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PSYCHE

A Journal of Entomology

Volume 80
1973

Editorial Board

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

E. O. Wilson
B. K. Holldobler

P. J. Darlington,
H. W. Levi
J. M. Burns
R. E. SlLBERGLIED

Published Quarterly by the Cambridge Entomological Club
Editorial Office: Biological Laboratories
1 6 Divinity Avenue
Cambridge, Massachusetts, U.S.A.

The numbers of Psyche issued during the past year were mailed on the
following idates:

Vol. 79, no. 4, December, 1972: April 25, 1973
Vol. 80, no. 1-2, March-June, 1973: September 7, 1973
Vol. 80, no. 3, September, 1973: December 16, 1973

L J

Q. L

^rL\

y'fv r

PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol. 80

March-June, 1973

Nos. 1-2

CONTENTS

Notes on the Life Cycle and Natural History of Parides areas mylotes
(Papilionidae) in Costa Rican Premontane Wet Forest. A. M. Young 1

Body, Web-building and Feeding Characteristics of Males of the
Spider Araneus diadematus (Araneae: Araneidae). R. Ramousse 23

Annotations on Two Species of Linyphiid Spiders Described by the
Late Wilton Ivie. P. J. van Helsdingen 48

Correlation Between Segment Length and Spine Counts in Two Spider
Species of Araneus (Araneae: Araneidae). L. D. Carmichael 62

Ant Larvae of Four Tribes: Second Supplement (Hymenoptera : For-
micidae: Myrmicinae). G. C. Wheeler and J. Wheeler 70

A New Species of Anacis from Northwest Argentina (Hymenoptera,
Ichneumonidae). C. C. Porter 83

Growth of the Orb Weaver, Araneus diadematus, and correlation with
Web Measurements. J. Benforado and K. H. Kistler 90

The Cockroach Genus Calolampra of Australia with Descriptions of
New Species (Blaberidae) . L. M. Roth and K. Princis 101

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1972-1973

President H. F. J. Nijhout, Harvard University

Vice-President T. F. H la vac, Harvard University

Secretary R. B. Swain, Harvard University

Treasurer F. M. Carpenter, Ilarvard University

Executive Committee A. F. Newton, Jr., Harvard University

J. W. Truman, Ilarvard University

EDITORIAL BOARD OF PSYCHE
F. M. Carpenter (Editor), Fisher Professor of Natural History ,
Harvard University

P. J. Darlington, Jr., Professor vf Zoology, Emeritus , Harvard
University

W. L. Brown, Jr., Professor of Entomology , Cornell University ;

Associate in Entomology , Museum of Comparative Zoology
E. O. Wilson, Professor of Zoology , Harvard University
H. W. Levi, Professor of Biology and Curator of Arachnology ,
Museum of Comparative Zoology

H. E. Evans, Alexander Agassiz Professor of Zoology, Harvard
University

J. M. Burns, Associate Professor of Zoology, Elarvard University

PSYCHE is published quarterly by the Cambridge Entomological Club, the
issues appearing in March, June, September and December. Subscription
price, per year, payable in advance: $4.50 to Club members, $6.00 to all other
subscribers. Single copies, $2.00.

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

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

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

Authors are expected to bear part of the printing costs, at the rate of
$13.50 per printed page. The actual cost of preparing cuts for all illustra-
tions 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, $20.00 each;
smaller sizes in proportion.

The December, 1972, Psyche (Vol. 79, no. 4) was mailed April 25, 1973

The Lexington Press. Inc.. Lexington. Massachusetts

PSYCHE

Vol. 80 March-June 1973 No. 1-2

NOTES ON THE LIFE CYCLE AND
NATURAL HISTORY OF

PA RIDES ARCAS MYLOTES (PAPILIONIDAE) IN
COSTA RICAN PREMONTANE WET FOREST*

By Allen M. Young
Department of Biology, Lawrence University
Appleton, Wisconsin 5491 1

The “ Aristolochia- feeding” swallowtails of the New World tropics
comprise a well-known group of butterflies famous for their roles in
mimicry complexes (Brower and Brower, 1964). Although the
adult stages of many congeneric species of notable genera such as
Battus and Parides have been known for some time (Godman and
Salvin, 1879-1901; Seitz, 1924), there is considerably less informa-
tion concerning the immature stages of these butterflies. This is
particularly the case for the Central American species of Parides , one
of the three genera ( Battus , Parides , and the Old World Troides)
of the Troidini, the tribe of pharmacophagous swallowtails (Ehrlich
and Raven, 1965). While the Troidini are most abundant in the
Old World tropics, it is apparent that New World genera in this
tribe, such as Battus and Parides , have undergone extensive speciation
in Central and South America. And with the exception of a few
studies such as the recent study of Battus polydamus in Costa Rica
(Young, 1971a) and another on the related Ornithoptera alexandrae
on New Guinea (Straatman, 1971), the life cycles, behavior, and
food plants of many species remain obscure. It is believed that the
primarily neotropical distribution of the Aristolochiaceae (Pfeifer,
1966) is a major factor in accounting for the extensive adaptive
radiation of Parides and Battus on these plants ( Brower and Brower,
1964; Ehrlich and Raven, 1965).

It is the close and perhaps coevolutionary association of genera
such as Parides with Aristolochia (in the Aristolochiaceae) and the
co-occurrence of several sympatric congeneric species in lowland

* Manuscript received by the editor March 26, 1973

I

2

Psyche

[March-June

tropical forests (Young, 1971b) that makes these butterflies suitable
candidates for the study of butterfly community structure in the
tropics. In the Caribbean premontane wet forests of Costa Rica,
there occur at least three species of Parides whose adults are often
found together on the same flowers in forests: P. areas mylotes, P.
childrenae, and P. sadyattes. Another subspecies of P. areas , namely
mycale , is also seen in association with these species. As an initial
approach to determining the ecological factors responsible for the
co-occurrence of these similar species as a functional Mullerian
mimicry complex (Young, 1971b), studies have been conducted on
the life cycle, food plants, and other aspects of butterfly biology, for
all of these species in Costa Rica. To date, the biological data for
P. areas mylotes (Bates) both in the laboratory (Young, 1972a)
and held (Young, 1971b; 1972a) has been the most extensive for
these species. This paper touches upon various aspects of biology in
this species not covered in the previous studies. Other reports will
subsequently appear concerning the biology of the remaining species.
Godman and Salvin (1879-1901) mention that P. areas mylotes is
common in the Pacific and Caribbean lowlands of Central America,
ranging from southern Mexico to Costa Rica. Thus the widespread
geographical distribution of the butterfly throughout Central America
makes it an even more attractive species to study from the standpoint
of the effects of local selection pressures on natural history and life
cycle.

Methods

The studies summarized here are: habitat selection, life cycle,
larval food plant acceptance, and behavior of immatures and adults.
Life cycle and larval food plant acceptance were examined in the
laboratory, while the other studies were conducted in the field at two
localities. At various times between late 1968 and mid- 1970, field
studies of P. areas mylotes were conducted at Finca la Selva, a region
of relatively undisturbed premontane tropical wet forest (elev. about
90 m) located on the confluence of the Rio Puerto Viejo and Rio
Sarapiqui. During the months of July and August 1972, the butter-
fly was studied at Finca Tirimbina, a forest site located about 8 km
west of Finca la Selva and at the basal belt transition zone (about
200 m. elev.) between montane and premontane tropical wet forest.

Habitat selection was studied by observing feeding and egg-laying
activities of adults at various places in the forest, both at “La Selva”
and “Tirimbina”. At La Selva, habitat selection was studied spo-

1973]

Young — Parides areas mylotes

3

radically several days each month over a 14-month period. At Tirim-
bina, it was studied systematically 14 days over a two-month period.

Life cycle studies consisted of the description of life stages and
the estimation of egg-adult developmental time under “laboratory”
conditions. These measurements were made on individuals reared
on a natural food plant, and eggs were obtained in one of two basic
ways. The first method was to collect eggs witnessed to be ovi-
posited in the wild ; this method was employed primarily in the
Tirimbina studies and to a lesser extent in the earlier La Selva
studies. The second method was to obtain eggs by hand-pairing
newly-emerged adults, using the technique of Clarke (1952) for
Papilio machaon, or allowing mating to occur in pairs of adults
confined to plastic bags. The latter technique is useful to obtain
estimates of fecundity in this species (Young, 1972a). Both methods,
obtaining eggs in the wild, and mating females in the laboratory with
subsequent induction of oviposition, are very successful for this
species, provide large numbers of eggs for rearing studies. Combining
both methods, a large number of individuals were reared from La
Selva (primarily through the laboratory mating method) and a lesser
number were reared from Tirimbina. The “laboratory” for the La
Selva studies consisted of a well-ventilated room in an apartment in
San Jose, while the “laboratory” for the Tirimbina studies was a
room in a different apartment, located about 1.5 km from the first.
In both cases, air temperature usually varied between 2i-23°C and
the humidity was about 45%.

The techniques for rearing immatures of this butterfly are given
in Young (1972a) for La Selva individuals, and essentially the same
methods were employed for the Tirimbina studies.

The larval food plant acceptance studies were conducted from
individuals obtained at Tirimbina during 1972. This study consisted
of offering first instar larvae immediately after hatching, in the
laboratory, fresh clippings of several species of Aristolochia from
various sources. The rationale was to offer separate small groups
of young larvae various species of Aristolochia including species
known to be natural food plants. Larvae on each food plant were
then scored for survival rate and body size. There were five species
of Aristolochia that were called “novel” food plants in addition to
the two natural food plant species. Two experiments were con-
ducted in San Jose: in each of these, 12 larvae were reared on the
natural food plant and 13 were reared on each of two “novel” food
plants collected from different localities in Costa Rica. The remain-
ing three food plants were tested at Lawrence University during

4

Psyche

[March-June

September and October 1972. The three species of Aristolochia
involved were already growing in a greenhouse tropical room for
about two years, and the Parides eggs were transported (by air)
from Costa Rica to Lawrence on September 6, 1972. Since the
natural food plant was not in culture at Lawrence, enough cuttings
of it were also brought to Wisconsin to sustain the larvae through
the earlier instars. At Lawrence, 10 larvae were on the natural food
plant, and 8 on each of the “novel” food plants.

Field studies of larval and adult behavior consisted of making
repeated observations on the feeding, resting, and defensive habits of
larvae in different instars, and on the oviposition behavior of adults.

Results

1 Habitat selection

Adults of both sexes of P. areas mylotes are most commonly en-
countered along paths, natural clearings, swamp edges, and other
exposed areas that either border forest or those which are found in
the forest interior. For example, the general study site at Tirimbina
where adults were most frequently seen is between the edge of forest
and a small river (Fig. 1). This small strip of dense secondary
growth vegetation is the result of forest being cut back from the
river edge for the original purpose of growing yucca and other veg-
etables that form the major diet of these people. Here, the adults
fly low over dense second-growth vegetation, seldom crossing the
small river, and frequently flying several meters into the shaded
forest understory and canopy. Excursions into the forest were most
frequently done by mated females in search of oviposition sites while
males and very fresh (presumably unmated) females generally lin-
gered in the sunlight second-growth. The strip of second-growth
between the forest and river is a major courtship area for this butter-
fly at Tirimbina and extensive growths of the larval food plant vines
are found hanging down from trees along the forest edge and grow-
ing horizontally in the canopy within a few meters from the edge.
A later paper (Young, et ah, in prep.) will demonstrate that mated
females of this species are far more prone to dispersal than either
males or unmated females. In the present paper, we can say that
mated females cruise along extensive tracts of cleared forest edge in
search of egg-laying sites, while males and unmated females remain
close to their eclosion sites. Courtship encounters are generally con-
fined to low sunlight vegetation very close to where the adults
emerged from their pupae. Males precede females in emergence.

1973]

Young — Parides areas mylotes

5

Fig. 1. A major habitat of adult Parides areas mylotes (Bates) at Finca
Tirimbina, near La Virgen, Heredia Province, Costa Rica. An adult popu-
lation is found along the forest edge, and males are active in the low
secondary growth vegetation between the forest and small river (Rio
Tirimbina) to the left. August, 1972.

Thus habitat selection, which can obviously be exercised only by the
adults (since eggs and larvae are relatively fixed through the oviposi-
tion strategy), is molded strongly in this species by two factors:
( i ) establishment of optimal courtship sites by males in sunlight
second-growth bordering forests or forest clearings, and (2) the
response by mated females to become more prone to disperse in search
for oviposition sites. Similar adult movement patterns have been seen
at La Selva, and the lesser vagility of individual males was mentioned
in Young (1971b). A courtship strategy in which males patrol an
area of the habitat consistently day after day (Young, et al., in prep.)
and mate with females as the latter emerge from their pupae, is
optimal for butterflies in which males are shorter-lived than females,
as is the case with Parides (Young, 1972a). But Cook et al.,
(1971) report a short life expectancy of about 10 days in P. anchises

6

Psyche

[March-June

and P. neophilus in a seasonal habitat on Trinidad where torrent
rains may kill off the adults of both sexes.

Although adult feeding preferences do not appear to be ,a major
factor in dispersal patterns at Tirimbina, it is interesting to note that
mimetic association with other species of Parides is most intense at
nectaries at La Selva (Young, 1971b). At Tirimbina, P. areas
mylotes is the only species of this genus seen consistently at the study
site, and flower specificity is not apparent. At La Selva., this butterfly
as a functional component of Mullerian mimicry complexes exercise
a strong preference to visit a single species of flower {I~I amelia
patens ) also visited by other Parides (Young, 1971b) ; judging from
the amount of time spent daily at Hamelia flowers, there appear to
be very few or no other preferred adult food sources of Parides at
La Selva. In the absence of the other Parides at the Tirimbina study
site, adult P. areas mylotes is found on a variety of flowers, usually
ranging from red to purple. Thus in the absence of strong selection
pressures favoring mimetic association, and where this mimicry is
potentially most effective at a food source, flower specificity may
break down for Parides in habitats where the species do not co-occur
on a regular basis. Similar diurnal patterns of visitation at flowers
between members of a tropical Battus mimicry complex in addition
to the co-occurrence of several Parides at flowers at La Selva suggest
strong selection pressures resulting in convergence of feeding habits
to enhance mimicry (Young, 1971b; 1972b).

Although courtship activity is generally limited to the sunniest
hours of the morning (Young, et ah, prep.), adults of both sexes
and various age-classes (distinguished by the extent of wing tatter-
ing) generally forage throughout the day, and they are relatively
unaffected by changes in local weather conditions. Even at a mon-
tane tropical forest locality (Cuesta Angel) where a cloud forest
occurs at about IOOO meters elevation, adults are seen foraging
throughout the day at the bright red flowers of Impatiens sultani
( Balsaminaceae — “Touch-me-nots”), a small herbaceous plant that
is imported from Africa and that grows in large numbers. As the
day becomes less bright in terms of illumination from the sun, these
flowers become even more conspicuous due to increased contrast of
the red coloration with the misty air; to the human observer, the
flowers are more conspicuous, and perhaps the butterflies respond in
a similar fashion. In both lowland and mountain localities, adult
activity drops off sharply after about 4:00 P.M. When there is
short succession of unusually dry days in both lowland and mountain

1973]

Young — Parides areas mylotes

7

localities, adults, especially males, are frequently seen visiting reced-
ing mud puddles and moist patches of ground.

Life cycle and developmental time

The egg (Fig. 2-A-C) is deep rusty-brown and slightly flattened
at the base. The diameter is i.i mm and the egg is covered with an
irregular thick layer of an orange-red sticky substance, which at-
taches it to the leaf surface and gives the entire surface of the egg
a rough appearance. This sticky substance forms thin threads which
hang down from the upper half of the egg and assist in attachment
(Fig, 2-A). It is not known if the sticky substance is also defensive
in function, in the sense of discouraging attack by ants and other
leaf-wandering predatory arthropods. The apical region of the egg
darkens considerably immediately before hatching. Eggs are gen-
erally laid on the ventral surface of older leaves and occasionally in
the crotches of small stems and petioles (Fig. 2-C). The amount
or thickness of the sticky substance covering the eggs is apparently
very variable, since other details of egg external morphology, such as
deep grooves (Fig. 2-B, C) can be seen on some eggs while com-
pletely obscured on others. Eggs are laid singly but usually in loose
clusters of 2-5 eggs on a single leaf.

At La Selva, the natural food plant is tf Aristolochia sp.” (this is
a new species from northeastern Costa Rica soon to be described by
H. W. Pfeifer based on my collection of it during March, 1970).
At Tirimbina, the natural food plant is Aristolochia constricta
Griseb. Both of these species occur in lowland forest on the Carib-
bean side of the central Cordillera in Costa Rica. Pfeifer (1966)
mentions that A. constricta is a forest species found from Costa
Rica to Panama, the Lesser Antilles, and probably northern South
America.

The first instar is about 3.2 mm long when it hatches, and the
ground color of the body is dark orange-brown. The head is shiny
black. After the young larva begins to feed on leaf tissue, the body
ground color becomes a deep wine red. All segments bear long
tubercles of the same color as the body, but the lateral pair on the
first segment are orange-white, and this color also characterizes the
dorsal pairs of tubercles on segments two, seven, ten, and twelve
(Fig. 2-D ) . The tubercles are fleshy for about one-third their
length, with the apical two-thirds being stiff and bearing numerous
tiny black spines (Fig. 2-D). The oSmeterium is bright orange-
yellow throughout larval life. By the time of the first molt, the larva
is about 9 mm long.

8

Psyche

[March-June

I ! ■ 1 jflj

IlllMMiiiMil

1973]

Young — Parides areas mylotes

9

The second instar is remarkably similar in appearance to the first
instar, with the only major difference being a loss of the spines seen
on tubercles in the previous instar. The larva (Fig. 2-E, F) retains
the six rows of spines of the first instar, in addition to the shiny
black head and true legs. The precise arrangement of the tubercles
is very noticeable in this instar. The first four thoracic segments
bear two pairs of lateral tubercles, and the uppermost pair disappears
until the tenth segment where it is resumed until the twelfth seg-
ment. The two pairs of lateral tubercles on these segments are not
precisely in line: the tubercle of thoracic segment i are juxtaposed
with those of thoracic segment 2 etc. The lateral tubercles of the
thoracic segment i are considerably shorter than these tubercles on
the remaining segments. The dorsal pair of tubercles on abdominal
segments i and 4 are white, while the upper lateral pair of the fourth
and fifth abdominal segments are also white. The highly reduced
dorsal pair of the abdominal segment 1 1 are also white. This pat-
tern of tubercle arrangement and coloration is retained throughout
the rest of larval life. By the second molt, the larva is about 14 mm
long.

The third instar is an exact replica of the second instar except that
the ground color of the body is a very deep purplish black. The third
instar is shown in Fig. 2-G. By the time of the third molt, the larva
is about 23 mm long. The fourth instar (Fig. 3-A) is identical to
the third instar except that the skin is very shiny and reflective. It
attains a length of 35 mm by the fourth molt.

A dramatic change in the ground color occurs with the molt to
the fifth instar (Fig. 3-B, C). The ground color is a dull, velvety
purplish-brown mottled with irregular blotches of black. The black
coloration is most extensive on the segments bearing white tubercles
(Fig. 3-B, C). The wfrite ridge along the anterior edge of the
osmeterial cuff behind the head is more prominent in this instar. As
this instar continues to grow, the ground color becomes even lighter
in coloration as extensive velvety grayish-tan areas replace the for-
merly purplish-brown areas of the body. The coloration of the dark
tubercles is also variegated during the fifth instar, with each tubercle
bearing lines of white in addition to the mottled coloration of the

Fig. 2. Life cycle and behavior of Parides areas mylotes (Bates).
(A) dorsal view of two eggs on a leaf; note the rough surface and sticky
strands on the eggs (B) single egg showing deep vertical grooves (C) sin-
gle egg in crotch of stems (D) first instar, lateral view (E) two second
instar larvae (one is feeding) (F) several second instar larvae living to-
gether (G) third instar, dorsal view.

IO

Psyche

[March-June

1973]

Young — Parides areas mylotes

1 1

body ground color. By the time of pupation, the larva is about
45 mm long. The coloration of the larva remains unchanged at the
time of pupation.

The pupa (Fig. 3-D) is about 25 mm long and the color pattern
consists of various light shades of green and yellow. The frontal
portions of the thorax and abdomen are yellow while the rest of the
body is light green.

Godman and Salvin (1879-1901) and Seitz (1924) give good il-
lustrations of wing color patterns of the adults (Fig. 3-E). The
single light area of the dorsal surface of the forewing in the female
is cream-colored while the dorsal bands on the hindwings are orange-
red. This color pattern is very consistent in both laboratory-reared
and wild-caught females of P. areas mylotes. Less stable is the fore-
wing dorsal coloration in the male within a single local population.
The large spot on each forewing (Fig. 3-E) is light green but with
the apical portion being cream-colored. Considerable variation is
apparent in this “two-component’ 5 spot on the dorsal surface of the
male’s forewing; this variability concerns the presence, absence, and
size of a second, very small two-component spot just inside the radial
cell of each forewing, and almost touching the major spot (Fig.

3-E). Similarly, there is considerable variation in the discal cell

spot. Godman and Salvin (1879-1901) mention the considerable
variation in the forewing spotting pattern of male in the closely
related species, P. iphidamas. Adults of both sexes of P. areas
mylotes can be distinguished from the subspecies mycale by the
presence of a thin light red marginal border of the wings in the
former subspecies, while these markings are white in the latter sub-
species. The red patch on the dorsal surface of the hindwings in

male P. areas mylotes is more intense than in the female, and the
distribution of the coloration is very different between the sexes
(Fig. 3-E). In bright sunlight, the red patches of the male’s hind-
wing are often iridescent, giving off a purple lustre; this is not seen
in the female. The mean length of the forewing in the female is
about 40 mm, while the same statistic of the male is about 38 mm.
Thus, not only is there a striking color sexual dimorphism in this
butterfly, but also a consistent wing length difference between the
sexes. In the absence of crowding, laboratory-reared individuals often

Fig. 3. Life cycle and behavior of Parides areas mylotes (Bates).
(A) fourth instar, lateral view (B) fifth instar, lateral view (C) fifth
instar, feeding on the tip of a young stem of Aristolochia (D) pupa, lateral
view (E) adults, female above, male below.

12

Psyche

[March-June

bear the same wing-length as wild-caught individuals from the same
locality.

The egg-adult developmental time for P. areas mylotes in the
laboratory for individuals reared on Aristolochia constricta is sum-
marized in Table i. In a previous study (Young, 1972a), the egg-
adult developmental time of this butterfly on Aristolochia sp. from
La Selva was about 42 days. The developmental time in that study
was measured on eggs obtained from La Selva adults. The develop-
mental time for eggs obtained at Tirimbina, and reared on A. con-
stricta is 53 days (Table 1). This difference in developmental time
between the two populations is apparent in eggs, larvae, and pupae:
the egg stage lasts 4 days in La Selva individuals as opposed to 6 days
in Tirimbina individuals ; the total larval period for La Selva in-
dividuals is 17 days as opposed to 33 days in Tirimbina individuals;
the pupal stage lasts 21 days in La Selva individuals as compared to
14 days in Tirimbina individuals.

Larval food plant acceptance

Development from the egg stage on natural food plants is suc-
cessfully completed in the laboratory (Young, 1972b; Table 1).
When other species of Aristolochia are tested, differences in food
plant acceptance by the larvae become apparent. Development is
successfully completed, and without a change from the Tirimbina
developmental time when larvae are reared from the egg stage on
Aristolochia labiata Willd. in Costa Rica. But larvae die during the
first instar when offered A. veraguensis Duchr. in Costa Rica. For

Table 1. The developmental time of Parides areas mylotes on a natural
food plant, Aristolochia constricta, under laboratory conditions.*

INSTAR

INSTAR

INSTAR

INSTAR

IN STAR

TOTAL

EGG

1

2

3

4

5

PUPA EGG-ADULT

MEAN

DURATION

(days)

6

5

5

6

6

11

14 53

± S.E.

± 0.1

± 0.3

± 0.5

± 0.3

± 0.2

± 0.8

± 0.2

N

46

46

42

42

40

37

37

^Laboratory conditions consisted of confining larvae to closed plastic bags
containing clippings of food plant. Physical conditions around the bags
were 21-23 °C and about 45% relative humidity. See text for further details
of rearing techniques, laboratory conditions, etc.

1973]

Young — Parides areas mylotes

13

the rearing studies at Lawrence, all the larvae died either in the
first or second instar when reared on A. ringens Vahl, A. littoralis
Parodi, and A. gigantea (Mart. & Zucc.). For the groups of larvae
offered these species, survivorship was 0%. Thus, in addition to the
two known natural food plants of P. areas mylotes , namely Aristo-
lochia sp. from La Selva and A. constricta from Tirimbina, the
butterfly only feeds successfully on A. labiata Willd. in Costa Rica.

Behavior of adults and larvae

Observations on adult behavior are limited to the oviposition strat-
egy of this species, since a later report (Young et al., in prep.) will
discuss other aspects of adult behavior, most notably, the spacing
patterns of males and females, and the courtship strategy.

Adults of both sexes generally cruise very low over second-growth
vegetation. Mated females in search of oviposition sites exhibit
extreme forms of cruising behavior in two ways : ( 1 ) they perform
sudden, almost vertical darts into the canopy where Aristolochia
lianas are found, and (2) they flutter through very dense second
growth within a few inches of the ground, and often being obscured
from view for several minutes.

Such patterns of cruising behavior by egg-laying females are con-
sistent with the observation of well-developed food plant specializa-
tion in this butterfly. The usual situation locally is that eggs are
laid on a single species of Aristolochia , and there is considerable site-
selectivity exercised in terms of placing the individual eggs securely
on the older leaves of an individual plant. The eggs are seldom
laid on young leaves and occasionally on stems at crotches between
two stems. Eggs are customarily laid on the dorsal surface of older,
well-shaded leaves of the vine, and anywhere from one to five eggs
may be laid in a loose cluster in this manner. Upon landing on a
leaf for oviposition, the female exhibits considerable wing fluttering
and drumming behavior with the antennae; an egg is usually laid
within 12 seconds. Oviposition is most commonly seen during sunny
hours throughout the day. While males may be cruising in the
general vicinity of egg-laying females, there is virtually no observable
interactions between the sexes. The less cohesive nature of the mated
female portion of a local breeding population of P. areas mylotes
(Young et al., in prep.) results in there usually being no more than
one or two ovipositing females at a larval food plant patch on a given
day. These individuals cover large tracts of habitat in searching for
oviposition sites, but usually return repeatedly on the same day to a
given food plant patch.

While clustering of eggs in the field is generally loose and vari-

14

Psyche

[March-June

able, when mated females of this butterfly lay eggs in the laboratory,
there is usually a tight clustering of eggs (Young, 1972a). Thus
tight clustering of eggs (the arrangement of eggs into a group where
the eggs touch each other) can be induced in the laboratory when
females are confined individually or in low numbers to plastic bags
containing clippings of the food plant. Such clustering, however, is
seldom found in the wild in this butterfly and other species of
Parides.

The larvae of P. areas mylotes exhibit several behavioral patterns
that warrant more intensive study. Upon hatching the larva in-
variably eats its emptied egg shell, and then moves a considerable
distance to the closest youngest leaves. Locomotor movement is ac-
companied by the production of silken treadwork on which the larva
crawls from one place to another. Although small groups of larvae
are frequently found in the field (Fig. 2-E, F) there is no evidence
for gregarious habits among the individuals in a group. All individ-
uals on an individual vine generally feed at the same times of day,
but there is no indication of coordinated locomotor movements among
the individuals. Furthermore, single or doublets of larvae are also
frequently encountered in the field. Larvae of all instars are gen-
erally inactive at night. The extent of larval dispersion when several
eggs are laid on a vine may be governed by the size of the vine. For
example, it is not uncommon to find one or two fourth or fifth instar
larvae present on a young vine ( 1-2 m tall) in the field, and in cases
where there are two present, these individuals are often found to-
gether on the same stem. Both in the field and laboratory, older
larvae eat the stems of young Aristolochia vines (Fig. 3-C). On
very large vines in which woody tissue is well-developed, older larvae
are generally confined to feeding on leaves and it is unusual to find
two or more individuals resting close together. Group formation is
frequently encountered only in the younger larvae (first and second
instars) and in cases where larger (older) larvae are clumped, this
is most likely due to the fact that they are feeding on a young vine
and the food supply is limited. It is not known if Parides larvae
crowded on young Aristolochia vines will leave the vine in response
to intense crowding. The osmeteria of the larvae of swallowtail
butterflies are functional defense organs. Predatory attack on the
larvae of P. areas mylotes in the wild has not been observed to date.
The defensive strategy of the larvae against predators includes
( 1 ) possession of conspicuous body coloration in which the dark
body and pattern of white tubercles stands out against the light green
coloration of Aristolochia leaves, (2) possession of an apparently

1973]

Young — Parides areas mylotes

15

functional and brightly-colored defensive organ, the osmeterium, and
(3) probably the possession of generally toxic or poisonous systemic
properties making the insect unpalatable, since they feed on vines
reputed to have very toxic properties.

Discussion

Young (1971a) reported a developmental time for Battus poly-
damus on Aristolochia veraguensis of about 14 days under similar
laboratory conditions to those employed in the present study. Straat-
man (1971) reported the developmental time of Ornithoptera alex-
andrae Rothschild to be 13 1 days on Aristolochia schlechteri and
107 days on A. tagala, where the difference occurred during the
larval period. The developmental time of Parides areas mylotes on
Aristolochia sp. from La Selva is 42 days (Young, 1972a) while
53 days on A. constricta (Table 1.). Furthermore, the develop-
mental time of Parides childrenae on Aristolochia pilosa at La Selva
is about 42 days (Young, 1972a). Thus different genera in the
Troidini have different developmental times on different species of
Aristolochia. At La Selva there has been ecological divergence be-
tween P. areas mylotes and P. childrenae with respect to the species
of Aristolochia used for oviposition and larval food-consumption.
Furthermore, two different strains of P. areas mylotes are evolving
between La Selva and Tirimbina: the duration of all immature life
cycle stages has been altered and the species feeds on a different
species of Aristolochia at each locality. If this difference in develop-
mental time was due solely to differences between the larval food
plant species, we would expect to find only a change in duration of
the larval period similar to that noted by Straatman (1971) in
Ornithoptera alexandrae on New Guinea. But in the case of P.
areas mylotes, there has been a change in the embryonic and post-
embryonic developmental time which suggests genetic alterations.
Strain-effect is not solely confined to food plant differences of the
type noted for Victorina epaphus on the Pacific and Caribbean slopes
of the central Cordillera in Costa Rica (Young, 1972c). Precisely
what sorts of ecological factors are reshaping the developmental
architecture of P. areas mylotes at different localities on the Carib-
bean drainage of the central Cordillera in Costa Rica remain obscure
at this time. One interesting hypothesis concerning this question
would focus on a higher level of predation pressure on eggs and
larvae in La Selva populations of the butterfly, which would favor
an accelerated developmental period for these life stages.

i6

Psyche

[March-June

The inability of young larvae of P. areas mylotes to survive on
Aristolochia ringens, A. littoralis, A. gigantea, and A. veraguensis
may be due to the lack of evolutionary contact (Ehrlich and Raven,
1965) with these plants. An alternative explanation is that extreme
food plant specialization in the butterfly has resulted in the narrow
restriction to only a few species of Aristolochia locally. Until more
is known about the regional and geographical distribution of various
species of Aristolochia in Central America, it will be difficult to
resolve the question of larval food plant adaptability in Parides.
Unfortunately eggs from La Selva have not been reared on A. con-
stricta from Tirimbina nor the converse, namely, eggs from Tirim-
bina reared on A ristolochia sp. from La Selva.

The question of unpalatability is of considerable ecological and
evolutionary interest. Brower and Brower (1964) have demon-
strated that freeze-killed adult Parides neophilus L., which feeds on
various species of Aristolochia on Trinidad, are very unpalatable to
Scrub Blue Jays in the laboratory. Brower and Brower (1964),
Ehrlich and Raven (1965) and Pfeifer (1966) cite previous studies
which illustrate the toxic properties of various compounds derived
from the vegetative portions of Aristolochiaceae. The question of
palatabifity in genera of the Troidini ( Parides , Battus , Ornithop-
tera, and Troides ) is of interest since the larvae are presumably
unpalatable in addition to possessing a defensive organ (Eisner et. al.,
1971). The larvae of these genera, as exemplified in the present
study by P. areas mylotes , are generally conspicuous in appearance
(Fig. 2, 3) to the human observer.

The possession of a dual system of defense by Parides larvae and
other troidines is related to the functional responses of each com-
ponent (unpalatability and chemical defense secretion) to different
kinds of predators that the larvae encounter in their habitats. Un~
palatability, as evidenced here by the conspicuous coloration of the
larvae and the toxic properties of their food plants, is an adaptation
for defense against vertebrate predators such as insectivorous birds,
mammals, and reptiles. Brower and Brower (1964) have demon-
strated that blue jays become ill after eating an unpalatable butterfly
and that there is a subsequent modification in prey-selection behavior
by such an experienced predator to avoid the prey on further visual
contact with it. Thus, the flexible learning abilities of vertebrate
predators makes unpalatability an effective defensive mechanism that
increases the likelihood of survival of individuals in a prey popula-
tion. An insectivorous bird foraging in forest edge second-growth or
forest canopy has daily opportunity for visual contact with the

1973]

Young — Parides areas mylotes

7

poisonous Parides larvae which stand out against the foliage back-
ground during the day time when they are feeding. This is an ideal
situation for unpalatability to be effective against vertebrate preda-
tors. The bird does not have to make tactile contact with the poten-
tial prey, but can recognize it from a distance. On the other hand,
the added possession of a defensive organ that produces a volatile
chemical secretion would be an adaptation primarily against inverte-
brate (arthropodan) predators that make tactile or very close visual
contact with Parides larvae and elicit a behavioral response. Such a
defensive mechanism would be essentially ineffective against verte-
brate predators since the larvae could not respond fast enough to the
strike of the predator, and the larva would invariably be killed.
This is especially true since lepidopterous larvae have low visual
sensing ability but quick discriminatory ability for tactile stimuli.
In a similar fashion, the generally instinctive nature of the be-
havioral repertoire of invertebrate predators would make unpalata-
bility an ineffective defense mechanism against these predators.
Under such conditions, there is strong selection for the evolution with
a dual system of defense, one adapted to vertebrate predators with
developed learning abilities (unpalatability), and the other adapted
to smaller invertebrate predators with instinctive behavior patterns
(defense glands). Furthermore, the larvae would probably survive
single attacks by invertebrates such as ants, even though the in-
stinctive nature of the predator’s behavior results in repeated attacks
on the prey. The small size of invertebrate predators and the ability
of Parides larvae to survive individual attacks (in the form of small
bites) reduces the threat of death from instinctive predatory be-
havior patterns. Thus, in the absence of conclusive evidence, I sug-
gest that the unpalatable properties of troidine butterfly larvae (Euw
et al., 1968) are an adaptation to potential large vertebrate preda-
tors, while their defensive organs comprise an adaptation to inverte-
brate predators. This effect is even more pronounced in the adults,
which are very unpalatable to birds (Brower and Brower, 1964),
since there are ample opportunities for foraging birds which catch
insects on the wing to recognize, at a distance, the butterflies through
conspicuous coloration. Therefore, adult butterflies should possess
unpalatability rather than defensive gland as an adaptive strategy
against vertebrate predators. The studies of Euw et al. (1968) and
Eisner et al. (1971) indicate that unpalatability and chemical de-
fense secretions in troidine butterflies are due to very different kinds
of chemical compounds.

The oviposition behavior varies greatly for different genera of

i8

Psyche

[March-June

troidine butterflies. Straatman (1971) found that Ornithoptera alex-
andrae lays eggs singly, and Cook et al. (1971) comment that single
oviposition also occurs in Parides neophilus and P. anchises on Trini-
dad. But Young (1971a) found tight cluster oviposition in the held
to prevail in Battus poly damns in Costa Rica. The oviposition in P.
areas mylotes is very variable since eggs may be laid singly or as
loose clusters of varying numbers of eggs per cluster. But oviposi-
tion in the wild is never tightly clustered as seen in Battus polydamus
(Young, 1971a). The P. areas mylotes pattern is basically single,
but with a behavioral tendency to lay several eggs close together on
a single leaf. This behavior results in first and second instar re-
maining together in small groups and dispersing later, which is very
different from the more well-defined gregarious behavior exhibited
by the larvae of Battus polydamus (Young, 1971a). Larvae in
the latter case are generally gregarious through all instars and
presumably fitness is increased as noted in other studies (Ghent,
i960). A similar oviposition pattern to that found in P. areas
mylotes also occurs in P. childrenae and P. sadyattes (Young,
in prep.). Thus the oviposition pattern of Parides in Costa Rica
(and perhaps for all of the Central American mainland) is a
variable one being basically single but typified by loose clusters of
a variable number of eggs, usually ranging between two and five
on a leaf. It is clearly not entirely single, nor is it the tight-cluster-
ing pattern seen in Battus. As might be predicted, the larvae are
semi-gregarious in P. areas mylotes (Fig. 2-E, F) as well as in P.
childrenae and P. sadyattes (Young, in prep.) and perhaps in most
Parides , while truly gregarious in Battus. These preliminary find-
ings in different species suggest that there may exist distinct phylo-
genetic patterns of type of oviposition and extent of larval gregari-
ousness at the generic level in the Troidini, and perhaps within other
tribes of Papilioninae. Superimposed upon evolutionary history will
be the prevailing ecological conditions (Birch and Ehrlich, 1967)
such as food plant specialization, patchiness of food plant populations,
predation pressure on immatures, adult population cohesiveness, and
several others, which mold the oviposition strategy in either direction
(single versus clustering) and the likelihood of larval gregarious
behavior.

Summary

In this paper concerning the life cycle and natural history of
Parides areas mylotes (Bates) on the Caribbean side of the central
Cordillera in Costa Rica, the following points were emphasized:

1973]

Young — Parides areas mylotes

19

( 1 ) The butterfly is a forest species which is most commonly
encountered along forest edges associated with extensive borders of
secondary growth vegetation or small forest clearings.

(2) Habitat selection by adults is governed primarily by two
factors: (a) the selection of optimal courtship sites by males ex-
hibiting home range behavior, and (b) the search pattern of mated
females for suitable oviposition on Aristolochia vines along forest
borders and in the canopy.

(33 The larvae of this species are probably warningly-colored,
since they contrast greatly with the light green leaves of the food
plant. The pupae are cryptically colored against the same back-
ground.

(4) The egg-adult developmental time varies on different natural
food plants in different localities: on Aristolochia sp. from Finca
La Selva the developmental time is about 42 days; on A . constricta
from Finca Tirimbina 53 days. This difference is due to more than
food plant difference since the egg stage is considerably shorter in
individuals reared on Aristolochia sp. There appears to have been
the evolution of different strains in different localities where different
food plants are also exploited.

(5) Development is successfully completed on A. labiata but
unsuccessful on A. veraguensis , A. ringens , A. littoralis, and A.
gigantea. The inability of young larvae to feed on these species
may be due to either (a) a lack of contact with those species, or
(b) the development of narrow food plant specialization.

(6) The conspicuous coloration (contrast) of the larvae against
the light green food plant leaves and the known toxic properties of
the Aristolochiaceae indicate that the larvae are unpalatable to verte-
brate predators with well developed learning abilities. The un-
palatability of the larvae is inferred from the known unpalatability
of the adults of a related species of Parides. The possession of an
osmeterial defensive organ is interpreted here, on the other hand, as
being primarily an adaptation of defense against invertebrate (ar-
thropodan) predators with rather inflexible (instinctive) learning
abilities.

(7) The variable oviposition strategy of P. areas mylotes in the
wild is not strictly single nor is it clustering. Eggs are generally
laid in loose clusters of two to five eggs on a leaf, and this pattern
appears to be a modified form of single oviposition. When mated
females are confined to plastic bags in the laboratory, tight clustering
of eggs can be induced. Previous studies show that at least one
tropical species of Battus lays eggs in tight clusters in the wild,

20

Psyche

[March-June

while some species of Parides undoubtedly lay eggs singly and
Ornithoptera lays eggs singly. It is suggested that there may exist
phylogenetic differences in oviposition patterns at the generic level
in the Troidini, and that secondary differences in these patterns are
molded by contemporary ecological factors.

Acknowledgements

The La Selva portion of these studies was financed by N.S.F.
Grant GB-7805, Daniel H. Janzen, principal investigator, with
logistic support through the Organization for Tropical Studies, Inc.
The Tirimbina studies were financed by Grant No. 12 1 of the
Bache Fund of the National Academy of Science and partially by
N.S.F. Grant GB-33060. Logistic support was provided by the
Costa Rican program of the Associated Colleges of the Midwest.
Roger Kimber and John Thomason assisted with rearing and food
plant acceptance studies. Howard W. Pfeifer identified the species
of Aristolochia and provided rooted cuttings of several species. Lee
D. Miller confirmed the identification of the butterfly. The manu-
script was read by Murray S. Blum.

Literature Cited

Birch, L. C., and P. R. Ehrlich.

1967. Evolutionary history and population biology. Nature 214: 349-
352.

Brower, L. P. and J. V. Z. Brower.

1964. Birds, butterflies, and plant poisons: a study in ecological chem-
istry. Zoologica 49: 137-159.

Clarke, C. A.

1952. Hand pairing of Papilio machaon in February. Entomol. Record
64: 98-100.

Cook, L. M., K. Frank, and L. P. Brower.

1971. Experiments on the demography of tropical butterflies. I. Sur-
vival rate and density in two species of Parides. Biotropica 3 :
17-20.

Ehrlich, P. R., and P. H. Raven.

1965. Butterflies and plants: a study in coevolution. Evolution 18:
586-608.

Eisner, T.

1970. Chemical defense against predation in arthropods. In Chemical
Ecology (ed. by Sondheimer, E. and Simeone, J. B.), pp. 157-218.
New York: Academic Press.

Eisner, T., A. F. Kluge, M. I. Ikeda, Y. C. Meinwald, and J. Meinwald.

1971. Sesquiterpenes in the osmeterial secretion of a papilionid butter-
fly, Battus polydamas. J. Insect Physiol. 17: 245-250.

1973]

Young — Parides areas mylotes

21

Euw, J. V., T. Reichstein, and M. Rothschild.

1968. Aristolochic acid-1 in the swallowtail butterfly Pachlioptera
aristolochiae (Fabr.) (Papilionidae) . Israel J. Chem. 6: 659-670.
Ghent, A. W.

1960. A study of the group-feeding behavior of larvae of the Jack pine
Sawfly, N eodiprion pratti banksianae Roh. Behavior 16: 110-148.
Godman, F. D. and O. Salvin.

1879-1901. Biologia centrali-americana, Insecta, Lepldoptera-Rhopalo-
cera. Vol. I.

Jordan, K.

1907-1908. Papilio. In The Macrolepidoptera of the world.
Vol. 5. The American Rhopalocera. (Ed. by A. Seitz), pp. 11-45.
Stuttgart: A. Kernan Verlag.

Pfeifer, H. W.

1966. Revision of the North and Central American hexandrous species
of Aristolochia (Aristolochiaceae) . Annals Missouri Botan. Gar-
den 53 : 115-196.

Straatman, R.

1971. The life history of Ornithoptera alexandrae Rothschild. J. Lepid.
Soc. 25: 58-64.

Young, A. M.

1971a. Mimetic associations in natural populations of tropical papilionid
butterflies. I. Life history and structure of a tropical dry forest
breeding population of Battus polydamus polydamus. Rev. Biol.
Trop. 19: 211-240.

1971b. Mimetic associations in natural populations of tropical papilionid
butterflies (Lepidoptera : Papilionidae). J. New York Entomol.
Soc. 79 210-224.

1972a. Breeding success and survivorship in some tropical butterflies.
Oikos 23: 318-326.

1972b. Mimetic associations in populations of tropical butterflies. II.
Mimetic interactions of Battus polydamus and Battus bellus.
Biotropica 4: 17-27.

1972c. The ecology and ethology of the tropical nymphaline butterfly,
Victorina epaphus . I. Life cycle and natural history. J. Lepid.
Soc. 26: 155-170.

BODY, WEB-BUILDING AND FEEDING
CHARACTERISTICS OF MALES OF
THE SPIDER ARANEUS DIADEM ATUS
(ARANEAE: ARANEIDAE)

By Raymond Ramousse*

Division of Research

North Carolina Department of Mental Health
P. O. Box 7532
Raleigh, North Carolina 27611

INTRODUCTION

Many investigators have observed female orb-web spiders in their
natural habitats (Enders, 1972; Eberhard, 1971), but there have
been relatively few scientific observations of males outdoors. A major
reason for this is because after maturation males discontinue web-
building and they seek mates and are difficult to follow in an un-
confined setting. Males have also attracted less attention in labora-
tory situations since they have shorter life spans than females and
because they stop building webs after reaching maturity. The activity
of spiders in laboratories has been observed primarily in relation to
their web-building behavior (LeGuelte, 1966, Witt, 1963a, b),
making the female a more frequent subject of study. Thus, with
the exception of maturation on web-building (Witt et al.f 1972),
only females have been comprehensively studied.

The focus of this research is to explore the activities of the males
of Araneus diadematus ’Clerck and their role in the female-male
relationship which ultimately determines the continuity of the species.
Two characteristics related to the females have already been identi-
fied as possibly playing a part in the survival of the species. These
include cocoon hatching and differential maturing. Cocoons have
been observed hatching at two different times for a single species of
spider — presumably providing an advantageous distribution of egg-
production over a period of time (Potzsch, 1963). Also, within a
set1 of spiderlings, different rates of maturation have been observed.
Some females grow rapidly and die early while others grow slowly

^Present address of author: Laboratoire d’Ethologie experimentale, 1 rue
Raulin, 69 Lyon 7e, France

Manuscript received by the editor March 26, 1973

To avoid confusion with the designation of “family” used in nomencla-
ture, offsprings from a single cocoo-n will be called a “set.”

22

1973]

Ra?mousse — A raneus diadematus

23

and live at least four months longer (Reed & Witt, 1972). The
related differential maturing rates may provide an advantageous
distribution of spiderlings over a period of time. Together, these
mechanisms would seem to help a species survive drastic or poten-
tially destructive changes in environmental conditions. This research
seeks to explore the male’s role in these phenomena. At what rate is
he growing, maturing and dying during the female’s life cycle?

This leads to the question of inbreeding. An observation of the
maturation rates of spiderlings of the same set was conducted in an
effort to determine if inbreeding is possible.

Also, if the rate of growth is a factor in the rate of maturation
(and spiders of the same set are known to present a considerable
variation in size even under apparently optimal conditions), (Witt
et al., 1968), is growth prenatally or genetically determined or a
function of external factors?

The effects of an even diet independent of manifest behavior (Witt
et al. , 1972) and differential force-feeding on various schedules
(Benforado & Kistler, 1972) have already been studied. What,
however, would happen to the growth rate of male and female
spiders if they could choose their food quantity through web-building
frequency?

The answers to some of these questions about the growth and
maturation of male spiders should provide clues about their role in
the reproductive cycle and, more generally, about their role in the
continuity of the species.

METHODS

Two Araneus diadematus cocoons collected in the field, were placed
in two different rearing boxes in the laboratory, where they hatched
(February 23, 1972, one cocoon and 14 days later, March 6, 1972,
the other). The offspring from the first cocoon will be called set I,
and the offspring from the second cocoon, set II. The laboratory
provided a cycle of long warm days and short cool nights throughout
the lifespan of the animals.

As the animals left the communal web to build individual webs,
they were put in glass tubes. Five weeks after hatching for set I and
three weeks after hatching for set II the spiderlings were caged in
individual labeled frames (50 X 50 X 10 cm) where they could
build webs without apparent limitation in size. All observations
began at this moment; however, some molts were noticed inside the
cocoon, and the spiderlings molted one or two times in the glass tubes.

24

Psyche

[March-June

In the rearing boxes as well as in the glass tubes they were pro-
vided water and gnats ad libitum. In the frames the spiderlings were
fed with de-winged houseflies. A weighed fly was given one time
every three days only when a web had been built, thus rewarding the
spiders for high frequency of building.

The individual weights of the spiders (accuracy o.i mg) were
recorded every week and web-building was recorded every day. Each
web was photographed then collapsed by the experimenter, and ana-
lyzed for size, shape, fine structure and regularity (Reed et al.,
1965). The dates of the molts of each spiderling were recorded and
the length of the first leg was measured on the molted limb (ac-
curacy in mm.).

In the following pages the initials FG and SG are used in place
of fast growing males and slow growing males. Statistical com-
parison between the two groups (SG & FG) where not specifically
mentioned was made with the Wilcoxon test, adapted by White for
unpaired measurements (White, 1952).

RESULTS

Of 31 spiderlings that reached maturity in set I, twelve were
identified as males. There were 14 males out of a total of 29 animals
in set II. The number of males in each set is significantly repre-
sentative of the expected 50% probability of males in a population
(Binomial test, p = 0.01 in each case).

Some characteristics of the male

The adult males of Araneus diadematus have enlarged black palps,
relatively narrow elongated abdomens, and weigh about a fifth of the
adult females. Adult females are characterized by long yellow palps
and a globulous abdomen (Figure 1). Other characteristics of the
males include banding of the legs that is generally darker, a lack of
humps on the abdomen, and a modified second tibia that is stronger
than in females and has short spines (Levi, 1971).

The enlarged palps appear at the end of the next-to-the-last molt,
whitish instead of black, and blacken between the two last molts.
One animal exhibited enlarged palps prematurely two molts before
the last one and four other animals after the last molt, but these
were exceptions.

After the last molt, when they reached sexual maturity and maxi-
mum weight, the males stopped building webs. Sekiguchi (i955)
reported that a male of Araneus ventricosus, in the laboratory, did

1973]

Ramousse — A raneus diadematus

25

Figure 1. Outline of a male (left) and a female (right). Note the
difference of size (female front leg: 16 mm, male front leg: 12 mm), of
weight (female: 144.1 mg, male: 47.0 mg) and the difference of form of
the palps (short and enlarged for the male, long and thin for the female).

not spin a web after its last molt, and that the aggregate glands
become vestigial in the adult males. Prior to this point the involve-
ment of the aggregate glands in the formation of the catching area
of a web was clearly shown (Peakall, 1964). We may suppose that
adult males are unable to spin webs because their aggregate glands
are no longer functional.

The males ate scarcely, even when we attempted to induce prey
catching by placing the flies in front of their mouths. While an
immature male transformed a fly into a small compact ball through
eating; the different parts of the body of a fly abandoned after eating
by a mature male were easily recognizable. Even when they ate, the
mature males used only a small amount of the food available. Males
of Linyphia triangularis Clerck did not require food in the adult
stage, and were still able to mate with females that later produced
fertile eggs. When these males were provided with food, the rate of
prey capture and the rate of food consumption dropped sharply

26

Psyche

[March-June

Figure 2. Body weight of four FG and seven SG littermate males in
set I of Araneus diadematus, hatched in the laboratory from one cocoon
on February 23, 1972. Dashed line: weekly mean body weights of the FG
males. Dotted line: weekly mean body weights of the SG males. Numerals
followed by an arrow indicate the number of animals molting for the last
time during a week. Numerals surmounted on black circles indicate the
number of animals dying during a week. The FG males reached their
maximum weight the 13th week of post-hatching, the SG males reached
their maximum the 29th week of post-hatching. Note that the SG animals
need twice as much time to mature as the FG.

(Turnbull, 1962). We may assume that the adult males, no longer
able to build a web, do not neeed food to fulfill their mating role.

Four males in this study continued to spin webs until they died;
they built webs for a few days after the last molt was recorded, then
stopped building for three or four weeks and generally built a final
web six or seven days before death. These facts suggest that these
four males were not able to go through an additional molt to com-
plete their development. Also, these males presented enlarged palps
only after the last molt recorded which is another confirmation of
thir inability to complete their development.

During the web building period the males are distinct from the
females only between the two last molts (about 3 weeks). This
explains why few studies have been made of the males either outdoors
or in the laboratory.

1973]

Ramousse — A raneus diadematus

27

W eight increase

In each set, the individual weight curves follow two distinct pat-
terns and no in between: a group with an early maximum (FG)
and a group with a late maximum (SG). In set I the course of the
growth of four males with early maxima (between 10th and 15th
week post hatching) was compared to seven males with late maxima
(between 22nd and 33rd week post hatching) (Figure 2). In the
second set the growth of 1 1 males which reached their maximum
weight between the 8th and 16th week post hatching was compared
to three males reaching their maximum weight between the 19th
and 23rd week post hatching (Figure 4). In both sets the SG
animals needed approximately twice as much time to complete the
last molt and to attain sexual maturity as did the FG animals. In
each set the females could be divided into fast and slow growth
groups in the same way as males. Figure 6 shows the body weight of
the FG and SG males and females. The data from the two sets
were combined forming four groups: FG and SG males and females.
The weight gain per day until maturation, in both sets, was sig-
nificantly higher for the FG males than for the SG males (set I:
T = 6,P :: - 0.05; set II: T = 7, P = 0.05).

The mean weight gain per day between the two last molts for
each group was :

set I set II

FG

2.09 mg/d

1.59 mg/d

SG

0.64 mg/d

1. 6 1 mg/d

In each set, every animal showed a weight gain per day significantly
higher between the two last molts than during the preceding period
of observation (Wilcoxon matched-pairs signed ranks test: set I:
N = 9, T = 3, P = 0.02; set II : N feft 13, T = o, P — 0.01).

Frequency of building

Th® mean of webs built per day to reach the last molt were:

set I set II

FG

0.57 web/day

0.49 web/day

SG

0.22 web/day

0.18 web/day

week percentage web building

28

Psyche

[March-June

The FG males had a higher rate of building while they grew
than did the SG males (set I: T = 6, P = 0.05; set II: T = 6,
P — 0.01 ) . The differences in the rate of building appear clearly
on the graphs (Figs. 3 and 5) obtained by plotting the mean fre-
quency of building per week for each group in each set.

The frequency of building is strongly correlated with the amount
of food eaten per day (Kendall rank coefficient; set I: 7 — 0.59,
P = 0.004; set II: 7 — 0.52, P = 0.005). This is the necessary
consequence of the feeding schedule. We might suppose that this
relation occurs in nature. A fresh snare probably increases the
chances of capturing prey.

F G males

'f?j set 1

F G males

[ l 1

- 1 1

1 1

1 1

1 1

f 1 1

1 1 I".

1 1 1 1
r •. ; ; ; .

rj— J -J ;

.... j

— 1 • :

••••••

.....

• ...... . .r"*

! .

8 12 16 20 24

weeks post hatching

Figure 3. Frequency of building of the FG and SG males of set I.
Dashed line: weekly mean of frequency of building for the four FG males.
Dotted line: weekly mean of frequency of building for the seven SG males.
Note the similarity in the pattern between weight increase and web building
frequency. (Compare with Fig. 2.)

1973]

Ranvousse — A raneus diadernatus

29

Figure 4. Body weight of 14 male littermates in set II of Araneus
diadernatus hatched in the laboratory on March 6, 1972. Dashed line:
weekly mean body weight for the 11 FG males. Dotted line: weekly mean
body weight for the three SG males. Numerals followed by an arrow
indicate the number of animals molting for the last time during a week.
The FG animals reached their maximum weight the 12th week post-hatch-
ing, the SG males reached their maximum the 27th week post-hatching,
when the FG males are dead. (Compare with Fig. 2.)

The rate of building :

set 1 set II

FG

0.57 w/d

0.64 w/d

SG

O.31 w/d

0.30 w/d

between the two last molts was significantly higher than the rate of
building during the previous stages of growth in both sets (set I:
N = 9, T = 2, P = 0.01 ; set II: N — 12, T = 1.5, P = 0.01
Wilcoxon test).

What explanations are there for differences in frequency of build-
ing? A multiplicity of factors have been found to have some in-
fluences on web-building: a change from dark to light, a steep rise

30

Psyche

[March-June

Figure 5. Frequency of building of the FG and SG males of set II.
Dashed line: weekly mean frequency of building for the 11 FG males.
Dotted line: weekly mean frequency of building for the three SG males.
Note similarity to Fig. 3.

in temperature following a temperature minimum, weather condi-
tions, barometric pressure, a full silk supply, hunger (Witt, et al .,
1968). In the laboratory, all the spiders were subjected to the same
environmental conditions, therefore the differences in rate of building
should be due to an internal state, such as hunger. There is a gen-
eral agreement in the literature that hunger is a strong drive for
web-building. Heavy feeding is followed by several days without
web-building (Koenig, 1951; Wolf & Hempel, 1951; Wiehle, 1927;
Peters, 1932). The interpretation is that the hunger drive is too low
for releasers like temperature and light to operate. On the other
hand, spiders deprived of food built almost every day (Peters, 1939)
and built webs even at the expense of other body constituents (Witt,
1963b). We may assume that the FG males have a higher level of
hunger than the SG males, which induces a higher rate of building.

Pood consumption

Each time a spider was fed, the fly was weighed before eating.
Since only one or two percent of a fly was rejected by a spider after

1973]

Ra?nousse — A raneus diadematus

3i

eating, we assume that a fly was eaten entirely. The mean quantity
of food consumed per day was :

set I set II

FG

2.44 mg/d

2.06 mg/d

SG

i.44mg/d

1.39 mg/d

The FG spiders ate a significantly higher quantity of food per day
than the SG ones (set I: T = 6, P ~ 0.05; set II : T = 8,
P — O.05 ) . There was a significant difference in the amount of
food consumed per day between FG males of the two sets (T = 8.5,
P = 0.05).

The mean quantity of food eaten between the last two molts was:

set I set II

FG

3-33 mg/d

2.61 mg/d

SG

2.85 mg/d

3.54 mg/d

In each set the mean quantity of food consumed per day between the
last two molts was significantly higher than the mean amount of
food eaten per day during the preceding observation period, (Wil-
coxon test: set I : N = 10, T = o, P = 0.0 1 ; set II: N = 13,
T = 1, P = 0.01).

A relationship exists between the amount of food eaten per day
and the growth rate in both sets, indicating that the growth rate is
a function of the amount of food consumed (Kendall rank coefficient;
set I: y = 0.55, P = 0.01 ; set II : y = 0.60, P = O.OOi). The
foot eaten was used to sustain the basal metabolism, to make silk,
and to build the body of the spiders. A rough estimate of the per-
centage of food transformed into spider tissues was obtained by
dividing the gain of body-weight per day by the quantity of food
consumed per day: the FG males used about 57 % (set I) and 47%
(set II) of the food they ate, while the SG males transformed only
33% (set I) or 32% (set II) of their food into spider tissues. The
FG groups transformed a greater amount of food consumed into
spider tissues than did the SG groups (set I : T = 6, P == 0.05;

32

Psyche

[March-J une

Figure 6. Body weight and number of molts of 25 males (15 FG, 10 SG),
and 25 females (15 FG, 10 SG) from the two sets cocoons of Araneus
diadematus studied. Each line connects mean body weights at one, two,
five, seven and nine months post-hatching. Large black circles: FG females,
large dashed line: early life of SG females; small black circles: FG males,
small dashed line: SG males. Arrows indicate the number of molts to the
time. Note the different growth rates and the related different speed of
maturation in FG and SG males and females, and the similarities for
both sets.

set II: T = 14, P = O.05). As a result of having more food
available for metabolism, an FG male was able to utilize more energy
for other metabolic processes than basal metabolism, such as synthesis
of silk, synthesis of body constituents, etc. This would assure a
larger supply of silk for the FG spiders than for the SG, which
could be an important drive for web-building (Peakall, 1967). The
increased frequency of building in the FG spiders leads to a greater
amount of food consumed which in time results in the rapid weight
gain and growth.

1973]

Ranvousse - — A ranens diadematus

33

Maturation

Between the start of the observations and the time of sexual ma-
turity (last molt) the mean number of molts recorded for each
group was:

set I set II

FG

3.25 molts

3.27 molts

SG

4.50 molts

3.66 molts

The SG males in set I went through a significantly higher number
of molts than did the FG males (T = 12, P = 0.05) and reached
a higher weight (see below). In set II, we had only three SG males
and one of them did not complete its development, this explains the
difficulty to obtain a significant difference between SG and FG ani-
mals in this set.

For set I the mean time of maturation was 81.6 days for the FG
spiders and 202.5 days for the SG spiders. In set II maturation was
reached in a mean time of 78.0 days for the FG males and 163.0
days for the SG ones. The time of maturation was significantly
longer for the SG animals (set I: T = 6, P — 0.05 ; set II:
T = 6, P = 0.01). In addition the time of maturation was sig-
nificantly longer for the SG in the first set than in the second set
(T = 6, P = 0.05) .

The rate of maturation, number of molts divided by the number
of days necessary to complete these transformations, was significantly
higher for the FG males than for the SG males (set I: T — 6,
P = 0.0 1 ; set II: T — 6, P = 0.05).

The Kendall rank coefficient between the gain of weight per day
and the number of molts per day was 0.61 for set I and 0.66 for
set II ( in both P — 0.001). A relationship exists between the rate
of growth and the rate of maturation which is in agreement with
the findings of Deevey (1949) with Latrodectus mactans (Fabri-
cius) and of Benforado and Kistler (1973) with Araneus diadema-
tus. We may assume that the maturation rate is correlated with the
growth rate. The mean length of time in days between two con-
secutive molts was determined. In 3 out of 4 groups, the last inter-
molt was longer than the other intermolts (table 1); for the FG
males, this last intermolt was significantly longer than the earlier
(N — 10, T = 1.5, P = 0.01 ).

34

Psyche
Table i

[March-June

1

2

3 4

set I FG

20.6

14.0

SG

63-3

37-5

46.1 29.6

set II FG

22.7

12.0

SG

26.5

54-0

55.0 17.0

Mean length of time

in days

separating two consecutive molts. The

numerals designate each intermolt and its order in relation to the
final one, i being the last.

Increase in leg-length

The mean length of the first leg as measured on the last molt was :

set I set II

FG

1 1 .0 mm

10.5 mm

SG

14.3 mm

12.3 mm

SG males, which were also heavier, had significantly longer first legs
than FG males after the last molt (set I: T — io, P = o.oi ;
set II : T = 7.5, P = 0.05 ) .

The rate of leg growtdi is given by the ratio of the length gain in
the number of days necessary to obtain this increase of length. The
mean rate of leg growth during the entire observation was:

set I set II

FG

0.183 rnm/d

0.176 mm/d

SG

0.069 rnm/d

0.069 mm/d

The rate of length increase was significantly higher for the FG
males than for the SG males (set I: T — 10, P = 0.01; set II:
T = 7, P = 0.05). This points out the relationship existing be-
tween rate of maturation and rate of lengthening. No correlation
was found between the leg-growth between molts and the length of
time of the intermolt.

Maximum weight

Body weight increased for all males to a maximum at the last
molt, declining from this point onwards. This is in contrast to

1973]

Ramousse — A raneus diadematus

35

female weight increase which continues after the last molt, possibly
due to egg formation. The mean weight was :

set I set II

FG

62.95 mg

57.64 mg

SG

88.52 mg

70.23 mg

The maximum weight reached by the FG males, in both sets, was
lower than for the SG males. This suggests that the maximum
weight may be a function of the duration of development. In that
case, rapid maturation would occur at the expense of weight growth.

No correlation between initial weight and final weight was found
in contrast to the findings of Benforado and Kistler (1972). A
relatively small difference in initial weights and the low accuracy of
the weights may explain the contrast.

Mating

The FG males of set I matured 81.6 days post-hatching (p-h.)
and those of set II 78.0 days p-h. The FG females reached ma-
turity 229 days p-h. in set I and 104 days p-h. in set II. In the
first set all the FG males were dead before any of the FG females
were mature, preventing them from mating. In the second set, the
FG males survived 71 days after the last molt, so that some of them
were still alive when the first FG females matured. But the females
accepted the advances of the males only 60 days or more after the
last molt, preventing the FG males from mating with their sisters.
In summary, the FG males of both sets were sexually mature too
early to mate with any of the females of the same set.

The SG males reached maturity 202 days p-h. in set I and 163
days p-h. in set II. At this time the FG females of set II were
already mature (104 days p-h.) and those of set I were almost ma-
ture (229 days p-h.), as well as the SG females of set II (223 p-h.).
In these conditions the SG males of both sets may have been able to
mate with the FG females of their own set or the other set.

Each of the nine SG males, when they were an average of 300
days p-h., were brought into the presence of three different females.
All the males seemed to behave in the same way, but only three of
them mated successfully with a single female, and one with two
different females. These successful males were the biggest of the
SG males. One male of each set was able to mate with a female of

3 6

Psyche

[March-June

his own set. Only the FG females of the set I accepted the males,
while both FG and SG females of set II accepted the males. How-
ever, the small number of males limited the number of trials and
did not permit us to know statistically which of the females (FG
or SG) were the most successful in mating.

The FG males cannot mate with females of their own set, but we
may assume that they can find females of other sets in a natural
habitat which are mature at the same time. The SG males can
mate with the FG females of their own set, permitting limited in-
breeding. This is merely a possibility since different sets in nature
may have mature females at the same time.

Comparison of the webs of SG and FG spiders

FFeb changes during development :

Webs for the FG and SG males were compared on the basis of
the spiral area, mesh size and thread length.

The spiral area of all webs built by males in both groups and both
sets showed a general increase until reaching a maximum area during
the period between the last two molts. The spiral area decreased in
size thereafter. Four males mentioned earlier, who did not follow
the general web-building pattern (they built webs until they died),
showed no decrease in spiral area in their final webs. This further
supports speculation that they died before achieving full development
through a complete series of molts. In contrast, the spiral area of
the females of Araneus diadematus and Neoscona vertebrata in-
creased until the last three months, after which time the catching
area does not change significantly (Witt & Baum, i960). The
catching area of the females of the golden garden spider, Argiope
aurantia , showed a growth and decline, the peak size coinciding
roughly to the time of last molt and sexual maturation (Reed, Witt
& Scarboro, 1969).

An essentially upward linear growth in mesh size throughout the
lifetime occurred for 11 of the males studied. For the 12 other
males the mesh size increased until reaching a plateau during the
last intermolt. Witt, Rawlings and Reed (1962) have pointed out
that the mesh size of the female webs of Araneus diade?natus show
also an increase until the last molt, and then reach a plateau. But
Argiope aurantia shows a linear growth in mesh size throughout the
lifetime (Reed, Witt & Scarboro, 1969).

The thread length fol’ows the same pattern as the two other pa-
rameters with a peak during the last intermolt and then a decrease.
Argiope aurantia and Araneus diadematus females have been shown

1973]

Ranvousse — A raneus diadematus

37

Figure 7. Web built by a young male of Araneus diadematus. Young
FG and SG males built webs with the same characteristics, small and fine-
meshed. The vertical white lines of the scale are spaced 20 mm apart.

38

Psyche

[March-June

to change according to the same pattern for thread length (Reed,
Witt & Scarboro, 1969; Witt, Rawlings & Reed, 1972). The de-
crease of the thread length may be due to the thickening of the
threads as the weight of the spider increases (Christiansen et al.,
1962). The males follow the same pattern of web-changes as the
females of the same species, except for the catching area. The gen-
eral effect is that young Araneus diademcitus males build small,
fine-meshed webs (Fig. 7) ; and during the onset of the last inter-
molt build large, wide-meshed webs; males at the end of the last
intermolt, without changing weight and leg-length significantly, build
medium meshed webs. Therefore it appears that web size cannot
simply be explained by the spider’s bodily dimensions (Witt, Rawl-
ings & Reed, 1972; Reed, Witt & Scarboro, 1969).

Comparison of the webs of the FG and SG males
at the same age

All the webs photographed between the 9th and 12th weeks post
hatching of five FG males of both sets were measured. So were the
webs of eight SG males of both sets during the same period of time.
All these spiders were at the same age, but the five FG spiders were
almost mature and the eight SG males were 100 days before reach-
ing maturation. Table 2 shows the figures for spiral area, mesh size,
thread length and a measure of the regularity of the spacing of the
threads (standard error of median mesh size North). The two
measures of the body, and the four measures of the web show sig-
nificant differences between the FG and SG males, with the excep-
tion of the variance of the mesh size: the webs of the two groups
show similar regularity.

Table 2

FG

SG

t

P

Body weight

50.1 ± 19.0 mg

22.6 ± 6.4 mg

3.58

0.005

Leg length

9.1 ± 1.70 mm

6.68 ± 1.48 m

2.49

0.05

Spiral area

42,138 ± 9,883 mm2

13,956 ± 8,178 mm2

4.88

0.001

Mesh size

58.60 ± 6.96 mm2

34.75 ± 10.08 mm2

4.05

0.005

Thread length

16,066 ± 4,111 mm

6,950 ± 3,282 mm

4.07

0.005

SE median mesh
size North

0.114 ± 0.030

0.162 ± 0.062

1.47

Measures of webs of the FG and SG males at the same age. Only the
regularity measures in the last line are not significantly different.

1973]

Ranvousse — A raneus diadematus

39

Table

3

FG

SG

t

P

Body weight

60.14 ± 11.56 mg

79.37 ± 18.07 mg

0.11

Leg size

10.75 ± 1.22 mm

13.60 ± 1.41 m

4.90

0.001

Spiral area

33,931 ± 10,131mm2

28,766 ± 6,559 mm2

1.15

0.400

Thread length

15,546 ± 2,768 mm

10,425 ± 1,715 mm

3.66

0.005

Mesh size

46.46 ± 9.64 mm2

70.73 ± 15.95 mm2

3.99

0.005

SE median mesh
size

0.900 ± 0.022

0.293 ± 0.068

8.78

0.001

Measure of webs of the FG and SG males at comparable stage of maturity.

It is neither possible to relate the regularity measures to leg length
— the FG’s legs were significantly longer than the SG’s legs — nor
to maturation since the FG spiders were at the last stages of matura-
tion and the SG males two or three stages before. We may assume
that the size of the males’ webs but not their regularity is a function
of the rate of growth and in consequence of the body dimensions.

Comparison of the FG and SG males’ webs
at the same stage of maturation

All the FG webs photographed during the last stage were com-
pared with all the SG webs photographed about ioo days later,
during their last stage; this compares webs built at different times
but comparable stages of maturity. Table 3 gives the figures for
body weight, leg size, spiral area, thread length, me^h size and the
variance of the mesh size. The webs of the FG (lighter) males had
a spiral area larger, a significantly longer thread, smaller mesh size,
and a higher regularity than the webs built by the SG males at
comparable maturity (Figs. 8 and 9). The longer length of thread
produced by the FG males than the SG males indicates that they
have a better supply of silk or thinner thread. This may be sup-
ported by the higher amount of food eaten per day and the higher
rate of utilization of the food by the FG than SG males. The larger
mesh size and irregularity of the SG webs is related to the larger
body dimensions of these animals. The difference between the dimen-
sions of the bodies of the FG and SG males coincides with a longer
duration of development for the SG than for the FG males. We
may assume that the regularity of spacing the spiral thread is related
to the duration of the development in the two groups of males with
different rate of growth, and is related to maturation within a group
having a homogenous growth rate.

40

Psyche

[March-June

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white lines being originally spaced 20 mm apart.

1973]

Ramousse — A raneus diadematus

4i

Mortality

The mean mortality in each group was :

set I set II

FG

1 6 1. 6 days post-hatching

146.5 days post-hatching

SG

3 dead (mean 329.0 days
post-hatching)

3 still living (342.0 days
post-hatching)

308-6 days post-hatching

The SG males lived significantly longer than the FG males (set I:
T = 10, P — 0.01 ; set II: T gg 4, P = 0.05). Rapid growth
occurs at the expense of endurance which is in agreement with the
findings of Bonnet (1935), who found that the spiders’ lives short-
ened with an increase in food supply, and the findings of Reed and
Witt (1972), who found that the FG females of A raneus diade-
matus lived shorter than the SG females.

In our laboratory, the males matured from July 1972 to January
1973. But in North America males of Araneus diadematus can only
be found in September and early October in the Boston area, (Levi,
1971) and in Southern France in August and September (Bonnet,
1935). Nevertheless some authors found Araneus diadematus in the
field during different seasons even in winter, Bertkau (1885) in
Germany and Termeyer (1791) in Italy. Millot (1926) also ob-
tained, in the laboratory, the survival of young Araneus diadematus
during the winter; they completed their development the following
spring. The biological cycle of Araneus diadematus varies according
to the environmental conditions it goes through, and a low rate of
feeding with the stable laboratory conditions could allow a longer
lifespan for the spiders reared in the laboratory than in the field.
In that case, the lengthening of the development merely emphasizes
the difference betwen FG and SG animals.

The FG males survived an average of 80 days after the last molt
in set I and 71.4 days in set II. These males grew and built webs
approximately half their lives, then sought out mates. We can note
equivalent facts for the SG males with their relatively long time
scale.

42

Psyche

[March-June

General Discussion

The males grow until the last molt, at which time they attain
their maximum weight; weight decreases slowly thereafter. The
females have a distinctly different course of growth; their weight
increased long after the last molt, generally until they lay a cocoon.
The males mature more rapidly than the females, but the females
grow bigger than the males and live longer.

The females and the males of each of the two sets of Araneus
diadematus studied are clearly divided into FG and SG. The FG
males are characterized by a higher frequency of building, rate of
food consumption, rate of weight increase, rate of leg growth, rate
of maturation as well as a smaller number of molts than the SG
males, significant only for set I in the last instance.

The positive correlation between the rate of building and the rate
of food consumption for all males must be expected, since the spiders
were fed only when they had built a web. The different frequency
of building may be explained as a lower threshold for web-building
through hunger in the FG than in the SG spiders. The number of
prey captured is a function of behavior mechanisms of the spider and
potential prey; among the former are the stimuli that induce the
spider to attack, the efficiency of this attack, and also a number of
other variables such as web-site, web-characteristics, and frequency of
building. The hunger stimulus which induces both, the attack and
web-building, has a lower threshold for the FG than for the SG
spiders, suggesting that in a natural habitat the FG males would be
able to capture and eat more food than the SG spiders. In addition,
the usual effect of genes on animals with rigid patterns is to alter
behavior in a quantitative, rather than a qualitative, fashion (Mann-
ing, 1967). The environmental conditions being the same for all
the animals, the difference in threshold of hunger may be the conse-
quence of different genotypes.

A relationship between the rate of food intake and the rate of
growth indicates that the food was converted into spider tissues, in
addition to maintain basal metabolism and to support the necessary
activities like prey catching. The percentage of food converted into
spider tissue was higher for the FG males than for the SG males,
explaining the different growth rates. The same mechanism could
provide a more ample supply of silk for the FG than for the SG
males, which is suggested by the analysis of the web dimensions of

1973]

Ramousse — A raneus diadejnatus

43

the two groups. Hunger is an important drive for web-building and
prey catching, which in turn increases the amount of food available
to the spider. As a consequence, a good supply of food permits the
spider to use more energy to metabolize tissues and silk, and a full
supply of silk lowers the threshold of web-building. So, the fre-
quency of building may be controlled by a changed feed-back between
hunger and amount of food eaten.

A strong relationship exists between the rate of growth and the
rate of maturation. But the number of molts was not constant, nor
was the time separating two successive molts. The FG males went
through fewer stages than the SG males (significant only in set I)
and in less time. The rapid increase of the body weight of the FG
spiders seems to force these spiders to change more often their rigid
skins. Ecdysis is a crisis that requires extra energy to overcome; the
heavy eating FG spiders could accumulate this extra energy in the
form of reserves more rapidly than the SG spiders. The differential
maturation may be attributed to nutrition. But nutrition is a func-
tion of the amount of food eaten controlled through appetite, pro-
ficiency in prey-catching, and web-building frequency. Similar rela-
tions must explain the differences in development between males and
females as well as between females of the same set.

Different schedules of feeding result in differential growth and
maturation for spiders of the same set (Benforado and Kistler,
1972), suggesting that the amount of food eaten is a determinant
factor. But with the same amount of food available, the spiders of
the same set show different growth rates and maturation rates (Reed
& Witt, 1972). This indicates that prenatal or genetic conditions
control the development and maturation. In our study, the spiders
could choose the food quantity they need through their behavior.
When the spiders are in identical environmental conditions, we may
assume that the difference in behavioral patterns present at hatching
time probably are genetically determined. One pattern induces some
spiders (FG) to capture and eat more food than other spiders, and
in turn this large amount of food eaten by these spiders, increases
their rate of development and maturation. The rapid growth in the
two sets occurs at the expense of endurance and maybe weight in-
crease, the FG males are short livers and small weighers. The short
life-span of the FG spiders prevent them from mating with females
of the same set, while some SG males live 'long enough to mate with
the FG females of their own set.

44

Psyche

[March-June

Poetsch has shown (1963), that cocoons of a. single species of
spider hatch at different times. This presumably provides an ad-
vantageous distribution of egg-production over a period of time. The
two cocoons studied hatched at different times, and the males of
set II, which hatched fifteen days later, grew faster than the males
of set I (significant only for the SG males). Between the sets the
rapid growth occurs also at the expense of endurance and maybe
weight increase. Differential growth occurs between the sets as well
as within the sets, favoring the distribution of mature animals over
a period of time. This suggests that during the favorable season
mature males and females can mate and produce cocoons at various
times, providing the species with a better chance to survive any
drastic crisis due to the environment.

The relative quick maturation of males favors mating between
animals of different sets and of different behavior instead of inbreed-
ing. This allows the species to conserve a genetic pool with high
selective potentialities, Dobshansky, 1951).

During development the last intermolt was distinct from the other
stages. During this period the males built more webs, ate more food
per day, and grew faster than during the other stages. The time
separating the last two molts was generally longer than the time
separating any other two successive molts. Sexual differentiation
also took place during this period, and we may assume gametogenesis
too. This would explain why the males need more food and a longer
time to complete the last stage of development. The importance of
the requirements during this time must make it the most difficult
for the males.

Summary

The offsprings from two cocoons of Araneus diadematus , hatched
at different times and placed in individual frames, were studied in
the laboratory during the life-span of the males. During this time,
the characteristics of the body (weight and size), the frequency and
the parameters of the webs, the number and date of the molts, and
the amount of food eaten were recorded for each animal. The
spiders could choose their feeding schedules through their building
behavior.

The males built and increased their weight only until the last
molt, in contrast to the females which continued both building
and increasing their weight long after the last molt. During the

1973]

Raiwousse — A rcineus diadematus

45

building period the males were distinctly different from the females
only during the last stage. The males lived shorter and grew less
than the females. The last intermolt was distinct from the other
stages: the males built more webs, ate more food, grew faster than
during the other stages.

Two different rates of development appeared among the males
of each set, determining a fast and slow growing group. The fre-
quency, the amount of food eaten, the rate of weight increase and
the rate of maturation were higher for the fast growers than for the
slow growers. As a consequence of the rapid growth, the life-span
of the fast growing males was shorter and the maximum weight was
lower (but not significantly) than for the slow growing males.
Hunger and amount of food eaten determined the different growth
rates and related maturation rates; a lower threshold is supposed
for the fast growing males than for the slow growing males, and
may be the consequence of a genetic difference. Maturation would
be controlled by different patterns of behavior determined on a
genetic level.

The differential maturation, which occurs within animals of a set
and between sets, results in a distribution of mature animals over
various times of year. The relative quick maturation prevents the
fast growing males from mating with a female of the same set, but
limited inbreeding is possible between the slow growing males and
the females of the same set. A potential high survival of the species
is assured by the dispersion of the individuals of one set and the dis-
persion of the sets over different seasons.

ACKNOWLEDGEMENTS

This work was carried out in the laboratories of the Division of
Research, North Carolina Department of Mental Health and was
supported by Grant Number GB-25274 from the National Science
Foundation to Dr. Peter N. Witt. The author gratefully ack-
nowledges the assistance of Dr. Witt during all stages, the assistance
of Mrs. Mabel Scarboro for all technical and laboratory work, Mrs.
Rubenia Daniels for her administrative assistance, and of Dr. John
O. Rawlings with whom the statistical tests used were discussed.

References cited
Benforado, J. and Kistler, K. H.

1973. Growth of the orb weaver, Araneus diadematus, and correlation
with web measurements. Psyche, 80: 90-100.

46

Psyche

[March-June

Bertkau, Ph.

1885. Ueber den Saisondimorphismus und einige andere Lebenser-
scheinungen bei Spinnen. Zool. Anz. 8 : 459-464.

Bonnet, P.

1935. La longevite chez les Araignees. Bull. Soc. Etomol. de France.
40: 272-277.

Christiansen, A., Baum, R. and Witt, P. N.

1962. Changes in spider webs brought about by mescaline, psilocybin
and an increase in body weight. J. Pharmac. ex. Ther. 136:
31-37.

Deevey, G. B.

1949. The developmental history of Latrodectus mactans (Fabr.) at
different rates of feeding. Amer. Midi. Nat. Notre Dame, 42:
189-219.

Dobzhansky, T.

1951. Genetics and the origin of species. Columbia University Press.

Eberhard, W. G.

1971. The ecology of the web of Uloborus dwersus (Aranea: Ulobori-
dae). Oecologia (Berlin), 6: 328-342.

Enders, R.

1972. Web site selection by Argiope aurantia Lucas and other orb
weaving spiders (Araneidae). Thesis, N. C. State University,
Raleigh.

Koenig, M.

1951. Beitrage zur Kenntnis des Netzbaus orbiteler Spinnen. Z. Tierp-
sychol. 8 : 462-493.

LeGuelte, L.

1966. Structure de la Toile de Zygiella-x-notata Cl. (araignees, Agri-
opidae) et quelques facteurs qui regissent le comportement de
1’araignee pendant la construction de la toile. These, Nancy.

Levi, H. W.

1971. The diadematus group of the orb-weaver genus Araneus north
of Mexico (Araneae: Araneidae), Bull. Mus. Comp. Zool.,
141 (4) : 131-179.

Manning, A.

1967. Genes and the evolution of insect behavior. Jerry Hirscb-
McGraw-Hill (Behavior-genetic analysis).

Millot, J.

1926. Contribution a 1’ histophysiologie des Araneides. Bull. Biol. Fr.
et Belg., Supp. 8: 1-238.

Peakall, D. B.

1964. Composition, function and glandular origin of the silk fibroions
of the spider Araneus diadematus Cl. J. Exp. Zool., 156: 345-350.

1969. Silk synthesis, mechanism and location. Amer. Zoologist, 9: 71-79.

Peters, H. M.

1939. Uber das Kreuzspinnennetz und seine Probleme. Naturwissen-
schaften 47: 776-786.

Potzsch, J.

1963. Von der Brutfiirsorge heimischer Spinnen. Wittenberg, Ziemsen.

Reed, C. F. and Witt, P. N.

1972. Growth rate and longevity in two species of orb-weavers. Bull.
Brit. Archnol. Soc. 2(6) : 111-112.

1973]

Raiwousse — A rcineus diadematus

47

Reed, C. F., Witt, P. N. and Jones, R. L.

1965. The measuring function of the first legs of Araneus diadematus
Cl. Behavior 25 : 98-119.

Reed, C. F., Witt, P. N. and Scarboro, M. B.

1969. The orb web during the life of Argiope aurantia (Lucas).
Devel. Psychobiology 2(2): 120-129.

Reed, C. F., Witt, P. N., Scarboro, M. B. and Peakall, D. B.

1970. Experience and the orb- web. Devel. Psychobiology 3 (4): 251-265.

Sekiguchi, K.

1955. Differences in the spinning organs between male and female
spiders. Sci. Rep. Tokyo Kyoiku Daigaku, 8: 23-32.

Termeyer, R. M. de

1791. Richerche e sperimenti sulla seta dei Ragni e sulla loro gen-
erazioni. Scelte d’opusculi interessanti 3 : 288.

Turnbull, A. L.

1962. Quantitative studies of the food of Linyphia triangularis Cl.
(Aranea: Linyphiidae) . Canad. Entomologist, 94(12) : 1233-1249.

White, C.

1952. The use of ranks in test significance for comparing two treat-
ments. Biometrics, 8: 33-41.

Wiehle, J.

1927. Beitrage zur Kenntnis des Radnetzbaues der Epeiriden, Tetrag-
nathiden and Uloboriden. Z. Morpholog. u. Okolog. der Tiere, 8 :
468-537.

Wolff, D. and Hempel, U.

1951. Versuche liber die Beeinflussung des Netzbaues von Zilla-x-
notata durch Pervitin, Scopolamin and Strychnin. Z. vergl.
Physiol., 33: 497-528.

Witt, P. N.

1963a. Interrelationships between web-building behavior and amount of
thread material in the spider Araneus diadematus Cl. Proceed,
of XVI Intern. Cong, of Zool.

1963b. Environment in relation to behavior of spiders. Arch of environ.
Hlth., 7: 4-12.

Witt, P. N. and Reed, C. F.

1965. Spider web-building. Measurements of web geometry identifies
components in a complex invertebrate behavior pattern. Science,
149: 1190-1197.

Witt, P. N., Reed, C. F. and Peakall, D. B.

1968. A spider’s web. Problems in regulatory biology. Springer-Verlag,
New York.

Witt, P. N., Rawlings, J. O. and Reed, C. F.

1972. Ontogeny of web building behavior in two orb-weaving spiders.
Am. Zoologist 12: 445-454.

ANNOTATIONS ON TWO SPECIES OF
LINYPHIID SPIDERS

DESCRIBED BY THE LATE WILTON IVIE*

By P. J. van Helsdingen

Rijksmuseum van Natuurlijke Histone, Leiden, Netherlands

In December 1966 a short paper by Wilton Ivie was published in
the Journal of the New York Entomological Society. In that paper
the author described and illustrated two new species of Linyphiidae,
Taranucnus durdenae from Pennsylvania, and Troglohyphantes ho->
koko, from Ontario. The latter was also recorded from the state of
New York. In the course of my study of the North American Liny-
phiidae it appeared that both were junior synonyms of North Amer-
ican species, as will be discussed below. In a vast faunal region as
North America such mistakes are quite understandable and not easily
avoided, especially when a species originally was described in a rather
ill-chosen genus, or when the older description of the species was
based on the opposite sex.

While correcting the names of the two species, I take the oppor-
tunity to discuss their distributions, habitats, and affinities wherever
I have something of presumed importance to add. To avoid con-
fusion the two species will be discussed separately under their correct
names.

T aranucnus ornithes (Barrows)

Figures 1-10

Lepthyphantes ornithes Barrows, 1940, Ohio J. Sci., 40: 134, figs. 7-7C
(descr. $ $ ; Ohio, Tennessee). — Vogel, 1967, Mem. Amer. Ent. Soc.,
23: 93 (catal.) .

Taranucnus durdenae Ivie, 1966, J. New York Ent. Soc., 74: 224, figs. 1-5
(descr. $ $ ; Pennsylvania). — Vogel, 1967, Mem. Amer. Ent. Soc.,
23: 100 (catal.). Vogel, 1968, J. New York Ent. Soc., 76: 102 (Penn-
sylvania). NEW SYNONYMY.

Types. — Lectotype cf of Lepthyphantes ornithes , by present desig-
nation, from Sugar Grove, Ohio. There are two $ paralectotypes,
from Sugar Grove, Ohio, and the Great Smoky Mountains National
Park, Tennessee. All three specimens are in the Barrows Collection
at the Ohio State University, Columbus, Ohio, and were examined.

*Based on research done at the Museum of Comparative Zoology and
published with a grant from the Museum of Comparative Zoology.

Manuscript received by the editor March 25, 1973

48

1973]

van Helsdingen — Linyphiid Spiders

49

Figure 1. Map showing distributions of Taranucnus ornithes (Barrows)
(•) and Oreonetides recurvatus (Emerton) (★).

The cf holotype and $ paratype of Taranucnus durdenae, from Rec-
tor, Pennsylvania, should be in the American Museum of Natural
History, New York, but they were not examined by me.

In the Barrows collection, through kind cooperation of Dr. C. A.
Triplehorn, one vial with Lepthyphantes ornithes was located. This
vial contained one and two ? specimens, together with a label
referring to the Ohio locality. A second vial, with the recorded ?
from the Smoky Mountains, could not be found. It appears, that
the single $ specimen from the latter locality has been added to the
Ohio material. The original description mentions one $ from each
locality (page 134, ninth line from bottom), while now two females
are present in the Ohio vial. The description does not give any clue
as to the sizes of the different specimens, and the mixed series there-
fore cannot be separated again in accordance with the origin of the
specimens.

Name. — I vie dedicated his new species to Beatrice Vogel Durden.
Barrows did not give any explanation of the derivation of the name
ornithes , but his remarks on the female epigyne gives us a key: “The
epigynum (Fig. 7A) appears as if made up of two gasteropod shells
with the large openings toward the abdomen. When seen from be-
hind the two parts appear as two bird heads placed beak to beak/'
The name therefore appears to refer to the two birds, revealed in a

50

Psyche

[March-June

posterior view of the epigyne (' ornis is Greek for bird, ornithes is the
plural form). It is, of course, pure coincidence that durdenae, de-
rived from the name Beatrice Vogel obtained through her marriage,
has to be discarded in favor of the older ornithes, a name that still
links the species with Beatrice Vogel, but now through the plural
form of her maiden name (Vogel is the Dutch and German word
for bird, ornithes is Greek for birds).

Distribution. — The number of available specimens has consider-
ably increased during the years. The distribution (see map, Fig. i)
now comprises, beside Pennsylvania, Tennessee and Ohio, also North
Carolina, W. Virginia and Virginia. The species is probably not
rare, but, in my own experience, is not easily collected because of its
concealed habitat. Barrow’s Ohio specimens came from “under a log
in a wooded ravine” (October), Ivies specimens from Pennsylvania
were collected in July, and so was Vogel’s specimen, which came
from the same locality in Pennsylvania. No other data, on the habitat
are available from literature. I have collected a fair series (21? 2cJ )
in the Great Smoky Mountains National Park along the trail to the
Alum Cave Bluffs. This trail leads off the road from Sugarlands to
Newfound Gap in northern direction to Mount Le Conte, and is on
the Tennessee side of the park, though close to the North Carolina
line.

At the end of nearly three weeks of fruitless search for this species
at both sides of the Smoky Mountains Range, we found a few speci-
mens on what was planned to be our last day in this area. The next
day we returned to the same spot and added a few more specimens
to our collection. The species was found to inhabit small cavities and
crevices in the steep rocky sides of the trail, and also in the dark hol-
lows under and between the roots of trees. The forest at this height,
between 1100 and 1300 m, has a heavy undergrowth of Rhododen-
dron spec, and Dog-hobble (Leucothoe fontanesiana). The spiders
were very difficult to collect. Even when we knew where to find
them, many escaped by retreating from the entrances of their little
caves into dark and impenetrable depths.

The distribution, as presented in Figure 1, is based on specimens
examined by myself as well as on data supplied by my colleague and
friend Dr. William A. Shear. The collecting dates of the various
samples range from May until October. There is only one other
mention of the exact circumstances of collecting: William Shear
collected i? by sweeping tall weeds near Athens in West Virginia,
a situation that does not agree with the habitat described above
(which may have been delimited too rigidly).

1973]

van Helsdingen — Linyphiid Spiders

5i

Figures 2-6. Taranucnus ornithes (Barrows). Figs. 2-3, epigyne, ven-
tral and dorsal aspect, respectively. Fig. 4, vulva, route to be followed by
embolus indicated by consecutively numbered arrows. Fig. 5, tegulum and
median apophysis ( ma ) of male palp. Fig. 6, median apophysis, dorsal
aspect.

52

Psyche

[March-June

The genus Taranucnus is represented in Europe by the type-spe-
cies, T. setosus (O.P.-Cambridge) , which I have collected from sev-
eral localities in The Netherlands, and by T. bihari Fage, a troglo-
biontic species from Rumania, which I do not know. A diagnosis of
the genus, as based on setosus and \ornithes, reads as follows.

Small spiders (3.2 mm or less). Cephalothorax not much longer
than wide. Eyes large, with PME closer to PLE than to each other.
Chelicerae with stridulating files, dorsal margins with three teeth.
Legs slender and very long (femur I ca. 2 times length cephalo-
thorax, tibia I even slightly longer) . Legs spinose, including femora
and metatarsi. Metatarsus IV without trichobothrium, Tm I 0.15-
O.25. Abdomen with pattern, composed of blackish bars and areas
but without white pigmented spots. Male palp with a short tibia, a
rather flat and broadly rounded cymbium, which has a strongly modi-
fied proximal part, and with a long, thin embolus; the embolus is
supported by a well-developed embolic membrane. Epigyne with
membraneous, coiled ducts leading to small receptacula; no socket or
semi-covered depression visible for the reception of an apophysis of
the male palp.

Both species prefer dark, protected places for a habitat: T. ornithes
in crevices and cavities, T. setosus under overhanging vegetation (e.g.
thick layers of heath) at the border of fens. The habitat of T. bihari
would very well fit into this picture.

T aranucnus clearly fits into the tribe Linyphieae and seems very
close to Labulla.

T aranucnus ornithes and T. setosus j beside their occurrence on dif-
ferent continents, differ from each other mainly in the genitalia. Also
ornithes seems to be the smaller species with more slender legs and
a slightly more proximal Tm I.

The descriptions of T. ornithes by Barrows and Ivie can be sup-
plied with the following data.

Measurements (in mm). Male, total length 2.0-2. 7, length cephalo-
thorax 1. 05-1. 3, width 0.9-1.05. Female, total length 2. 2-3.0, length
cephalothorax 1.05- 1.2, width 0.85-1.0.

Ratio length to width of cephalothorax ca. 0.85, which is high if
compared with other, related genera (cf. Linyphia 0.65-0.75, Neriene
°-55~o.75). Eyes large (diameter PME 0.09 mm) and close to-
gether, the PME one-half diam. apart, and closer to each other than
to PLE. AME not much smaller than PME. Chelicerae with dis-
tinct stridulating files; three teeth in dorsal and three in ventral row.

Legs very long and slender: femur I 2 times as long as cephalo-

1973] van Helsdingen — Linyphiid Spiders 53

Figures 7-10. T aranucnus ornithes (Barrows), male palp. Fig. 7, lateral
aspect. Fig. 8, radix (r) with embolus (e). Fig. 9, paracymbium (pc)
and basal section of cymbium, showing modifications, lateral aspect. Fig.
10, cymbium, dorsal aspect.

54

Psyche

[March-June

thorax ( cf ) or slightly less ($), tibia I 22-24 ( 6 ) or 20-22 ($)
diameters of segment long. Tibia I (without patella) slightly longer
than femur, metatarsus shorter than tibia. Femora I-III with a d-
spine on one-fourth of length, femur I with an additional l'-spine.
All tibiae with the usual d-spines (basal d'^-spine1 at 0.3-0.35) ;
tibia I with a pair of v-spines, a 1' and a l"-spine, and a whorl of
apical spines; tibia II with a v" and a l"-spine only, III and IV with-
out ventrals or laterals, but apical spines present on all tibiae. Meta-
tarsi all with a single d-spine on O.20-0.25. All spines long (3 times
diameter of segment or more) and thin. Tm I 0.16-0.20.

Abdomen as depicted by Ivie (1966: fig. 5), characterized by the
inverted heart-shaped dark grey spot dorsally, followed by chevron
and cross-bars of the same colour.

The malp palp was depicted and described at some length by Ivie
(1966: 224, figs. 3-4). I add the following remarks to his observa-
tions (Figs. 5-10). Patella and tibia short and simple, the patella
bearing a strong dorsal spine, the tibia with a number of hardly
thickened spine-hairs. Cymbium (Figs. 9-10) with complexly modi-
fied basal parts, but with a simple, elbow-shaped paracymbium with
the tip of the free arm widened into a flat, rounded plate; cymbium
proper with two sclerites, a mesal and a lateral one, which are more
sclerotized than the distal part of the element, and which enclose,
together with the distal part, a deep dorsal depression of the cym-
bium ; mesal sclerite distinctly connected without seam along the
sclerotized mesal brim of the cymbium, lateral sclerite apparently
with membraneous connections only. Median apophysis (Figs. 5-6,
ma) with slender base but broadening into a flat and thin plate-like
structure; no tooth or sharp tip; spermduct leaving element at the
inside of the curvature. Radix (Fig. 8, r) a short, centrally situated
element, spermduct running through the element and showing a dila-
tion in the middle (Fickert’s gland?). Embolus (Fig 8, e) with firm
base, long and tapering to a thin, thread-like tip. Embolic membrane
a flat but twisted membraneous structure with narrowly upturned
brim, arising from connecting membrane between median apophysis
and radix.

Epigyne (Figs. 2-3) showing at either side a chitinous roof over
the entrance of the duct, mesally separated by a narrow incision.
Vulva (Fig. 4) showing the spirally coiled ducts, which run in loops
in anterior direction but then turn backward again toward the thick-

JAs usual, the directions of the individual spines are noted by means of
single or double accents, e.g. d' for pro-dorsal, d" for retro-dorsal, l' for
pro-lateral, l" for retro-lateral, etc.

1973]

van Helsdingen — Linyphiid Spiders

55

walled portions close to the receptacula, the latter situated against the
posterior wall at the outside of the entrances. Receptacula small.

It is clear that the embolus is supported by the embolic membrane
in the unexpanded palp, -and it is not unlikely that it also guides and
supports the long embolus during the difficult process of introducing
the long and flexible element into the vulva. The whole palp may
find firm support against the epigyne by means of the intricate dor-
sal modification of the cymbium. The median apophysis does not
show any hook-shaped parts (cf. Linyphia Neriene , etc.) and, at
the most, may serve the purpose of supporting the functioning palp
during copulation by being pressed with its broad and flattened
apical portion against the epigyne. The embolus of T. ornithes does
not have the small, toothed apophysis at the base of the embolus as
shown in the figures of T. setosus by Merrett (1963: 382, fig. 39).

Oreonetides recurvatus (Emerton, 1913)

Figures 1, 11-17

Bathyphantes recurvatus Emerton, 1913, Trans. Connecticut Acad. Arts Sci.,
18: 218, pi. 2 fig. 8 (descr. $, Vermont). — Ivie, 1969, Amer. Mus.
Novit., 2364: 7 (= Oreonetides r.) .

Aigola recurvata ; Crosby, 1937, Proc. Biol. Soc. Washington, 50: 40, pi. 1
fig. 10 (descr. $, New York).

Troglohyphantes kokoko Ivie, 1966, J. New York Ent. Soc., 74: 226, figs.
6-7 (descr. $, Ontario; New York). NEW SYNONYMY.

Types. — cf holotype of Bathyphantes recurvatus from Gore
Mountain, Norton, Vermont, in MCZ (examined). ? holotype and
9 paratype of Troglohyphantes kokoko from Ko-ko-ko Bay, Lake
Timagami, Ontario, reported to be in the AMNH (not seen).

Additional material. — The specimens from Mt. Whiteface, New
York, mentioned by Crosby (1937) have been examined in the
AMNH. More recently collected specimens ( i9 3 cf ) come from
George Lake, Alberta, Canada (CNC), 20.IX-1.X.1966 (Fig. 1).

Oreonetides recurvatus is a small species with long, slender legs,
which are well provided with spines on femora, tibiae, and meta-
tarsi. Tibia I has, beside the dorsal spines, a 1', a 1" and 2 v-spines.
The chelicerae have three teeth in the dorsal row. The abdomen
shows a dorsal pattern of grey cross-bars. Taking all together, it
is not very likely that the present species belongs to Oreonetides , but
it is maintained there for the time being. The genus is due for re-
vision, and it does not seem appropriate to create a new genus for
recurvatus here without having studied the other species presently
residing in Oreonetides.

56

Psyche

[March-June

Figures 11-14. Oreonetides recurvatus (Emerton). Fig. 11, epigyne, dor-
sal aspect. Figs. 12-13, vulva, ventral (12) and dorsal (13) aspects. Fig.
14, radix (r), embolus ( e ) and embolic membrane (em) of male palp,
mesal aspect.

57

1973] van Helsdingen — Linyphiid Spiders

A first attempt to revise Oreonetides has been published recently
(Saaristo, 1972). The paper contains a good characterization with
excellent figures of the type-species, O. vaginatus (Thorell), but the
many other, mostly Nearctic, species are not included. The diagnosis
of the genus, therefore, might be too narrow to fit the other species,
though it very well may be necessary to divide the genus into a num-
ber of smaller units. The creation of Montitextrix by Denis (1963)
for O. glacialis (L. Koch) was a first step in that direction. Oreone-
tides flavus (Emerton) and O. rotundas (Emerton), both from the
Nearctic region, are very close to M. glacialis in their genital struc-
tures, but differ in the positions of the Tm I (0.65-0.70 in glacialis ,
0.30-0.40 in flavus and rotundus) and the presence of a trichoboth-
rium on metatarsus IV in glacialis (absent in all others).

A few species are more closely related to — i.e. more closely re-
semble— the type-species vaginatus, viz., filicatus, firmus and abnor-
mis, and possibly also rectangulatus. I do not see the principal dif-
ference between vaginatus on the one hand, and fir?nus and abnormis
on the other (cf. Saaristo, 1972: 70). Oreonetides might constitute
an example of transition from the still folded scapes (but how un-
der-developed in comparison with the flexible and elaborately built
scapes of Lepthyphantes species!) of vaginatus and filicatus to the
rigid and unfolded scape of firmus and abnormis, where the narrow
portion of the scape with the socket or semi-covered depression is not
present. The absence of a well-developed median apophysis (Saaristo:
suprategulum ) is, of course, correlated with this simple build of the
epigyne, and should not be used as an independent character.

Oreonetides filicatus is a good species and not a synonym of
vaginatus as suggested by Saaristo (1972: 72). It is smaller than
vaginatus, but both male palp and epigyne are resembling each other,
though they differ in detail. Without question the species is con-
generic with vaginatus. However, the anterior tibia does not bear
the T-spine, which is one of the characters mentioned in the diagnosis
of the genus by Saaristo (1972: 69). I put this forward here as a
demonstration of my above remark on the too narrow delimination
of the genus.

Oreonetides rectangulatus (Emerton), of which I only know the
male, is different in several respects. Most striking are the pecul-
iarly shaped chelicerae, which bear a strong conical protrusion on
their dorsal surface for three-fifths of their length. The palp also
deviates from the true Oreonetides type.

Oreonetides flavescens (Crosby), described in Aigola (Crosby,
1937: 39, figs. 7-8) from New York I have not seen, but judging

58

Psyche

[March-June

17

0-5 mm

Figures 15-17. Oreonetides recurvatus (Emerton), male palp. Fig. 15,
lateral aspect. Fig. 16, cymbium, dorsal aspect. Fig. 17, tegulum with me-
dian apophysis (ma) .

1973]

van Helsdingen — Linyphiid Spiders

59

from the figures and description it was correctly placed with vagina-
tus and related species.

To the descriptions of O. recurvatus given by the various authors

I add the following remarks.

Measurements (in mm). Male, total length 2. 7-3.0, length cephalo-
thorax 1.25- 1.43, width 1.05- 1.20. Female, total length 2.7 (Ivie,
1966: 2.8 mm), length cephalothorax 1.22 (1.3), width 1.05 (1.1).

Ratio length to width of cephalothorax 0.85. PME large (0.09
mm), separated by their diameter, same distance to AME which
are barely smaller; lateral eyes of posterior and anterior rows con-
tiguous. Chelicerae showing sexual dimorphism; c? with three “teeth”
on dorsal margin: a basal, round tubercle with a more slender and
sharp tooth just dorsally of its tip, a second tooth at some distance
of the basal tubercle, and with a small tubercle at the end of the
row; ventral row with two small teeth; 9 also with three dorsal
“teeth”, ventral row with five small teeth (three ventral teeth ac-
cording to Ivie, 1966). Stridulating files well-developed, ridges fine
and close together.

Legs long and slender, femur I 1.8- 1.9 times length cephalothorax,
tibia I 20-23 ( c? ) or 18 (9) diams. of segment long. Tibia I longer
than femur I, metatarsus as long as femur ( cf ) or slightly shorter
(9). Femora I-III with a d-spine (on 0.5), femur I with an addi-
tional l'-spine. All tibiae with 2 d-spines (position basalmost d-
spine 0.25-0.35), tibia I also with one 1', one 1", and 2 v-spines, tibia

II with one v-spine and a 1 "-spine; apical spines on segments hardly
developed; all metatarsi with a single d-spine (position ca. 0.20 ).
Tm I 0.15-0.20, metatarsus IV without trichobothrium.

Male palp (Figs. 14-17). Characteristic is the meso-proximal
hook-like projection of the cymbium, which points laterad (Fig. 16).
The paracymbium lacks the isolated ventral hair which occurs in O.
vaginatus (see Saaristo, 1972), and also in O. filicatus and rotundas;
only a small tubercle present on this spot. Median apophysis (Fig.
17, ma) well-developed, with slightly curved tip. Embolic section
(Fig. 14) in general structure resembling vaginatus , but without
lamella; ventro-lateral branch rounded-truncated, a slender, chiti-
nous projection present between this branch and base of embolus, and
a second, equally narrow, slightly larger, membraneous projection,
arising from the dorsal surface of the radix. Embolus {e) short and
squat, curved, with a sharp spermduct tooth, and well-protected by
the embolic membrane {em) , which arises from the dorsal surface of
the radix from the membraneous connection between median apophy-
sis and radix (r).

6o

Psyche

[March-June

Epigyne and vulva (Figs. 11-13). Epigyne as depicted by Ivie
(1966: figs. 6-7). Scape folded and reappearing from below the
main body of the scape with a rounded, membraneous tongue, which
possesses a semi-covered depression. Entrances of ducts situated in
the main body of the epigyne, laterally and behind the bend of the
scape; ducts converging in forward direction, then curving outwards
to turning-points and to the receptacula seminis, which lie close to
the turning-points. The short fertilization-ducts curving to dorsal
side and ending as open gutters.

From the structures of palp and epigyne it is clear that the me-
dian apophysis, in connection with the depression at the tip of the
scape, serves as an important means of support during copulation.
The scape of the epigyne looks rather rigid and probably cannot be
pulled out of its resting position very far (cf. Lepthyphantes) , though
the ventralmost part may get pushed away from the main body so as
to allow the embolus to reach the entrance of the duct of the epigyne.
The exact functions of the hook-like projection of the cymbium, the
proximal, roughened extension of the paracymbium, and the lateral
arm of the radix are not easily understood without the aid of actual
observation of the pairing in one of the species of this group.

Acknowledgements

The present study is part of a general survey of North American
Linyphiidae, supported by Public Health Service Research Grant
AI-01944, from the National Institute of Allergy and Infectious
Diseases, to H. W. Levi.

The help with types and other specimens by the following institu-
tions and their curators is thankfully acknowledged : Dr. Herbert
W. Levi, Museum of Comparative Zoology, Harvard University,
Cambridge, U.S.A. (MCZ) ; Dr. John A. L. Cooke, American
Museum of Natural History, New York (AMNH); Dr. C. A.
Triplehorn, Ohio State University, Columbus, Ohio; Dr. Robin E.
Leech, Canadian National Collection, Ottawa, Canada (CNC).
Thanks are also due to Dr. William A. Shear, Concord College,
Athens, W. Virginia, for information on his collection.

References

Barrows, W. M.

1940. New and rare spiders from the Great Smoky Mountains National
Park region. Ohio J. Sci., 40: 130-138, figs. 1-12.

Crosby, C. R.

1937. Studies in American spiders: the genus Aigola Chamberlin. Proc.
Biol. Soc. Washington, 50: 35-42, pi. 1.

1973]

van Helsdingen — Linyphiid Spiders

61

Denis, J.

1963. Araignees des Dolomites. Atti 1st. Veneto Sci. Lett. Arti, 121:
253-271, figs. 1-16.

Emerton, J. H.

1913. New England spiders identified since 1910. Trans. Connecticut
Acad. Arts Sci., 18: 209-224, pis. 1-2.

I VIE, W.

1966. Two new North American spiders (Araneae: Linyphiidae). J.
New York Ent. Soc., 74: 224-227, figs. 1-7.

1969. North American spiders of the genus Bathyphantes (Araneae,
Linyphiidae). Amer. Mus. Novit., 2364: 1-70, figs. 1-121.
Merrett, P.

1963. The palpus of male spiders of the family Linyphiidae. Proc.
Zool. Soc. London, 140: 347-467, figs. 1-127.

Saaristo, M. I.

1972. Redelimitation of the genus Oreonetides Strand, 1901 (Araneae,
Linyphiidae) based on an analysis of the genital organs. Ann.
Zool. Fennici, 9: 69-74, figs. 1-17.

Vogel, B. R.

1967. A list of new North American spiders (1940-1966). Mem. Amer.
Ent. Soc., 23 : 1-186.

1968. Additional records of spiders from Western Pennsylvania. J.
New York Ent. Soc., 76: 101-105.

CORRELATION BETWEEN SEGMENT LENGTH AND
SPINE COUNTS IN TWO SPIDER SPECIES OF
ARANEUS (ARANEAE: ARANEIDAE)*

By L. David Carmichael
Museum of Comparative Zoology

Abstract

Observations made on several hundred adult male spiders of
two species of Araneus indicate a highly significant correlation
(p<0.ooi) between the length of a segment (tibia of the second
leg) and the number of macrosetae (“spines”) present on the seg-
ment. This result is further supported by observations on the first
tibiae of about twenty male A. trifolium, one of the two species, and
by a few observations on immatures of the two species. A short
summary of the methods used in taking the measurements and mak-
ing the calculations is followed by discussion of the implications of
this correlation with reference to species determination and geo-
graphic variation.

Methods

The study was done on two species of common North American
spiders, Araneus trifolium (Hentz) and A . marmoreus Clerck. 185
male specimens of A. trifolium yielded 347 tibiae of the second leg
(some specimens had lost one leg) ; the length of this segment was
measured, and the number of spines on the segment was counted.
In addition, lengths and spine counts were taken for the first tibiae
of 23 of the spiders, yielding 41 observations. Similarly, 120 speci-
mens of A. marmoreus yielded 210 second tibiae; the length was
measured, and two spine counts were made: the total number on
the tibia, and the number of modified, “dentiform”1 spines (see
Figure 3). The samples of both species were museum collections,
and represented almost the entire known range of each in North
America, extending from coast to coast and roughly from the 35th
to the 55th parallels.

The spines of the second tibia, like all the spines of these spiders,
are actually setae in the entomological sense ; they are set in a socket

* Manuscript received by the editor January 10, 1973

Term used by Locket & Millidge (1953), pp. 120, 121.

62

1973]

Carmichael — A raneus

63

Figures 1 and 2. A. trifolium tibia II, right leg. 1, anterior (prolat-
eral) surface; 2, posterior (retrolateral) surface.

Figures 3 and 4. A. marmoreus tibia II, right leg. The dentiform spines
are blackened. 3, anterior (prolateral) surface; 4, posterior (retrolateral)
surface.

Note the difference in scale between 1, 2 and 3, 4.

64

Psyche

[March-June

of the cuticle, and are movable. Thus even when the spine itself
becomes detached and lost, as is common with preserved specimens,
its presence or absence can be determined unequivocally by the pres-
ence or absence of the setal socket.

In the two species examined, the spines are uniformly larger than
the hairs which are also present, the former having a diameter at
the base of roughly 0.03 to 0.06 mm, while the latter are five to
ten times smaller. In A . marmoreus, there is a second type of spine,
described as dentiform, which is roughly 0.07 to 0.10 mm at the
base. Since this constituted a distinct group, it was counted sep-
arately.

Finally, the spines in both species are arranged in fairly constant
patterns, particular to the species (see Figs. 1-4). This makes it
possible to recognize each spine, which further eliminates any un-
certainty as to the number of spines. These three factors, the clear
difference between spines and hairs, the presence of a socket whether
or not the spine itself has been lost, and the possibility of recogniz-
ing each spine, make the spine counts unambiguous and as accurate
as possible within the limits of observor error.

The length of the tibia was measured along the dorsal midline
of the segment, between points “a” and “b” as shown in Figures 1
and 3. A grid in the microscope eyepiece, each cell of which meas-
ured 0.325 mm on a side (as determined with a stage micrometer),
permitted accuracy to ± 0.02 mm, or about =+= 1%.

The treatment of the data followed standard statistics texts; the
actual calculations were performed by the Harvard SDS 940 digital
computer.

Results

Table I presents the correlation coefficient for the number of
spines versus the segment length in the four different samples, as
well as the coefficient of regression (b) and the results of the t-test
for b = o. These values indicate that in all four cases the correla-
tion is highly significant (i.e. significant at the 0.1% level), b, the
coefficient of regression, quantifies the relationship established here;
it is the slope of the estimated regression line drawn in Graphs I
and II. The line is:

(number of spines) = a + b X (length of tibia)

Table II gives the mean and variance for the two variables in
both species, and the mean absolute difference between right and
left legs of individual specimens. In general the results are very

1973]

Carmichael — A raneus

65

Graph I. Scatter diagram and estimated regression line for A. trifolium.
measurements, b = 2.75, a — 15.24, (expected number of spines) — a +
b X (tibia length). Open circle, adult specimens; closed circle, immature
specimens; triangles, mean spine counts for given tibia! length.

66

Psyche

[March-June

similar to those found by Beatty (1967) for Adriana : the spine
count is quite constant within each species, though few specimens
are actually identical. Furthermore, in the two species of Araneus j
as in Ariadna , almost no individuals are completely symmetrical in
pattern, and most are asymmetric in actual spine counts. Beatty
attributed such differences within and between individuals to develop-
mental “accidents”; it is clear, however, that in the case of Araneus
some of the variation between individuals is specifically related to
difference in size. But for any single specimen the difference in
spine count (between left and right legs) seems not to be correlated
with the difference in segment length. (This correlation is cal-
culated as r2 in Table in II; the values, though positive, are not sig-
nificant at the 5% level.) Thus Beatty’s assertion is correct for
individual spiders; differences between left and right legs do seem
to be due to developmental accidents, and quite independent of each
other. This point will be important in the following discussion.

Discussion

It is important first to note that the above correlation does not in
itself imply cause and effect; this is clear from the fact that for any
individual, segment length and spine count are not correlated. It is
likely that both the length of the segment and the number of spines
are dependent on some other factor, such as general body size, etc.

One obvious possibility is that both measurements are related to
the degree of development, that is, to the number of molts the spider
has undergone. In most spiders raised there is some variation in the
number of preadult instars within a species. Furthermore, it is not
known with certainty at what stage these spiders mature or how
many molts occur after maturation, so this possibility cannot be ex-
amined with the data available. All the calculations here are based
on sexually mature specimens, but their ages cannot be determined
more precisely. Consequently, part of the spine count variation may
be dependent on this unmeasured variable; of course, size is some-
what dependent on this variable too.

On the basis of the data presented here, the best statement is
simply that spine count is very significantly correlated with segment
length, in these two species of Araneus.

Then there is the question of geographic variation. The samples
studied represent a pool of many local populations in North America,
and it is possible that the relationship between segment length and
spine count is different in different regions. (Preliminary examina-
tion of the data with regard to this question indicate that this is in

1973]

Carmichael — A raneus

67

Graph II. Scatter diagram and estimated regression lines for A. mar-
moreus measurements. Upper part of graph (open circles) shows total
spine counts: b = 4.43, a = 16.52. Lower part of graph (closed circles)
shows dentiform spine counts only: b=1.44, a = 8.12. Triangles show
mean spine counts for given tibial length.

68

Psyche

[March-June

TABLE I

A. trifolium

II

Tibiae
347
0.54

2.75 ±0.60
12.0

Tibiae I
41
0.61

2-39 ± 1.36
4.8

A
Total
count
210
0.63

4.43 ± 1.00
11.6

marmoreus

Dentiform
only
210
0.42

1.44± 0.56
6.7

N
n
b

t-test
(for b — 0)

Calculation of correlation and regression coefficients for spine counts
versus lengths of first and second tibiae of A. trifolium and for total and
dentiform spine counts versus length of second tibia of A. marmoreus. ri
is the correlation coefficient for spine counts versus length ; b is the slope
of the regression line, presented with its 99% confidence intervals. The
significance of the correlation may be found either from the value of the
coefficient r or from the t-test on the null-hypothesis b — 0.

fact the case.) While this does not affect the validity of the results
as they have been presented, it would modify the quantitative rela-
tionship (expressed by b) significantly in separated areas. This ques-
tion is open to further study; its significance will be mentioned
below.

Conclusion

The correlation between the number of spines on a segment and
the length of the segment is important to at least two aspects of
Araneology: taxonomy and the study of geographic variation. Mac-
rosetal counts have often been used to distinguish between different
genera of spiders2, as well as between species of one genus such as
Araneus. If the situation described in this paper is a general one,
then clearly any character based on setal counts should be used for
taxonomic purposes only after careful study. In general, it would
seem from observations on these two species of Araneus that the
number of spines alone is not highly reliable, but the pattern is quite
constant within a species (or at least recognizable, though spines
may be missing, or present in “unusual” locations). This is sup-
ported by observations made on species of the genus Neoscona (Ber-
man and Levi (1971), p. 467 ) .

Secondly, in studying geographic variation, it is necessary at least
in this case to consider the mean dimensions of local populations as
well as the spine counts. A marked variation in spine count between

2For examples, see Kaston (1948), with reference to: Gnaphosidae

(Drassodidae) pp. 347, 354; Clubionidae, pp. 367, 382; Thomisidae, pp.
410, 440; and Salticidae, p. 445.

1973]

Carmichael — A raneus

69

TABLE II

A. trifolium (N=166) A, marmoreus (N=98)

tibial

spine

tibial

total

dentiform

length

(mm)

count

length

(mm)

spine

count

spine

count

mean

2.31 ± 0.06

21.59 ± 0.31

3.39 ± 0.09

3 1. 54 ± 0.66

13.01 ± 0.32

standard

deviation

0.44

2.22

0.52

3.66

1.76

mean

difference

(absolute)

*

1.25 ±0.23

1.82 ±0.43

r2

0.134

0.127

. —

Self-explanatory: the means are presented with their 99% confidence

intervals. See text for details.

two regions might be obscured by the fact that specimens from one
of the regions are generally smaller than those from the other (which
itself might be due to significant geographic variation in size, or to
differences in sampling techniques, etc.).

Acknowledgments

I wish to thank Dr. H. W. Levi, my advisor at the time this
study was done, for his advice and patient help; Dr. S. J. Gould,
who advised me on the interpretation of statistical data; and the
late Mr. Ivie of the American Museum of Natural History, who
loaned me specimens from that museum. While doing the research
for this paper, I was supported until June, 1969 by a scholarship
from the National Merit Foundation, and subsequently by an
NDEA title IV fellowship.

References

Beatty, J. A.

1967. The Spider Genus Ariadna in the Americas. Doctoral Thesis,
Harvard University, Dept, of Biology.

Berman, J. D. and H. W. Levi

1971. The Orb Weaver Genus Neoscona in North America (Araneae:
Araneidae). Bull. Mus. Comp, Zoo-1. 141 (8) : 465-500.

Kaston, B. J.

1948. Spiders of Connecticut. Bull. Conn. Geol. Natur. Surv. Vol. 70 :
1-874.

Locket, G. H., and A. F. Millidge

1953. British Spiders, 2. Ray Society (London).

Simpson, G. G., A. Roe, and R. C. Lewontin

1960. Quantitative Zoology (2nd ed.). Harcourt, Brace & World, Inc.
Wetherill, G. B,

1967. Elementary Statistical Methods. Methuen & Co.

ANT LARVAE OF FOUR TRIBES:

SECOND SUPPLEMENT

(HYMENOPTERA: FORMICIDAE: MYRMICINAE)*

By George C. Wheeler and Jeanette Wheeler
Laboratory of Desert Biology
Desert Research Institute
University of Nevada System
Reno, Nevada 89507

Subsequent to the publication of our first supplement on the larvae
of the subfamily Myrmicinae (1960a)1 we have received from other
myrmecologists so much additional material that it has become neces-
sary to publish additional supplements.

Tribe Leptothoracini
Genus Macromischa Roger
Machromischa subditivci Wheeler

Creighton 1965 — Life cycle: egg 30 days, larva 2 3 days, pupa
19 days.

Genus Leptothorax Mayr2

Kempf 1959: 393 — “The morphological distinctness of the im-

aginal stages and the distribution of the species may even suggest to
accord Nesomyrmex full generic status. The larvae, however, are
quite close to the holarctic subgenus Leptothorax s. str., according
to G. C. & J. Wheeler (1955), who studied those of echinatinodis’1

Leptothorax carinatus Cole

semipupa: Length (through spiracles) about 2.2 mm. Profile

probably similar to L. amhiguus (1955: 22), otherwise differing in
the following details. Body hairs (1) about 0.006 mm long; (2)
0.006-0.087 mm long; (3) about 0.14 mm long, four on the dorsum
of each AI-AIII. Cranium transversely subelliptical. Head hairs
0.012-0.03 mm long, simple or bifid. Ventral border of each lobe of
labrum with one isolated and three contiguous sensilla and a few
minute spinules. Each labial palp a cluster of five sensilla. (Mate-

*Manuscript received by the editor January 30, 1973

To save space we cite our own papers by year and page; the complete
references are in Literature Cited.

2In 1950, M. R. Smith changed the well established subgeneric names
Leptothorax and Mychothorax to Myrafant and Leptothorax respectively.
Could more confusion be generated in less than two pages? The established
names should have been conserved. We refuse to accept these changes.

70

1973]

Wheeler Wheeler — Ant Larvae

71

rial studied: three semipupae from Texas, courtesy of Dr. A. C.
Cole.)

Leptothorax hispidus Cole

worker semipupa. Length (through spiracles) about 3 mm.
Similar to L. ambiguus (1955: 22) except as follows. No spinules
on integument. Body hairs: (1) 0.013-0.025 mm long; (2) 0.025-
0.068 mm long; (3) about 0.2 mm long, four each on AI-AIIL
Head hairs 0.01 5-0.038 mm long, with tip bifid. Labrum with about
ten hairs, about 0.025 mm long, on the anterior surface; posterior
surface with eight sensilla. Each maxillary palp a raised cluster of
four sensilla; each galea a very short peg with two sensilla. Each
labial palp with five sensilla.

young sexual larva. Length (through spiracles) about 3.2
mm. Similar to the above larva except as follows. Body sac-like.
Dorsal surface of posterior somites with a few minute spinules.
Cranium transversely subelliptical. Anterior surface of labrum with
four hairs.

Material studied: four worker semipupae and two young sexual

larvae from Texas, courtesy of Dr. A. C. Cole.

Leptothorax nevadensis Wheeler

Length (through spiracles) about 2.8 mm. Similar to L. ambiguus
(1955: 22) except as follows. Integument with a few minute spi-
nules on the venter of anterior somites and the dorsa of posterior
somites. Body hairs: (2) 0.025-0.19 mm long; (3) about 0.25 mm
long, on AI-AIV. Head hairs 0.013-0.03 mm long, generally dis-
tributed. Each mandible with narrow blade and two medial teeth.
Each maxilla with a ventral projection on the lateral surface; each
palp a cluster of five sensilla. Each labial palp a cluster of five sen-
silla. (Material studied: six larvae from Oregon, G. C. and J.
Wheeler #8)

Leptothorax niger fplendidiceps Urbani

Urbani (1968: 460-464) described the larva and figured young
and mature larvae and the head of the latter.

Leptothorax nitens Emery

Length (through spiracles) about 2.6 mm. Similar to L. ambiguus
(1955: 22) except as follows. Thorax slightly more constricted and
arched ventrally. Integument of venter of anterior somites and dorsa
of posterior somites with a few minute spinules, isolated or in short
rows. Body hairs: (1) 0.006-0.012 mm long; (2) 0.025-0.125 mm
long; (3) about 0.2 mm long. Head hairs 0.013-0.05 mm long,

72

Psyche

[March-June

Fig. 1. Leptothorax (M.) provancheri. a, larva in side view, Xl7;
b, head in anterior view, X67; c, very young larva in side view, Xl7;
d, simple and branched body hairs, X169; e, surface view of dendritically
branched body hair, X169; f, anchor-tipped body hair, Xl69; g, left
mandible in anterior view, X163. Fig. 2. Rogeria procera . a, left mandible
in anterior view, X169; b, head in anterior view, X 67 ; c and d, branched
body hairs, X264; e, anchor-tipped body hair, X264; f, larva in side
view,, X1B*

1973]

Wheeler & Wheeler — Ant Larvae

73

simple or bifid. Labrum with eight hairs on the anterior surface;
posterior surface with eight sensilla. Each mandible with the blade
narrow and bearing two teeth. Each maxillary palp a cluster of
four sensilla. Each labial palp a cluster of five sensilla.

very young larva. Length (through spiracles) about 1.3 mm.
Similar to very young larva of L. ambiguus (1955: 23) in shape,
otherwise similar to mature larva above except as follows. Cranium
more rounded. Head hairs 0.013-0.038 mm long, all simple. Labrum
more narrowed ventrally. Mandibles with narrower blade and
sharper teeth. Maxillae very small ; each galea a slight elevation with
two sensilla.

Material studied: 15 larvae from Oregon, G. C. and J. Wheeler

# 14.

Leptothorax (Mychothorax) provancheri Emery
(Fig. 1)

Length (through spiracles) about 3.7 mm. Paraponeriform (i.e.,
shaped somewhat like a crookneck squash ; neck short and stout ; body
elongate, stouter, straight and subcylindrical) ; posterior end rounded;
a ventrally projecting boss on each lateral surface of T r; each thoracic
somite and AI-AIII with a hairless midventral boss. Anus postero-
ventral. Leg, wing and gonopod vestiges present. Diameter of spira-
cles decreasing posteriorly. Integument of posterior somites with
minute spinules in short rows dorsally and isolated ventrally. Body
hairs rather sparse. Of three types: (1) 0.025-0.45 mm long, with
straight or kinked shaft and branched (bifid to dendritic) all branches
with denticulate tip, on all surfaces of all somites except venter of
Ti; (2) 0.025-0.1 mm long, simple, on all surfaces of Ti, fewer
on T2 and T3; (3) 0.25-0.375 mm long, anchor-tipped, with curled
to kinked shaft, four in a row across the dorsum of each AI-AV
(sometimes also one on AVI). Cranium subhexagonal, longer than
broad ; sides of head nearly straight. Each antenna on a teardrop-
shaped base; each a slight dome with three sensilla, each of which
bears a spinule. Head hairs numerous, minute (0.003-0.019 mm
long). Labrum paraboloidal in anterior view; anterior surface with
12 hairs (about 0.012 mm long), and two sensilla; ventral border
with two isolated and two clusters of three sensilla each; posterior
surface with about eight sensilla and a few minute spinules in short
rows. Mandibles leptothoraciform (i.e., moderately narrow; taper-
ing gradually and curving gradually to an apical tooth; anterior
surface produced medially into a blade bearing two subapical teeth) ;
all teeth subequal. Each maxilla with the apex conoidal ; each palp

74

Psyche

[March-June

a short skewed peg with five sensilla; each galea a short frustum
with two apical sensilla. Labium narrowly paraboloidal ; sparsely
spinulose, the spinules minute and in short transverse rows ; each palp
represented by a cluster of five sensilla; an isolated sensillum be-
tween each palp and the opening of the sericteries, the latter a short
transverse slit. No spinules on hypopharynx.

very young larva. Length (through spiracles) about 1.6 mm.
Abdomen sac-like, thorax forming a stout neck. Integument of ven-
ter of anterior somites and entire surface of posterior somites with
minute spinules. Body hairs sparse. Of three types: (i) o.oi 3-0.1
mm long, on dorsal and lateral surfaces of thorax and on dorsa of
AI-AIV; (2) about 0.006 mm long, on venter of Ti, few on AV :
(3) about 0.1 mm long, four each on AI-AIV. Antennae minute,
each with three sensilla. Head hairs about 1/3 as numerous, minute
(0.002-0.005 mm long). Labium subrectangular ; anterior surface
with about ten sensilla. Mandibles subtriangular in anterior view,
with all teeth sharp-pointed. Each maxillary palp represented by a
cluster of five sensilla; each galea represented by two contiguous
sensilla. Otherwise as in the mature larva.

Material studied : numerous larvae from Colorado, G. C. and

J. Wheeler #16.

The larva of L. provancheri resembles our other species of Mycho-
thorax in profile and in mandible shape, but it differs markedly in
the shape of the dominant type of body hair, in the shape of the head,
in the shape of the labrum, and in the abundance and size of head
hairs.

Genus Rogeria Emery

Because we had inadequate material previously (1955: 28) we
are giving a complete description below.

revised description. Profile solenopsidiform. Body hairs mod-
erately abundant ; of two types — ( 1 ) short, generally distributed
and variously branched and (2) anchor-tipped, on mesothorax, meta-
thorax and first three abdominal somites. Head hairs moderately
numerous, moderately long and bifid. Mandibles leptothoraciform.

In our 1960b key Rogeria would go to “ Monomorium antarcti-
cum” ( = Chelaner) , from which it cannot be distinguished generic-
ally.

Rogeria pro c era Emery
(Fig. 2)

Length (through spiracles) about 4 mm. Solenopsidiform (i.e.,
short and stout; head ventral, near the anterior end; prothorax bent

1973]

Wheeler & Wheeler — Ant Larvae

75

ventrally to form a very short stout neck ; remainder of body straight ;
both ends broadly rounded. Anus ventral.) Dorsal profile long and
C-shaped; ventral feebly sigmoid. Leg vestiges present. Spiracles
small; diameter diminishing posteriorly. Integument of venter of
thorax with relatively coarse spinules in transverse rows ; a few min-
ute spinules on dorsa of posterior somites. Body hairs moderately
abundant. Of two types: (i) 0.044-0.138 mm long, with tip short-
bifid to long-branched, the branches variously denticulate or branched ;
(2) about 0.25 mm long, anchor-tipped with flexuous shaft, six on
the dorsum of each T2, T3, AI, All and four on AIII. Cranium
subtrapezoidal, broadest dorsally; occiput nearly flat; clypeus bulg-
ing. Antennae each with three sensilla, each of which bears a rather
long spinule. Head hairs moderately numerous, 0.05-0.075 mm long,
bifid with the branches short to long. Lab rum bilobed, narrowed
dorsally ; each lobe with two hairs on the anterior surface about 0.006
mm long, ventral border with three isolated and two contiguous
sensilla, posterior surface with six isolated and a cluster of three
sensilla; entire posterior surface with a few short rows of minute
spinules dorsally and with coarse isolated spinules ventrally. Mandi-
bles leptothoraciform (i.e., moderately stout, tapering gradually and
curving gradually to an apical tooth, anterior surface produced me-
dially into a blade, which bears two medial teeth and a few denticles).
Each maxilla paraboloidal, apex with coarse isolated spinules; each
palp a cylinder with four apical and one subapical sensilla; each
galea a short stout cylinder with two apical sensilla. Labium nar-
row, anterior surface with coarse spinules, which are isolated or in
short rows near each lateral surface; each palp a slight elevation
with five sensilla; an isolated sensillum between each palp and the
opening of the sericteries, the latter a transverse slit. (Material
studied: 12 larvae from Brazil, courtesy of Drs. W. L. Brown and
K. Lenko.)

Tribe Ocymyrmecini
Genus Ocymyrmex Emery

Profile aphaenogastriform. Prothorax narrowed rapidly to the
diameter of the head. Head small. Anus with a prominent poste-
rior lip. Body hairs numerous, short, with frayed tip. Cranium
subcircular. Antennae high on cranium, minute and in pits. Head
hairs few, short, with bifid tip. Labrum paraboloidal, as long as
broad. Mandibles vollenhoviform but with only one medial tooth.

In our 1960b key this genus would fit under Group D but would
require a new rubric: 6. Body aphaenogastriform (Di) ; mandibles
vollenhoviform (Ilf).

76

Psyche

[March-J une

Ocymyrmex arnoldi Forel

(Fig. 3)

Length (through spiracles) about 5.9 mm. Aphaenogastriform
(i.e., stout and rather elongate; diameter greatest at AIV and AV ;
slightly constricted at AI ; thorax stout and arched ventrally, but
not differentiated into a neck; posterior end broadly rounded, anus
ventral). Prothorax narrowed rapidly to diameter of head. Head
small. Anus with a prominent posterior lip. Leg and wing vestiges
present. About six differentiated somites. Spiracles small, dimin-
ishing slightly posteriorly. Entire integument spinulose, the spinules
minute and in short transverse rows, the rows longer and closer
together on the venter of the anterior somites. Body hairs abundant,
uniformly distributed, all short (0.025-0.075 mm long), with stout
shaft and frayed tip. Cranium subhexagonal ; mouth parts rather
large. Antennae minute, each in a small pit bounded medially by a
high rim, three sensilla each bearing a tall spinule. Head hairs few,
short (0.019-0.038 mm long), slightly curved, with short-bifid tip.
Labrum paraboloidal, with three small ventral projections; anterior
surface with six minute hairs; ventral border with two small sensilla
on each ventrolateral projection and two groups of two larger con-
tiguous sensilla ; posterior surface with 16 sensilla and scattered min-
ute spinules. Each mandible vollenhoviform, but with only one
medial tooth (i.e., slender, rather long and nearly straight, apex form-
ing a moderately long slender tooth which is slightly curved medially ;
with a narrow medial blade, from the edge of which arises one in-
conspicuous medial tooth). Each maxilla with the apex paraboloidal
and with minute isolated spinules; each palp a narrow frustum with
four apical and one lateral sensilla ; galea digitiform with two apical
sensilla. Labium paraboloidal, with a few short rows of minute
spinules; each palp a skewed peg with five sensilla; an isolated sensil-
lum between each palp and the opening of the sericteries, the latter
a transverse slit. Hypopharynx with minute spinules in short rows.
(Material studied: three larvae and one semipupa from Rhodesia,
courtesy of Dr. W. L. Brown.)

Trire Tetramoriini
Genus Tetramorium Mayr
T etramorium caespitum (Linnaeus)

Bruder and Gupta 1972 — Description p. 366; photographs of
first, second and third instars and semipupa; drawings of mandibles
and maxillae of first, second and third instars. Life cycle in incipient
colony: egg 9-12 days, larva 8-14 days, semipupa 5 days, pupa 12-18

1973]

Wheeler & Wheeler — Ant Larvae

77

Fig. 3. 0 cymyrmex arnoldi. a, body hair, X167; b, larva in side view,

Xl6; c, head in anterior view, X81; d, right antenna in anterior view,
X376; e, left mandible in anterior view, X133. Fig. 4. Procry ptocerus
adlerzi. a, head in anterior view, X67; b-d, three types of body hairs,
X267; e, left mandible in anterior view, X185; f, larva in side view,
X 14.

7§

Psyche

[March-June

days, total 36-45 days. Life cycle in mature colonies: egg 8-12
days, first instar 2-7 days, second instar 3-7 days, third instar 10-19
days, semipupa 5 days, pupa 12-18 days, total 43-63 days.

Genus Triglyphothrxx Forel

Triglyphothrix striatidens Emery

immature larva. Length (through spiracles) about 2.2 mm.
Dorsal profile C-shaped ; ventral feebly sigmoid ; thorax stout and
curved ventrally but not differentiated from abdomen in diameter;
abdomen bag-like. Spiracles small; diameter diminishing posteriorly.
Integument of venter of anterior somites sparsely spinulose, the spi-
nules minute and in short transverse rows. Body hairs very few:
6 on Ti, 2 each on T2-AIV; 0.008-0.033 mm long, longest and
with multifid-tip on Ti, becoming shorter and simple posteriorly.
Head large; cranium subpyriform. Each antenna with three sensilla,
each of which bears a spinule. Head hairs minute (about 0.006 mm
long) , simple, six only, near mouth parts. Labrum twice as broad
as long, bilobed, lateral borders curved; each lobe with five sensilla
on the anterior surface, two contiguous sensilla on the ventral bor-
der, and one isolated and three contiguous sensilla on the posterior
surface; entire posterior surface moderately spinulose, the spinules
minute and in short rows which are arranged in longer subtransverse
rows medially, laterally the spinules are coarser and isolated. Each
mandible heavily sclerotized, narrowly subtriangular in anterior
view; of two portions: lateral thick and terminating in a long

slender apical tooth ; medial blade arising from the anterior surface
and bearing two sharp-pointed medial teeth. Each maxilla with the
apex conoidal and bearing a few spinules; palp a low rounded knob
with five sensilla; galea a short frustum with two apical sensilla.
Labium feebly bilobed, with minute spinules in subparallel rows;
each palp a low rounded knob with five sensilla ; an isolated sensillum
between each palp and the opening of the sericteries, the latter a
short transverse slit. Hypopharynx densely spinulose, the spinules
minute and in short arcuate rows which are arranged in long sub-
parallel transverse rows, base with numerous heavily sclerotized long-
itudinal ridges. (Material studied: numerous immature larvae from
New South Wales, courtesy of Rev. B. B. Lowery.)

Tribe Cryptocerini3

revision : Posterior surface of labrum usually without spinules.

Hypopharynx usually without spinules.

3In 1949, M. R. Smith concluded that the well established name Crypto-
cerus, which had been in use for 146 years, was a synonym of Cephalotes ;
he changed Cryptocerini to Cephalotini, Cryptocerus to Paracryptocerus
and subgenus Cryptocerus to Harnedia. In four pages could anyone intro-
duce more confusion into stable nomenclature? The old names should
have been conserved. We refuse to accept any of these changes.

1973]

Wheeler & Wheeler — Ant Larvae

79

Genus Cryptocerus Fabricius
Cryptocerus rohweri Wheeler

Creighton and Nutting 1965: 63 — “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).” Eggs de-
veloped in about a month into male larvae, which overwintered.

Cryptocerus ( Cyathomyrmex) pallens Klug

immature larva. Length (through spiracles) about 3.4 mm.
Similar to C. minutus (called Paracryptocerus rninutus in 1954:
155) except as follows. Head very large and covering approximately
half of the anterior end. Integument of venter of anterior somites and
all surfaces of posterior somites with a few minute spinules in short
transverse rows. Body hairs: (1) 0.006-0.038 mm long, most nu-
merous on the prothorax; (2) about 0.225 mm long. Head hairs very
numerous, slightly longer (0.013-0.05 mm long). Anterior surface
of labrum with eight hairs and/or sensilla. Mandibles moderately to
feebly sclerotized. An isolated sensillum between each palp and the
opening of the sericteries.

very young larva. Length (through spiracles) about 1.6 mm.
Body subellipsoidal, head on the anterior end (i.e., similar to 1.9 mm
larva of C. minutus, 1954: 155). Otherwise similar to mature larva
except in the following details. Entire integument spinulose, the
spinules minute and in short transverse rows. Head hairs shorter
(0.013-0.038 mm long). Labrum with ten hairs on the anterior
surface. Mandibles sickle-shaped, no blade.

Material studied: five larvae from Brazil, courtesy of Dr. K.

Lenko.

Genus Procryptocerus Emery

revision. Profile cataulaciform. Body hairs moderately numer-
ous. Of three types: (1) simple with flexuous tip; (2) tip short-
branched, multifid; (3) anchor-tipped. Head hairs of two types: (1)
simple, with long flexuous tip; (2) with short-branched (multifid)
tip. Mandibles cryptoceriform. Posterior surface of labrum with or
without spinules. Maxillae adnate and rounded. Hypopharynx with
or without spinules.

Procryptocerus adlerzi (Mayr)

(Fig. 4)

Length (through spiracles) about 4.6 mm. Profile cataulaciform
(i.e., straight, elongate-subelliptical ; segmentation indistinct; head
applied to the ventral surface near the anterior end) ; no neck. Anus
ventral, with a posterior lip. Leg, wing and gonopod vestiges present.

8o

Psyche

[March-J une

Six feebly differentiated somites. Spiracles small ; decreasing in diam-
eter posteriorly. Integument of ventral surface of anterior somites
with minute spinules in short transverse rows, entire surface of poste-
rior somites with scattered minute spinules. Body hairs moderately nu-
merous. Of three types: (i) 0.013-0.036 mm long, simple, tip very
fine and flexuous; (2) 0.022-0. 13 mm long, tip denticulate, a few on
each somite; (3) about 0.2 mm long, with flexuous shaft and anchor-
tip, four on each AI-AIV. Cranium subcircular in anterior view.
Antennae moderately large; just below middle of cranium; each with
three small sensilla, bearing a spinule each. Head hairs moderately
numerous, short to moderately long. Of two types: (1) 0.01 9-0.03
mm long, simple, with long fine flexuous tip, the more numerous
type; (2) 0.028-0.075 mm long, with short-branched tip. Labrum
arcuate; with eight hairs 0.025-0.038 mm long; ventral border with
four isolated and two clusters of three sensilla each ; posterior sur-
face with four isolated and two clusters of three sensilla each; spi-
nules lacking. Mandibles cryptoceriform (i.e., stout, subtriangular in
anterior view; lateral portion thick and terminating in a sharp-pointed
apical tooth ; medial blade arising from the anterior surface and bear-
ing two subapical teeth, which are subequal to apical tooth). Max-
illae rounded and adnate; each palp a short peg with five sensilla,
larger than the galea, the latter a short cylinder with two apical
sensilla. Labium small ; each palp a short stout peg with five sensilla ;
an isolated sensillum between each palp and the opening of the seric-
teries, the latter a short transverse slit. No spinules on hypopharynx.

immature larva. Length (through spiracles) about 3.1 mm.
Body relatively stouter. Head on anterior end. Integument of dorsal
surface of posterior somites with rather coarse spinules. Body hairs
(1) 0.025-0.05 mm long; (2) 0.0 19-0. 125 mm long; (3) about
O.225 mm long, four on the dorsal surface of each AI-AIV. Other-
wise similar to mature larva.

very young larva. Estimated length about 1.4 mm. Similar
to the immature larva except as follows. Body hairs sparse: (1)
0.002-0.025 mm long; (2) 0.024-0.05 mm long, with tip denticulate
to flattened; (3) about 0.086 mm long, on AI-AVI. Head hairs
sparse, all short spikes (about 0.003 mm long). Anterior surface of
labrum with six hairs about 0.006 mm long. Each mandible with the
apex turned medially; all teeth narrowly sharp-pointed. Each labial
palp represented by a cluster of five sensilla.

Material studied: 14 larvae from Brazil, courtesy of Dr. K.

Lenko.

1973]

Wheeler & Wheeler — Ant Larvae

81

Procryptocerus regular if Emery

Length (through spiracles) about 5.4 mm. Similar to P. adlerzi
except as follows. Body stouter; head relatively larger. Integument
with few spinules on the anterior somites. Body hairs (1) 0.013-
0.05 mm long; (2) 0.038-0.1 13 mm long; (3) four on the dorsa
of each AI-AIV. Cranium transversely subelliptical. Antennae
larger. Head hairs numerous; (1) 0.007-0.05 mm long; (2) 0.038-
0.087 mm long. Labrum feebly bilobed, with eight hairs 0.01 3-0.05
mm long; each lobe with six isolated and two contiguous sensilla on
and near the ventral border; entire posterior surface with a few
transverse rows of minute spinules. Each mandible with apical tooth
longer and basal teeth shorter. Each galea represented by two con-
tiguous sensilla. Each labial palp represented by a cluster of five
sensilla. Hypopharynx with a few transverse rows of minute spi-
nules.

young larva. Length (through spiracles) about 1.8 mm. Sim-
ilar to mature larva except as follows. Entire integument with min-
ute spinules, in short rows anteroventrally, elsewhere isolated,
coarsest posteriorly. Body hairs sparse; (1) 0.003-0.044 mm long;
(2) 0.013-0.088 mm long; (3) 0.125-0.188 mm long. Head hairs
moderately numerous, 0.003-0.044 mm long, simple. Hairs on labrum
about 0.003 mm long.

youngest larva. Length (through spiracles) about 1 mm long.
Similar to young larva except as follows. Body egg-shaped. Spiracles
very small. Integument with spinules on the dorsal surface of abdo-
men from spiracle to spiracle, more extensive on AIX and AX.
Body hairs very sparse; (1) 0.003-0.025 mm long, few, none on
AIX and AX: (2) 0.025-0.075 mm long, few, with short-bifid to
short-multifid tip, on lateral surfaces; (3) 0.09-0.18 mm long, four
on the dorsum of each AI-AVII, with straight shaft and smooth
anchor-tip. Head hairs very few, about 0.003 mm long. Posterior
surface of labrum lacking spinules. Each mandible with one apical
and one medial tooth. Each maxillary palp represented by a cluster
of sensilla. Labium and hypopharynx without spinules.

MALE LARVA. Length (through spiracles) about 5.8 mm. Sim-
ilar to worker larva except as follows. Thirteen feebly differentiated
somites. Body hairs (3) about 0.15 m m long, four on the dorsal
surface of each AI-AIV. Cranium subrectangular. Head hairs
(1) 0.013 mm long; (2) about 0.38 mm long, with short-bifid tip,
about eight on the cranium. Mandibles heavily sclerotized.

Material studied : numerous larvae from Brazil, courtesy of Dr.

K. Lenko.

82

Psyche

[March-J une

Procryptocerus striata scabriuscula Emery
Length (through spiracles) about 5.2 mm. Similar to P. adlerzi
except as follows. Body stouter. Integument of venter of anterior
somites and dorsa of posterior somites with minute spinules in short
transverse rows. Body hairs (1) 0.003-0. 19 mm long, spike-like;
(2) 0.013-0.05 mm long, very few on each somite. Cranium trans-
versely subelliptical. Head hairs few. Lab rum with six hairs and
six isolated and two clusters of three sensilla each on the anterior
surface; posterior surface with six isolated and two clusters of two
or three sensilla each and with minute spinules in short arcuate rows,
the rows forming a reticulate pattern. Mandibles quadrilateral,
heavily sclerotized, with all teeth straight and round-pointed. Each
maxillary palp an ungula, with two apical and three lateral sensilla;
each galea a very short stout peg with two apical sensilla. (Material
studied: nine larvae from Mexico, courtesy of Roy R. Snelling.)

Literature Cited
Bruder, K. W., and A. P. Gupta

1972. Biology of the pavement ant, T etramorium caespitum . Ann.
Entomol. Soc. Amer. 68: 358-367.

Creighton, W. S.

1965. The habits and distribution of Macromischa subditiva Wheeler.
Psyche 72: 282-286.

Creighton, W. S., and W. L. Nutting

1965. The habits and distribution of Cryptocerus roh'weri Wheeler.
Psyche 72: 59-64.

Kempf, W. W.

1959. A synopsis of the New World species belonging to the Nesomyr-
mex-group of the ant genus Leptothorax Mayr. Studia Entomol.
(Rio de Janeiro) 2: 291-432.

Smith, M. R.

1949. On the status of Cryptocerus Latreille and Cephalotes Latreille.
Psyche 56: 18-21.

1950. On the status of Leptothorax Mayr and some of its subgenera.
Psyche 57: 29-30.

Urbani, C. B.

1968. Studi sulla mirmecofauna d’ltalia. IV. La fauna mirmecologico
delle isole maltesi ed il suo significato ecologico e biogeografico.
Ann. Mus. Civ. Stor. Nat. Genova 77: 408-559.

Wheeler, G. C., and Jeanette Wheeler

1954. The ant larvae of the myrmicine tribes Cataulacini and Cepha-
lotini. J. Washington Acad. Sci. 44: 149-157.

1955. The ant larvae of the myrmicine tribe Leptothoracini. Ann.
Entomol. Soc. Amer. 48: 17-29.

1960a. Supplementary studies on the larvae of the Myrmicinae. Proc.
Entomol. Soc. Washington 62: 1-32.

1960b. The ant larvae of the subfamily Myrmicinae. Ann. Entomol.
Soc. Amer. 53: 98-110.

A NEW SPECIES OF ANACIS
FROM NORTHWEST ARGENTINA
(HYMENOPTERA, ICHNEUMONIDAE)

By Charles C. Porter*

Department of Biological Sciences, Fordham University,

Bronx, New York 10458

In two previous contributions (Porter 1967, 1970), the author
characterized the mesostenine ichneumonid genus Anacis, assigning
to it four species from Chile and contiguous regions of southwestern
Argentina. Meanwhile, Townes (1969, p. 176-177), as a result of
his study of the world Mesostenini, enlarged the definition of A nacis
to include also Cryptus exul (Turner, 1919, Ann. & Mag. Nat.
Hist. (9)3: 558) from Tasmania. Consequently, Anacis seemed to
emerge as pertaining to that zoogeographic category comprised of
taxa restricted at the present time to the Nothofagus zone of south-
ern South America and to similar areas of the Australian region.

Now, however, discovery of a fifth Neotropic Anacis from sub-
tropical wet forest in northwestern Argentina obliges us to modify
our distributional concept of the genus. Thus, in South America
Anacis appears to be of Andean rather than of strictly Neantarctic
or Araucanian range and, quite possibly, extends to other areas on
the continent. Its New World distribution, therefore, may be com-
pared to that of several other ichneumonid genera — such as Macro -
grotea , Trachysphyrus (sensu Townes), Picrocryptvides, Dotocryp-
tus, Deleboea } Alophophion , and Thymebatis — all of which are
well represented in Andean and temperate South America, including
Chile, but which concurrently have a greater or lesser number of
species on the peripheries of the lowland tropics. Taxa of this same
distributional type which moreover have species in the Australian
region are, of course, much rarer, but the ichneumonid genus Labena
(two species also reach North America) and the seolioid family
Thynnidae constitute approximate parallels.

The present study offers a description of this new Argentine
Anacis and a revised key to all known South American species of
the genus.

KEY TO THE SOUTH AMERICAN SPECIES OF ANACIS

(Based on females)

1. Mesoscutum mat, finely granular; setae of second gastric ter-
gite dense, mostly approaching or exceeding the length of their

83

84 Psyche [March-June

interspaces; mesosoma pale red or reddish brown with white
markings and black areas of variable extent 2

Mesoscutum silky shining with more or less fine punctuation;
setae of second gastric tergite sparser, mostly shorter to much
shorter than the length of their interspaces; mesosoma black
with sparse to profuse white markings 3

2. Gaster black with white apical bands on tergites; flagellum with

a white annulus; hind-tarsomeres 2-4 white; first flagellomere
8.5-10.0 as long as deep at apex; postpetiole weakly expanded,
O.8-O.9 as wide apically as long from spiracle to apex; ovi-
positor tip 0.18-0.20 as high at notch as long from notch to

apex 1. A. f estiva Porter

Gaster bright reddish brown with more or less prominent white
apical bands on tergites; flagellum without a white annulus;
hind-tarsus without white markings; first flagellomere 6. 2-7. 3
as long as deep at apex; postpetiole more strongly expanded,
1. 3-1. 4 as wide apically as long from spiracle to apex; ovipos-
itor tip 0.26-0.31 as high at notch as long from notch to apex
2. A. tucumana n. sp.

3. Thorax and propodeum with profuse white markings; all gastric

tergites with a complete white apical band ; apical margin of
clypeus with a small subdentate median projection; dorsal mar-
gin of pronotum without a definite submarginal groove

3. A. stangeorum Porter

Mesosoma with white at most on anterior margin of pronotum,
tegula, and subalarum; not all gastric tergites with a com-
plete white apical band; no median projection on apical margin
of clypeus; dorsal margin of pronotum with a conspicuous
submarginal groove 4

4. Legs mostly black with white markings; gaster with the follow-

ing white: median subapical mark on tergite 1, sometimes

marks on 2 and 3, and broad apical bands on 4-7 ; sheathed
portion of ovipositor 0.5-0.6 as long as fore-wing; nodus of

ovipositor tip with an unusually large and deep notch

4. A. varipes Porter

Legs mostly orange; gaster with white at most on tergites 6-8
and only on 7 sometimes with a complete white apical band;
sheathed portion of ovipositor O.3-0.4 as long as fore-wing;
nodus with a small but distinct notch 5. A. rubripes Spinola

1973]

Porter — A nacis

85

2. Anacis tucumana new species

( fig. 0

Holotype: female, ARGENTINA ( Tucuman : Horco Molle,

Dto. Tafi, October 24, 1970, C. C. Porter). (Tucuman). Para-
types: 4 females, ARGENTINA ( Jujuy : Post a de Lozano, Octo-
ber 26, 1969, December 8, 1969, C. C. Porter; Tucuman: Horco
Molle, Dto. Tafi, September 9-1 1 & September 30, 1969, C. C.
Porter). (Gainesville, Porter, Tucuman) .

Female: Color: flagellum pale reddish brown with slight dusky

staining, especially above on first segment; pedicel largely blackish
brown above and pale reddish brown to yellowish below; scape
largely blackish brown above and grading through reddish brown
into yellowish below; head black with some brown staining toward
apex on mandibles, sometimes in malar space, often around clypeus
above, often irregularly on face, and on antennal sockets, as well
as with the following white: most of basal 1/2 of mandible; very
large transverse blotch on clypeus; and a complete or sometimes
ventrally interrupted orbital ring, which is broadest below where it
extends 1/2 to 3/4 or more the distance into malar space; mesosoma
bright reddish brown with black on areas of variable extent includ-
ing most of prothorax (which at most is irregularly reddish stained),
mesoscutum slightly to entirely, mesopleuron slightly to on as much
as dorsal 1/3 plus most of prepectus, many margins and sutures
more or less broadly, as well as with the following white: broad
anterior margin of pronotum except toward apex ; broad dorsal mar-
gin of pronotum except for about median 1/4-1/3; tegula; most of
subalarum; sometimes large spot toward lower anterior corner of
mesopleuron; small area in upper hind corner of mesopleuron on
mesepimeron; most of apical 3/4-778 of scutellum; and sometimes
most of postscutellum ; gaster bright reddish brown with a prominent
to dull and little contrasting white apical band on tergites 1 or 2-
8 ; fore-leg with coxa mostly white with a conspicuous dorso-apical
brown blotch and some brown staining postero-basally ; trochanter
white with a large pale brown blotch dorsally; trochantellus white
with dusky to black staining on apex and above; femur bright red-
dish brown grading into whitish below and with a little dusky to
black staining on base; and tibia and tarsus somewhat duller brown
with some faint dusky staining and with fifth tarsomere mostly
dusky to blackish; mid-leg with coxa pale reddish brown with a
large, pale to dark brown dorso-apical blotch and sometimes a less
well defined ventral brownish area basad, as well as with white on

86

Psyche

[March-June

Fig. 1. Anacis tucumana Porter, female holotype. Lateral view of
mesosoma and gaster, showing color pattern.

most of dorso-basal 3/5 and on an extensive ventro-apical area;
trochanter white with a large brown area dorsally; trochantellus
white basally and below and blackish brown apically and above;
femur bright reddish brown with base narrowly blackish; and tibia
and tarsus a little duller reddish brown with last tarsomere dusky
to blackish; hind-leg with coxa uniformly bright red brown, except
sometimes for an obscure whitish area above near base; trochanter
varying from white with reddish brown staining to almost uniformly
reddish brown with white narrowly on apex, and sometimes with
blackish l rown staining on about basal 3/4 above; trochantellus
more or less blackish brown above and mostly reddish to brownish
white or white below; femur bright red-brown with base narrowly
blackish; and tibia and tarsus a little more dully red-brown with
last tarsomere dusky to blackish; wings hyaline with very slight
dusky staining apicad. Length of fore-wing: 5. 3-5. 7 mm. 1st fiagel-
lomere: 6. 2-7. 3 as long as deep at apex. Clypeus: in profile high
and asymmetrically convex to bluntly subpyramidal, with the apical
face shorter than the basal and a little concavely declivous ; the apical
margin practically straight, not produced or dentate medially. Malar
space : 0.75-0.85 as long as basal width of mandible. Temple: 0.30-
0.36 as long as eye in dorsal view; gently rounded-off and strongly

1973]

Porter — A Jiacis

87

receding. Fore-tibia: moderately stout but scarcely swollen. Pro-
notum: dorsal margin a little swollen, especially anteriad, and with-
out a submarginal groove; epomia sharp in scrobe but only faintly
prolonged below; anterior margin not angled at mid-height below.
Mesoscutum: notauli traceable about 1/2-2/3 the length of meso-
scutum, well defined but not very sharp anteriad and becoming much
weaker behind ; surface mat and finely granular with abundant,
small, faint, subadjacent to confluent punctures and an area of con-
trastingly coarser puncto- reticulation apicad between and beyond
notauli. Mesopleuron: subalarum not unusually swollen or expanded;
speculum swollen, mostly smooth and polished; surface otherwise
more or less strongly shining with fine but moderately strong, rather
uniform, obliquely longitudinal wrinkling, which becomes only slight-
ly more irregular on lower 1/2, and with medium-sized, variably
distinct intercalated punctures which are best defined on lower 1/2.
Wing venation: radial cell 2. 8-3. 2 as long as broad; areolet large
and broad, intercubiti strongly convergent above, 2nd abscissa of
radius 0.6-0.7 as long as 1st intercubitus; 2nd recurrent vertical, at
most weakly outcurved on upper 1/2; disco-cubitus broadly angled
or simply arched, without or rarely with a vestige of a ramellus;
nervulus at least slightly antefurcal ; mediella strongly arched ; axil-
lus close to hind-margin of wing. Propodeum: moderately short and
high in profile, basal face gently arched and sloping rearward to join
the much more steeply declivous but not sharply discrete, subequal
apical face ; spiracle round ; basal trans-carina sharp throughout,
weakly to moderately bowed forward medially, rather far from base
of propodeum; apical trans-carina more or less traceable throughout,
but becoming a little to, often, very weak and irregular on its broad-
ly bowed forward median portion, laterally well defined, forming
very low, sub-crescentic cristae and continuing ventrad to pleural
Carina; areola not defined; without lateral longitudinal carinae;
surface basad of basal trans-carina shining with considerable fine
wrinkling, especially mesad, and abundant, rather large, shallow,
subadjacent or sparser to confluent punctures but distad of basal
trans-carina uniformly mat with stronger, granularly reticulate
wrinkling and puncto-reticulation. 1st gastric segment: with a low
and weakly crescentic lateral flange at base; petiole moderately
broad and flat; postpetiole strongly expanded and 1.3-1.4 as wide
apically as long from spiracle to apex; ventro-lateral carina sharp on
postpetiole and about apical 1/2 of petiole but sometimes more or
less fading out toward base on petiole, or continuing sharp to base;
dorso-lateral carina fine and sharp for a short distance near spiracle

88

Psyche

[March-June

and again toward base of petiole but otherwise faint or absent; dor-
sal carinae absent or at most faintly suggested above spiracle. Cas-
ter: moderately elongate fusiform; 2nd tergite dully shining to mat
with fine, granularly reticulate wrinkling and abundant, medium
sized to large, mostly obscure, densely intercalated punctures which
emit numerous short setae that in great part approach or equal the
length of their interspaces; the following tergites with somewhat
longer and denser setae that mostly equal or exceed the length of
their interspaces. Ovipositor: sheathed portion 0.2 as long as fore-
wing; straight, moderately stout, strongly compressed; nodus high,
with a small but sharp notch ; dorsal valve in profile with a straight
to slightly concave taper between notch and apex; ventral valve on
tip with fine, inclivously oblique ridges; tip 0.26-0.3 1 as high at
notch as long from notch to apex.

Male*, unknown.

Types: The holotypes and two paratypes are deposited in the

collection of the Institute Miguel Lillo, San Miguel de Tucuman,
Republica Argentina. One paratype has been donated to the Florida
State Arthropod Collection (Gainesville, Florida, USA) and a
fourth paratype is in the collection of Charles C. Porter (RFD 3,
Cambridge, Maryland, USA).

Discussion: Among South American species of its genus, Attach

tucumana comes closest to the Araucanian A. f estiva, as shown by
several common characters emphasized in the key. Nonetheless, that
relationship is comparatively remote and tucumana has some features
which set it apart from the other South American Anacis ; for ex-
ample, its shorter notauli, shorter second radial abscissa, less elongate
propodeum, weaker dorso-lateral and dorsal carinae of the first gas-
tric tergite, and slightly shorter ovipositor. Indeed, the northwest
Argentine and Araucanian populations of Anacis probably have been
out of contact since the late Tertiary and early Pleistocene Andean
uplift, although it may be surmised that this is an old genus which
ranged throughout the climatically and biotically more uniform
South America of Pre-Andean times.

Worth noting also is the difference in abundance between the
southern and northern Anacis. At least two of the southern species
(A. f estiva and A. ruhripes ) are very common insects likely to be
encountered in numbers almost any day during the growing season;
whereas, A. tucumana only has been collected on five occasions. This
circumstance coincides with the probable relict status of Anacis.
Populations isolated in the Araucanian zone, where almost none of

1973]

Porter — A nacis

89

the modern Neotropic ichneumonid fauna has penetrated, would be
expected to flourish in the absence of aggressive competitors; whereas,
populations in an area such as the Selva Tucumano-Boliviana, where
occur scores of the Neotropic genera represented by hundreds of
species, would form a much more inconspicuous part of the fauna
and might have difficulty surviving at all.

Field notes: All localities for this species belong to the wet

subtropical forest community commonly designated Selva T ucumano-
Boliviana. Horco Molle in Tucuman Province is located in the
lower stratum of this forest (about 700 m.), while Posta, de Lozano
in Jujuy Province is at 1600 m. in an area of transition between
several types of environment, including many components of the
Selva and some Chaco elements, as well as stands of alders ( Alnus
joruliensis) .

Specimens of A. tucumana were collected by sweeping in weedy
areas both at the edge of the forest and in partially shaded places
within the forest.

Acknowledgements

Part of the material covered in this study was collected while the
author was working as Associate Investigator under a United States
National Science Foundation Grant (GB-6925) awarded to Dr.
Howard E. Evans of the Museum of Comparative Zoology at Har=
vard University.

The figure was inked by Miss Alicia Sandoval of the Instituto
Miguel Lillo.

References

Porter, C. C.

1967. A Review of the Chilean genera of the tribe Mesostenini. Studia
Ent. 10: 369-418.

1970. The Genus Anacis in Argentina. Acta Zoologica Lilloana
26(2) : 9-22.

Townes, H. K.

1969. Genera of Ichneumonidae, Part 2, Gelinae. Mem. Amer. Ent.
Inst. 12.

GROWTH OF THE ORB WEAVER,
ARANEUS DIADEM A TVS, AND
CORRELATION WITH WEB MEASUREMENTS*

By Jay Benforado and Kent H. Kistler
Division of Research, North Carolina Department of Mental Health
Raleigh, North Carolina 27611

Introduction

It is a well-known fact that within any population of spiders of
similar age there is considerable variation in the size of individual
spiders of the same species. In literature as early as 1890, McCook
has observed this variation and repeated observations (Comstock,
1940; Savory, 1928) have verified this phenomenon. Although ob-
servations are frequent, explanations are few. Bristowe (1958)
cites differences in feeding as a reason for differential size, but the
reference is made merely in passing and to the authors’ knowledge
is not elaborated upon elsewhere. This paucity of explanation lends
itself to further analysis of the factors contributing to the phenom-
enon of differential size.

Our purpose in this paper is to isolate some of the factors which
contribute to differential size in Araneus diadematus Clerck (for
identification of species, see Levi, 1971), and to elaborate upon cer-
tain of these factors as we are able.

Corresponding with differential size, in an orb-weaver such as
Araneus diadematus , differential growth is also manifested in chang-
ing dimensions of the web. That large differences in dimensions
exist betwen the individual webs of spiders is also a well-known
fact. An attempt to clarify some of the factors influencing web
changes is also made.

Method

environment: The spiders used were from two cocoons of Ara-

neus diadematus , obtained from Canastota, New York, which hatched
on April 26, 1972. From the time of hatching and throughout the
experiment, the spiders were kept in a room which was lighted 16
hours per day aand kept cool during the eight dark hours by an air
conditioner. (See Witt, 1971).

early rearing: At the time of hatching, the offspring from each

* Manuscript received by the editor March 1 , 1973

90

1973] Benforado & Kistler — Araneus diadematus 91

cocoon were placed in a separate rearing box. The spiderlings were
kept in these boxes, living on a communal web with a constant sup-
ply of loose gnats in the box, until they began to build individual
webs approximately three weeks after hatching. As each animal built
her first web she was removed from the rearing box and placed in
an individual glass tube, approximately 1X7 cm, with the ends
of the tube stoppered with cotton. From the time the animals were
placed in the tubes until onset of the experiment they were fed ap-
proximately 10-15 gnats per week, by placing the gnats in the tube
with the spiderling. The animals were watered by wetting the cot-
ton with water daily.

distribution: Seven weeks after hatching the two sets of spider-

lings were each separated into three equal groups by means of a
random numbers chart. No attempt was made to distribute males
and females evenly. Although the growth (body weight) of males
and females differs, it has been shown that the early growth of both
sexes is alike (Witt et al., 1972). Because of the short duration of
the experiment and the difficulty in identifying male spiders before
the last molt, distribution of males and females was left to chance.

At the time of initial grouping the two sets of spiderlings num-
bered twenty and thirty respectively. It was decided to feed each
of the three groups of each set according to a different feeding sched-
ule: one group every day, one group twice weekly, and one group
every ten days. Thus there were six groups, one for each set of
offspring on each feeding schedule. After one week of this procedure,
however, it was decided because of the small size of the groups to
reduce the number of schedules to two, and the middle /schedule was
dropped and its members distributed randomly between the lighter
and heavier-fed groups.

Data for animals that died or escaped during the course of the
experiment were removed, so figures represent only animals observed
for the duration of the experiment.

weighing: Each spider in the heavy-fed groups was weighed once

a week, to 0.1 mg, while animals in the light-fed groups were
weighed on the day of feeding and the day after feeding.

web analysis: After eleven days of controlled differential feeding

in the tubes, the spiders were transferred to aluminum and glass
laboratory cages, 50 X 50 X 10 cm. At this time the animals were
eight weeks old. From this time on the spiders began to build webs.
Photographs of webs were taken daily and analyzed (Reed et al.,
1965). Daily records of web building were kept and the webs were

92

Psyche

[March-June

Figure 1. Mean weights of 19 light-fed and 15 heavy-fed Araneus
diadematus with standard errors (vertical lines). Figures are for both
sets of spiderlings combined. Sharp increases in weight in the light-fed
group are due to the animals being weighed before and after feeding.
Note the increasing difference in weight between the light-fed and heavy-
fed group.

1973]

Benforado & Kistler — Araneus diadematus

93

destroyed daily with the thread left in the cage for the spider to
digest.

feeding: While in the tubes, the spiders were fed by placing a

previously weighed de-winged housefly in the tube daily or every
ten days. Those spiders that would not eat a housefly had three to
seven unweighed gnats placed in their tube. By visual inspection the
following day it was determined whether the fly had been eaten. The
remains of the eaten flies were then weighed to obtain an approxi-
mation of the amount eaten by each spider. The spiders were
watered by wetting the cotton every other day.

After being placed in the cages, if the spider had a web, feeding
was by means of placing the housefly in the web ; if there was no
web, we attempted to induce the animal to eat by placing the fly
in front of its mouth. The heavy-fed spiders were offered at least
one fly per day and more, if they would accept it. The light-fed
group was fed one fly once every ten days. If on the day of feeding
of the light-fed group a spider would not eat, a note was made and
the attempt repeated until successful. All spiders were watered on
Mondays, Wednesdays and Fridays by spraying a small amount of
water in each cage.

molts: From the onset of the experiment molts were recorded by

date of the molt to give an indication of the maturation of the
animal.

Results

feeding AND weight increase: At the end of a period of five

weeks the two feeding schedules resulted in two significantly differ-
ent weight groups. This development is shown in Figure i, which
illustrates the increasing difference in weight between the two groups.
At the onset of differential feeding the mean weights of the two
groups were alike, however, a T-test between the mean weights at
the end of the experiment is significant at the o.i % level.

An analysis of covariance was performed on the data.. Because
the original data was non-homogeneous, a transformation [log (x +
io) ] was made (Winer, 1962). The initial observation was used
as a covariate in the analysis of covariance. Because the analysis
of covariance indicated no significant difference in the behavior
(growth) of the two families, all figures are for both families com-
bined. For the heavy-fed group the mean weight changed from 7.93
mg db 1.04 on June 12 to 74.28 mg ± 10.93 on July 17. The
mean weight of the light-fed group changed from 6.40 mg dz 0.98

94

Psyche

[March-June

on June 12 to 17.91 mg ± 2.56 on July 13; there was a significant
interaction between time and feeding schedule below the 1% level.
FEEDING AND Maturation: If the number of molts over time is

taken as an indication of speed of maturation, then a relationship
between feeding and rate of maturation can be seen. During the
period of differential feeding the number of molts of the heavy-fed
and light-fed groups differed significantly at the 5% level. The
heavy-fed group had a mean number of 3.0 molts while the light-fed
group had a mean number of 2.3 molts. These results are in agree-
ment with the findings reported by Deevey (1949) with Latrodec-
tus mactcins (Fabricius) and indicate that in the laboratory with
only food quantity as a variable, a relationship exists between the
rate of weight increase and the rate of maturation.

INITIAL WEIGHT and rate of growth: From the beginning of

the experiment we noted a wide variation of weights of the individ-
ual animals. At the onset of differencial feeding individual weights
ranged from 1.1 mg to 16.2 mg. In both the light-fed and heavy-
fed groups there existed a positive correlation between initial weight
and final weight. For the light-fed group r = 0.7713 and for the

TABLE 1

Measurement

Light-fed
Early Late

Heavy-fed
Early Late

Mean wt. (mg) of spiders

12.52

22.30

20. 1 4

59-34

Spiral area (cm2)

1 18.92

119.51

1 19.88

138.32

Center area (mm2)

7 1 1 .00

877.53

920.30

1424.30

Thread length (m)

7-35

7-47

7.56

8.47

Mesh width (mm2)

20.16

22.34

21.79

27.48

Angle regularity

4-25

4.16

5.52

4.62

# of oversized angles
Relative deviation of

1.67

1.87

2.50

1.80

spiral turns (South)

0.34

0-33

0.41

0.35

Selected measurements of webs built by a group of light-fed and heavy-fed
spiders. Because not all spiders built on the same day, early and late webs
of both groups were chosen from two five day periods two weeks apart.
Measurements are divided into those which measure size (above the broken
line) and those which measure regularity. Note the difference between the
light-fed and heavy-fed animals in measures of web size at the late date.
While the heavy-fed group increased in all size measures (see Fig. 2), no
web regularity measures changed. For an explanation of web measure-
ments see Witt et al., A Spider’s Web.

1973]

Benforado & Kistler — Araneus diadematus

95

Figure 2. Selected web samples from two spiders: one heavily-fed spi-
der and one light-fed spider. Webs are from the periods measured in
Table 1 and are all reproduced to the same scale. Note that while both
the heavy-fed and light-fed animals began with webs of similar size, after
two weeks of differential feeding the large, heavy-fed spiders’ webs had
increased in size while the webs of the small, light-fed spiders remained
the same size.

96

Psyche

[March-June

heavy-fed group r = 0.9m; both correlations are significant at
the 0.1% level. In most instances those animals with the extreme
weights at the beginning of feeding remained the extremes in their
group. Reasons for the variation in initial weight are unknown.
Different rates of growth for light and heavy hatchlings have re-
cently been shown to occur in several species of spiders, apparently
independent of food available, and appear correlated with different
lengths of life (Reed and Witt, 1972).

AMOUNT EATEN : An approximation of the amount eaten was ob-

tained for a three week period. For nine heavy-fed animals the
mean amount eaten during the three week period was 115.8 mg and
for fifteen light-fed animals the mean amount for the same period
was 35.O mg. Within each group, however, there was an enormous
variation in the amount consumed : in the heavy-fed group the

amount eaten by individual animals ranged from 209.0 mg to 42.6
mg while in the light-fed group the amount eaten ranged from
49.6 mg to 3 gnats weighing 18 mg.

feeding and web changes: Table i gives a summary of web

changes that accompanied the growth of the animals. In measure-
ments of web size both groups increased, with the heavy-fed group
having a much larger increase as illustrated in Figure 2. In measure-
ments of web regularity both heavy-fed and light-fed groups re-
mained constant, as shown in figures of Table 1.

Discussion

The observed differential growth and development in Araneus
diadematus seems to be a function of several factors. Although an
exposure to a greater than normal supply of food generally results
in faster than normal growth and development, even within a group
exposed to the same food supply there seems to be a great variation
in growth rates. Evidence of these differences is expressed in the
increasing standard errors in Figure 1, and seems to be dependent
upon individual factors in the animals rather than environmental
variations. Large differences in the amount eaten by individual
animals in the laboratory existed and presumably exist in nature.
These differences seem to correspond to differences in the rate of
growth in agreement with the findings of Turnbull in other species
of spiders (Turnbull, i960, 1965). However, whether these dif-
ferences in the amount of food eaten are due to differences in pro-
ficiency in prey-catching or to differences in appetite or some other
factor in the animal is not clarified by our findings.

1973]

Benforado & Kistler — - Araneus diadematus

97

Another important factor influencing differential growth is the
initial weight of the animal. Variations in initial weights within a
family are generally retained during the course of development. Al-
though several possible reasons for different initial weights within a
family have been given by others, the authors are reluctant to offer
any explanations.

In an orb web weaving spider such as Araneus diadematus the
amount of food available to the animal is roughly equivalent to the
number of prey which become entrapped in the web. The number
of prey entrapped in the web is in turn determined by a number of
variables such as web-site, size and fine structure of the web, and
frequency of web building. Thus, it can be seen that the interaction
of the variables resulting in differential size and growth is complex
and can be divided into those factors which influence the amount of
food available to the spider and those factors which influence the
spider’s use of the food available to it.

Repeated attempts have been made to explain web characteristics
in terms of characteristics of the individual spider (Peters, 1936).
More common, however, has been the notation of changes in the
form of the orb web during the life of the spider (Tilquin, 1942;
Savory, 1952) and the attempt to relate these changes to changes in
the animal (Witt and Baum, i960; Witt, 1963; Reed et al.> 1969).
Because influencing factors vary concurrently, it is frequently diffi-
cult to assess the causes of changes in the form of the web.

In our experiment we attempted to isolate the effect of one vari-
able (weight) while minimizing the effect of a variable which nor-
mally changes concurrently (time). All animals used hatched on
the same date, however, one group (the heavy-fed) gained consid-
erable weight over the period measured. The web changes accom-
panying these weight increases are summarized in Table 1. Be-
cause all of the animals were hatched on the same date, we conclude
that increases in web size are due to differences in size of the ani-
mals resulting from differential feeding rather than differences in
age. If appetite were a factor influencing web size, it would appear
that the hungrier, light-fed animals would build a larger web in
an attempt to catch more food ; however, this is not the case.

The relationship between food and the web of a spider is a deli-
cate one. Without food, the spider’s web-building ability diminishes,
but without a web there is no food (Peakall, 1968). Thus, like a
businessman, the spider faces the law of diminishing returns. It
appears that the hungry spider chooses to conserve its resources

98

Psyche

[March-June

rather than gamble on a larger web trapping more food. Early food
deprivation experiments (Witt, 1963) show that the spider con-
tinues to build the same size web when deprived of food, but with
less thread until finally a decreasing in web size occurs. Because
our hungry (light-fed) animals were kept on a diet closer to a main-
tenance level than a deprivation level, we observed no decreases in
web dimensions.

Feeding conditions in a natural environment vary more than those
imposed in a controlled laboratory. Yet the spider is able to survive
in these naturally diverse conditions because of its adaptability. In
situations where there is little food available, the spider is able to
survive by growing at a slow rate and maintaining the same size
web. Where food is abundant, the spider takes advantage of the
situation, growing at a fast rate and increasing the size of its web.

The spider has developed a method for coping with a wide range
of feeding conditions. By varying its body and web growth, the
spider can survive under the diverse conditions imposed by nature,
thus minimizing the necessity of seeking new food supplies and re-
locating the web. Our findings provide new insight into the spider
as an example of an animal that adapts itself successfully to its en-
vironment.

Summary

Spiders from two cocoons of Araneus diadematus were exposed to
five weeks of two different feeding schedules: one group was of-
fered large amounts (one housefly per day) of food, the other group
scarce (one fly every ten days) amounts. Although both groups in-
creased in weight, weight gains of the heavy-fed group were signifi-
cantly greater than those of the light-fed group, regardless of cocoon
origin. Within each group there was a wide variation in the growth
of individual animals, indicating the presence of factors other than
food supply; i.e. animals with extreme weights within a group at
the onset remained the extremes.

In conjunction with increases in weight, over the three week period
of observation, webs of the heavy-fed spiders showed an increase in
size but not in regularity and shape in comparison to webs of the
smaller, light-fed animals of the same age which did not change.
Such data suggest an increased chance of survival of the species
through variations in rate of growth and maturation dependent on
environmental factors.

1973]

Benforado & Kistler — Araneus diadematus

99

Acknowledgements

This work was carried out in the laboratories of the North Caro-
lina Department of Mental Health and was supported by Grant
Number GB 25274 from the National Science Foundation to Peter
N. Witt. The authors gratefully acknowledge the assistance of Dr.
Peter N. Witt during all stages and the assistance of Mrs. Mabel B.

Scarboro during the period of laboratory work.

References Cited

Bristowe, W. S.

1958. The World of Spiders. Collins, London.

Comstock, J. H.

1940. The Spider Book. Revised and edited by W. J. Gertsch. Corn-
stock, Ithaca, N.Y.

Deevey, G. B.

1949. The development history of Latrodectus mactans (Fabr.) at dif-
ferent rates of feeding. Amer. Midland Naturalist. 42: 189-218.

Levi, H. W.

1971. The Diadematus group of the orb-weaver genus Araneus north
of Mexico (Araneae: Araneidae). Bull. Mus. Comp. Zool., 141:
131-179.

McCook, H. C.

1890. American Spiders and Their Spinningwork. Vol. 2, Published
by the author, Philadelphia.

Peakall, D. B.

1968. The spider’s dilemma. New Scientist, pp. 28-29.

Peters, H. M.

1936. Studien am Netz der Kreuzspinne (Aranea diadema.) 1. Die
Grundstruktur des Netzes und Beziehungen zum Bauplan des
Spinnenkorpers. Z. Morphol. Okol Tiere, 32: 613-649.

Reed, C. F. and P. N. Witt

1972. Growth rate and longevity in two species of orb-weavers. Bull.
Brit. Arach. Soc., 2: 111-112.

Reed, C. F., P. N. Witt and R. L. Jones

1965. The measuring function of the first legs of Araneus diadematus
Cl. Behavior, 25 : 98-119.

Reed, C. F., P. N. Witt and M. B. Scarboro

1969. The orb web during the life of Argiope aurantia (Lucas).

Develop. Psychobiology, 2: 120-129.

Savory, T. H.

1928. The Biology of Spiders. Sidgwick and Jackson, London.

1952. The Spider’s Web. Frederick Warne and Co., London and N.Y.

Tilquin, Andre

1942. La Toile Geometrique des Araignees. Presses Universitaires de
France, Paris.

Turnbull, A. L.

1960. Quantitative studies of the food of Linyphia triangularis (Clerck)
(Araneae: Linyphiidae) . Canad. Ent. 94: 1233-1249.

I <30

Psyche

[March-June

Turnbull, A. L.

1965. Effects of prey abundance on the development of the spider
Agelenopsis potteri (Blackwell) (Araneae: Agelenidae). Canad.
Ent. 97: 141-147.

Winer, B. J.

1962. Statistical Principles in Experimental Design. McGraw-Hill,
N.Y. pp. 606-615.

Witt, P. N.

1963. Environment in relation to behavior of spiders. Arch. Environ.
Health, 7 : 4-12.

1971. Instructions for working with web-building spiders in the lab-
oratory. BioScience, 21: 23-25.

Witt, P. N. and Ricarda Baum

1960. Changes in orb webs of spiders during growth. Behavior, 16:
309-318.

Witt, P. N., J. O. Rawlings and C. F. Reed

1972. Ontogeny of web-building behavior in two orb-weaving spiders.
Am. Zoologist, 12: 445-454.

Witt, P. N., C. F. Reed and D. B. Peakall

1968. A Spider’s Web. Springer Verlag, Berlin.

THE COCKROACH GENUS CALOLAMPRA OF
AUSTRALIA WITH DESCRIPTIONS OF
NEW SPECIES (BLABERIDAE)

By Louis M. Roth1 and Karl Princis2

Princis (1963) listed 23 species of Calolampra, thirteen of them
from Australia and the others from Africa, India, Burma, China,
Sarawak and the Philippines. Calolampra simlansis from the Hima-
layas was described by Baijal and Kapoor (1966). Recently, the
African species were placed in a new genus Pseudocalolampra (Roth
and Princis, 1971) and Calolampra laevis (Brunner v. W. ) from
Burma and China was assigned to the new genus Calolamprodes
(Bey-Bienko, 1969). Of the other species, ‘‘Calolampra” brunneri
(Brancsik) is a mislabeled Derocalymma cruralis, and “C.” punctosa
(Walker) belongs to the genus Laxta (Princis, 1967, p. 7°8)-
Calolampra truncata (Brunner v. W.) was listed by Princis (1967,
p. 627) under the genus Rhabdoblatta. In this paper we describe
26 species of Australian Calolampra , of which 12 are new. Five
species, C. gracilis (Brunner v. W.), C. atomifera (Walker), C.
notabilis (Walker), C. signatura (Walker), and C. propria
(Walker) have been considered to be synonyms of C. irrorata
(Fabr. ) (Princis, 1963). Of these, propria is a synonym of C.
atomifera (Walker) ; the other 4, we believe, are valid species.

The genitalia of Calolampra consist of 3 major phallomeres
(Fig. 1). An L2d (dorsal sclerite of the second left phallomere)
is apparently absent. The prepuce is covered with microtrichia,
curves upward and is fused to the L2vm. When well developed the
prepuce is cup-shaped and with L2vm as a handle looks like a ladle;
rarely is the prepuce unmodified (e.g. Fig. 82). Li (first sclerite
of left phallomere) and R2 (hooked sclerite of the right phallomere)
is well developed and lacks a subapical incision. The hypandrium
(subgenital plate) is symmetrical and bears 2 small equal sized
styli. All photographs of genitalia, supra-anal plate, and hypandrium
are from specimens treated with 10% KOH, dehydrated, and
mounted in Permount. Supra-anal plates were mounted dorsal sur-
face upward and subgenital plates ventral side uppermost. One or

’Pioneering Research Laboratory, U.S. Army Natick Laboratories, Natick,
Massachusetts 01760.

Zoological Institute, Lund University, S-223. 62 Lund, Sweden.

Manuscript received by the editor March 26, 1973.

IOI

102

Psyche

[March-June

Figure 1. Calolampra confusa. Male genitalia, dorsal view (115 MCZ),
A. (without exact locality). LI — first sclerite of left phallomere; L2vm —
ventromedial sclerite of second sclerite of left phallomere; P — prepuce;
R2 — hooked sclerite of right phallomere.

R2

0.5 mm

1973]

Roth & Princis — Genus Galolampra

103

both cerci or styli may be missing from the figures, but it is the
shape of the hind margins of the supra-anal and subgenital plates
which are of some diagnostic value. Because the supra-anal and sub-
genital plates were mounted on slides, their flattened shapes may
differ from that seen in dried pinned specimens. For example, Rehn
and Hebard (1927, p. 239, and plate XVIII, figs. 8, 9) described
the supra-anal plate of Ccilolampra aliena as “. . . transverse, sub-
rectangulate, distal margin weakly arcuate, . . .” and the subgenital
plate (p. 240) as “. . . slightly asymmetrical distal margin mesad
arcuato-emarginate . . .”. The shapes of these structures (from the
same specimen) when cleared and mounted on a slide do not agree
with the above description (Figs. 87, 88).

The first tergite of some male Calolampra may have elevations or
ridges which appear to be tergal glands (Figs. 15, 199) probably
involved in sexual behavior (Roth, 1969). However, some males
have slight ridges which do not appear to be glandular; regardless of
the nature of these modifications, the pattern formed by them has
some diagnostic value (Figs. 260-279).

The following abbreviations are used for localities, and source
(museums) of the specimens examined: A = Australia; (BMNH)
== British Museum (Natural History), London; (L) = Lund
Museum, Zoological Institution, Lund, Sweden; (MCZ) = Mu-
seum of Comparative Zoology, Harvard University, Cambridge,
Mass., U.S.A. ; N.S.W. = New South Wales, Australia; N.T. —
Northern Territory, Australia; Q = Queensland, Australia;
(QM) = Queensland Museum, Brisbane, Australia; S.A. —
South Australia; (SAM) — South Australian Museum, Adelaide,
South Australia; (SM) — Stockholm Museum, Stockholm, Sweden;
(USNM) = United States National Museum, Washington, D.C. ;
V. = Victoria, Australia; (VM) = Vienna Natural History Mu-
seum, Vienna, Austria; W.A. = Western Australia; (WAM)
Western Australian Museum, Perth, Western Australia. The num-
ber perceding some of the museum abbreviations refers to the num-
ber assigned the specimen and its corresponding genitalia ( cf ) on
a slide, which are deposited in their respective museums. The male
genitalia of the Lund material were mounted between small cover
slips and are attached to the pinned specimens.

1. C. elegans sp. n.

(Figs. 2-5)

cf (Fig. 2). Apterous, nitid. Head piceous, nitid, with transverse
yellow band between eyes on vertex. Antennae with their basal
10-15 segments nitid, piceous, remainder fuscous. Pro-, meso-, and

104

Psyche

[March-June

Figures 2-5. Calolampra elegans. $ (269 L, in SM), holotype, Peak

Downs, Q. 2. Adult. 3-5. Genitalia. Scale: Fig. 2 — 5 mm; Figs. 3-5 =
0.25 mm.

1973]

Roth & Pr'incis — Genus Calolampra

105

metanotum bicolored, with evenly scattered impressed punctures.
Pronotum with piceous disk on reddish-yellow ground, bordered all
around with piceous. Meso- and metantotum piceous, with yellow
maculae bordered with piceous and imitating vestigial tegmina and
wings. First tergite unmodified piceous, the others mostly brown
with yellowish flecks laterally on each side. Supra-anal plate sordid-
yellowish, transverse, its posterior margin slightly emarginate mes-
ally. Cerci brownish, hardly surpassing the posterior margin of
supra-anal plate. Venter of abdomen piceous laterally, brownish
mesally. Hypandrium transverse, symmetrical, provided with 2 short
styles; posterior margin slightly emarginate mesally. Legs piceous
to castaneous, only middle and posterior coxae margined posteriorly
with yellow. Lower anterior marign of front femur armed with
1 distal, and 4 additional spines at the proximal half; lower posterior
margin of same bearing 1 distal and 1 additional spine. Lower an-
terior margin of middle femur with 1 distal and 3 additional spines;
lower anterior margin of hind femur with 1 distal and 3 or 4 addi-
tional spines. Lower posterior margin of mid femur with 1 distal
spine only; same margin of hind femur unarmed. Caudal basitarsus
decidedly longer than the succeeding tarsal joints together, with
small apical pulvillus. Tarsal claws symmetrical, arolia very small.
Genitalia as figured (Figs. 3-5) ; prepuce poorly developed (Fig. 3).
Length of body 23 mm; length of pronotum 7 mm; width of pn>
notum 10 mm.

This new species differs from all known species of Calolampra by
the absence of tegmina and wings in the male sex.

9- Unknown, but it is probably apterous.

Material examined: cT (269 L in SM) (holotype) Peak Downs,
Q. The type specimen is unique.

2. C. darlingtoni sp. n.

(Figs. 6-13)

cf (Fig. 8). Pronotum triangular, widest at posterior margin;
disk speckled with black and with small reddish dots placed among
the black speckles; lateral margins yellow with small reddish dots
and larger dark ones ; the narrow yellowish hind margin with a series
of black longitudinal striae. Head pale yellowish with a blackish
patch in interocular-ocellar area. Antennae proximally yellowish,
distally dull ferrugineous. Tegmina reduced to lateral pads hardly
exceeding hind margin of mesonotum ; outer as well as sutural margin
convex; sutural half speckled with dark brown (Fig. 9). Wings
absent. Mesonotum, metanotum, and tergites speckled with black

io6

Psyche

[March-June

Figures 6-13. Calolampra darlingtoni. 6. $ (136 MCZ), paratype.

Alagate, Mt. Lofty Range, S. A. 7. Tegmen of 9 shown in figure 6.
8. $ (140 MCZ), holotype, The Creel, Mt. Kosciusko, N.S.W. 9. Tegmen
of $ shown in figure 8. 10-11. Genitalia. 12-13. Supra-anal and subgenital
plates. Scale: Figs. 6, 8 — 3 mm; Figs. 7, 9, 12-13 — 0.5 mm; Figs. 10-11
— 0.2 mm.

1973]

Roth & Princis — Genus Calolampra

107

and dotted with reddish in the same fashion as pronotum; longi-
tudinal striae present on their hind margins. First tergite unmodi-
fied. Weakly emarginate supra-anal plate (Fig. 12) and cerci dull
yellowish. Sternites yellowish with reddish dots and speckles; black
stigmatic spots decided. Hypandrium dull yellowish with mesally
incised hind margin (Fig. 13). Genitalia shown in figures 10-11.
Legs yellow with deep brown spines; tarsal claws symmetrical, well
developed arolia present. Length of body 17 mm; length of pro-
notum 4.5 mm; width of pronotum 7.2 mm; length of tegmina
3 mm.

$ (Fig. 6). Somewhat larger than male. Head pale brownish
with a wide black band between eyes, extending decidedly beyond
interocular space. Tegmina obliquely truncate (Fig. 7). Supra-anal
plate and cerci speckled with dark. Subgenital plate almost entirely
black except the reddish middle part. Very small arolia present.
Length of body 18.5-20 mm; length of pronotum 4.5 mm; width
of pronotum 7.5-8 mm; length of tegmina 3-4.5 mm.

Material examined: cf (140 MCZ) (holotype, herewith desig-
nated). The 'Creel, Mt. Kosciusko (3000 ft.), N.S.W. 14, 15.
XII. 1 93 1, Darlington leg; $ (136 MCZ) (paratype), Alagate, Mt.
Lofty Range, S.A., 29. XI. 1931, Darlington leg.; $ (SAM) S.A.
(without exact locality).

The species is named in honor of Dr. Philip Darlington, who
collected the holotype.

3. C. truncata (Brunner v. W.)

(Figs. 14-20)

Epilampra truncata Brunner v. W., Nouv. Syst. Bl., 1865, p. 178, 9.

c? (Fig. 14). Tegmina truncate, reduced, and extending only to
about the hind margin of second tergite; sutural half of tegmen
covered with large irregular shaped dark dots (Fig. 14) ; wings
reduced to rounded “leathery” lappets which reach only to hind
margin of first tergite (Fig. 15), and are completely hidden under
the tegmina. Abdomen pale, speckled with black, each tergite and
sternite with a pair of large sublateral dark blots, anteriorly. Ter-
gite 1 modified (Fig. 15). Supra-anal (Fig. 18) and subgenital
(Fig. 19) plates rounded. Male genitalia as figured (Figs. 16-17).
Legs pale brown, arolia well developed. Length of body 16 mm;
length of pronotum 4.6 mm; width of pronotum 7.2 mm; length of
anterior margin of tegmen 5.6 mm; length of wing 3 mm.

? (Fig. 20). Head black, vertex brown. Pronotum testaceous

io8

Psyche

[March-June

Figures 14-20. Calolampra truncata. 14-19. $ (85 MCZ), Salisbury Ct.,
N.S.W. 14. Male. 15. Modified (arrow) first abdominal tergite and wings
(w). 16-17. Male genitalia. 18-19. Supra-anal plate and hypandrium.

20. $ (2VM), holotype, Sydney, N.S.W. Scale: Fig. 14 =: 2 mm; Figs. 16-
17 — 0.2 mm; Figs. 18-19 = 0.5 mm. Fig. 20 = 3 mm.

1973]

Roth & Princis — Genus Calolampra

109

with disc fuscous; whole surface of pronotum furnished with small
impressed punctures and among them larger fuscous flecks. Tegmina
obliquely truncate, with sutural margin decidedly shorter than an-
terior one; veins distinct, wings reduced to lateral pads hidden
under tegmina. Abdomen brown speckled with black. Tergites pro-
vided on their hind margins with longitudinal striae, distance be-
tween them about 1 mm. Supra-anal plate rounded. Legs brown.
Length of body 19 mm; length of pronotum 5.5 mm; width of pro-
notum 8 mm; length of anterior margin of tegmina 5.3 mm; length
of sutural margin of tegmina 3.8 mm.

Material examined: 9 (2 VM) (holotype), Sydney, N.S.W. ; cf
(85 MCZ) , Salisbury Ct., N.S.W., ex Wheeler Collection.

4. C. ignota sp. n.

(Figs. 21-26, 278)

Syn: C. fraserensis Princis (nec Tepper 1893), Lund Univ.
Arsskr., N. F. Avd. 2, Vol. 50:13 (also: Kungl. Fysiogr. Salsk. i
Lund, Handl., N. F. Vol. 65:13, 1954, p. 31, cf ?•)

cf (Fig. 21). Head lacking (probably reddish). Pronotum el-
liptical with yellowish lateral margins which are connected anteriorly
and bear scattered blackish dots; disk with symmetrically arranged
black and reddish-brown speckles and streaks; hind margin provided
with black longitudinal striae. Tegmina somewhat exceeding apex
of abdomen, brown with whitish costal margins and black humeral
streaks; the usual dark speckling is lacking and tegmina appear
solidly colored throughout (the normally covered part of the right
tegmen is somewhat darker). Wings as long as tegmina, brown with
deep brown veins. First tergite as figured (Fig. 278). Hind margins
of supra-anal plate (Fig. 22) and hypandrium (Fig. 23) weakly
emarginate mesally. Venter and legs yellowish. R2 of genitalia
apically bifurcate (Fig. 24) (prepuce and Li accidentally lost).
Arolia lacking. Length of body 18 mm; length of pronotum 4.8 mm;
width of pronotum 6 mm; length of tegmina 18 mm.

$ (Fig. 25). Head yellowish, with reddish-brown band between
eyes; face regularly arcuate with occiput (no angulation). Pronotum
transverse, but less than 2 times broader than long; disk provided
with several impressed punctures (not numerous fine dots) ; hind
margin of pronotum convex, at least in its middle part ; latero-caudal
angles not produced backwards; dark longitudinal raised striae on
hind margin of pronotum. Tegmina reduced to lateral lappets, their
apices subacuminate (Fig. 26) ; edging of outer margin brown.

I IO

Psyche

[March-June

Figures 21-26. Calolampra ignota . 21-24. $ (1 WAM), holotype, Bu-

long, W. A. 21. Adult. 22-23. Supra-anal plate and hypandrium. 24. R2 of
genitalia. 25. $ (179 L), paratype, Norseman, W. A. 26. Tegmen of female
shown in figure 25. Scale: Figs. 21, 25 — 3 mm; Figs. 22-23 — 0.5 mm;
Fig. 24 — 0.1 mm; Fig. 26 — 1 mm.

1973]

Roth & Princis — Genus Calolampra

1 1 1

Wings absent. Dorsum of abdomen with dark maculation on pale
ground, underside reddish, laterally somewhat darker; the dark
striae on hind margins of tergites usually reduced to raised acuminate
tubercles, especially on the distal tergites. Legs yellowish; lower
posterior margin of hind femora completely unarmed. Length of
body 17.5-23 mm; length of pronotum 4.5-6 mm; width of pronotum
8.5-10 mm; length of tegmina 3.6-5 mm.

Material examined: cf (1 WAM) (holotype, herewith desig-
nated), Bulong, W. A. 1931. This specimen was recorded by Prin-
cis as C. fraserensis; 9 (:79 L) (paratype), Norseman, W. A.
1940; 9 (SAM), Camp 23, 1894, Horn Explor. Exped. ; 9 (SAM)
Murray Bridge, S. A., 12.XI.1909 (killed by ants), Tepper leg;
$ (SAM) Adelaide, S. A. 8.VI.1895.

5. C. paula (Tepper)

(Figs. 27-41, 261)

Epilampra paula Tepper, Trans. R. Soc. S. Austral. XVII, 1893, p. 60,
$ only.

cf (Fig. 27). Pronotum elliptical, whitish to yellowish with
scattered black or reddish impressed dots; disk of pronotum with
black symmetrical figure; hind margin with a row of dark rather
long longitudinal striae. Tegmina with yellowish costal field and
black humeral stripe, which beyond middle is divided into separate
oblique streaks; remainder is either nearly uniform brownish or
speckled with dark. Wings as long as tegmina, darkened and with
brownish veins. First tergite as figured (Fig. 261). Hind margins
of supra-anal plate (Figs. 28, 33, 38) and hypandrium (Figs. 29, 34,
39) mesally emarginate. Genitalia as shown in figures 30-32, 35-37,
40, 41; R2 and prepuce somewhat variable (cf. Figs. 31, 36, 41).
Legs yellowish; lower posterior margin of hind femora unarmed.
Arolia present. Length of body 13-14 mm; length of pronotum 3.2-
3.6 mm; greatest width of pronotum 4.6-6 mm; length of tegmina
13-15 mm.

9. Clouded dull brown. Pronotum provided with numerous fine,
impressed dots. Dorsal thoracic segments with black irregular mark-
ings and black striae on their hind margins. Abdominal tergites
likewise with raised longitudinal striae on hind margins; these striae
are relatively long, equaling about Y to x/i of tergal length. Teg-
mina reduced to lateral lappets, their apices subacuminate. Wings
absent. Length of body 17-18 mm; length of pronotum 4.5 mm;
width of pronotum 6.8-7 mm; length of tegmina 3 mm.

1 12

Psyche

[March-June

Figures 27-32. Calolampra Paula. 27-32. $ (2 SAM), lectotype, Ardros-
san, S. A. 27. Adult. 28-29. Supra-anal and subgenital plates. 30-32. Geni-
talia (Fig. 32 is a ventral view). Scale: Fig. 27 — 3 mm; Figs. 28-29 —
0.5 mm; Figs. 30-32 — 0.2 mm.

1973]

Roth & Princis — Genus Calolampra

Figures 33-41. Calolampra paula. Male supra-anal plates, subgenital
plates, and genitalia. 33-37. (21 SAM), Rudy Hole, N. T. 38-41. (20 SAM),
Cunnamulla, Q. Scale: Figs. 33-34, 38-39 = 0.5 mm; Figs. 35-37, 40-41 —
0.1 mm.

Psyche

[March-June

114

Material examined: cT (2 SAM), (lectotype, herewith desig-
nated), Ardrossan, S. A., II.1879, J. G. O. Tepper leg.; c? (20
SAM), Cunnamulla, Q., H. Hardcastle leg.; cf (21 SAM), Rudy
Hole, N. T., 1894, Horn Explor. Exped.; 2 9$ (SAM) Rudy
Hole, N.T., 1894, Horn Explor. Exped.; nymph (SAM), Para-
ehilna Hale, Flinders Range, S. A.; nymph (SAM) Port Lincoln,
S. A., Lea leg.;; 2 nymphs (SAM), Rudy Hole, N. T., 1894, Horn
Explor. Exped.; nymph (SAM), Everard Rgs., S. A. to Warberton
Rgs., W. A., A, Brumley leg.; nymph (SAM), near Fraser’s Hul,
III. 1 950, 'C. G. Gross leg.

6. C. candidula Shaw
(Figs. 42-49, 266)

Calolampra candidula Shaw, Proc. Linn. Soc. N. S. Wales 50, 1925, p. 175,
Fig. 3, $.

cf (Fig. 42). Head yellowish with a transverse row of four
black maculae between eyes. Antennae with the basal one-fourth
pale yellowish and remainder ferrugineous. Pronotum elliptical,
yellowish with a symmetrical black macula on disk and with hind
margin black lined. Tegmina considerably exceeding the apex of
abdomen, diaphanous with whitish veins which bear scattered brown-
ish dots; black humeral stripe present. Wings as long as tegmina,
pellucid with whitish veins. Abdomen above and below yellow.
First tergite as figured (Fig. 266). Hind margin of supra-anal
plate very slightly emarginate mesally (Fig. 43). Hind margin of
hypandrium distinctly emarginate (Fig. 44). Genitalia shown in
figures 45-47. Legs yellowish. Well developed arolia present.
Length of body 22-23 mm; length of pronotum 5.3 mm; width of
pronotum 7.5 mm; length of tegmina 23.5-28 mm.

9 (Fig. 48). Head yellowish with a black, medially interrupted,
transverse band between eyes. Pronotum with disk thickly speckled
with black and reddish; lateral margins yellow with small reddish
dots and darker dots among the smaller ones ; the narow yellow hind
margin provided with a series of black longitudinal striae. Tegmina
lobiform, apex rounded (Fig. 49) reaching °f their length beyond
hind margin of mesonotum; yellow with scattered reddish and dark
brown dots which are concentrated in the sutural half. Mesonotum,
metanotum and tergites speckled with black and reddish ; hind
margins provided with series of black longitudinal striae. Cerci and
supra-anal plate yellow, the latter dotted with small reddish and
several larger dark brown dots among the smaller ones. Sternites

1973]

Roth & Princis — Genus Calolampra

115

Figures 42-49. Calolampra candidula. 42-46. $ (1 QM), holotype,

Bellevue, Q. 42. Adult. 43-44. Supra-anal plate and hypandrium. 45-47.
Genitalia. 48-49. 9 (116 MCZ), Cairns, Q. 48. Female. 49. Tegmen of

female shown in figure 48. Scale: Fig. 42 — 4 mm; Figs. 43-44 — 0.5 mm;
Figs. 45-47 = 0.1 mm; Fig. 48 — 3 mm; Fig. 49 = 1 mm.

Psyche

[March-June

1 16

yellowish brown, subgenital plate deep brown. Legs yellowish with
brown spines; tarsal claws symmetrical; very small arolia present.
Length of body 21 mm; length of pronotum 7 mm; width of pro-
notum 12 mm; length of tegmina 5 mm.

Distribution: Aramac, Q. and northeast corner of S. A. (Shaw,

1925, p. 175).

Material examined: c? (1 QM) (holotype), Bellevue, Q., 1917,
E. C. Hurtridge leg.; ? (116 MCZ), Cairns, Q., Wheeler leg.

7. C. as per a (Tepper)

(Figs. 50-68, 262)

Epilampra aspera Tepper, Trans. R. Soc. S. Australia XVII, 1893, p. 62

$ 9.

cf (Fig. 50). Head with a dark transverse interocular band
which is continued on both sides towards clypeus. Pronotum almost
elliptical, but with the greatest width beyond middle; whitish to
yellowish with scattered brown impressed punctures; disk with dark
symmetrical figure; the dark longitudinal striae of hind margin very
short, mostly present on the edge of margin. Tegmina whitish or
yellowish more or less distinctly speckled with brown (this speckling
becoming indistinct in distal half of tegmina as well as in the nor-
mally covered portion of the right tegmen) ; radius deep brown
beyond middle breaking up into separate oblique lines. Wings as
long as tegmina, pellucid with whitish to yellowish veins. First
tergite as figured (Fig. 262). Supra-anal plate (Figs. 51, 57, 59,
64) as well as hypandrium (Figs. 52, 58, 60, 65) mesally emargi-
nate. Genitalia shown in figures 53-56, 61-63, 66-68. Legs yellow-
ish; lower posterior margin of hind femora unarmed. Well de-
veloped arolia present. Length of body 1 7-20.5 mm; length of
pronotum 4.5-5 mm ; width of pronotum 6-7 mm ; length of tegmina
19-20 mm.

9. Pale ochraceous to dull brown. Head with a dark band be-
tween eyes. Pronotum provided with numerous, fine, impressed dots.
Tergites asperous or rugose, distally with short raised tubercles which
are generally provided with recurved sharp points. Tegmina lobi-
form, subacuminate. Wings absent. Length of body 19.5-25 mm;
length of pronotum 4.5~5.2 mm; width of pronotum 7. 5-8. 5 mm;
length of tegmina 3. 8-4. 5 mm.

Material examined: cf (3 SAM) (lectotype, herewith desig-
nated), Western Plains, S. A., 29.XI.1888, A. G. Percy leg.; cf
(10 SAM), Clayton Crossing, S. A., 13.XI.1955 (at light), E. T.

1973]

Roth & Princis — Genus Calolampra

1 17

Figures 50-58. Calolampra aspera. 50-54. $ (3 SAM), lectotype,

Western Plains, S. A. 50. Adult. 51-52. Supra-anal and subgenital plates.
53-54. Genitalia. 55-58. $ (11 SAM), Kingoonya, S. A. 55-56. Genitalia.
57-58. Supra-anal and subgenital plates. Scale: Fig. 50 — 5 mm; Figs. 51-
52, 57-58 — 0.5 mm; Figs. 53-56 = 0.1 mm.

1 1 8

Psyche

[March-June

Figures 59-68. Calolampra aspera. Male supra-anal and subgenital
plates, and genitalia. 59-63. (10 SAM), Clayton Crossing, S. A. (R2 in
figure 62 damaged). 64-68. (12 SAM), Muloorina Stn., S. A. Scale: Figs.
59-60, 64-65 — 0.5 mm; Figs. 61-63, 66-68 — 0.1 mm.

1973]

Roth & Princis — Genus Calolampra

1 19

Giles leg.; cf (n SAM), Kingoonya, S. A., R. Harvey leg.; cf
(12-19 SAM), Muloorina Stn., S. A., 18. II. 1956 (at light), G. F.
Gross leg.; 2 cf cf (SAM) Muloorina Stn., S. A., 17. II. 1956 (at
light), G. F. Gross leg.; 20 cf cf (SAM) Muloorina Stn., S. A.,
1 8.II.1956 (at light), G. F. Gross leg.; cf (SAM) Ooldea, S. A.,
Tepper leg.; cf and 9 (SAM) Hermannsburg (McDonnell
Ranges), Capt. S. A. White; cf (SAM), north western S. A.,
H. Basedow leg.; (SAM) Herrgott Springs, S. A., 1. XII. 1898,
Blackburn leg.; 9 (28 SAM) ( paralectotype, herewith designated),
Algebuckina, S. A., 23.V.1887, Driffield leg.; this specimen was
labeled by Tepper in 1915 as type but he neither published nor
designated it in the description in 1893; 9 (SAM), Lake Calla-
bonna, S. A., A. Zietz leg.; 9 (SAM), Painta Swamp, Burt Plains,
1894, Horn Explor. Exped.; 9 and nymph (SAM) Oodnadatta to
Todmorton, Capt. S. A. White leg.; 9 (SAM) Eyre Pen., S. A.,
XII. 1954, G. F. Gross leg.; nymph (SAM), Callington, S. A.
19.I.1886, Tepper leg.; Tepper labeled this specimen as the 9 type
of C. paula in 1915, but his action is not valid; 3 nymphs (SAM),
Ardrossan, S. A., 26, 27. XI. 1885, Tepper leg.; nymph (SAM),
Oodnadatta, S. A., 1894, Horn Explor. Exped.; nymph (SAM),
north western S. A., H. Basedow leg.; nymph (SAM), Eyre Pen.,
S. A., 4.IX.1889, W. Graham leg.; 2 nymphs (SAM), Sedan,
S. A., 23.VIII.1888, Rothe leg.; 2 nymphs (SAM), Ooldea, S. A.,
R. T. Maurice leg.; nymph (SAM), Ellery’s Creek (McDonnell
Ranges), Capt. S. A. White leg.; nymph (SAM), Hermannsburg
(McDonnell Ranges), Capt. S. A. White; nymph (SAM), Finke
Gorge, S. A., 1894, Horn Explor. Exped.

8. C. subgracilis sp. n.

(Figs. 69-73, 275)

cf. Pronotum triangular with lateral angles broadly rounded and
with the greatest width at about the middle; lateral margins yellow-
ish dotted with black; disk very thickly speckled with black. Teg-
mina well developed, reaching distinctly beyond apex of abdomen,
covered with dark speckles, distally becoming larger and paler; the
normally covered part of the right tegmen solidly colored. Wings
as long as tegmina, brownish with deep brown veins. Abdomen
above deep brown, below yellowish. First tergite as figured (Fig.
275). Supra-anal plate with posterior margin entire (Fig. 73) ;
hypandrium damaged. Genitalia shown in figures 71, 72. Legs yel-
lowish with brown spines; lower posterior margin of hind femora

120

Psyche

[March-June

Figures 69-78. Calolampra suhgracilis and C. confusa. 69-73. C. sub-
gracilis. 69-70. $ (150 MCZ), paratype, A. (without exact locality). 69.

Adult. 70. Tegmen of female shown in figure 69. 71-73. $ (86 MCZ),

holotype, Melbourne, V. 71-72. Genitalia. 73. Supra-anal plate. 74-78. Calo-
lampra confusa. $ (115 MCZ), holotype, A. (without exact location). 74-
75. Supra-anal plate and hypandrium. 76-78. Genitalia (arrow indicates
lateral spur near base of L2vm in figure 76). Scale: Fig. 69 — 5 mm;
Fig. 70 — 1 mm; Figs. 71-72 — 0.1 mm; Figs. 73-75 — 0.5 mm; Figs. 76-
78 — 0.2 mm.

1973]

Roth & Princis — Genus Calolampra

121

unarmed; well developed arolia present. Length of body 16 mm;
length of pronotum 4.2 mm; width of pronotum 6 mm; length of
tegmina 17 mm.

9 (Fig. 69). Head brownish testaceous, with a blackish trans-
verse band between eyes; face continuously passing into occiput (no
angulation present). Antennae proximally brownish becoming dis-
tally blackish. Pronotum edged with brown, feebly so in front, but
strongly laterally; front margin regularly arcuate with undiffer-
entiated lateral margins; disk of pronotum with scattered large im-
pressed punctures and so thickly speckled with black that the brown-
ish ground color almost disappears; hind margin convex in dorsal
aspect and provided with the usual black striae which are rather
short and indistinct. Tegmina lateral, lobiform, reaching beyond
hind margin of mesonotum (Fig. 70), their edging of outer margin
brown. Dorsum of abdomen thickly speckled with black. Lower
posterior margin of anterior femora armed only with 1 distal spine;
same margin of posterior femora completely unarmed. Length of
body 17-19. 5 mm; length of pronotum 5-5.2 mm; width of pronotum
7. 5-8. 3 mm;; length of tegmina 3. 5-4.2 mm.

Material examined: c? (86 MCZ) (holotype herewith desig-
nated), Melbourne, V., H. Edwards leg.; 3 9? (145, 147, and 149
MCZ) (paratypes), same data as above; 9 (146 M'CZ) (paratype),
A. (without exact locality); 9 (T5° MCZ) (paratype), A. (with-
out exact locality), H. Edwards leg.; 9 (SAM), Ardrossan, S. A.,
26. XI. 1885. Tepper leg.; Tepper designated this specimen as the
type of Walker’s notabilis but his action is invalid.

9. C. confusa sp. n.

(Figs. 1, 74-78, 269)

cf. Pronotum nearly elliptical, with lateral margins sparsely
dotted with dark; disk symmetrically speckled with black. Antennae
damaged. Tegmina distinctly reaching beyond apex of abdomen,
speckled with black, distally becoming brown; the normally covered
part of the right tegmen solidly colored. Wings as long as tegmina,
infumate. Abdomen above brown, below yellowish. First tergite as
figured (Fig. 269). Posterior margin of supra- anal plate entire,
truncate except for lateral corners (Fig. 74). Hypandrium very
shallowly emarginate mesally (Fig. 75). Genitalia shown in figures
1, 76-78; L2vm with a lateral spur near its base (Fig. 76), a char-
acter not found in any other Calolampra examined. The spur is
reminiscent of the structure we called L2d in the male of Pseudo-

122

Psyche

[March-June

Figures 79-85. Calolampra fenestrata. 79. $ (298 L in SM), holotype,
Mt. Tambourine, Q. 80. $ (178 L), paratype, Cabramatta, N.S.W. 81.

Tegmen of 9 shown in figure 80. 82-85. Genitalia, supra-anal plate, and
hypandrium of $ shown in figure 79. Scale: Fig. 79 — 6 mm; Fig. 80 —
4 mm; Fig. 81 — 1 mm; Figs. 82-83 — 0.1 mm; Figs. 84-85 — 0.5 mm.

1973]

Roth & Princis — Genus Calolampra

123

calolampra pardalina (Roth and Princis, 1971, Figs. 9, 12, 15).
Legs dull yellowish with pale brown spines; lower posterior margin
of hind femora unarmed. Tarsal claws symmetrical; well developed
arolia present. Length of body 20 mm ; length of pronotum 5 mm ;
width of pronotum 7 mm ; length of tegmina 20 mm.

9. Unknown.

Material examined : cf ( 1 1 5 MCZ) (holotype, herewith desig-
nated), A. (without exact locality).

10. C. fenestrata sp. n.

(Figs. 79-85)

cf (Fig. 79). Head yellowish with brown interocular-ocellar
area; the former area somewhat paler. Antennae pale brownish.
Pronotum elliptical, encircled all around by narrow yellow margin ;
thickly dotted and speckled with black ; hind margin provided with
black longitudinal striae which are generally marked only by color
and do not extend to the narrow outer edging. Tegmina reaching
somewhat beyond apex of abdomen, brown with translucid fenestra,
which however are lacking on the normally solidly (castaneous)
colored part of the right tegmen. Wings as long as tegmina brown-
ish with deep-brown veins and whitish cross-veinlets. Abdomen
above dark brown, below yellow. Hind margin of supra-anal plate
entire (Fig. 84). Hind margin of hypandrium notched mesally
(Fig. 85). L2vm without a modified preputial attachment (Fig.
82) ; R2 shown in figure 83. Legs yellowish, lower posterior margin
of hind femora usually armed with 3 or 4 spines. Length of body
20 ( ?) mm; length of pronotum 4.5 mm; width of pronotum 6 mm;
length of tegmina 20 mm.

9 (Fig. 80). Head yellowish, with blackish interocular-ocellar
area; another blackish patch reaches to clypeus. Pronotum, thickly
speckled with black ; several impressed punctures present ; hind
margin convex in dorsal aspect; the dark longitudinal striae of hind
margin distinctly marked by darker color and only weakly raised.
Tegmina lobiform (Fig. 81), edging of outer margins brown. Dor-
sum of abdomen thickly speckled with black on pale ground color ;
venter almost throughout solidly black; only several reddish patches
present. Legs as in male. Length of body 23 mm ; length of pro-
notum 5.8 mm; width of pronotum 9 mm; length of tegmina 4 mm.

Material examined: cf (298 L in SM) (holotype, herewith desig-
nated), Mt. Tambourine, Q., Mjoberg leg.; 9 (178 L) (paratype).
Cabramatta, N. S. W., 19.XI.1961, M. Nikitin leg.

124

Psyche

[March-June

Figures 86-94. Calolampra signatura. 86-91. $ (61 BMNH), holotype,

St. Helena. 86. Adult. 87-88. Supra-anal plate and hypandrium. 89-91.
Genitalia (R2 in figure 90, is damaged). 92-94. $ (67 MCZ). Genitalia
of holotype of Calolampra aliena, Haiti. Scale: Fig. 86 = 5 mm; Figs. 87-
88 — 0.5 mm; Figs. 89-91 % 0.1 mm; Figs. 92-94 — 0.2 mm.

1973]

Roth & Princis — Genus Calolampra

125

11. C. signatura (Walker)

(Figs. 86-94, 277)

Pattchlora signatura Walker, Cat. Derm. Salt. Brit. Mus. 5, 1871, Suppl.
Blatt. p. 13, $ (St. Helena).

Calolampra aliena Rehn and Hebard, Bull. Amer. Mus. nat. Hist. 54-, 1927,
p. 238, PI. XVIII, figs. 5, 7.-9, $.

Calolampra signatura (Walker), Ann. Mus. Roy. Afr. Centr, in — 8°,
Zool., 181, 1970, p. 170, $.

cf (Fig. 86). Interocular-ocellar space with a solid dark area
which is almost divided in two parts anteriorly by a triangular
emargination. Antenna with basal joint yellow and the remainder
fuscous. Pronotum thickly speckled and dotted with blackish and
reddish; edge of lateral margins uniform yellow; hind margin with
dark, radially arranged striae. Tegmina markedly extending beyond
the apex of abdomen, covered by numerous dark small dots; costal
margin yellow ; humeral streak broad and black, beyond middle
breaking into separate maculations; the normally covered part of
the right tegmen solidly brownish colored. Wings infumate, as long
as tegmina, with deep brown veins. Dorsum of abdomen pale,
banded transversely with brown; first tergite as figured (Fig. 277).
Hind margins of supra-anal plate (Fig. 87) and hypandrium (Fig.
88) not emarginate. Venter yellowish with brown stigmata. Geni-
talia shown in figures 89-94. Legs yellowish with brown spines;
lower posterior margin of hind femora unarmed; arolia normally
developed. Length of body 16 mm; length of pronotum 5 mm;
width of pronotum 6.6 mm; length of tegmina 21.7 mm.

9. Unknown.

Material investigated : cf (61 BMNH) (holotype of signatura) ,
St. Helena; cf (67 MCZ) (holotype of aliena ), Haiti, P. R. Uhler
leg.

This species is probably an introduction from Australia to St.
Helena and Haiti. St. Helena was an important station for slave
ships as well as ships coming from Australia. C. signatura was pos-
sibly introduced to St. Helena with Australian plants (there are
many introduced Australian plants in St. Helena) . C. aliena is
possibly a secondary introduction by slave ships from St. Helena to
Haiti (Princis, 1970).

12. C. atra (Tepper)

(Figs. 95-109, 279)

Epilampra atra Tepper, Trans. R. Soc. S. Austral. XVII, 1893, p. 65,
only $ (not $ $ as given).

Epilampra propria Tepper (nec Walker 1868), Ibid. XVII, 1893, p. 64,
only o from Mannum (nec 2 = Calolampra teppcri Kirby).

126

Psyche

[March-June

cf. Head with broad black band in interocular-ocellar area.
Antennae pale brownish. Pronotum elliptical in outline, with lateral
margins pale yellowish dotted with brown; disk thickly speckled
with black sometimes almost uniform black. Tegmina dull brown
with somewhat darker dots which are sometimes obscured ; coastal
area yellowish; humeral streak black, broad and short. Wings as
long as tegmina, smoky brown with brown veins. First tergite as
figured (Fig. 279). Abdomen above deep brown, below yellowish.
Supra- anal plate, cerci, and hypandrium yellowish. Hind margin of
supra-anal plate almost straight (Fig. 95) or slightly emarginate
(Figs. 100, 104). Hind margin of hypandrium weakly (Fig. 96) or
distinctly (Figs. 101, 105) mesally emarginate. Genitalia shown in
figures 97-99, 102, 103, 106, 107; R2 with apex beaklike (Figs. 98,
103, 107). Legs yellow with brown spines; lower posterior margin
of hind femora usually unarmed (distal spine is always lacking).
Length of body 19-23 mm; length of pronotum 4.5-6 mm; width
of pronotum 7-8 mm; length of tegmina 20.5-23 mm.

9 (Fig. 109). Shining blackish. Pronotum with lateral margins
pale yellowish dotted with brown; latero-caudal angles not produced
backwards. Tegmina lobiform bicolored, the outer part yellow, the
inner solidly dark; extending to about the hind margin of mesonotum
(Fig. 108). Length of body 23-27 mm; length of pronotum 6 mm ;
width of pronotum 10-12 mm; length of tegmina 3-5 mm.

Material investigated: 9 (1 SAM) (lectotype, herewith desig-
nated), Sedan, S. A., XII. 1885, F. Rothe leg.; Tepper erroneously
labeled this specimen as male; 9 (SAM) (paralectotype) , same data
as above; cf (22 SAM), Mannum, S. A., 24.V.1890, R. Schroeder
leg. ; this specimen was labeled by Shaw as the type of Kirby’s tepperi
but the action is invalid; cf (29 SAM), Adelaide, S. A.; N. B.
Tindale leg.; cf (30 SAM), Lucindale, S. A., Feuerheerdt leg.

13. C. gracilis (Brunner v. W.)

(Figs. 110-136, 273)

Epilampra gracilis Brunner v. W., Nouv. Syst. Blatt. 1865, p. 170, $ 2,
Fig. 20 (2).

cf (Figs, no, 130). Head yellow;; interocular-ocellar area with
a transverse dark brown band which is interrupted in the latter area.
Pronotum in dorsal aspect triangular, with the greatest width at its
middle, thickly speckled and dotted with dark brown ; hind margin
bearing a series of dark brown longitudinal striae. Tegmina some-
what exceeding apex of abdomen, with scattered brown maculation

1973]

Roth & Princis - — Genus Calolampra

127

Figures 95-109. Calolampra atra. 95-99. $ (22 SAM), Mannum, S. A.
95-96. Supra-anal plate and hypandrium. 97-99. Genitalia. 100-103. $

(29 SAM), Adelaide, S. A. 100-101. Supra-anal plate and hypandrium.
102-103. Genitalia. 104-107 $ (30 SAM), Lucindale. 104-105. Supra-anal
plate and hypandrium. 106-107. Genitalia. 108-109. 9 (1 SAM), lectotype,
Sedan, S. A. 108. Lateral view of pronotum, mesonotum, and tegmen. 109.
Dorsal view. Scale: Figs. 95-96, 100-101, 104-105 — 0.5 mm; Figs. 97-99,
102-103, 106-107 = 0.2 mm; Fig. 108 — 3 mm; Fig. 109 = 4 mm.

128

Psyche

[March-J une

Figures 110-117. Calolampra gracilis. 110. $ (1 VM), lectotype, Ade-
laide, S. A. 111-112. $ (139 MCZ) , Blue Mts., Hartley Vale, N.S.W.
111. Dorsal view. 112. Tegmen. 113-117. Supra-anal plate, hypandrium,,
and genitalia of lectotype shown in figure 110. Scale: Fig. 110 = 5 mm;
Fig. Ill — 4 mm; Fig. 112 — 1 mm; Figs. 113-114 — 0.5 mm; Figs. 115-
117 — 0.2 mm.

1973]

Roth & Princis — Genus Calolampra

129

Figures 118-136. Calolampra gracilis. 118-121. $ (25 SAM), Mt. Bryan
E., S. A. 118-119. Supra-anal plate and hypandrium. 120-121. Genitalia.
122-125. $ (26 SAM), Norwood, S. A. 122-123. Supra-anal plate and hy-
pandrium. 124-125. Genitalia. 126-129. $ (27 SAM), Yardea, S. A. 126-
127. Supra-anal plate and hypandrium. 128-129. Genitalia. 130. $ (261 L),
Cabramatta, N.S.W. 131-132. Genitalia of specimen shown in figure 130.
133-134. $ (260 L), same locality as 261 L. Genitalia. 135-136. $ (264 L),
same locality as 261 L. Genitalia. Scale: Figs. 118-119, 122-123, 126-127 —
0.5 mm ;Figs.l20-121, 124-125, 128-129, 131-136 = 0.2 mm; Fig. 130 = 4 mm.

130

Psyche

[March-June

on yellowish ground ; broad dark brown humeral stripe present ; the
normally covered part of right tegmen solidly brown colored. Wings
as long as tegmina, infumated and with brown veins. Dorsum of
abdomen yellowish brown, distally becoming darker; first tergite as
figured (Fig. 273). Cerci and supra-anal plate yellowish, the latter
with hind margin very weakly emarginate (Figs. 113, 118, 122,
126). Venter of abdomen yellow with dark brown stigmatic macu-
lae. Hypandrium distinctly (Figs. 114, 123, 127) or weakly (Fig.
1 19) emarginate. Genitalia shown in figures 115-117, 120-12 1, 124-
125, 128-129, 131-136. Legs yellow with brown spines; lower pos-
terior margin of front femora usually unarmed, the distal spine ex-
cepted, only occasionally one additional spine present; same margin
of hind femora totally unarmed. Length of body 19-22 mm; length
of pronotum 5-6 mm; width of pronotum 7-7.5 mm; length of
tegmina 21 mm.

$ ( Fig. hi). Head generally as in male, but sometimes in dark
individuals a blackish additional macula between interocellar area
and clypeus present; face passing into occiput without angulation.
Pronotum at most twice as broad as long, thickly speckled and
dotted with dark brown (in dark individuals with blackish) ; disk
with scattered impressed punctures; hind margin bearing a series of
dark brown to blackish longitudinal striae; middle part of the hind
margin weakly convex; lateral margins not differentiated, but regu-
larly arcuate with anterior margin; edging of lateral margins in
dorsal aspect not lined brown inside; latero-caudal angles slightly
produced backwards. Tegmina reduced to lateral lappets (Fig. 112) ;
edging of their outer margins dark brown (even in dark individuals
not black). Dorsum of abdomen thickly speckled and dotted with
dark brown to blackish on yellowish ground. Supra-anal plate yel-
lowish, only weakly dotted with dark brown. Venter of abdomen
dark brown to blackish with several reddish maculations. Legs yel-
lowish brown with dark brown spines; lower posterior margin of
front femora and same margin of posterior femora as in male. Length
of body 18-21 mm; length of pronotum 5-5.5 mm; width of pro-
notum 8. 5-8. 8 mm; length of tegmina 3-4 mm.

Material examined: cT ( 1 VM) (lectotype, herewith designated),

Figures 137-145. Calolampra marginalis. 137. $ (967 L), Rottnest Isl.,
W. A. 138. $ (180 L), Jandakot, W. A. 139. Tegmen of female shown in
figure 138. 140-142. $ (45 BMNH), Yanchep (32 miles north of Perth),

W. A. Genitalia (prepuce in figure 140 torn away from L2vm). 143-145.
Genitalia of $ shown in figure 137. Scale: Fig. 137 = 4 mm; Fig. 138 —

3 mm; Fig. 139 — 1 mm; Figs. 140-145 = 0.2 mm.

1973]

Roth & Princis — Genus Ccilolcunpra

I3i

145

132

Psyche

[March-June

Adelaide, S. A. ; 5 c? c? (260-264 L), Cabramatta, N.S.W., M.
Nikitin leg.; c? (114 MCZ), A. (without exact locality); cf (59
MCZ), A. (without exact locality) ; cf (25 SAM), Mt. Bryan E.,
S. A., 13.XII.1886, Best leg.; cf (26 SAM), Norwood, S. A.,
7. XII. 1885 (at light), Tepper leg.; cf (27 SAM), Yardea., S. A.,
16.XII.1952, G. F. Gross leg.; 9 (L), Cabramatta, N.S.W., 12.
XII. i960 (on trunk of Casuarina glauca) , M. Nikitin leg.; 9 (i39
MCZ), Blue Mts., Hartley Vale, N.S.W., 30.I.1932, Darlington
leg.

14. C. mcirginalis (Walker)

(Figs. 137-145, 274)

Ischnoptera marginalis Walker, Cat. Blatt. Brit. Mus. 1868, p. 119, $.

cf (Fig. 137). Head yellow with partly suffused reddish mark-
ings; face passing into occiput with an angulation between eyes. Pro-
notum triangular in dorsal aspect; disk almost uniformly reddish
brown, with suffused lyrate pattern which is indicated only by its
darker color; lateral margins transparent and their outer edging
yellow; the usual series of striae on hind margin almost completely
obscured by base color. Tegmina somewhat exceeding apex of ab-
domen, uniformly brownish (the darker maculation so usual in
other species is here completely lacking); marginal field whitish;
humeral stripe reddish brown, distally breaking up into branches
directed to anterior margins of tegmina. Dorsum of abdomen brown-
ish, distally becoming darker; first tergite as figured (Fig. 274).
Supra- anal plate whitish. Venter of abdomen yellowish distally
darker. Genitalia as shown in figures 140- 145; R2 beaklike (Figs.
1 41, 144). Legs yellowish; lower posterior margin of front femora
with a distal and 3 or 4 additional spines; same margin of posterior
femora with 0-2 spines. Length of body 17 mm; length of pronotum
4.5 mm; width of pronotum 5.8 mm; length of tegmina 17 mm.

9 (Fig. 138). Head as in male. Pronotum speckled with brown
and dotted with reddish ; disk with suffused lyrate pattern ; hind
margin convex in its middle part; la.tero-caudal angles not at all
produced backwards. Tegmina reduced to lateral lappets (Fig. 139),
their outer margins yellow. Dorsum of abdomen thickly speckled
with dark brown and sparsely dotted with reddish among the dark
speckles. Supra-anal plate with irregularly dentate hind margin.
Cerci reddish brown with yellowish apices. Venter of abdomen red-
dish brown with darker stigmatic maculae. Legs as in male. Length

1973 J

Roth & Princis — Genus Calolampra

Figures 146-152. Calolampra solida. 1+6. S (268 L in SM), holotype,
Peak Downs, Q. 147-148. 9 (4 L in SM), paratype, Peak Downs, Q. 147.
Dorsal view. 148. Tegmen of $. 149-152. Genitalia, supra-anal plate, and
hypandrium of $ holotype shown in figure 1+6. Scale: Fig. 146 — 6 mm;
Fig. 147 — 4 mm; Figs. 148, 151-152 = 1 mm; Figs. 149-150 = 0.2 mm.

134

Psyche

[March-June

of body 17.5-22.5 mm; length of pronotum 4.2-5 mm; width of pro-
notum 7. 5-8. 3 mm; length of tegmina 3-4.5 mm.

Material examined: c? (BMNH) (holotype), Swan River, W.
A., ex. Dr. Bacon’s coll.; cf (45 BMNH), Yanchep (32 miles
north of Perth), W. A. 20-31.XII.1935. R. E. Turner leg.; cf
(967 L), Rottnest Isl., W. A., 1940; 9 (180 L) Jandakot, W. A.,
1950; P (137 MCZ), Kings Park, Perth, W. A., 7. IX. 1931, W.
M. Wheeler leg.; 9 (L) Rottnest Isl., W. A., 1933; 9 (L), Kings
Park, Perth, W. A., XII.1951, T. Gislen leg.

15. C. solida sp. n.

(Figs. 146-152, 265)

cf (Fig, 146). Head yellow; face black from interocular area to
clypeus. Antennae with basal joint yellow, following 10-12 joints
brownish, remainder blackish. Pronotum triangular in dorsal aspect;
disk shining black, lateral margins yellow; the usual longitudinal
striae on hind margin only indicated by their color, not raised;
greatest width beyond the middle. Tegmina somewhat exceeding
apex of abdomen, semitransparent, rather sparsely furnished with
small brown speckles which are present even on the normally cov-
ered portion of right tegmen and are absent only in marginal field;
solidly black humeral stripe present. Wings as long as tegmina,
weakly infuscated and with brown veins. Dorsum of abdomen yel-
lowish; first tergite as figured (Fig. 265). Supra-anal plate (Fig.
15 1 ) with rounded latero-caudal angles and almost straight hind
margin. Hind margin of hypandrium weakly emarginate (Fig. 152).
Venter of abdomen yellow with blackish stigmatic markings. R2
and Li of genitalia shown in figures 149- 150 (L2vm and prepuce
accidentally lost). Legs pale brown to yellowish; lower posterior
margin of front femora with 1 additional spine; same margin of
hind femora unarmed or with one spine. Length of pronotum
6.6 mm; width of pronotum 8.8 mm; length of tegmina 25 mm.

9 (Fig. 147). Head and antennae as in male. Pronotum trans-
verse; disk shining black, provided with several scattered impressed
puntures; lateral margins yellow with blackish edging; the usual
longitudinal striae of hind margin suffused, nearly completely cov-
ered by black; middle part of hind margin convex; latero-caudal
angles not produced backwards. Tegmina (Fig. 148) reduced to
lateral vestiges; their outer margins with black edging. Wing ves-
tiges absent. Mesonotum black with a yellow marginal spot on each
lateral margin. Dorsum and venter of abdomen almost uniform

1973]

Roth & Princis — Genus Calolampra

135

Figures 153-164. Calolampra irrorata. 153-157. $ (59 BMNH), lecto-

type, A. (without exact location). 153. Adult. 154. Hypandrium. 155-157.
Genitalia. 158-162. $ (192 USNM), Cairns, Q. 158-159. Supra-anal and
subgenital plates. 160-162. Genitalia. 163. $ (2 L), Gayndah, Q. 164.

Tegmen of female shown in figure 163. Scale: Figs. 153, 163 — 4 mm;
Figs. 154, 158-159 = 0.5 mm; Figs. 155-157, 160-162 - 0.1 mm; Fig. 164
~ 1 mm.

136

Psyche

[March-J une

black. Legs as in male. Length of body 28 mm ; length of pronotum
7.5 mm; width of pronotum 10.5 mm; length of tegmina 4.5 mm.

Material examined: cf (268 L in SM) (holotype, herewith desig-
nated), Peak Downs, Q., ex Mus. Godeffroy; 9 (4 L in SM)
(paratype), same locality.

16. C. irrorata (Fabr.)

(Figs. 153-164, 268)

Blatta irrorata Fabricius, Syst. Ent. 1775, p. 272, $.

cf (Fig. 153). Pronotum triangular in dorsal aspect, with the
greatest width beyond its middle; lateral margins hyaline with their
edging lined dark brown inside; disk speckled with dark. Tegmina
well developed, somewhat exceeding apex of abdomen, covered with
rather small brown speckles which are partly present also in the
normally covered part of right tegmen. Wings as long as tegmina,
infumated and with brown veins. Abdomen above and below yellow-
ish. First tergite as figured (Fig. 268). Hind margins of supra-
anal plate (Fig. 158), and hypandrium (Figs. 154, 159) shallowly
emarginate mesally. Genitalia shown in figures 1 55- 1 57, 160-162.
Legs yellowish with brown spines; lower posterior margin of hind
femora armed with O to 2 spines; arolia well developed. Length of
body 23 mm; length of pronotum 5.5 mm; width of pronotum
8 mm ; length of tegmina 20 mm.

9 (Fig. 163). Pronotum transverse; disk thickly speckled with
black forming a lyrate pattern; lateral margins yellow, their edging
lined dark brown inside; hind margin straight in dorsal aspect and
provided with the usual series of black longitudinal striae which are
not raised but only marked by their color; latero-caudal angles dis-
tinctly produced backwards. Tegmina reduced to lateral vestiges
(Fig. 164) which are almost throughout yellow. Dorsum of abdo-
men thickly speckled with black and reddish brown. Supra-anal
plate yellow sparsely speckled and dotted with reddish. Venter of
abdomen almost uniform reddish brown, distally becoming darker.
Legs as in male but arolia very small. Length of body 23 mm;
length of pronotum 6 mm ; width of pronotum 9 mm ; length of
tegmina 4-4.5 mm.

Material examined: cf (59 BMNH), (lectotype, designated by
Roth and Princis, 1971, Figs. 27-29), A. (without exact locality),
ex Banks’ Coll.; cf (192 USNM), Cairns, Q., at light, J. F.
Illingworth leg.; 9 (2 L in SM), Gayndah, Q., ex. Mus. Godeffroy.

1973]

Roth & Princis — Genus Calolampra

137

Figures 165-175. Calolampra mjoebergi. 165. $ (296 L in SM), holo-

type, Colosseum, Q. 166. $ (143 MCZ), paratype, Ravenshoe, Atherton

Tab., Q. 167. Tegmen of $ shown in figure 166. 168-172. Supra-anal

plate, hypandrium, and genitalia of holotype shown in figure 165. 173-

174. $ (297 L), Malanda, Q. Genitalia. 175. $ (105 MCZ), paratype,

Ravenshoe, Atherton Tab., Q. R2 of genitalia. Scale: Fig. 165 — 5 mm;
Fig. 166 — 3 mm; Fig. 167 = 1 mm; Figs. 168-169 — 0.5 mm; Figs. 170-
175 — 0.2 mm.

138

Psyche

[March-June

17. C. mjoebergi sp. n.

(Figs. 165-175, 260)

cf (Fig. 165). Pronotum triangular in dorsal aspect, with great-
est width beyond its middle; disk speckled with black; lateral mar-
gins hyaline with their outer edging lined brown inside. Tegmina
somewhat exceeding apex of abdomen, furnished with scattered small
dark speckles which are lacking in the normally covered portion of
right tegmen. Wings as long as tegmina, weakly infumated and
with brownish veins. Dorsum of abdomen brownish, venter yellow-
ish. First tergite as figured (Fig. 260). Hind margin of supra-anal
plate convex, entire (Fig. 168). Hind margin of hypandrium
mesally very shallowly emarginate (Fig. 169). Genitalia shown
in figures 1 70-1 75. R2 relatively uniform in width and slender
throughout with rounded apex (Figs. 17 1, 1 74-1 75). Legs yellowish
with brown spines; lower posterior margin of hind femora armed
with 1-2 spines; well developed arolia present. Length of body
20 mm; length of pronotum 5.5 mm; width of pronotum 6.8 mm;
length of tegmina 19 mm.

9 (Fig. 166). Head pale brownish with a transverse black band
between eyes. Disk of pronotum as well as mesonotum, metanotum,
and tergites so thickly speckled with dark that they appear com-
pletely black (only dose examination shows that there are spots of
yellow ground color present among the dark speckles). Lateral
margins of pronotum yellow with numerous small reddish dots which
are crowded toward the disk; among these dots are several larger
blackish spots; outer edging of the lateral margins lined brown in-
side. Tegmina lobiform, yellowish with blackish dots, their apices
exceeding hind margin of mesonotum (Fig. 167). Posterior mar-
gins of pro- meso-, and metanotum as well as those of tergites pro-
vided each with a transverse row of somewhat raised black striae.
Supra-anal plate entire, yellow with numerous small reddish dots
and several larger blackish spots among the former. Legs as in male
but with very small arolia. Length of body 21-22 mm; length of
pronotum 6 mm; width of pronotum 9-9.5 mm; length of tegmina
4-4.5 mm.

Material examined: cf (296 L in SM) (holotype, herewith
designated), Colosseum, Q., Mjoberg leg. ; cf (297 L) (paratype).
Malanda, Q., Mjoberg leg.; cf (105 MCZ) (paratype), Raven-
shoe, Atherton Tab., Q., 8000 ft., IV.1932, Darlington leg.; 2 99
(142-143 MCZ) (paratypes), same data.

1973]

Roth & Princis — Genus Calolampra

139

Figures 176-181. Calolampra propinqua. 176. $ (266 L), holotype,

Wee Waa, N.S.W. 177. $ (267 L), paratype, Cabramatta, Blue Mts.,

N.S.W. 178. Tegmen of $ shown in figure 177. 179-181. Genitalia of $
holotype shown in figure 176. Scale: Fig. 176 — 4 mm; Fig. 177 — 3 mm;
Fig 178 — 1 mm; Figs. 179-181 = 0.2 mm.

140

Psyche

[March-June

1 8. C. propinqua sp. n.

(Figs. 176-181, 271)

cT (Fig. 176). Head yellowish, with a black patch in interocular-
ocellar area. Antennae yellowish distally becoming somewhat darker.
Pronotum triangular in dorsal aspect, widest beyond its middle ; disk
speckled and dotted with black ; edging of lateral margins lined
brown inside; hind margin with a series of black striae. Tegmina
somewhat exceeding apex of abdomen, covered with small blackish
spots; costal field uniform pale yellow; humeral streak black, dis-
tally broken up into separate flecks; the normally covered portion
of right tegmen solidly colored. Wings brownish, as long as tegmina,
with deep brown veins. Dorsum of abdomen dark brown; first ter-
gite as figured (Fig. 271). Supra-anal plate transverse, distally
weakly convex. Genitalia shown in figures 179-181. Venter of
abdomen yellow with black stigmatic maculae. Legs yellow with
brown spines ; posterior margin of hind femora unarmed ; well de-
veloped arolia present. Length of body 16 mm; length of pronotum
4 mm; width of pronotum 5.4 mm; length of tegmina 15.5-16 mm.

9 (Fig. 177). Head and antennae as in male. Pronotum speckled
and dotted with blackish and reddish; edging of lateral margins
yellow, lined brown inside; hind margin in dorsal aspect slightly
convex, with a series of black longitudinal striae; latero-caudal
angles weakly produced backwards. Tegmina reduced to lobiform
vestiges somewhat obliquely truncate (Fig. 178), edging of their
outer margins yellow. Dorsam of abdomen thickly speckled with
blackish. Supra-anal plate yellowish, provided with some dark dots.
Venter of abdomen castaneous. Legs yellowish with brown spines;
arolia absent. Length of body 16-18 mm; length of pronotum
44-5.2 mm; width of pronotum 6. 8-8. 2 mm; length of tegmina
3.5-4 mm.

Material examined: cf (266 L) (holotype, herewith designated),
Wee-Waa (at light), N.S.W., 21.I.1960, M. Nikitin leg.; $
(267 L) (paratype), Cabramatta, Blue Mts., N.S.W., 14.XII.1960,
M. Nikitin leg.; 9 (144 M'CZ) (paratype), Melbourne, V., H.
Edwards leg. $ (148 MCZ) (paratype), A. (without exact local-
ity), H. Edwards leg.

19. C. frctserensis (Tepper)

(Figs, 182-192, 263)

Epilampra fraserensis Tepper, Trans. R. Soc. S. Austral. XVII, 1893,
p. 59, $. '

<$ (Fig. 182). Pronotum broadly triangular in dorsal aspect, with

1973]

Roth & Princis — Genus Calolampra

141

Figures 182-192. Calolampra fraserensis. 182. $ (5 SAM), lectotype,
Fraser Range, W. A. 183. $ (138 MCZ), Rottnest Island, W. A. 184.
Tegmen of $ shown in figure 183. 185-188. Supra-anal plate, hypandrium,
and genitalia of $ lectotype shown in figure 182. 189-192. $ (SAM), Mt.
Bryan East, S. A. 189-190. Supra-anal plate and hypandrium. 191-192.
Genitalia. Scale: Figs. 182, 183 — 5 mm; Figs. 184 — 1 mm; Figs. 185-186,
189-190 = 0.5 mm; Figs. 187-188, 191-192 = 0.2 mm.

142

Psyche

[March-June

greatest width beyond its middle; disk speckled with dark forming
a lyrate pattern; lateral margins pale yellowish. Tegmina covered
with brown blotches which are partly confluent, especially in anal
area; the normally covered portion of right tegmen solidly colored.
Wings blackish, about as long as tegmina. First tergite as figured
(Fig. 263). Hind margin of supra-anal plate nearly straight (Figs.
185, 189), same margin of hypandrium emarginate (Figs. 186,
190). Genitalia shown in figures 187-188, 191-192. Venter of
abdomen as well as legs pale yellowish ; lower posterior margin of
hind femora unarmed; well developed arolia present. Length of
body 19 mm; length of pronotum 5 mm; width of pronotum 7.5
mm ; length of tegmina 24 mm.

$ (Fig. 183). Head reddish; face passing into occiput with a
distinct transverse angulation between eyes. Pronotum nearly smooth,
at most several large impressed punctures present; hind margin
straight in dorsal aspect. Tegmina reduced to lateral vestiges, apices
subacuminate (Fig. 184), edging of their outer margins dark brown.
Dorsum of abdomen thickly speckled with blackish. Venter of ab-
domen reddish in the middle, blackish laterally. Legs as in male.
Length of body 22 mm; length of pronotum 5.1 mm; width of pro-
notum 8 mm; length of tegmina 4.5 mm.

Material examined : cf (5 SAM) (lectotype, herewith designated).
Fraser Range, W. A., X.1891, R. Helms leg.; cf (24 SAM),
Mt. Bryan East, S. A., 13.XII.1886, J. Best leg.; $ (138 MCZ),
Rottnest Island, W. A., 23. X. 1931, P. J. Darlington leg.

20. C. notahilis (Walker)

(Figs. 193-215)

Epllampra notabilis Walker, Cat. Blatt. Brit. Mus. 1868, p. 202, $.

cf (Figs. 193, 200). Head yellowish, with a broad black trans-
verse band between eyes extending to the imaginary line connecting
antennae. Antennae with basal joint yellow and remainder fuscous.
Pronotum thickly speckled and dotted with blackish brown and
reddish; edging of lateral margins yellow; hind margin with dark,
radially arranged striae. Tegmina decidedly exceeding apex of ab-
domen, covered by numerous small dark dots; costal field yellow;
humeral streak broad, but beyond middle is broken up into separate
flecks; the normally covered portion of right tegmen solidly brown-
ish. Wings as long as tegmina, brownish, with deep brown veins.
Dorsum of abdomen pale, obscurely banded with brownish; first
tergite modified (Fig. 199). Supra-anal plate pale, with convex

1973]

Roth & Princis — Genus Calola?npra

14 3

Figures 193-199. Calolampra notabilis. $ (60 BMNH), holotype, A.

(without exact locality). 193. Adult. 194-196. Genitalia. 197-198. Supra-
anal and subgenital plates. 199. Modification of first tergite. Scale: Fig.
193 — .5 mm; Figs. 194-196 — 0.2 mm; Figs. 197-199 — 1 mm.

Psyche

[March-June

' a!

■

Figures 200-205. Calolampra notahilis. $ (295 L), V. (without exact

locality). 200. Adult. 201-202. Supra-anal plate and hypandrium. 202-205.
Genitalia. Scale: Fig. 200 = 4 mm; Figs. 201-202 = 1 mm; Figs. 203-
205 — 0.2 mm.

1973] Roth & Princis — Genus Calolampra 145

Figures 206-215. Calolampra notabilis. 206. $ (181 L), V. (without

exact locality). 207. Tegmen of $ shown in figure 206. 208-211. $ (23

SAM), Warunda, Eyrie Pen., S. A. 208-209. Supra-anal plate and hy-
pandrium. 210-211. Genitalia. 212-215. $ (151 MCZ), A. (without exact
locality). 212-213. Genitalia. 214-215. Supra-anal plate and hypandrium.
Scale: Fig. 206 = 4 mm; Fig. 207 = 2 mm; Figs. 208-209, 214-215 =

1 mm; Figs. 210-213 - 0.2 mm.

146

Psyche

[March-June

hind margin (Figs. 197, 201, 214) or very weakly emarginate (Fig.
208). Venter of abdomen yellowish with black stigmata. Hypan-
drium weakly emarginate (Figs. 198, 202, 209, 215), slightly ex-
ceeding supra-anal plate. Genitalia as figured (Figs. 194-196, 203-
205, 210-213). Legs yellowish with brown spines; lower posterior
margin of hind femora usually unarmed. Length of body 16.5 mm;
length of pronotum 5 mm; width of pronotum 6.5-7 mm;; length of
tegmina 21 mm.

$ (Fig. 206). Head generally as in male. Pronotum transverse,
its hind margin slightly convex in dorsal aspect; edging at lateral
margins uniform brown, not lined darker inside; latero-caudal angles
not produced backwards. Tegmina vestigial (Fig. 207) edging of
their outer margins yellow. Latero-caudal angles of metanotum
rounded. Legs as in male with very small arolia. Length of body
22.5 mm; length of pronotum 6 mm; width of pronotum 9-10 mm;
length of tegmina 4.5-5 mm.

Material examined: c? (60 BMNH) (holotype), A. (without
exact locality) ex Mr. Lambert’s Coll.; c? (295 L), V. (without
exact locality); cf (151 MCZ), A. (without exact locality), ex
S. H. Scudder’s Coll.; c? (23 SAM), Warunda, Eyrie Penin.,
S. A., 1 8.X. 1909, B. Zietz leg.; $ (181 L), V. (without exact lo-
cality).

21. C. queenslandica sp. n.

(Figs. 216-221, 270)

cf (Fig. 216). Head yellow; interocular-ocellar area with a
brown transverse band which in the latter area is medially inter-
rupted. Pronotum in dorsal aspect broadly triangular with greatest
width beyond its middle; disk with lyrate pattern formed by brown
speckles and dots; lateral margins yellow rather sparsely dotted with
brown; cephalic margin continuously arcuate with nondifferentiated
lateral margins ; hind margin provided with the usual series of brown
longitudinal striae. Tegmina somewhat exceeding apex of abdomen,
sparsely dotted with pale brownish; the normally covered portion of
right tegmen solidly colored. Dorsum of abdomen pale brownish;
first tergite as figured (Fig. 270). Venter of abdomen yellow with
brown stigmatic maculae. Hind margin of supra-anal plate (Fig.
217) and hypandrium (Fig. 218) weakly emarginate mesally. Gen-
italia shown in figures 2 19-221. Legs yellow with brown spines;
lower posterior margin of hind femora unarmed. Length of body

1973]

Roth & Princis — Genus Calolampra

147

Figures 216-221. Calolampra queenslandica. $ (271 L in SM), holo-

type, Q. (without exact locality). 216. Adult. 217-218. Supra-anal plate
and hypandrium. 219-221. Genitalia. Scale: Fig. 216 = 5 mm; Figs. 217-
218 — 1 mm; Figs. 219-221 = 0.1 mm.

Psyche

148

[March-J une

20 (?) mm; length of pronotum 5.4 mm; width of pronotum
8 mm ; length of tegmina 20 mm.

?. Unknown.

Material examined: cf (271 L in SM) (holotype, herewith desig-
nated), Q. (without exact locality).

22. C. atomiferci (Walker)

(Figs. 222-248, 272)

Epilampra atomifera Walker, Cat. Blatt. Brit. Mus. 1868, p. 69, $.

Polyzosteria propria Walker, Cat. Blatt. Brit. Mus. 1868, p. 161, $.

Epilampra notabilis Tepper (nec Walker 1868), Trans. R. Soc. S. Aus-
tral. XVII, 1893, p. 59, part. $ $.

cf (Fig. 222). Pronotum broadly triangular in dorsal aspect,
with greatest width beyond its middle ; disk speckled and dotted with
reddish to deep brown. Antennae ferrugineous, except the pale
yellowish basal joint. Tegmina somewhat exceeding apex of abdo-
men, transparent, but sparsely dotted with reddish brown; the nor-
mally covered portion of right tegmen solidly colored. Wings as
long as tegmina, moderately infumate and with deep brown veins.
First tergite as figured (Fig. 272). Supra-anal plate pale yellowish
with slightly emarginate (Figs. 225, 233, 237, 245), or entire (Figs.
229, 241) hind margin. Cerci and hypandrium pale yellowish; hind
margin of the latter almost straight (Fig. 226) or weakly concave
(Figs. 230, 234, 238, 242, 246). Genitalia shown in figures 227-
228, 231-232, 235-236, 239-240, 243-244, 247-248. Legs yellowish
with brown spines; lower posterior margin of hind femora unarmed.
Length of body 17-19. 5 mm; length of pronotum 4-5.2 mm; width
of pronotum 6-7.5 mm; length of tegmina 17-19. 5 mm.

9 (Fig. 223). Pronotum at most provided with several scattered
large impressed punctures (not with numerous, fine, impressed dots) ;
hind margin straight in dorsal aspect, at least in its middle part.
Tegmina reduced to lateral lappets; edging of outer margins of
lappets brown (not black) ; apex of tegminal vestiges rounded (Fig.
224). Latero-caudal angles of pronotum slightly produced back-
wards. Face regularly arcuate with occiput (no angulation present).
Abdomen above thickly speckled with dark. Underside of abdomen
reddish in the middle, black laterally. Legs yellowish; lower pos-
terior margin of front femora armed only with 1 distal spine; same
margin of hind femora usually completely unarmed. Length of body
17.3-20 mm, length of pronotum 5-5.2 mm, greatest width of pro-
notum 8-9 mm; length of tegmina 3.5 mm.

1973]

Roth & Princis — Genus Calolampra

149

Figures 222-232. Calolampra atomifera. 222. $ (64 BMNH), holotype,
Nova Hollandia (without exact locality). 223. $ (65 BMNH), lectotype of
Polyzosteria propria Walker, S. A. (without exact locality). 224. Tegmen
of female shown in figure 223. 225-228. Supra-anal plate, hypandrium, and
genitalia of $ holotype shown in figure 222. 229-232. $ (7 SAM), Ade-
laide, S. A. Supra-anal plate, hypandrium, and genitalia. Scale: Figs. 222-
223 = 4 mm; Figs. 224, 225-226, 229-230 = 1 mm ; Figs. 227-228, 231-
232 — 0.1 mm.

[March-June

150

Psycht

Figures 233-248. Calolampra atomifera. Male supra-anal plates, sub-
genital plates, and genitalia. 233-236. (6a SAM), Ardrossan Cliffs, S. A.
237-240. (8 SAM), Bordertown, S. A. 241-244. (9 SAM), Happy Valley
(near Adelaide), S. A. 245-248. (4 SAM), Ardrossan. S. A. Scale: Figs.
233-234, 237-238, 241-242, 245-246 = 1 mm ; Figs. 235-236, 239-240, 243-
244, 247-248 = 0.2 mm.

1973]

Roth & Princis — Genus Calolampra

I5i

Material examined: cf (64 BMNH), (holotype of atomiferd) ,
Nova Hollandia (without exact locality), ex Mr. Hunter’s Coll. ;
9 (65 BMNH), (lectotype of propria ), S. A. (without exact lo-
cality), ex R. Bakewell’s Coll.; cf (4 SAM), Ardrossan, S. A.,
I.1884, Cadd leg.; this specimen was labelled by Tepper as type of
notabilis, but his action is invalid because the specimen did not be-
long to the original material used by Walker; cf (6a SAM), Ar-
drossan Cliffs (in moist dust), S. A., 26.XI.1885, Tepper leg.;. this
specimen was labelled by Tepper as cotype of notabilis , but his action
is invalid (see above) ; cf (7 SAM), Adelaide, S. A., Botan. Park
(under bark) 17. XII. 1885, Tepper leg.; 9 (SAM) Callington,
S. A., (under rubbish) 19.I.1886, Tepper leg.; cf (8 SAM), Bor-
dertown, S. A., (under bark of Eucalyptus) , 8.1. 1878, Tepper leg.;
9 (SAM), Adelaide, S. A., 29.VI.1892, A. Zietz leg.; cf (9
SAM), Happy Valley (near Adelaide), S. A., 27.VIII.1903, Burs-
lem leg.

23. C. obscura (Tepper)

(Figs. 249-252, 264)

Epilampra obscura Tepper, Trans. R. Soc. S. Austral. XVII, 1893,
p. 64, 9.

Calolampra submarginalis Princis, Lund Univ. Arsskr., N. F. Avd. 2,
Vol. 50, No. 13, (also K. Fysiogr. Sallsk. i Lund Handl. N. F. Vol. 65,
No. 13, 1954, p. 32, $ and 9 nymph (not $ and $ as given), syn. nov.

cf (Fig. 249). Head yellowish, interocular space piceous. An-
tennae fuscous with basal joint yellowish. Pronotum thickly speckled
with blackish on pale yellowish ground ; disk with symmetrical mark-
ing of dark brown; hind margin furnished with dark longitudinal
striae. Tegmina considerably exceeding apex of abdomen, thickly
speckled with dark; costal margin pale yellowish. Wings blackish,
as long as tegmina. First tergite shown in figure 264. Supra-anal
plate subquadrate, with latero-caudal angles rounded. Venter pale
yellowish with dark stigmatic dots on sternites. Hypandrium emar-
ginate. Genitalia shown in figures 250-252. Legs pale yellowish;
lower posterior margin of hind femora unarmed. Length of body
18-19 mm; length of pronotum 5 mm; width of pronotum 6.8-
7.2 mm; length of tegmina 18-18.5 mm.

9. Dull reddish brown. Pronotum furnished with many fine,
impressed dots. Tegmina reduced to lateral lappets with rounded
apices. Wings absent. Length of body 23 mm ; length of pronotum
5 mm; width of pronotum 10 mm; length of tegmina! vestiges

152

Psyche

[March-June

Figures 249-259. Calolampra spp. 249-252. C. ohscura. 249. $ (270 L),
West Kimberly, W. A. 250-252. Male genitalia of specimen shown in figure
249. 253-256. $ C. insularis. (107 MCZ), holotype, Murray Island, Torres
Strait. 253-254. Genitalia. 255-256. Supra-anal and subgenital plates. 257-
259. C. pernotabilis. (1 L in SM), $ holotype, Mt. Tambourine, Q. Geni-
talia. Scale: Fig. 249 — 6 mm; Fig. 250 — 0.1 mm; Figs. 251-254, 257-
259 = 0.2 mm; Figs. 255-256 — 1 mm.

1973]

Roth & Princis — Genus Calolampra

153

3 mm. The specimen described by Princis (1954, p. 33) as a female
is in reality an immature individual (probably of last nymphal
stage) .

Material examined: 9 (SAM) (lectotype of obscura, herewith
designated), N. T., 3.VIII.1886, Dr. Magarey leg.; 2 nymphs
(SAM), same data; cf (L) (paratype of sugmarginalis) , West
Kimberley, northern W. A., 1946; 2 cf cf (270 L and L in SM),
Kimberley District, northern W. A., Mjoberg leg.; 4 nymphs, same
data; nymph (SM), Noonkangah, northern W. A., Mjoberg leg.;
nymph (SM), Derby, northern W. A., Mjoberg leg.; cf (SAM),
Port Hedland, northern W. A., 8.VII.1953, N. B. Tindale leg.

24. C. insularis sp. n.

(Figs. 253-256, 267)

cf. Pronotum triangular in dorsal aspect with greatest width
beyond its middle; disk with black lyrate pattern; lateral margins
yellow speckled with some larger black punctures and dotted with
many small reddish ones; lateral angles broadly rounded. Tegmina
well developed, somewhat exceeding apex of abdomen, speckled with
deep brown, however, distally becoming paler; the normally covered
portion of right tegmen solidly colored. Wings as long as tegmina,
infumate with brown veins. Abdomen above brown, below yellow-
ish. First tergite as in figure 267. Genitalia shown in figures 253-
254. Hind margin of supra-anal plate (Fig. 255) and hypandrium
(Fig. 256) distinctly incised mesally. Legs yellowish with brown
spines; lower posterior margin of hind femora with one spine. Well
developed arolia present. Length of body 20 mm ; length of pro-
notum 5 mm; width of pronotum 7.5 mm; length of tegmina 20 mm.

9. Unknown.

Material examined: cf (107 MCZ) (holotype, herewith desig-
nated), Murray Island, Torres Strait, H. L. Clark leg.

25. C. pernotabilis sp. n.

(Figs. 257-259, 276, 280-282)

cf. Head yellow; interocular-ocellar area blackish, parted by a
narrow pale median line into two parts. Pronotum in dorsal aspect
triangular, with the greatest width at its middle; disk speckled with
blackish and dotted with reddish brown, the blackish speckles form-
ing a lyrate pattern ; lateral margins lined brown inside ; hind mar-
gin bearing a series of dark brown longitudinal striae. Tegmina

154

Psyche

[March-June

Figures 260-279. Schematic outlines (not drawn to scale) of ridges found
on the first abdominal tergites of Calolampra males. 260. C. mjoebergi .
261. C. paula. 262. C. aspera. 263. C. fraserensis. 264. C. obscura.

265. C. solida. 266. C . candidula. 267. C. insularis. 268. C. irrorata.

269. C. confusa. 270. C. queenslandica. 271. C. propinqua. 272. C. atomi-
fera. 273. C. gracilis. 274. C. marginalis. 275. C. subgracilis. 276. C.
pernotabilis. 277. C. signatura (67 MCZ, holotype of C. aliena) . 278. C.
ignota. 279. C. atra. (drawings by K. Princis).

1973]

Roth & Princis — Genus Calolampra

155

decidedly exceeding apex of abdomen, with scattered brown macula-
tion on yellowish ground; broad dark brown humeral stripe present;
the normally covered part of right tegmen solidly brown colored.
Wings as long as tegmina, weakly infuscated and with brown veins.
Dorsum of abdomen yellowsh-brown. First tergite as in figure 276,
very similar to the modification of notabilis (cf. fig. 199). Genitalia
shown in figures 257-259. Venter of abdomen yellow with dark
brown stigmatic maculae. Legs yellow with brownish spines; lower
posterior margin of front femora with 1 distal and 1 additional spint,
same margin of posterior femora completely unarmed. Length of
body 20 (?) mm; length of pronotum 5.1 mm; width of pronotum
6.6 mm; length of tegmina 21 mm.

9 (Fig. 280). Head generally as in male, but with an additional
blackish brown macula between interocellar area and clypeus (Fig.
282). Pronotum less than twice as broad as long; disk bearing a
blackish lyrate pattern and several scattered impressed punctures;
lateral margins lined brown inside; hind margin straight in its mid-
dle part and provided with a series of blackish longitudinal striae;
latero-caudal angles slightly produced backwards. Tegmina reduced
to lateral lappets (Fig. 281) ; edging of their outer margins yellow.
Dorsum of abdomen thickly speckled with dark brown and dotted
with reddish. Supra-anal plate yellowish, sparsely dotted with red-
dish brown. Venter of abdomen dark to blackish brown with several
included reddish maculations. Legs as in male. Length of body
16.5-17.5 mm; length of pronotum 3. 8-4. 5 mm; width of pronotum
7 mm; length of tegmina 3-3.5 mm.

Material examined: c? ( 1 L in SM) (holotype, herewith desig-
nated), Mt. Tambourine, Q., Mjoberg leg.; 9 (SM) (paratype,
herewith designated) and nymph (SM), Gayndah, Q.; 9 (L)
(paratype, herewith designated).

26. C. tepperi Kirby

Epilampra propria Tepper (nec Walker 1868), Trans. R. Soc. S. Austral.
XVII, 1893, p. 64, part., only $ from Kangaroo Island (nec $ — Calo-
lampra atra ).

Calolampra tepperi Kirby, Ann. Mag. Nat. Hist. (7) XII, 1903, p. 275,
$ from Kangaroo Island.

cf. Unknown.

9- Head yellowish with three reddish transverse bands between
eyes, antennae, and clypeus. The band between the eyes is broken
up into 4 indistinctly limited parts. Disk of pronotum is largely
clouded with dark including some pale spots and lines; the usual

Psyche

[March-June

156

Figures 280-282. Calolampra pernotabilis. $ (SM), paratype, Gayndah,
Q. 280. Dorsal view. 281. Tegmen. 282. Head, frontal view. Scale:.
Fig. 280 — 3 mm; Figs. 281-282 — 1 mm.

1973]

Roth & Princis — Genus Calolampra

157

lyrate pattern is vaguely indicated ; lateral and anterior margins are
brown, contrastingly different from the yellow color which encircles
the disk; hind margin in dorsal aspect weakly convex, bearing a
series of dark striae which are not raised but only indicated by their
darker color. Tegminal pads lateral, their apices somewhat exceeding
the weakly concave hind margin of mesonotum. Meso-and metano-
tum as well as abdominal tergites much the same color as disk of
pronotum; their hind margins also bear similar longitudinal striae
but their distal ends are generally raised and only the remaining
portions are indicated by color. Hind margin of supra-anal plate
arcuate and mesally weakly emarginate. Venter of abdomen brown
with yellow maculations. Lower posterior margin of front femora
armed with one distal and one additional spine, the same margin of
hind femora completely unarmed. Length of body 19 mm; length
of pronotum 5 mm ; width of pronotum 8 mm ; length of tegmina
4 mm.

This species may prove to be identical with Brancsik’s Calolampra
depolita (Jheft Naturw. Ver. Trencs. Comit. XIX-XX, 1898,
P. 57, ?)•

Material examined: $ (SAM) (lectotype of tepperi, herewith
designated), Kangaroo Island, S. A., 1-6.III.1886, Tepper leg.
Kirby proposed the name Calolampra tepperi for Tepper’s propria,
however, without lectotype designation. Shaw selected a male and a
female from Tepper’s material of propria and attached his type labels
to these specimens using the name Calolampra tepperi. However,
his action cannot be accepted as proper lectotype designation because
he did not publish his designation; 2 nymphs (SAM), Kangaroo
Island, 1-6.III. 1 886, Tepper leg.; nymph (SAM), Kangaroo Island
(southern coast), 16. 1. 1906, O. Rail Leg.

Acknowledgements

We thank the following for the loan of museum material: Dr.
David R. Ragge, British Museum (Natural History) ; Dr. E. C.
Dahms, Queensland Museum ; Dr. Howard Evans, Museum of
Comparative Zoology, Harvard University; Dr. Ashley Gurney,
United States National Museum; Dr. L. E. Koch, Western Aus-
tralian Museum; Dr. G. F. Gross, South Australian Museum;
Dr. A. Kaltenbach, Vienna Museum of Natural History.

We thank Mr. Samuel Cohen for taking most of the photographs.

158

Psyche

[March-June

References

Baijal, H. N. and Kapoor, V. C.

1966. On a new species of cockroach from northwest Himalayas. Agra
Univ. J. Res. 15 (2) :5-8.

Bey-Bienko, G. Y.

1969. New genera and species of cockroaches (Blattoptera) from

tropical and subtropical Asia. Entomol. obozr. 48:831-862.

(Russian: English trans. in Entomological Review, 84:528-548:
1970).

Princis, K.

1963. Orthopterorum Catalogus, Edit. M. Beier. 4. ’s» — Gravenhage

pp. 76-172.

1967. Orthopterorum Catalogus, Edit. M. Beier. 11. ’s-Gravenhage,

pp. 616-710.

1970. La faune terrestre de l’lle de Sainte-Helene (Premiere partie).
Ann. Mus. Roy. Afr., in -8°, Zool., 181: 167-176.

Rehn, J. A. G. and Hebard, M.

1927. The Orthoptera of the West Indies 1. Blattidae. Bull. Amer.
Mus. Nat. Hist., 54:1-320.

Roth, L. M.

1969. The evolution of male tergal glands in the Blattaria. Ann.
Entomol. Soc. Amer., 6 2:176-208.

Roth, L. M. and Princis, K.

1971. Pseudocalolampra, a new genus of cockroach from Africa (Dic-
tyoptera :Blaberidae) . Proc. Entomol. Soc. Washington, 73:329-
336.

Shaw, E.

1925. New genera and species (mostly Australasian) of Blattidae,
with notes, and some remarks on Tepper’s types. Proc. Linn.
Soc. New South Wales, 1:171-213.

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PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol. 80 September, 1973 No. 3

CONTENTS

The Evolution of Cryptic Postures in Insects, with Special Reference

to Some New Guinea Tettigoniids (Orthoptera) .

Michael H. Robinson 159

Cretaceous Aculeate Wasps from Taimyr, Siberia (Hymenoptera).

Howard E. Evans 166

Paramuzoa (Nyctiborinae) , a New Cockroach Genus Previously Con-
fused with Parasphaeria (Epilamprinae) . Louis M. Roth 179

The Milliped Family Rhiscosomididae (Diplopoda: Chordeumida:

Striarioidea). William A. Shear 189

Supplementary Studies on Ant Larvae: Cerapachyinae, Pseudomyrme-
cinae and Myrmecinae. George C . Wheeler and Jeanette Wheeler 204
Studies on Neotropical Pompilidae (Hymenoptera). IX. The Genera

of Auplopodini. Howard E. Evans 212

Notes on Heteroonops and Triaeris (Araneae; Oonopidae).

Arthur M. Chickering 227

The Utilization of Various Asclepias Species by Larvae of the

Monarch Butterfly, Danaus plexippus. James M. Erickson 230

Copulatory Pattern Supports Generic Placement of Schizocosa avida

(Walckenaer) (Araneae: Lycosidae). Jerome S. Rovner 245

The Male Genitalia of Blattaria. X. Blaberidae. Pycnoscelus, Stilpno-
blatta, Proscratea (Pycnoscelinae) , and Diploptera (Diplopterinae) .

Louis M. Roth 249

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1973-1974

President .
Vice-President .
Secretary .

Treasurer .

Executive Committee

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EDITORIAL BOARD OF PSYCHE
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University

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The March-June, 1973 Psyche (Vol. 80, nos. 1-2) was mailed September 7,
1973

The Lexington Press. Inc.. Lexington. Massachusetts

PSYCHE

Vol. 80

September, 1973

No. 3

THE EVOLUTION OF CRYPTIC POSTURES IN
INSECTS, WITH SPECIAL REFERENCE TO SOME
NEW GUINEA TETTIGONIIDS (ORTHOPTERA)*

By Michael H. Robinson

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

Introduction

A number of authors (see, for instance, Cott 1940, and de Ruiter
1952) have suggested that the structural adaptations to defense by
concealment that are found in insects can only function efficiently
if they are accompanied by appropriate behavior patterns. These
behavior patterns include diurnal immobility and the adoption of
complex resting positions. A resting position may involve both the
selection of an appropriate location (background selection) and the
assumption of a special resting posture. The latter may entail sys-
tems that function to suppress signals that predators could use to
locate the insect, and in addition may give rise to signals that convey
false information about the edibility of the insect. The first category
involves the strategy of concealment, the second category involves
the strategy of mimicry.

I have suggested (Robinson 1968, 1969a, 1969b, 1973) that the
protraction of the anterior legs of stick-mimicking insects (particu-
larly phasmids and mantids, but including insects of other orders)
in line with the long axis of the body has, at the very least , a dual
function with respect to the signals that are potentially detectable by
the predator. Thus, this behavior may :

(a). Conceal the legs, head and antennae of the insect, thereby
suppressing signals that could be used as prey-detection cues by a
predator.

* Manuscript received, hy the editor June 6, 1973.

159

i6o

Psyche

[September

(b). Enhance the general resemblance of the insect to a stick by
increasing the apparent length of the body and providing it with a
long tapering termination, thereby adding to the plant-part mimicry
and signalling false information about edibility to (insectivorous)
predators. Experiments that show that some predators can use the
presence of heads and legs as prey-detection cues are described by
Robinson (1973).

It seems probable that the behavioral and structural devices that
serve to conceal prey-detection cues and also have a mimetic function
evolved in the first place as part of a strategy of concealment and
then constituted important steps towards the specializations involved
in stick- or leaf-mimicry. Thus, for instance, insects that rest against
a substrate can achieve maximum concealment by suppressing ‘relief’
or profile. This can be achieved by flattening or elongation, or both.
Flattening could be a starting point for leaf-mimicry and elongation
a starting point for stick-mimicry (examples in Robinson 1969b, but
see also the recent careful study of Ghanian praying mantids by
Edmunds 1972). Two examples of cryptic postures in tettigoniids
from New Guinea are detailed in this paper. Both involve adapta-
tions that are clearly related to concealment and at the same time
dead-ends in the sense that they do not lie on the path to leaf-mimi-
cry as it has been achieved in the orthoptera. Both adaptations are
complex and interesting in themselves. Both involve the concealment
of cue-structures and both involve profile reduction.

Materials and Methods

The insects were observed at the Wau Ecology Institute, Wau,
Morobe District, New Guinea during the period April 1970 to
April 1971, as part of a comprehensive study of insect anti-predator
adaptations. This mainly involved the rich phasmid fauna of the
area.1 Both species were collected at night at the Institute and also
at other localities in the Wau region. They were identified by Mrs.
Judith Marshall of the British Museum (N.H.) London to whom
the author is most grateful. Specimens are deposited with the Mu-
seum. Behavioral observations were carried out both in the field
and in a large screened insectary. More than ten specimens of each
species were examined.

Observations made on more than thirty species of phasmid will be pub-
lished as soon as the insects can be identified.

1973]

Robinson — Cryptic Postures

1 6 1

Psyche

[September

162

Description of Defensive Behavior
1. A cauloplacella immunis Brunner (Fam: Pseudophyllinae)

Specimens of this insect collected at Wau were a more-or-less
uniform bright green in color. At night when actively moving about
vegetation the insect looked like a ‘typical’ unspecialized tettigoniid.
However by day the insect assumed a resting attitude on leaves (both
upper and lower surfaces) that is shown in Figure 1.

This posture involves fairly complex changes in the orientation of
the tegmina, the alignment of the legs and also in the relationship of
the head, thorax and abdomen to the substrate. The change in the
orientation of the tegmina is the most striking. Note, from Figure 1.,
that the tegmina are kept together at their posterior margins (which
lie approximately in the midline of the body) and that their anterior
margins (which lie lateral to the insect) are closely applied to the
substrate. In the attitude of the active insect the angle between the
contiguous tegmina is less than 90° (i.e. between their internal
surfaces) while it becomes very obtuse (closer to 180°) in the flat-
tened cryptic posture. In the cryptic posture the tegmina form a
carapace-like structure that covers the second and third leg pairs.
This change in the orientation of the tegmina is achieved by slow
transition. In effect as the insect moves from a locomotory stance
into its resting posture it lowers the body against the substrate, re-
orients the limbs, tucks the ventral part of the head beneath the
prothorax and ‘feathers’ the tegmina outwards. In the resting pos-
ture the insect has a very low profile and the anterior legs are pro-
tracted side-by-side enclosing the antennae. The second and third
leg pairs lie beneath the expanded tegmina: concealed completely or
with part of the tarsi projecting.

It seems probable that in order to achieve this position the mus-
culature of the tegminal base must be modified in some way and that
each tegmen would exhibit some structural modification at its base.
Perhaps some of the movements involved in stridulation require
muscles and a form of tegminal articulation that facilitate the step
to this form of profile concealment.

Figure 2. Stages A, B, and C in the assumption of the full cryptic posture
(C) of Phyllophora sp. In A the major elements of the third leg are ap-
posed and the leg is being moved towards the anterior margin of the left
tegmen. In B the leg is about to be moved under the edge of the tegmen.
In C the leg has been rotated at its base, moved under the tegmen and the
tibia now lies closer to the midline of the body than the femur. Note the
position of the head and antenna in the final stage.

1973]

Robinson — Cryptic Postures

163

164

Psyche

[September

2. Phyllophora sp. * (Fam: Phyllophorinae)

Specimens of this robust dark-green insect assumed diurnal resting
attitudes on small branches. In the process the long third legs were
folded at the femorotibial joint so that the tibia was closely apposed
to the femur (inside edge to inside edge) and the apparent unit
formed in this way was then tucked beneath the lower edge of the
tented tegmen (its anatomically anterior margin). On one occasion
when we filmed the process the insect brought first one tarsus for-
wards to the jaws, beneath the body, then the other. The tarsal
region was groomed, in each case, and then the folding process was
finished and the leg fitted into its cryptic stance. As the long hind
legs are fitted into position beneath the tegmina the insect settles
down so that its ventral surface is in contact with the substrate. At
this stage legs I & II brace the resting insect. Interestingly enough
the coloration of the outer margin of the tibia III was much paler
(with pink overtones) than the rest of the joint. This coloration
closely matches that of the ventral surface of the abdomen against
which the folded unit is apposed. This coloration is visible only
when the insect is viewed from below. Figure 2 shows the process
of leg-folding and the final cryptic posture. We observed similar
behavior in a very much larger phyllophorine that we did not collect.

Discussion

The Phyllophora sp. device can be regarded as primarily an adap-
tation for concealing the large (“characteristic”) jumping legs of the
orthopteran. This conclusion is based on comparison with function-
ally similar devices in other insects and is supported by the fact that
the posture is adopted during the period of diurnal immobility when
the insect is presumably at risk from visually hunting predators. On
the other hand it is not a form of cryptic behavior consistent with
the main line of evolution of leaf-mimicry in the Tettigoniidae. In
that line leaf-mimicry has been achieved by flattening in the sagittal
plane and reduction in the length of the tegmina (examples in
Chopard 1938, Robinson 1969b).

The cryptic posture adopted by Acauloplacella immunis is a most
interesting one in that it reduces profile and affords leg-concealment
at the same time. It does not involve any marked dorso-ventral
flattening in the active insect. It is an adaptation that is essentially
more similar to the postural flattening of bark-living frogs and geckos
(see Cott 1940) than other forms of crypsis found in the Orthoptera.

♦Close to P. cheesmanae and P. similis de Jong.

1973]

Robinson — Cryptic Postures

165

Neither of the insects showed any of the forms of secondary de-
fense that Robinson (1969a) suggested were consequences of escape-
inhibiting cryptic postures. Both postures could be regarded as in-
hibiting the possibility of immediate escape following the penetration
of the first line of defense. Thus in the Phyllophora position the
jumping legs are in such a position that immediate escape by jump-
ing is not possible although the animal can push itself off the sub-
strate and drop. Similarly with Acauloplacella. Despite this neither
animal had a startle display, chemical secretion or was armed with
defensive spines.

Many of the orthopteroid insects that occur in this region of
New Guinea have complex secondary defenses and in particular use
strong spines in defense. This may be correlated with the fact that
most of the mammalian predators of insects (marsupials) are noc-
turnal and handle their prey. They may thus be less susceptible to
visual defenses and more affected by mechanical counter-attack.

References

Chopard, L.

1938. La biologie des Orthopteres. Paris: Lechavalier.

Cott, H. B.

1940. Adaptive coloration in animals . London: Methuen.

Edmunds, M.

1972. Defensive behaviour in Ghanian praying mantids. Zool. Journal
Linnean Soc. London. 51:1-32.

Robinson, M. H.

1968. The defensive behavior of Pterinoxylus spinulosus Redtenbacher,
a winged stick insect from Panama. Psyche. 75: 195-207.

1969a. The defensive behaviour of some orthopteroid insects from Pan-
ama. Trans. Royal Entomological Soc. London. 121: 281-303.
1969b. Defenses against visually hunting predators. In Dobzhansky
et al. Evolutionary Biology. 3 : 225-259.

1973. Insect anti-predator adaptations and the behavior of predatory
primates. Actas del IV Congresso Latinamericano de Zoologia
II. 811-836.

Ruiter, L. de.

1952. Some experiments on the camouflage of stick caterpillars. Be-
haviour. 4: 222-232.

CRETACEOUS ACULEATE WASPS FROM
TAIMYR, SIBERIA ( HYMENOPTERA) *

By Howard E. Evans
Mliseum of Comparative Zoology1

It seems incredible that only 16 years ago no aculeate Hymenop-
tera were known from the Mesozoic. In 1957 Sharov described
Cretavus from the Upper Cretaceous, an enigmatic form almost
certainly belonging to the Scolioidea. Ten years later Wilson,
Carpenter, and Brown described a beautifully preserved worker ant
from Upper Cretaceous amber, and in 1969 I described two dis-
similar wasps from the Upper Cretaceous and an undoubted aculeate
wing from the Lower Cretaceous. Thus within a few years it be-
came evident that the Aculeata were well diversified by the end of
the Mesozoic. Recent finds in the U.S.S.R. have more than con-
firmed that impression, and When fully described will document a
radiation so pronounced that events in the Tertiary will appear
anticlimactic.

It is the purpose of this brief report to provide names and de-
scriptions for several striking wasps occurring in Upper Cretaceous
amber from the Taimyr Peninsula, Siberia. These specimens were
kindly made available to me for study by Dr. A. Rasnitsyn of the
Palaeontological Institute of the U.S.S.R. Academy of Sciences,
Moscow. All specimens have been returned for deposit in that in-
stitution. All specimens bear the following data, as supplied by
Dr. Rasnitsyn: North Siberia, Taimyr Peninsula, Maimetcha River
[a branch of the Kheta River, Khatanga Basin], Yantardakh Hill
[3 km up from mouth of the Maimetcha] ; amber of Upper Cre-
taceous age, Coniacian-Santonian stage, Kheta formation.

This material contains several relatively well-preserved, more or
less complete specimens. As might be anticipated, some of them are
quite generalized and difficult to place in the commonly accepted
superfamilies. One of these I tentatively place in the Scolebythidae,
a recently-described family that I consider annectant between the
Scolioidea and Bethyloidea. Another I interpret as a probable gen-

®Published with the aid of a grant from the Museum of Comparative
Zoology.

Present address: Dept, of Entomology and Zoology, Colorado State Univ.,
Fort Collins, Colorado 80521.

Manuscript received by the editor , June 7 , 1973.

1973]

Evans — Cretaceous A culeate W asps

167

eralized sphecoid wasp. One specimen is quite clearly a sphecoid, a
pemphredonine probably related to Lisponema, described from Cre-
taceous Canadian amber (Evans, 1969). The bulk of the material
consists of Bethyloidea (26 of 29 specimens), including the first
records of Bethylidae from prior to the Oligocene.

As might be expected, all specimens are small (under 5 mm) and
all represent forms likely to be associated with trees. Present-day
Bethylidae attack larvae of Microlepidoptera and small Coleoptera,
while Cleptidae (the most abundant Aculeata in Taimyrian amber)
are parasites of sawfly larvae or the eggs of Phasmida. Many living
Pemphredoninae are associated with woody plants, and since the
Cretaceous forms lack spinose legs it seems a safe assumption that
they were xylicolous. The absence of larger wasps and of fossorial
forms is, I believe, merely an artifact, as such insects are unlikely to
become fossilized in small pieces of resin. But the diversity of non-
fossorial forms leads one to believe that the total aculeate fauna may
have been surprisingly rich.

? FAMILY SPHECIDAE

Taimyrisphex, new genus

Known from a single male approximately 4 mm in length, fully
alate [legs missing except coxae, front and middle trochanters, and
front femur] (Figs. 1, 2). Head about as wide as thorax; lower
part of front roundly prominent, overhanging bases of antennae,
the latter 13-segmented, very short, approximately capable of reach-
ing apices of front coxae, scape barely longer than thick, flagellar
segments about as long as thick (except ultimate segment 1.5 X as
long as thick) ; eyes large, reaching from close to top of vertex to
base of mandibles, inner margins weakly emarginate; ocelli large,
lateral ocelli removed from eye margins by less than their own
diameters; occipital carina present at least laterally; labial palpi
short, 4-segmented ; maxillary palpi slightly longer [probably 6-
segmented; details of mandibles and clypeus not clearly visible].
Pronotum sloping smoothly to collar, with small, rounded posterior
lobes which nearly reach the tegulae; dorsal and lateral faces of
pronotum separated by a subcarinate ridge; posterior margin of
pronotum forming a smooth arc between posterior lobes; meso-
scutum long, weakly convex, the notauli deeply and broadly im-
pressed on the posterior fourth, extending as weak lines almost
to anterior margin of scutum ; parapsidal furrows present, linear,
nearly complete; scutellum convex, with a transverse basal impres-

Psyche

[September

1 68

sion; metanotum rather long, but postnotum not visible; propodeum
with smooth contours, posterior rim well developed; mesopleurum
strongly convex, undivided by grooves; mesosternum simple; middle
coxae contiguous; coxae subconical; front femur simple, elongate.
Wing venation as figured. Metasoma sessile; tergite i convex,
forming a weak constriction at junction with tergite 2; sternite I
nearly flat in profile, hind margin nearly straight, thin, slightly
overlapping base of sternite 2, which has a narrow basal constriction
beyond which it is strongly convex [apical third of metasoma miss-

ing]-

Type-species. — - T aimyrisphex pristinus, new species.

Remarks. — • This is an exceedingly generalized aculeate, without
noteworthy features that would assign it unequivocally to any ma-
jor group. I assign it tentatively to the Sphecidae (in the broad
sense), largely on the basis of the rounded posterior lobes of the
pronotum, which lie slightly below the tegulae and do not quite
reach them. The wing venation is not inconsistent with that of a
generalized sphecid, and is more generalized than that of Archisphex
(Evans, 1969) with respect to the position of the recurrent veins.
The venation might also be that of a generalized scolioid, although
the lack of a constriction between the first two metasomal segments
and of a crease beneath the stigma (1 r) would exclude it from
most living families of Scolioidea. The general form of the prono-
tum is suggestive of a pompilid, and the venation would not exclude
it from that family, but there is no evidence of a transverse suture
on the mesopleurum. Since we know from other evidence that the
family Sphecidae was well represented in the Cretaceous, it seems
best to assign T aimyrisphex to that family, at least tentatively, until
such time as further pieces can be added to the puzzle.

Taimyrisphex pristinus, new species

Length about 4 mm; fore wing about 2.5 mm. Color dark brown,
pronotum apparently with a pair of dorsal pale spots, first metasomal

Fig. 1. T aimyrisphex pristinus, n. sp., wings of type. Fig. 2. Same speci-
men, oblique-dorsal view of body, wings mostly omitted. Fig. 3. Mandible
of Cretabythus sibiricus, n. sp. type. Fig. 4. Same specimen, body and wings.
Fig. 5. Pittoecus pauper, n. sp., type, portion of fore wing. Fig. 6. Same
specimen, front leg. Fig. 7. Same specimen, hind tibia. Fig. 8. Same speci-
men, ventral surface of abdomen. Fig. 9. Protamisega khatanga, n. sp.,
type, lateral view of head. Dashed lines in Figures 4 and 5 indicate parts
not clearly visible in specimens.

1973]

Evans — Cretaceous A culeate Wasps

169

70

Psyche

[September

tergite with a pair of large pale spots barely separated medially
[these maculations may be artifacts] ; antennae dark brown except
scape paler; preserved parts of legs stramineous except hind coxae
mostly fuscous; wings hyaline, with brown veins and stigma; wing
membrane covered with microtrichiae, but body without visible setae
or pubescence. Body surface smooth, without noticeable sculpturing.
Front angle of ocellar triangle exceeding a right angle; lateral ocelli
removed from eye margin by slightly less than half their own
diameters, removed from the rounded vertex crest by somewhat more
than their own diameters. Front femur 3 X as long as its maximum
width.

Holotype . • — ■ <$, Taimyr, N. Siberia, 1970, amber specimen no.
3130-16.

FAMILY SPHECIDAE: SUBFAMILY PEMPHREDONINAE

Pittoecus, new genus

Known from a single female approximately 5 mm in length, fully
alate but with a reduced wing venation resembling that of certain
living Pemphredonini (Figs. 5-8). Head broad, with large eyes
extending from base of mandibles to or close to top of head ; temples
broad; antennae short, 12-segmented, somewhat coiled, arising from
small elevations of the front; mandibles straight, tapered, without
visible teeth on upper or lower margin [clypeus and other mouth-
parts not clearly visible, top of head (including ocelli) missing].
Pronotum with rounded posterior lobes similar to those of living
Pemphredoninae [thoracic dorsum largely missing] ; mesopleura
somewhat convex, without visible grooves; legs fully preserved, with
only a few small spines, front tarsus without a pecten, hind tibia
with a group of very short spines basally and a few along the shaft ;
tibia! spur formula 1-1-2; claws dentate. Wings imperfectly pre-
served, fore wing apparently with two submarginal cells, as figured.
Metasoma slender and tapered, sessile basally, with six distinct seg-
ments and a well developed sting and sting-palps [dorsum of abdo-
men missing].

Type-species . — Pittoecus pauper , new species.

Remarks . — Although the only available specimen is incomplete,
consisting of a hollow mold of the ventral half of the body (with
complete legs and partially complete wings), the form is so similar
to that of living pemphredonine wasps such as Passaloecus that it
seems worthy of description. The form of the second submarginal
cell and the unusual distance between the origin of the basal and

1973]

Evans — ■ Cretaceous A culeate W asps

171

transverse median veins are suggestive of the Australian genus Har-
pactophilus.

Pittoecus pauper, new species

Length 4.8 mm; estimated length of fore wing 2.5 mm. Colora-
tion not preserved, specimen the color of amber. Scape barrel-
shaped, about twice as long as wide; antennal segments 2-12 slightly
longer than thick, 13 somewhat pointed, about 1.7 X as long as
thick. Front femur 2.7 X as long as wide; segments of front tarsus
in a ratio of 25:8:7:6:13, comparable measurements of hind tarsus
45:18:14:8:13; front basitarsus with two minute lateral spines as
well as some small apical ones; hind tibia with short spines as fig-
ured, hind tarsi also weakly spinose. Apical metasomal segment
bearing numerous short bristles adjacent to base of sting.

Holotype. — $, Taimyr, N. Siberia, 1970, amber specimen no.
3130-18.

? FAMILY SCOLEBYTHIDAE

Cretabythus, new genus

Known from a single male approximately 2.5 mm in length, fully
alate but with a reduced venation, hind wing without closed cells
(Figs. 3, 4). Antennae elongate, 13-segmented, all segments some-
what longer than wide; maxillary palpi elongate [probably 6-seg-
mented] ; mandibles short, with 4 sharp apical teeth; clypeus, short,
with a low median keel ; malar space short, about one fourth as long
as width of mandibles at their base; eyes and ocelli of moderate size,
ocelli in a broad triangle close to the broadly rounded vertex; head
contracted immediately behind eyes; occipital carina complete. Pro-
notum short, its posterior margin broadly arched between the pos-
terior lobes, which are small, rounded, slightly below and touching
the tegulae; pronotum with a short, anterior collar; notauli and
parapsidal furrows complete; scutellum large, rather flat; base of
propodeum with a slightly elevated, transverse band ( ?metanotum) ,.
disc with a basal, semicircular area with strong surface sculpturing,
otherwise smooth, with a strong transverse carina margining the
abrupt, posterior declivity. Propleura well developed, somewhat
prolonged anteriorly, but posternum and proepimeron (as described
for Scolebythus ) not evident; mesopleura rather smooth, without
evident pits, ridges, or sutures; front and hind coxae contiguous,
middle coxae slightly separated. Fore wings as figured, with one

172

Psyche

[September

closed submarginal cell and two closed discoidal cells (superficially
resembling certain pemphredonine Sphecidae) ; hind wing not fully
visible, but evidently with a strong vein on the basal, anterior mar-
gin, without closed cells. Legs not spinose; tibial spur formula
1-2-2; claws dentate. Metasoma slender, sessile, with 7 visible seg-
ments, without a constriction between first two segments and without
unusual modifications.

Type-species. — Cretabythus sibiricus, new species.

Remarks. — This specimen is reasonably well preserved and nearly
complete, but it presents a puzzling combination of characters. I
had at first supposed it was a pemphredonine sphecid related to
Pittoecus , but despite similarities in the wing venation there are
several reasons for excluding it from this group : the broad, 4-toothed
mandibles, pronotal lobes reaching the tegulae, two mid-tibial spurs,
lack of closed cells in the hind wing, and so forth. The wing vena-
tion appears closest to that of the Scolebythidae (though unfortu-
nately one cannot be sure whether the anal lobe is developed) and
there are other features in common with that group.2 However,
there are also several major differences: e.g. lack of a distinct pros-
ternum and proepimera, presence of an anterior pronotal collar and
of a transverse carina on the propodeum, and differences in the form
of the first metasomal segment. It is probable that modern Scole-
bythidae are specialized for life beneath bark or in holes in wood,
and the absence of such specializations in Cretabythus should not
necessarily exclude it from this family. For the present I can suggest
no better placement for this unusual wasp.

Cretabythus sibiricus, new species

Length 2.5 mm; fore wing about 1.9 mm. Fuscous, except pro-
notum apparently somewhat lighter than remainder of body; legs
stramineous; antennae stramineous basally, slightly infuscated toward
apex; wings hyaline, stigma brown, veins light brown. Body with-
out strong surface sculpturing and without noticeable setae. Anten-
nal segments in the following ratio : 5 :4 .*5 :5 : 5 :5 :5 :5 :5 :5 :5 :5 :5 ;
segment three about 1.7 X as long as wide. Segments of front
tarsus in a ratio of 8 14. :3 :2 :5 ; longer spur of hind tibia 0.4 X
length of hind basitarsus.

2When I described the Scolebythidae (Evans, 1963) I had only females.
I have since discovered a male Cly stops enella longiventris Kieffer and can
state that there is no marked sexual dimorphism in this group. The family
currently has a broadly discontinuous distribution in Brazil, Madagascar,
and Australia (the Australian element has yet to be described).

1973]

Evans — Cretaceous A culeate JVasps

73

10

II

Fig. 10. Protamisega khatanga, n. sp., type, fore wing. Fig. 11. Hypo-
cleptes rasnitsyni, n. sp., type, fore wing. Fig. 12. Archaepyris minutus,
n. sp., type, fore wing. Fig. 13. Hypocleptes rasnitsyni , n. sp., lateral view
of type. Fig. 14. Celonophamia taimyria, n. sp., head of type. Fig. 15.
Archaepyris minutus, n. sp., head of type. Fig. 16. Celonophamia taimyria,
n. sp., dorsal view of body and wings as preserved. Broadly dashed lines
in Figures 10 and 12 indicate pale streaks in wing membrane, while finely
stippled lines indicate weak veins.

174

Psyche

[September

Holotype. — cf, Taimyr, N. Siberia, 1970, amber specimen no.
3130-17.

FAMILY BETHYLIDAE

Archaepyris, new genus

Wasps approximately 2 mm in length, fully alate, of fuscous
coloration (known only from males) (Figs. 12, 15). Antennae
simple, with 13 segments; eyes large, slightly protruding from sides
of head, not noticeably hairy; mandibles short, broad, with several
sharp apical teeth ; clypeus short, median lobe only slightly produced ;
ocelli of moderate size, situated close to vertex crest. Pronotum
moderately long, disc only slightly shorter than mesoscutum along
midline; notauli and basal scutellar pits or groove not visible; pro-
podeum short, with a more or less flat dorsal surface that is slightly
wider than long, also with an almost vertical posterior declivity.
Legs not spinose ; claws weakly dentate. Fore wing as figured, with
a short vein arising from basal vein, the discoidal cell outlined by
weak veins and outer part of wing with a series of fine creases that
may mark the course of former veins. Metasoma sessile.

Type-species. — Archaepyris minutus, new species.

Remarks. — Since the classification of Bethylidae is based on struc-
tural minutiae that require perfect preservation and close study under
high power, placement of fossil material involves a large measure of
guess work. Archaepyris I would judge to be a very generalized
bethylid, combining features of the usually recognized subfamilies.
Although Epyris-Wke in general form, the venation is suggestive of
some Pristocerinae, while the vein arising from the basal vein sug-
gests Bethylinae.

Archaepyris minutus, new species

Length 2 mm; fore wing 1.4 mm. Fuscous; wings hyaline, with
brown veins and stigma. Front slightly narrower than eye height,
with a weak median groove; sculpturing of front rather weak;
antennae arising close to base of clypeus, the segments in the follow-
ing ratio: 15:10:9:10:9:9:9:9:9:9:9:9:15. Segments of middle tar-
sus in the following ratio: 16:7:7:5:9. Subgenital plate simple;
parameres hirsute, somewhat rounded apically.

'Holotype. — ■ cf, Taimyr, N. Siberia, 1971, amber specimen no.
331 1-7. Paratype. — cf in fair condition, in amber from same
source, no. 3130-21.

973]

Evans — Cretaceous A culeate W asps

175

Celonophamia, new genus

Wasps approximately 2 mm in length, fully alate, of fuscous
coloration (known only from females) (Figs. 14, 16). Antennae
simple, with 12 segments; eyes large, slightly protruding from sides
of head, not noticeably hairy; palpi short; mandibles short, the apex
broad, with several sharp teeth; front with a deep median groove.
Pronotum short, its posterior margin broadly arched ; mesoscutum
broad, notauli distinct. Legs simple, not spinose; front femur not
swollen, about 3 X as long as wide ; claws simple. Wings extending
well beyond middle of abdomen, rather evenly covered with micro-
trichiae ; stigma and radial vein well developed ; basal vein reaching
subcosta a short distance basad of stigma, not giving rise to a vein ;
discoidal and subdiscoidal veins absent or weakly defined. Metasoma
sessile, slightly flattened, bearing short bristles.

Type-species. — Celonophamia taimyria , new species.

Remarks. — This genus also occurs in Upper Cretaceous Canadian
amber (specimen to be described later). The 12-segmented anten-
nae suggest Cephalonomia and its allies (the generic name is ana-
gram of Cephalonomia) . However, the wings are broader and have
a fuller venation than in that genus. Possibly this genus is close to
the ancestral stock of the Cephalonomiini, subfamily Epyrinae.

Celonophamia taimyria, new species

Length 2 mm; fore wing about 1.3 mm. Fuscous; wings hyaline,
with brown veins and stigma. Front slightly wider than height of
an eye, with a deep median sulcus reaching to anterior ocellus;
ocelli of moderate size; sculpturing of front weak. Antennal seg-
ments in the following ratio: 15:8:7:6:6:6:6:6:6:6:6:10. Segments
of hind tarsus in the following ratio: 20:9:7:6:11.

Holotype. — 9, Taimyr, N. Siberia, 1971, amber specimen no.
331 1-5. Paratypes. — Two specimens of indeterminate sex, in rather
poor condition, nos. 3311-11 and 3311- 12.

FAMILY CLEPTIDAE
Hypocleptes, new genus

Small wasps bearing much resemblance to living members of the
genus Cleptes, although with a very simple wing venation (known
only from females) (Figs. 11, 13). Antennae short, scape only
about 2.5 X as long as wide, arising slightly below bottoms of

176

Psyche

[September

eyes; malar space moderately long; palpi simple, slender. Pronotum
rather long, crossed by a deep transverse furrow, posterior lobes
projecting well beneath sides of mesoscutum, the latter elongate,
with strong, complete notauli ; propodeum strongly foveolate, some-
what angulate but not really dentate on each side. Fore wing with
very simple venation, as figured. Legs simple, slender, non-spinose;
tibia! spur formula 1-2-2; claws apparently simple. Metasoma
robust except apical three segments in the form of a slender (prob-
ably extensible) tube. Body without strong surface sculpturing ex-
cept for the large foveae on the propodeum.

Type-species. — Hypocleptes rasnitsyni , new species.

Remarks. — This genus differs from Procleptes (Evans, 1969) in
lacking dentiform processes on the propodeum and apical processes
on the front coxae; also, the scapes are much shorter. The wings
of Procleptes are mostly missing, so this feature cannot be compared.

Hypocleptes rasnitsyni, new species

Length 2.8 mm; fore wing about 1.8 mm. Head, thorax, and
appendages brown to black, abdomen much paler, rufotestaceous ;
wings hyaline, veins and stigma brown. Antennae short, middle seg-
ments very slightly longer than wide. Mesoscutum, measured along
midline, about twice as long as pronotum, surface sparsely punctate;
mesoscutum and scutellum separated by a transverse groove. Seg-
ments of front tarsus in the following ratio: 22:5:5:4:10; middle
and hind tarsi more elongate than front tarsi.

Holotype. — ?, Taimyr, N. Siberia, 1971, amber specimen no.
3311-6. Paratypes. — 5 9$ in fair condition in other pieces of amber
from same source, nos. 3130-20, 3130-22, 3311-8, 33 1 1 _9, and
3311-10. 4 additional 9? in Poor condition are also tentatively placed
in this species.

Protamisega, new genus

Known from a single female only 1.9 mm in length, bearing much
resemblance to living members of the genus Amisega (Figs. 9, 10).
Eyes large, somewhat protuberant; antennae inserted far below bot-
tom of eyes, scape very long, about 4 X as long as wide, about half
as long as remainder of antenna. Pronotum moderately long, crossed
by a transverse carina anteriorly, midline not impressed ; notauli
deeply impressed, complete; scutellum with a transverse basal sul-
cus; propodeum with a steep posterior slope, surface with a number

1973]

Evans — Cretaceous A culeate TV asps

177

of ridges describing large foveae, ridges on sides converging to form
a small, spinose projection. Legs simple, not spinose; claws appar-
ently simple. Fore wing as figured. Metasoma robust on basal four
segments, terminal segments forming a tube very much as in Hypo-
cleptes. Body without noticeable setae and without strong surface
sculpturing except for propodeum.

Type-species. — Protamisega khatanga , new species.

Remarks. — This genus differs from the preceding in having the
wing venation less simplified and in having the scape longer and in-
serted lower on the head. In the last feature it resembles Procleptes
of Canadian amber (Evans, 1969), but the latter has stronger pro-
cesses on the propodeum and much more obvious surface sculpturing.

Protamisega khatanga, new species

Length 1.9 mm; fore wing 1.2 mm. Body dark brown, scape and
basal parts of legs also fuscous, but appendages fading to lighter
brown apically; wings hyaline, veins and stigma light brown. An-
tennae prominent, middle segments about as wide as long. Segments
of hind tarsus in the following ratio: 17:6:6:5:9. First four meta-
somal tergites in approximately the following ratio, measured from
the side: 13:13:8:4. The terminal tubular portion appears to be
made up of three segments, although the details are obscure.

Holotype. — Taimyr, N. Siberia, 1970, amber specimen no.

3130-19.

Discussion

Ten additional pieces of amber from the same source contain un-
identifiable specimens, all of which I would judge to be Bethyloidea;
some of these probably represent additional specimens of species
described above. The preponderance of Bethyloidea in this material
doubtless reflects the fact that many are associated with trees and are
small enough to be preserved in small pieces of amber, as mentioned
earlier. Nevertheless it is difficult to escape the impression that
Cleptidae, at least, were much more abundant than they are today,
for the family is now represented by only a few genera containing
species that are only rarely encountered. Since the Cretaceous
Cleptidae have the general habitus of contemporary forms, it is per-
haps a safe assumption that they, too, attacked Symphyta and Phas-
mida, groups that may well have been more important parts of the
fauna in the Cretaceous than they are today.

178

Psyche

[September

All of the Bethyloidea are clearly identifiable as to family, but
more difficult to place in present-day tribes and subfamilies. The
bethylid genus Archaepyris appears to have some features of each
of the major subfamilies, while Celonophamia appears annectant
between the Epyrini and Cephalonomiini. Cretabythus is a truly
enigmatic form, with many bethylid-like features (particularly the
mandibles) but with an unusual wing venation suggesting the Scole-
bythidae. Taimyrisphex is perhaps the most generalized wasp yet
described from the Cretaceous. Although I have placed it in the
Sphecidae, one might argue that it is a scolioid or even a prototype
of the Pompilidae. We do know that the Sphecidae underwent
considerable evolution before the end of the Cretaceous, for two
specialized forms of the subfamily Pemphredoninae have now been
described.

In conclusion, it may be useful to list the Aculeata now known
from the Cretaceous, bearing in mind that this list will be con-
siderably augmented when all recently collected material has been
described :

SCOLIOIDEA

Cretavidae: Cretavus

?SCOLIOIDEA-BETHYLOIDEA
PScolebythidae : Cretabythus

BETHYLOIDEA

Bethylidae : A rchaepyris

Celonophamia
Cleptidae: Procleptes
Hypocleptes
P rot amis ega

References

Evans, H. E.

1963. A new family of wasps. Psyche, 70: 7-16.

1969. Three new Cretaceous aculeate wasps (Hymenoptera) . Psyche,
76: 251-261.

Sharov, A. G.

1957. First discovery of a Cretaceous stinging hymenopteron (Acu-
leata). Dokl. Akad. Nauk., 112: 943-944. (In Russian).

Wilson, E. O., F. M. Carpenter, and W. L. Brown, Jr.

1967. The first Mesozoic ants, with the description of a new subfamily.
Psyche, 74: 1-19.

FORMICOIDEA

Formicidae: Sphecomyrrna

?SPHECOIDEA

PSphecidae : A rchisphex

Taimyrisphex

SPHECOIDEA

Sphecidae

Pemphredoninae: Lisponema
Pit to ecus

PARAMUZOA (NYCTIBORINAE),

A NEW COCKROACH GENUS
PREVIOUSLY CONFUSED WITH
PARASPHAERIA (EPILAMPRINAE)*

By Louis M. Roth
Pioneering Research Laboratory
U.S. Army Natick Laboratories
Natick, Massachusetts 01760

Kirby (1904, p. 194) placed Parasphaeria Brunner in the Peri-
sphaeriinae, probably because Brunner (1865) stated that the genus
was near Perisphcieria Biirm. Princis (1964) placed the genus in
the Perisphaeriidae. While studying the male genitalia of genera
which Princis assigned to the Perisphaeriidae, I found the phallo-
meres of Parasphaeria ovata (Blanchard) (type of genus, Princis,
1964, p. 240) to be more typical of the genitalia of genera which
belong to the Epilamprinae and that Parasphaeria linearis (Serville)
is not a member of ovoviviparous Parasphaeria , but is an oviparous
species.

All ovoviviparous cockroaches belong to one family, the Blaberidae
(McKittrick, 1964) and the arrangement of their male genital phal-
lomeres are similar with the hook (R2) always on the right side
(Fig. 10). The Plectopterinae ( Blattellidae) also have the hook on
the right, but in the other 4 blattellid subfamilies ( Anaplectinae,
Blattellinae, Ectobiinae, and Nyctiborinae) , the male genitalia are
the mirror image of those of the Plectopterinae and Blaberidae, hav-
ing the retractable hook on the left side (McKittrick, 1964). The
hooked phallomere (L3) of Parasphaeria linearis (Fig. 9) is on the
left and the phallomere L2d clearly shows a close relationship to
members of the Nyctiborinae, in which subfamily I place this species.

In this paper, I shall redescribe Parasphaeria ovata , and erect a
new genus for Parasphaeria linearis, in part from specimens used by
Brunner (1865) when he described the genus. I have examined
the S and 9 types of Blatta ovata Blanchard, and Brunner was
correct in his determination of his specimens of this species. Complete
synonyms for these 2 species can be found in Princis (1964).

Paramuzoa, n. gen.

Type species: Blatta linearis Serville (present designation)

Paramuzoa appears to be close to Muzoa Hebard, having sym-

*Manuscript received by the editor October 2, 1973

179

8o

Psyche

[September

Figs. 1-5. Adults of Paramuzoa linearis. 1. Male (Brazil). 2. Head
( cf ) (From specimen shown in Fig. 1). 3. Female (Brazil). 4. Left side
of female mesonotum showing the deep, incomplete lateral incision (arrow)
resulting in a tegmenlike lobe. 5. Supra-anal plate and cerci (dorsal) of £.
(Figs. 4 and 5 from female shown in Fig. 3. (Both adults from Brunner s
collection at the Natural History Museum, Vienna; scale: figs. 1, 3 = 5 mm,
figs. 2, 4-5 = 1 mm) .

1973]

Roth — Paramuzoa

1 8 1

metrical tarsal claws; in other genera of Nyctiborinae symmetrical
claws are found only in Megaloblatta (Hebard, 1921, p. 132).
Although the shape of L2d (Fig. 9) of Paramuzoa differs from that
of Muzoa madida Rehn, both have this phallomere tapering to a
sharp point directed to the left. In addition to specific differences
which distinguish it from Muzoa , the male of Paramuzoa has an
asymmetrical subgenital plate and the right and left styles differ
markedly in size (Fig. 8) ; in Muzoa the subgenital plate is sym-
metrical and the styles are equal. Femoral armament also differs
between the 2 genera. The female of Paramuzoa is wingless and the
mesonotum is modified laterally to form lobes which appear to be
movable tegmina (Fig. 3) but anteriorly are actually part of the
mesonotum (Fig. 4). The females of Muzoa are fully winged.

Paramuzoa linearis (Serville), n. comb.

(Figs. 1-9)

Syn. Blatta linearis Serv. (Serville, 1831, Ann. Sci. nat. 22,
P- 4F cf )

Parasphaeria linearis (Serv.) (Brunner, 1865; 314, cf and $)

cf (Figs. 1, 6). — Violaceous black. Head black, except for a
broad testaceous band above the labrum; surface pitted (Fig. 2).
Last segment of labial and maxillary palps black, the others testa-
ceous. Eyes wide apart, the interocular space larger than the width
of one eye. Antennae black, except for about 6 distal yellowish
segments terminating in several apical black segments (Fig. 6).
Pronotum densely pitted, piliferous, anterior and posterior margins
rounded, widest at posterior border, metallic blackish except for 2
smooth orange spots below the middle. Tegmina piliferous, the an-
terior portion distinctly pitted. Wings infuscated (Fig. 6). Legs
black. Ventral femoral margins: Front femur; anterior margin
lined with small piliform setae, those on the distal half closer to-
gether; distal spine present, genicular spine absent. Mid femur:
piliform spines on both margins; hind margin with 1 large spine,
absent on anterior margin, distal spines on both margins, the anterior
larger than the one on the posterior margin, genicular spine present.
Hind femur: anterior margin with piliform spines only, plus a distal
spine; posterior margin with piliform spines and one large spine,
distal spine absent, genicular spine present. Tarsi with large pulvilli
on all segments; tarsal claws simple, equal, with large arolia. Supra-
anal plate with sides straight, hind margin unevenly convex; cerci
about 4 times as long as wide, piliferous (Fig. 7). Subgenital plate

Psyche

[September

Figs. 6-8. Paramuzoa linearis. 6. Adult $ and left wing (Sao Paulo,
Cipo, Brazil, 25-XI-1967, leg. V. N. Alin, U.S. National Museum; det.
Gurney). 7. Supra-anal plate and cerci (dorsal). 8. Subgenital plate and
styles (ventral). (Figs. 7 and 8 KOH preparations from specimen shown
in Fig. 6; scale: fig. 6 = 5 mm, figs. 7-8 = 1 mm).

973]

Roth — P aramuzoa

183

asymmetrical, the hind margin rounded ; styles slender, the left about
2.5X longer than the right (Fig. 8). Genitalia shown in figure 9;
R2 with a subapical incision; L2d touching, but not attached to L2
vm, tapering to a sharp point directed to the left.

Measurements (mm.) of Brunner’s specimen (Brazil) as follows:
width and length of pronotum — 5.4 and 4.2 respectively; length
and width of tegmen 17 and 4 respectively; body length 15.

$ (Fig. 3) — Larger than and differs from male as follows: Head
hidden under anterior margin of pronotum. Pronotum with hind
margin slightly convex, the lateral corners curved backwards. What
appear to be movable tegmina are actually a modification of the
mesonotum which is deeply incised laterally forming a small lobe
having a broad longitudinal convex swelling; anteriorly the meso-
notum is entire and the mesad margin of the lobe continues as a
groove in the mesonotal surface (Fig. 4) enhancing the impression
of a freely moving tegmen (Fig. 3). Wings absent. Ventral hind
margin of mid femur with 2 large spines. Tergites with fine recum-
bent hairs, their insertions giving a finely pitted appearance. Sternites
somewhat similar to tergites. Supra-anal plate piliferous, the hind
margin strongly rounded with a shallow mesad indentation; cerci
flattened dorsally, convex ventrally, about twice as long as wide,
extending well beyond the hind margin of the supra-anal plate
(Fig. 5). Mid portion of subgenital plate weakly convex, laterally
strongly curved dorsad, hind margin rounded, laterally undulate.

Measurements (mm.) of Brunner’s specimen (Brazil) as follows:
Width and length of pronotum 8 and 5.6 respectively; width and
length of tegmen 2.5 and 2.8 respectively; length of body about
19 mm.

Parasphaeria ovata (Blanchard)

(Figs. 10-19)

cf (Figs. 11, 14). — Head brown, exposed beyond anterior mar-
gin of pronotum; interocular space about i.6x width of an eye;
interocellar space less than distance between eyes; labrum and clypeus
testaceous; antennae brown, longer than body. Pronotum widest at
about the middle, hind border convex, disk chestnut brown, impressed,
resulting in an uneven surface; anterior and lateral borders testa-
ceous. Tegmina very long, basally relatively narrow and expanding
beyond, testaceous, nearly transparent; veins testaceous, prominent,
especially on the basal half; Wings as long as tegmina, hyaline, veins
testaceous. Front femur: basal half of ventro-anterior margin with

Figs. 9-10. Male genitalia (dorsal). 9. Paramuzoa linearis (From speci-
men shown in Fig. 6). 10. Parasphaeria ovata (From specimen shown in

Fig. 14). Phallomeres numbered according to McKittrick (1964). (KOH
preparations; the phallomeres have been separated somewhat, and flat-
tened; Figs. 9 and 10 to same scale).

Psyche

[September

Figs. 11-16. Adult males of Parasphaeria ovata. 11. Chile (U.S. Na-
tional Museum, det. Gurney). 12. Supra-anal plate, cerci, and paraprocts
(ventral). 13. Subgenital plate (ventral). (Figs. 12-13 from $ shown in
Fig. 11). Chile (Brunner’s specimen from the Natural History Museum,
Vienna). 15. Supra-anal plate (dorsal). 16. Subgenital plate (ventral).
(Figs. 15-16, from $ shown in Fig. 14; scale: figs. 11, 14=5 mm, figs. 12-
13, 15-16=0.5 mm).

1973]

Roth — P aramuzoa

86

Psyche

[September

widely spaced piliform spines followed by a row of smaller more
closely spaced fine spines, small distal spine present, genicular spine
absent. Mid and hind femora: ventro-anterior and posterior mar-
gins unarmed (except for some small piliform spines), distal spines
absent, small genicular spines present. Tarsal claws simple, equal,
arolia and pulvilli well developed. Abdomen testaceous, lacking
tergal glands; hind margin of supra-anal plate rounded and pilifer-
ous; surface covered with small setae, more numerous on the pos-
terior half (Fig. 15); right and left paraprocts bulbous (Fig. 12) ;
subgenital plate asymmetrical, slightly indented on the right; styles
absent (Figs. 13, 16). Genitalia shown in figure 10; R2 on right
side, with a subapical incision; L2d separated from L2vm and with
a lateral lobe which is directed dorsad; Li with a deep upturned
cleft, heavily sclerotized along the margins; lower lobe (below cleft)
without setae (what appears to be setae are actually on a membrane
which overlaps the lobe).

Table 1. Measurements (mm) of Parasphaeria ovata

Overall Pronotum Tegmen

body length length width length width

$

Chile, Type (Paris Museum)

21

4.3

5.7

25.5

—

Chile (Brunner’s specimen ;
Vienna Museum)

0

22

4.4

5.8

26

Chile, Type (Paris Museum)

26

6.1

9

4.2

3.1

Peru (Brunner’s specimen;
Vienna Museum) a

28

6.1

8.3

4.1

2.9

Ancud, Chiloe Island, Chile

(2-7 April 1920, Cornell Univ.
Exp.; Phila. Acad. Natural Sci.,
det. Hebard)

25

6.5

9.1

4.3

3.2

Araucana, S. Chile (R. M.
Middleton, 1907-337; British
Museum; det. Hebard, 1928)

23

5.6

8.3

4.2

2.9

Unlabeled specimen in British
Museum

28

6.5

10.4

4.5

3.4

“Brunner (1865) lists Chile as

the

locality for

his $

specimen,

but th<

label indicated it was from Peru. Rehn (1933) believed that P. ovata is
limited in distribution to the southern half of Chile and he questioned the
determination of Shelford’s record of a male from Ecuador. Two small
nymphs, apparently of this species, at the British Museum had the following
collection data: 1) L. Nahuel Huapi, Puerto Blest, 2-3. XII. 1926, Argen-
tina, Terr. Rio Negro, F. and M. Edwards; 2) Casa Pangue, 4-10. XII.
1926, S. Chile, Llanquihue prov., F. and M. Edwards.

1973]

Roth — Paramuzoa

187

Figs. 17-19. Adult females of Parasphaeria ovata. 17. Thorax and part
of abdomen (Chile, U.S. National Museum; det. Gurney). 18. Peru (Brun-
ner’s specimen from the Natural History Museum, Vienna). 19. Terminal
segments of female shown in Fig. 18. (scale: tig. 17 — 4 mm, fig. 1 8 ~
5 mm, 19 = 2 mm) .

Male measurements are shown in Table 1.

$ (Figs. 17, 18). — Dark brown, metallic. Head exposed beyond
front margin of pronotum, largely black, shiny, with widely spaced
fine dot-like impressions; labrum, lower border of clypeus, and area
above and below antennal insertions testaceous; lower part of frons
weakly wrinkled; antennae black, not quite length of body; last
segments of maxillary palps darker than the others. Pronotum con-
vex, widest at posterior border; uneven testaceous band borders the
pronotum to the hind margin; posterior angles black. Tegmen lobi-
form, lateral, reaching slightly beyond hind margin of mesonotum,
testaceous except for a dark incomplete humeral stripe. Metanotum
black with a testaceous spot at each of the 2 anterior angles and 2
other similar markings on the posterior border within the posterior

1 88

Psyche

[September

angles. Legs stout, short, testaceous, margined with brown; arma-
ment as in cT. Abdomen convex, shiny black, slightly roughened;
segments 2 to 7 with testaceous spots on the posterior angles, these
spots decreasing in size from the anterior to the posterior tergites; a
broad median testaceous band on sternites 1-5, the remainder blackish;
supra-anal plate rounded, entire, cerci small, triangular (Fig. 19).
Female measurements are shown in Table 1.

Summary

The Brazilian cockroach originally described as Blatta linearis by
Serville has been referred to the genus P arasphaeria by Brunner ; it is
not a member of this ovoviviparous genus ( Blaberidae) . P. linearis
is oviparous with characters placing it in the subfamily Nyctiborinae
(Blattellidae) , and Pararnuzoa n. gen. is erected for this species.
The male genitalia, of P arasphaeria ovata (Blanchard) indicate that
it is a member of the Epilamprinae and is not close to Perisphaeria
(Perisphaeriinae) as suggested by Brunner.

Acknowledgements

I thank the following for the loan of specimens: Dr. M. Descamps,
Museum of Natural History, Paris, for Blanchard’s types of P ara-
sphaeria ovata ; Dr. A. Kaltenbach, Natural History Museum,
Vienna, for Brunner’s specimens of P. ovata and P. linearis ; Dr.
Ashley Gurney, U. S. National Museum, Dr. David Rentz, Phila-
delphia Academy of Natural Sciences, and Dr. David R. Ragge,
British Museum (Natural History), London, for additional speci-
mens. I also thank Mr. Samuel Cohen for taking the photographs.

References

Brunner von Wattenwyl, C.

1865. Nouveau systeme des Blattaires. Vienna, 426 pp.

Hebard, M.

1921. Studies in the Dermaptera and Orthoptera of Colombia. Second
paper. Trans. Am. Entomol. Soc. 47: 107-169.

Kirby, W. F.

1904. A synonymic catalogue of Orthoptera. 1 : 61-205. London.
McKittrick, F. A.

1964. Evolutionary studies of cockroaches. Cornell Univ. Agric. Exp.
St. Mem. 389, 197 pp.

Princis, K.

1964. Orthopterorum Catalogus. Pars 6: 174-281. ’s — Gravenhage.
Rehn, J. A. G.

1933. On the Dermaptera and Orthoptera of Chile. Trans. Am. Ento-
mol. Soc. 59: 159-190.

THE MILLIPED FAMILY RHISCOSOMIDIDAE
(DIPLOPODA: CHORDEUMIDA: STRIARIOIDEA) *

By William A. Shear
Department of Zoology, University of Florida,
Gainesville, Florida 32601

Silvestri (1909) established the Family Rhiscosomididae for the
single species Rhiscosomides miner i, from Oregon. Aside from the
description of a second Oregon species by Chamberlin (R. josephi)
from a female (Chamberlin, 1941), nothing further had been
learned about the relationships or ecology of the millipeds of the
family until my general review (Shear, 1972) of the North Ameri-
can families of the Order Chordeumida. In that paper, I described
a third species, R . acovescorJ from California, transferred Tingupa
monterea Chamberlin to Rhiscosomides/ and established the relation-
ship of the Family Rhiscosomididae to the Families Caseyidae, Uro-
chordeumidae and Striariidae. Together, these four families make
up the Superfamily Striarioidea.

Beginning in late 1971, I received nearly 600 unsorted Berlese
samples from Ellen M. Benedict, of the Department of Biology,
Portland State University, Portland, Oregon, and about 50 similar
samples from Dr. David Malcolm, Pacific University, Forest Grove,
Oregon. These samples were taken pursuant to studies of pseudo-
scorpions, but also contained a large number of millipeds. The milli-
peds from this material add enormously to our knowledge of the
fauna of the northern Pacific coast of the United States, and I am
extremely grateful to Mrs. Benedict and Dr. Malcolm for allowing
me to examine them. This paper represents the first report based
largely on the Oregon Berlese material, which now allows a more
or less comprehensive revision of several little-known milliped fam-
ilies. I am also grateful to Dr. Paul Arnaud, California Academy
of Sciences, San Francisco, California, for allowing me to borrow
that institution’s collection of unidentified millipeds.

However, despite the rich material now available, a number of
questions remain to be answered concerning the rhiscosomidids.
( 1 ) Chamberlin’s species R. montereum , the southernmost known
representative of the genus, remains unstudied, since the types (the
only known material) are no longer in existence. It appears to be a
species distinct from R. acovescor , of Marin County. (2) The re-

*Manuscript received by the editor July 5, 1973

89

190

Psyche

[September

lationship of the rhiscosomidids to the striariids is even more obvious
than before. I think that when the striariids from the Benedict and
Malcolm collections are thoroughly studied, it might prove desirable
to even consider the Rhiscosomididae a subfamily of the Striariidae,
despite the numerous differences in gross body form. (3) More
material from California is needed, as there are probably several
additional species occurring there. (4) Rhiscosomidids have not
been collected in the state of Washington, where they may also
occur, though numbers of Berlese samples from suitable habitats in
that state contained no rhiscosomidids. The southern coastal region
of Washington needs further exploration for millipeds.

Ecologically, there do not appear to be any really significant dif-
ferences in the habitats of the several known species. All have been
collected most frequently from rotted wood, from conifer duff, and
less frequently from deciduous duff and litter. Collections where
elevational data is available are from 1100 ft. elevation or less.
Nearly all were taken between November and March. However, the
holotypes of R. montereum and R. trinitarium were collected in
June and July respectively, and the latter was taken above 3400 ft.
elevation. It should be emphasized that this data represents negative
evidence from many samples from suitable habitats taken by Mrs.
Benedict at much higher elevations and at other times of the year.
Summer and early fall samples were poor in all types of millipeds;
perhaps we are dealing here with a fauna adapted to low to nioderate
temperatures and high humidity, individuals of which burrow deeper
into the soil during unfavorable seasons.

All type material for new species described below has been de-
posited in the Museum of Comparative Zoology, Cambridge, Massa-
chusetts, except for the holotype of R. trinitarium , which is the
property of the California Academy of Sciences, San Francisco,
California.

Family Rhiscosomididae Silvestri

Rhiscosomididae Silvestri, 1909, Rend. R. Accad. Lincei 18: 232; 1913, Boll.

Lab. Zool. Portici 7: 307; Shear, 1972, Bull. Mus. Comp. Zool. 144(4):

261.

Type Genus: Rhisoosomides Silvestri, 1909. The family is mono-
basic.

Diagnosis: Distinct from species of Caseyidae in having broad

segmental paranota, from species of Urochordeumidae in having the
collum wider than the head, and from species of Striariidae in the

1973J Shear — Rhiscosomididae 191

body ornamentation (Fig. 4) : tiny, sharp, seta-tipped tubercles,
rather than longitudinal ridges. Rhiscosomidids also resemble some-
what the larger species of the genus Tingupa (Tingupidae) , but
may be distinguished from them by the body ornamentation as well;
tingupids are covered with short, longitudinal carinae.

Description: Small, striarioid millipeds (Fig. 4) with 30 post-

cephalic segments. Collum broader than head, only slightly reflexed
ventrad laterally. Antennae clavate, short. Mentum of gnathochi-
larium divided. Postcollum segments with strong paranota extending
laterad, posterior lateral corners becoming strongly reflexed posteriad,
segmental setae long, rather blunt. Surfaces of metazonites covered
with closely set, sharply pointed tubercles bearing tiny branched
setae. Sixth segment of males enlarged in some species. Epiproct
trilobed. Legs normal, pregonopodal legs of males somewhat more
crassate than postgonopodal legs. Gonopods of males (Figs. 2, 7,
10, 18) with sternum strongly sclerotized, two prominent groups of
coxal processes. Anterior coxal processes partially fused in some
species to form anterior plate. Posterior coxal processes usually fur-
nished with fimbriate, membranous, or flagelliform branches and
areas. Telopodites irregular, lobelike. Ninth legs reduced in size,
with blunt coxal process, flattened, granular telopodite of one seg-
ment (Figs. 9, 14, 16). Coxae of legs 10 with glands opening on
anterior faces. Legs 1 1 normal. Cyphopods embraced by expansions
of sternites and coxae of second and third legs, with postgenital
structures of uncertain origin (Fig. 3).

Distribution: Pacific coast region of the United States from the

Monterey Penninsula north to the Columbia River, usually at ele-
vations below 1 100 ft.

Genus Rhiscosomides Silvestri

Rhiscosomides Silvestri, 1909, Rend. R. Accad. Lincei 18: 232; 1919, Bull.

Lab. Zool. Portici 7: 308; Shear, 1972, Bull. Mus. Comp Zool. 144(4) : 261.

Type Species: Rhiscosomides mineri Silvestri, by monotypy.

Description : The genus and family are coextensive, but the fol-
lowing additional characters may be noted. Body generally dark
brown in color, collum usually cream-white, bases of segmental setae
marked with light spots. In species in which sixth segment of males
is enlarged, that segment lighter in color dorsally than the others.
General appearance is of parallel-sided polydesmiform animals,
squared off anteriorly at collum, tapering abruptly to blunt epiproct
from segment 25. Ocelli vary in number from 5 to 7, variable within
species.

92

Psyche

[September

Gonopod Anatomy of Rhiscosomides Species

The gonopods of Rhiscosomides species males conform well to the
striarioid pattern. The sternum (Fig. 2, S) is heavily sclerotized
and anteriorly margined, with deeply depressed openings from the
tracheal spiracles. Laterally, the sternum is broadly expanded, con-
cealing the bases of the coxal processes. The anterior coxal processes
(Fig. i, AC) are closely appressed, and in the Acovescor Group
of species are wholly or partially fused to form a broad plate. A
strong lateral branch is usually present (Fig. 2, LB), but may be
a broad flange (Fig. 7), or the largest part of the process (Fig. 18).
The length of the anterior coxal process and the form of its terminal
branches are of excellent taxonomic and diagnostic value. The pos-
terior coxal process usually has three branches: the anterior branch
(Fig. 7, AB), the mesal branch (Fig. 7, MB), usually the largest,
and fhe generally much smaller posterior branch (Fig. 7, PB). The
mesal and posterior branches are usually connected by a membranous
or fimbriate area. There is also a flabelliform median structure
arising from the heavily sclerotized part of the sternum that extends
between the gonopods. This sternal flap (Fig. 10, SF) usually
comes off with one or the other of the gonopods when they are sep-
arated. The telopodites (Figs. 2, 10, T) are amorphous, lobelike
structures that are usually displaced laterally, but may interlock
basally with the lateral extensions of the sternum or of the anterior
coxal processes. They are of little taxonomic value. The posterior
gonopods are rather uniform throughout the genus (Figs. 9, 14, 16).
In my 1972 description, I erred in calling the narrow anterior mesal
coxal lobe the telopodite. The actual telopodite is flattened and
irregular in outline and bears a more or less pointed process on the
posteriomesal margin, which extends posteriad. This telopodite pro-
cess and the coxal lobe protect and partially support the anterior
gonopods, and in some animals, clasp between them the coxae of
legs 10.

In terms of gonopod anatomy and a few other characters, the six
species known from males fall into two groups. In the Acovescor
Group ( R . acovescor, R. trinit ariurn) , the anterior coxal processes
tend to be fused or broadly contiguous mesally, and are rather short.
Areas of highly branched, fine cuticular fibers are well developed on
the posterior coxal processes. The sixth segment of the males is only
slightly or not at all enlarged ; in R. acovescor the collum is colored
like the other segments, instead of being white. Females have not
been collected. Both species occur in California, and R. montereum
will probably also prove to belong to the group.

1973]

Shear — Rhiscoso?nididae

193

The Mineri Group includes R. mineri , R. josephi , R. malcolmi
and R. benedictae. In these species, the anterior coxal processes are
usually more rodlike, not at all fused mesally, and are more or less
sharply curved anteriad. The fimbriate or membranous areas on the
posterior coxal processes are of limited extent, and there is an area
on the mesal branch that appears to be glandular. The sixth segment
of males is enlarged; the collum is white. Females have postgenital
bodies (Fig. 3) of uncertain origin immediately posterior to the
cyphopods. These are of limited utility in diagnosis, though they
cannot be used to separate some species. The four species occur
along the Oregon coast and in the foothills of the Coast Ranges.

Certainly, by the standards that have been applied in the past,
these two species groups might have been recognized as genera, and
I suggested (Shear, 1972) that R. acovescor might not be congeneric
with R. mineri. However, R. mineri, the type species of Rhiscoso-
mides , shows a degree of intermediacy in the form of the anterior
coxal processes, particularly when compared to R. trinitarium, which,
in turn, is intermediate between R. mineri and R. acovescor . In the
light of these facts, and because of the small number of species in
the family, it seems pointless to recognize a second genus at this time.

Key to Species of Rhiscosomides
(excluding R. montereum)

1 a. Sixth segment of males conspicuously enlarged (Fig. 4), lighter
in color than other segments; anterior coxal processes of go no-
pods usually rather rodlike (Figs. 7, 10, 15), touching mesally

but not fused. 3

ib. Sixth segment of males not much larger, if at all, than other
segments; anterior coxal processes of gonopods somewhat flat-
tened to platelike, more or less contiguous mesally, or fused. 2
2a. Anterior coxal processes without lateral branches, fused into a

broad plate (Fig. 17) ; Marin Co., Calif. acovescor

2b. Anterior coxal processes with elaborate branches; Trinity Co.,

Calif trinitarium

3a. Apical teeth or processes of anterior coxal process small, not

directed posteriad (Figs. 7, 10, 15). 4

3b. Anterior coxal process with large apical branch directed ventro-

posteriad (Fig. 2) mineri

4a. Anterior coxal processes bent sharply anteriad at nearly a right

angle (Figs. 7, 15) 5

4b. Anterior coxal processes long, rodlike, evenly curved (Fig. 10)
benedictae

194

Psyche

[September

5a. Anterior branch of posterior coxal process broad, bladelike (Fig.

15) male ol mi

5b. Anterior branch of posterior coxal process narrow, more rod-
like (Fig. 7) josephi

Rhiscosomides montereum (Chamberlin)

Tingupa monterea Chamberlin, 1910, Ann. Ent. Soc. Amer. 3: 240-241,
figs. 3-5, sex not specified.

Rhiscosomides monterea, Shear, 1972, Bull. Mus. Comp. Zool. 144(4): 262.

Type: Holotype of unspecified sex from Pacific Grove, California,
collected June, 1902, lost, presumed destroyed.

Notes: Little more can be said about this form until males are
discovered, but as I earlier pointed out (Shear, 1972), the detailed
description of the nonsexual characters leaves no doubt that this
species belongs to Rhiscosomides and not to Tingupa. There are
eight ocelli. An immature Rhiscosomides female from San Mateo
County, California, also has eight ocelli, and may be an example of
R. montereum .

The change in spelling of the specific epithet, which I did not ob-
serve in my 1972 report, is made necessary by the gender of the
generic name.

Rhiscosomides mineri Silvestri
Figs. 1-3

Rhiscosomides mineri Silvestri, 1913, Boll. Lab. Zool. Portici 7: 308-310,
figs. 4-7, $ ; Shear, 1972, Bull. Mus. Comp. Zool. 141(4): 261-262.

Type: Male holotype from a rotting log, Lebanon, Linn Co.,
Oregon; whereabout of specimen unknown, not examined. Silvestri’s
excellent figures (Silvestri, 1913) leave no doubt about the identity
of this species.

Description: Male from 5 mi east of Yamhill, Yamhill Co., Ore-
gon; length, 7.1 mm, width, 1.12 mm. Body of typical form, head
broad, front somewhat flattened, depressed in anterior midline, su-
prantennal swellings moderate. Antennae short, strongly clavate,
reflexed along sides of head, reaching anterior margin of segment 4
when fully extended. Ocelli seven, in two rows of three and four.
Collum broader than head, lateral margins only a little deflexed
ventrad, posteriolateral corners rounded, curved anteriad, anterior
margin sinuous, posterior margin arcuate. Segments with rather
narrow, polydesmiform paranota at first curved forward, posterio-
lateral corners becoming acute, reflexed posteriad, anterior and pos-

1973]

Shear — Rhis cosomididae

195

Figs. 1-3. Rhiscosomides mineri. Fig. 1. Anterior coxal processes of
anterior gonopods, anterior view. Fig. 2. Right anterior gonopod, lateral
view. Fig. 3. Cyphopods, posterior view. Figs. 4-6. R. josephi. Fig. 4.
Body of male, lateral view of anterior end. Fig. 5. Sternum and coxae of
legs 2 of female, posterior view. Fig. 6. Left postgenital structure of female,
posterior view.

g6

Psyche

[September

terior margins of paranota evenly curved. Prozonites of segments
with small, rounded granules becoming larger, acute on metazonites,
bearing minute branched setae. Segmental setae along anterior mar-
gin of metazonite, outermost at midpoint in lateral margins of para-
nota. Epiproct trilobed. Legs short, femora clavate.

Legs 7 with coxae somewhat enlarged. Anterior gonopods (Figs,
i, 2) typical. Anterior coxal processes closely appressed in midline
(Fig. 1), with large posteriorly directed apical branch (Fig. 2),
blunt, spatulate lateral branch embracing posterior coxal process.
Anterior branch of posterior coxal process long, acute, curved, sword-
like; mesal branch nearly sigmoid, conforming to lateral branch of
anterior coxal process; posterior branch apparently absent. Telo-
podite lobelike. Posterior gonopod (ninth leg) typical, flattened,
setose. 'Coxae of legs 10 enlarged, gland opening on anterior face.
Other postgonopodal legs normal.

Coloration: dark brown, prozonites and metazonites of sixth seg-
ment lighter tan, collum cream-white, legs and venter white. Bases
of segmental setae marked by light spots.

Female from same locality: Size and body form much as in male,
but sixth segment of normal size. Ocelli of 3 females: 2 specimens
have 5 ocelli, one has 6. Cyphopods and postgenital structures as
in Fig. 3.

Distribution: OREGON: Yamhill Co., 5 mi east of Yamhill on Hwy
240, Berlese of litter and grass, 2 October 1971, E. Benedict, 5 $ $ ;
Washington Co., 2 mi north of Helvetia on Bishop Road, Berlese
of mixed conifer and deciduous duff, 21 January 1968, D. Malcolm,
9; Tillamook Co., 4 mi south of Blaine, elev. 50c/, Berlese of rotten
wood, 15 March 1972, E. Benedict, $.

Rhiscosomides josephi Chamberlin
Figs. 4-9

Rhiscosomides josephi Chamberlin, 1941, Bull. Univ. Utah Biol. Ser. 6:

16-17, no figs.; Shear, 1972, Bull. Mus. Comp. Zool. 141(4): 262.

Type: Female holotype from “John Day Creek,” Douglas Co.,
Oregon, collected 18 November 1941, by J. C. Chamberlin; speci-
men in Chamberlin Collection, now at U. S. National Museum.
There is no “John Day Creek” in Douglas Co., but there is a town
of Day Creek, and a small stream of that name flowing into the
South Umpqua River. It is presumed that this is the type locality,
and not the region of the John Day River to the northeast, semiarid
country from which few millipeds have been collected. All subse-

1973]

Shear — Rhiscosomididae

197

quent collections of R. josephi are from the Douglas Co. region.
Admittedly, the assignment of this species name is somewhat arbi-
trary, but no harm is done by using it for the commonest species of
southwestern Oregon.

Description: Male from Canyonville County Park, Douglas Co.,
Oregon; length, 7.0 mm, width, 1.10 mm. Body of typical form,
nonsexual characters as described for R. mineri .

Anterior gonopods (Figs. 7, 8): anterior coxal processes short,
sharply curved anteriad, termination complex, somewhat variable
(compare Figs. 7 and 8), usually with large lateral tooth, smaller
mesal teeth, small anterior tooth; lateral branch of process a broad
flange embracing posterior coxal processes. Posterior coxal processes
with anterior and mesal branches bent sharply posteriad at right
angles, posterior branch much reduced. Telopodites typical. Ninth
legs (posterior gonopods) typical of genus (Fig. 9). Coloration as
usual.

Female from same locality; size and structure much as in male,
sixth segment not enlarged. Sternite of second legs with blunt exten-
sion between coxae (Fig. 5), postgenital structure similar to that of
R. mineri (Fig. 6).

Distribution: Oregon: Coos Co., 8 mi east, 2 mi south of Alle-
gany, Weyerhauser Co. Millicoma Tree Farm, company road 5000,
Berlese of Pseudotsuga bark flakes on clear-cut slope, 20 November

1971, E. M. Benedict, 2 ; Curry Co., 13 mi east of Gold Beach on
road to Agness, elev. 600', Berlese of tan oak duff, 10 March 1972,
E. M. Benedict, $22; Douglas Co., Canyonville County Park, 2
mi east of Canyonville off Rt. 227, Berlese of duff, moss, wood, soil,
elev. 1000', 6 November 1971, E. M. Benedict, c? cT?9, 2 mi north
of Melrose, elev. 400', berlese of rotted wood and duff, 7 February

1972, E. M. Benedict, $ , 0.7 mi west of Scottsburg, near Umpqua
River, elev. 300', Berlese of rotted myrtle heartwood, 1 1 December
1971, E. M. Benedict, 8 2, Elliot State Forest, 1 mi south, 2 mi
west of Ash, elev. 1 100', Berlese of mixed duff from conifers, bigleaf
maples, 11 December 1971, E. M. Benedict, 2 , 2 mi southeast of
Day Creek on Rt. 227, elev. 1000', berlese of oak and mad rone
litter, 6 November 1971, E. M. Benedict, $ .

Notes: Fig. 8, of the male from 2 mi southeast of Day Creek, and
Fig. 7, of the described male from Canyonville County Park, though
fairly close geographically, represent the extremes of variation in the
termination of the anterior coxal process. Females are difficult to
distinguish, except by locality (see Map 1) from those of R. mineri,
as the postgenital structures are very similar (cf. Figs. 3 and 6).

98

Psyche

[September

Figs. 7-9. Rhiscosomides josephi. Fig. 7. Right anterior gonopod, lateral
view. Fig. 8. Termination of gonopod of variant specimen, lateral view.
Fig. 9. Left posterior gonopod, anterior view. Figs. 10-13. R. benedictae.
Fig. 10. Right anterior gonopod, lateral view. Fig. 11. Termination of
gonopod of variant specimen, lateral view. Fig. 12. Sternum and coxae of
legs 2 of female, posterior view. Fig. 13. Left cyphopod and postgenital
structure, posterior view.

1973]

Shear — Rhiscosomididae

199

Map 1. Coastal Oregon, showing distribution of species of Rhiscosomides.
Dots, R. mineri. Squares, R. josephi. Triangles, R. henedictae. Circles, R.
malcolmi.

200

Psyche

[September

Rhiscosomides benedictae n. sp.

Figs. 10-14

Types: Male holotype and female paratype from woods behind
Marine Biological Institute, Charleston, Coos Co., Oregon, col-
lected from a Berlese sample of spruce, alder and cedar duff by
E. M. Benedict, 30 April 1967.

Description : Male holotype; size and nonsexual characters as
described for R. mineri.

Anterior gonopods: anterior coxal processes long (Fig. 10),
slightly curved, terminating in lateral and dorsal teeth and blunt
mesal lobe; lateral branch small lamella. Posterior coxal processes
with anterior branch small, rodlike, mesal branch upright, of mod-
erate size, posterior branch relatively large, membranous anterior
face. Telopodites lobed, interlocking with lateral extensions of an-
terior coxal processes. Ninth legs (Fig. 14) of usual form.

Female paratype typical of genus; process from second sternite
(Fig. 12) thin, postgenital structures as in Fig. 13.

Distribution: Oregon: Lincoln Co., State Forest Camp east of
Waldport, 30 October i960, D. R. Malcolm, $ ; Benton Co., Rt.
34 at Benton Co. line, Berlese of maple and alder duff, 30 October
i960, Malcolm, $$99; Douglas Co., 3.2 mi northeast of Scotts-
burg, elev. 400', Berlese of rotted wood and bark, 1 1 December
1971, E. M. Benedict, c?¥$; Coos Co., 4 mi east, 2 mi south of
Allegany, Weyerhauser Co. Millicoma Tree Farm, company road
6000, Berlese of rotted wood from riparian zone of Fall Creek,
between steep canyon walls, 21 November 1971, E. M. Benedict,
$ $99.

Notes: Fig. 11 illustrates a slight variation in the form of the
termination of the anterior coxal process of the anterior gonopod of
the Benton Co. specimen. Those from the Millicoma Tree Farm
are similar to this; at that place R. benedictae is nearly syntopic with
R. josephi, which was taken from bark chips on a clear-cut slope,
while R. benedictae was taken from litter in a riparian zone. It
would be interesting to further explore the ecological situation be-
tween these two species.

Rhiscosomides malcolmi n. sp.

Figs. 15, 16

Types: Male holotype, female paratypes from 13 mi north, 5 mi
west of Brookings, Curry Co., Oregon, collected 10 March 1972

1973]

Shear — Rhiscosomididae

201

from Sitka spruce duff on bluff overlooking ocean, by E. M. Bene-
dict.

Description: Male holotype; length, 7.0 mm, width, 1.10 mm.
Body form typical, as described for R. mineri.

Anterior gonopods (Fig. 15) robust, with anterior coxal processes
bent anteriad at right angle, lateral branch strong, irregular in form.
Posterior coxal processes with anterior branch bent posteriad at
right angle, blade-like, mesal branch large, upright, posterior branch
relatively large, with extensive fimbriate anterior edge. Telopodites
as usual. Posterior gonopods (Fig. 16) as usual for genus.

Female paratype typical. Postgenital structures not distinguishable
from those of R. bene diet ae.

Distribution: OREGON: Curry Co ., 14 mi east of Gold Beach,
elev. 600', berlese of rotted wood and fir duff, 10 March 1972,
E. M. Benedict, $ S99.

Notes: Just as R. josephi and R. mineri females are difficult to
separate, so are those of R. malcolmi and R. benedictae. At the
locality near Gold Beach, R. malcolmi is nearly syntopic with R.
josephi , which was taken there from tan oak duff. A mile away,
R. malcolmi was collected from rotted wood and fir duff.

Rhiscosomides acovescor Shear

Fig. 17

Rhiscosomides acovescor Shear, 1972, Bull. Mus. Comp. Zool. 144(4): 262-

263, figs. 451-458, $.

Types: Male holotype from Sequoia duff, S. P. Taylor State Park,
Marin Co., California, collected 7 January 1962 by C. W. O’Brien,
deposited in Museum of Comparative Zoology, examined.

Description: Male paratype; length, 6.0 mm, width, 1.10 mm
(specimen broken). Body form as described for R. mineri, except as
follows: 5 ocelli; sixth segment not at all enlarged, collum pig-
mented as other segments, not cream-white.

Anterior gonopods : anterior coxal processes of each side completely
fused distally, separated by slight suture proximally, forming broad
anterior plate, simple, not branched (Fig. 17). Posterior oxal pro-
cesses with anterior branch small, weak, mesal branch thick, heavy,
posterior branch thin, laciniate. Telopodites relatively large. Pos-
terior gonopods (ninth legs) typical for genus.

Females unknown.

Distribution: Known only from the type locality.

Notes: The female paratype I designated in 1972 was not dis-

[September

202 Psyche

Fig. 14. Left posterior gonopod of Rhiscosotnides benedictae, anterior
view. Figs. 15, 16. R. malcolmi. Fig. 15. Right anterior gonopod, lateral
view. Fig. 16. Left posterior gonopod, anterior view. Fig. 17. Right
anterior gonopod of R. acovescor, lateral view. Fig. 18. Right anterior
gonopod of R. trinitarium, mesal view.

1973]

Shear — Rhiscosornididae

203

sected at that time and turned out to be immature. My interpreta-
tion of the posterior gonopods in that paper was likewise in error;
see above. I have examined the tiny gonopods at high magnification
under phase contrast, but cannot determine for certain if the large
branch on the anterior gonopod made of closely appressed cuticular
fibers is attached to the anterior or posterior coxal process. In R.
trinitarium (see below) this branch is definitely a part of the pos-
terior coxal process, while when gonopods of R. acovescor are dis-
sected, it always seems to go with the anterior coxal process.

Rhiscosomides trinitarium n. sp.

Fig. 18

Type: Male holotype from Butter Creek, elev. 3450', 12 mi south-
east of Hyampom, Trinity Co., California, collected 22 July 1968,
by H. Leech. Deposited in California Academy of Sciences.

Description: Male holotype; length 7.1 mm, width 1.12 mm.
Body form as described for R. acovescor , but sixth segment slightly
larger than seventh, collum cream-white.

Anterior gonopods: Anterior coxal process highly complex (Fig.
18), branches closely appressed in midline, but not fused; lateral
branch large, blunt. Posterior coxal process with small, sharp an-
terior branch, posterior and mesal branches fused (?), complexly
laciniate. Telopodites bilobed. Posterior gonopods (ninth legs)
typical.

Female unknown.

Distribution: Known only from the type locality.

Notes: This species is clearly intermediate between R. acovescor
and the more typical northern group of species.

Literature Cited

Chamberlin, R. V.

1941. New southern millipeds. Bull. Univ. Utah. 32, Biol. Ser. 6: 1-19.
Shear, W. A.

1972 Studies in the milliped Order Chordeumida (Diplopoda) : A re-
vision of the Family Cleidogonidae and a reclassification of the
Order Chordeumida in the New World. Bull. Mus. Comp. Zool.
144(4) : 151-352.

SlLVESTRI, F.

1909. Descrizioni preliminari di vari artropodi, specialmente d’America.
Rend. R. Accad. Lincei 18: 229-233.

1913. Ulustrazione di due famiglie di Chordeumidea (Diplopoda) del
Nord America. Boll. Lab. Zool. Portici 7: 303-310.

SUPPLEMENTARY STUDIES ON ANT LARVAE:
CERAPACHYINAE, PSEUDOMYRMECINAE
AND MYRMICINAE*

By George C. Wheeler and Jeanette Wheeler
Laboratory of Desert Biology, Desert Research Institute,
University of Nevada System, Reno 89507

Subsequent to the publication of our first supplement on the ant
larvae of the subfamily Cerapachyinae (1964), our first paper on
Pseudomyrmecinae (1956) and several supplements on Myrmicinae
(i960, 1972, 1973) we have received from other myrmecologists so
much additional material that it has now become necessary to pub-
lish a supplement.

Genus Phyracaces Emery

revision : The last sentence of our generic characterization

(1964: 69) should read: Hypopharynx usually spinulose dorsally.

Phyracaces elegans Wheeler (Fig. 2). Length (through spiracles)
about 4.7 mm. Very similar to Ph. larvatus (1964: 69) except as
follows. Body more slender. A pair of bosses on lateral surfaces of
venter of AI-AVI. Spiracles small, AI largest, diameter decreasing
posteriorly. Integument densely spinulose, spinules in short to long,
subtransverse to arcuate rows. Body hairs less numerous and shorter
(0.025-0.05 mm long). Head hairs shorter (0.009-0.019 mm long).
Posterior surface of labrum with a ventrally directed medial boss
bearing 6 sensilla, about 5 sensilla on each lateral surface. Mandi-
bles with narrower base. No spinules seen on hypopharynx.

young larva: Length (through spiracles) about 2.1 mm. Simi-

lar to mature larva above except as follows. Neck curved, abdomen
with straight ventral profile and C-shaped dorsal profile. Body hairs
shorter (0.01-0.033 mm long). Integument spinulose, spinules mi-
nute, isolated laterally and in short rows dorsally and ventrally.
Antennae less distinct. Maxillae lacking spinules; galea a slightly
raised pair of sensilla. No spinules seen on labium; opening of
sericteries a short slit.

very young larva: Length (through spiracles) about 1.5 mm.

Entire body arcuate ventrally. Otherwise similar to young larva.

Material studied: numerous larvae from New South Wales,

courtesy of Rev. B. B. Lowery.

* Manuscript received by the editor September 28, 1973

204

1973]

Wheeler & Wheeler — Ant Larvae

205

Fig. 1. Cerapachys {Sycia) australis: a, head in anterior view, X95;
b, left mandible in anterior view, X314; c, larva in side view, X 22 ;
d and e, two types of body hairs, X 444. Fig. 2. Phyracaces elegans: head
in anterior view, X 74. Fig. 3. Tetraponera natalensis: left antenna in
lateral view, X339.

Phyracaces ficosus Wheeler. Length (through spiracles) about
4.4 mm. Very similar to Ph. larvatus (1964: 69) except in the
following details. Spiracles on first abdominal somite slightly larger,
remainder small and subequal. Body hairs shorter (0.013-0.063 mm
long). Antennae with 2 sensilla each. Head hairs shorter (0.008-
0.025 mm long). Galeae digitiform. (Material studied: 14 larvae
from New South Wales, courtesy of Rev. B. B. Lowery.)

Genus Cerapachys F. Smith

revision: Our generic characterization (1964: 67) should be

replaced with the following: Leg vestiges small paraboloidal papillae.
Body hairs usually simple. Head (including mouth parts) subpyri-
form in anterior view. Head hairs usually short. Mandibles long,
slender and with median border erose. Maxillary palp short; galea
long and digitiform.

Cerapachys (Syscia) australis Forel (Fig. 1). Length (through
spiracles) about 3.2 mm. Body long and subcylindrical ; about 12

206

Psyche

[September

differentiated somites; head on anterior end; a small posteriorly
projecting boss on AX. Anus ventral. Spiracles small. Entire
integument densely spinulose, spinules minute and in short to long
straight or arcuate rows. Body hairs short, uniformly distributed
and moderately numerous. Of two types: (i) 0.025-0.063 mm long,
mostly bifid, sometimes with one or both branches rebranched, on all
somites; (2) 0.037-0.05 mm long, simple, a few on each somite.
Cranium subhexagonal in anterior view, slightly longer than wide.
Antennae large, each a low mound with 3 minute sensilla, each
bearing a minute spinule. Head hairs few, 0.025-0.05 mm long,
simple or bifid. Labrum subarcuate, about twice as wide as long;
anterior surface with 8 sensilla on and near ventral border; posterior
surface with about 6 sensilla ventromedially and with a few oblique
arcuate rows of minute spinules. Mandibles narrowly subtriangular
in anterior view; apex rather long, narrow and heavily sclerotized;
medial border with 6-8 small denticles. Maxillae with apex para-
boloidal and sparsely spinulose, spinules minute to short and in a
few arcuate rows; palp a peg with 4 (2 encapsulated and 2 bearing
a spinule each) apical and one lateral sensilla; galea digitiform with
2 apical sensilla, each bearing a minute spinule. Labium subtrape-
zoidal, widest distally, anterior surface densely spinulose, spinules
minute and in numerous short arcuate rows; palp a rounded elevation
with 5 (2 encapsulated and 3 bearing a spinule each) sensilla; an
isolated sensillum between each palp and opening of sericteries; the
latter a slit in a shallow depression on anterior surface. Hypopharynx
with minute spinules in long transverse sub-parallel rows. (Material
studied: 10 larvae from Queensland, courtesy of Rev. B. B. Lowery.)

Subfamily Pseudomyrmecinae

We have never been able to key the genera of this subfamily.
Except for head shape, where the difference in the species of Pachy-
sima is greater than that between any two genera, some of the vari-
ants of any character in any genus can be found in other genera.

Bernard (1951: 1053) included larval characters in his character-
ization of the subfamily, which he called family Promyrmicidae.

Sudd (1967: 123) discussed the feeding of the larvae. He stated
(erroneously) that the trophothylax was formed by the bases of the
rudimentary legs; we have shown (1956: 375, 383) that it is
“formed from the depressed ventral surface of the thorax and elab-
oration of the first and second abdominal somites.”

1973]

W heeler & Wheeler — Ant Larvae

207

Genus Pseudomyrmex Lund

Janzen (1967: 344). Beltian bodies are cut up by the workers
and fed to the larvae.

The following species of Pseudomyrmex are compared with Ps.
alliodorae 1956: 379); only differences are given here.

Pseudomyrmex adustus Borgmeier. Length (through spiracles)
about 4.8 mm; straight length about 4.6 mm. Body hairs: (2)
0.05-0.25 mm long, longest of AI-AV; (3) 0.2-0.25 mm long, 2
only on each T1-3 and AI-AIII. Head hairs more numerous and
0.05-0.25 mm long. Posterior surface of labrum with a cluster of 3
sensilla in the middle of each half. (Material studied: 7 larvae from
Brazil, courtesy of Dr. K. Lenko.)

Pseudomyrmex belti fulvescens Emery (= Ps. ferrugineus F.
Smith). Janzen 1967: Description p. 394; feeding of larvae p. 416-
417; handling of larvae p. 418. Similar information in Janzen
1966.

Pseudomyrmex elongatus Borgmeier. Length (through spiracles)
about 3.8 mm; straight length about 3.6 mm. Body hairs longer:
(1) 0.006-0.018 mm long; (2) 0.018-0.2 mm long; (3) 0.175-0.22
mm long, 4 in a row across the dorsum of each T1-3 and AI-AIV.
Head hairs more numerous and slightly longer (0.01-0.05 mm long).
(Material studied: 15 larvae from Brazil, courtesy of Dr. K. Lenko.)

Pseudomyrmex schuppi Forel. Length (through spiracles) about
5.9 mm; straight length about 5.7 mm. Largest spiracles on AI.
Body hairs longer: (1) 0.013-0.025 mm long; (2) 0.038-0.25 mm
long; (3) 0.25-0.33 mm long, 4 in a row across the dorsum of each
T1-3 and AI-AIII. (Material studied: numerous larvae from Brazil,
courtesy of Dr. K. Lenko.)

Pseudomyrmex subtilissimus Emery. Length (through spiracles)
about 4.3 mm; straight length about 3.9 mm. Body hairs ( 1) 0.006-
0.018 mm long; (2) 0.018-0.15 mm long; (3) about 0.15 mm long,

4 in a row across the dorsum of each T1-3 and AI-AIV. Head hairs
slightly longer (0.025-0.05 mm long). (Material studied: 6 larvae
from Brazil, courtesy of Dr. K. Lenko.)

Pseudomyrmex termitarius F. Smith. Length (through spiracles)
about 5.1 mm; straight length about 4.7 mm. Body stouter. Body
hairs about twice as numerous: (1) 0.013-0.05 mm long; (2) 0.05-
0.275 mm long; (3) 0.25-0.35 mm long, 4 in a row across dorsum
of each T1-3 and AI-AIV. Head hairs more numerous, longer
(0.013-0.05 mm long) and finely denticulate. Labrum with width
twice the length, with anterior lobes more prominent and with 2

208

Psyche

[September

minute hairs on anterior surface. Mandibles with teeth stouter and
blunter; lateral outline less curved; denticles on anterior surface
more numerous. Maxillary apex less constricted and with spinules
longer and covering a greater portion of the surface. Labium with
more numerous spinules. (Material studied: 9 larvae from Brazil,
courtesy of Dr. K. Lenko.)

Genus Tetraponera F. Smith

Tetraponera natalensis F. Smith (Fig. 3). Length (through spira-
cles) about 8.2 mm; straight length about 6.2 mm. Similar to T.
aitkeni (1956: 388) except as follows. Body slightly stouter at AV
and AVI. Integument of AIX and AX with minute spinules. Body
hairs: (1) 0.008-0.075 mm long; (2) 0.025-0.15 mm long, longest
with tip branched or denticulate; (3) 0.175-0.3 mm long, 4 in a
row across the dorsum of each T1-3 and AI-AVI. Each antenna
represented by 3 individually raised sensilla on a small base. Head
hairs longer (0.013-0.11 mm long) and less numerous, with or
without alveolus and articular membrane, some with denticles near
the tip. Labrum with breadth less than twice length ; borders sinu-
ate; anterior surface with 6 sensilla and 2 hairs on each half; pos-
terior surface with 9 sensilla on each half; spinules as in T. aitkeni.
Anteromedial surface of mandibles with large spinules, which are
isolated or in short rows of 2 or 3. Maxillae with rather numerous
long spinules in short arcuate rows; palp represented by a cluster of
5 sensilla on a slight elevation. (Material studied: numerous larvae
from South Africa., courtesy of Dr. W. L. Brown.)

Genus Pachysima Emery

Pachysima latifrons Emery: Bernard ( 1 95 1 : 1054-1057) de-
scribed and figured the young (after W. M. Wheeler).

Genus V iticicola Wheeler

Viticicola tessmanni (Stitz) : Bernard (1951: 1054) described
and figured the larva (after W. M. Wheeler).

Subfamily Myrmicinae

Ettershank (1966: 161, 162): “The larvae of the Formicidae

have not been used to any extent in taxonomic studies, although
numerous descriptions and figures of scattered genera and species
occur in the literature. The only wide-scale comparative larval study

1973]

Wheeler & Wheeler — Ant Larvae

209

that has been attempted is the series of papers by G. C. Wheeler
(later with J. Wheeler), which constitute a fundamental contribu-
tion to the subject that will be used for a long time.” “Reference to
all the publications by the Wheelers on myrmicine ants are contained
in a summary article (G. C. and J. Wheeler i960). In this paper,
the authors conclude that three characters are of major importance:
body profile, mandible shape, and setal form. They recognize 22
body profiles and 30 mandibular shape categories all of which are
explained and illustrated.”

Genus Messor Forel

Messor capitatus Latreille: Delage (1968a) gave in a table the
sizes and abundance of larvae throughout the year. She stated that
only small larvae overwinter. She (1968b) discussed larval enzymes
and digestion.

Genus Pheidole Westwood

Kempf (1972: 457): Ph. vallifica is the host of the eucharitid
Orasema costaricensis Wheeler and Wheeler.

Genus Melissotarsus Emery

Delage-Darchen (1972a) : Hairs few, long, with bifid tips. Crude
sketch of a larva on p. 219.

Genus Crematogaster Lund

Delage-Darchen (1972b) found only three larval stages in C.
(N ematocrema) stadelmanni Mayr. Fig. 1 hairs enlarged; Fig. 2
and 3 larvae of various stages in side view; Fig. 4 head in anterior
view. Pilosity is taxonomically worthless because of extreme vari-
ation between colonies and even in the same colony.

Genus Monomorium Mayr

Cloudsley-Thompson (1962: 179): The calliphorid flies Ben-

galia peuhi Vil. and B. minor Malloch fed on the larvae of M.
salomonis (Linnaeus) in the central Sudan.

Van Pelt and Van Pelt (1972: 978): Larvae of the syrphid
Microdon baliopterus Loew fed upon the larvae of M. minimum
( Buckley) .

210

Psyche

[September

Genus Solenopsis Westwood

Markin et al. (1972: 1053): Life cycle of Solenopsis invicta
Buren in an incipient colony: egg 6-8 days, larva 14-15 days, pupa
20-24 days.

Genus Tetramorium Mayr

Tetramorium caespitum (Linnaeus). Donisthorpe (1927: 197):
“The larvae were fed with disgorged liquid food as long as they
were young and gathered together in groups, but when they grew
older and were separated, the workers fed them with solid sub-
stances.” Many larvae were hung on to the plaster walls of the nest
by their anchor-tipped hairs.

Literature Cited

Bernard, F.

1951. Super-famille des Formicoidea. Traite de Zoologie, Tome X,
Fasc. II: 907-1119, 1258-1263, 1272-1275.

Cloudsley-Thompson, J. L.

1962. A note on the association between Bengalia spp. (Dipt., Cal-
liphoridae) and ants in the Sudan. Entomol. Monthly Mag. 98:
177-179.

Delage, Bernadette.

1968a. Recherches sur la fourmis moissoneuses du Bassin Aquitain:
ecologie et biologie. Bull. Biol. 100: 315-367.

1968b. Recherches sur les fourmis moissoneuses du Bassin Aquitain:
ethologie. Physiologie de l’alimentation. Ann. Sci. Nat., Zool.
Biol. Anim. (12) 10: 197-265.

1972a. Une fourmi de Cote-d’Ivoire : M elissotarsus titubans Del., n. sp.
Insectes Sociaux 19: 213-226.

1972b. Le polymorphisme larvaire chez les fourmis N ematocrema
d’Afrique. Insectes Sociaux 19: 257-277.

Donisthorpe, H. St. J. K.

1927. British ants. Geo. Routledge & Sons, London. 436 pp.

Ettershank, G.

1966. A generic revision of the world Myrmicinae related to Solenopsis
and Pheidologeton. Australian J. Zool. 14: 73-171.

Janzen, D. H.

1966. Coevolution of mutualism between ants and acacias in Central
America. Evolution 20: 249-275.

1967. Interaction of the bull’s-horn acacia (Acacia cornigera L. ) with
an ant inhabitant (Pseudomyrmex ferruginea F. Smith) in eastern
Mexico. Univ. Kansas Sci. Bull. 47: 315-558.

Kempf, W. W.

1972. A study of some Neotropical ants of genus Pheidole Westwood. I.
Studia Entomol. 15: 449-464.

1973]

Wheeler & Wheeler — Ant Larvae

2i i

Markin, G. P., H. L. Collins and J. H. Dillier.

1972. Colony founding by queens of the red imported fire ant, Solenop-
sis invicta. Ann. Entomol. Soc. Amer. 65: 1053-1058.

Sudd, J. M.

1967. An introduction to the behavior of ants. St. Martin’s Press, New
York. 200 pp.

Van Pelt, A. F., and S. A. Van Pelt.

1972. Microdon (Diptera: Syrphidae) in nests of Monomorium in
Texas. Ann. Entomol. Soc. Amer. 65: 977-979.

Wheeler, G. C., and Jeanette Wheeler.

1956. The ant larvae of the subfamily Pseudomyrmecinae. Ann. Ento-
mol. Soc. Amer. 49: 374-398.

1960. Supplementary studies on the larvae of the Myrmicinae. Proc.
Entomol. Soc. Washington 62: 1-32.

1964. The ant larvae of the subfamily Cerapachyinae : supplement.
Proc. Entomol. Soc. Washington 66: 65-71.

1972. Ant larvae of the subfamily Myrmicinae: second supplement on
tribes Myrmicini and Pheidolini. J. Georgia Entomol. Soc. 7:
233-246.

1973a. The ant larvae of six tribes: second supplement. J. Georgia
Entomol. Soc. 8: 27-39.

1973b. Ant larvae of four tribes: second supplement. Psyche 80: 70-82.

1973c. Ant larvae of the myrmicine tribe Attini: second supplement.
Entomol. Soc. Washington. (In press.)

1973d. The ant larvae of the tribes Basicerotini and Dacetini : second
supplement. Pan-Pacific Entomol. (In press.)

STUDIES ON NEOTROPICAL POMPILIDAE
( HYMENOPTERA ) . IX.

THE GENERA OF AUPLOPODINI*

By Howard E. Evans

Museum of Comparative Zoology, Harvard University,
Cambridge, Mass. 02138, U.S.A.1

The posthumous paper of Hermann Haupt on the classification
of the Macromerinae (Haupt, 1959) is an unworthy memorial to
its author and an unfortunate step backward in the systematics of
spider wasps. Working from very little material and a lack of
awareness of research in other parts of the world, Haupt erected
12 new genera, few if any of which are likely to stand the test of
time. Two of them actually belong in the subfamily Pompilinae, as
synonyms of Priochilus Banks (Evans, 1966), while others will fall
in the Pepsinae in most classifications ( Compsagenia , Anapriocnemis) .
H is inclusion of such diverse elements in the Macromerinae suggests
the difficulty in defining the group, which I would rank as a rather
weakly characterized tribe of Pepsinae and call the Auplopodini,
after the first genus to be used in a suprageneric sense, Pseudagcnia ,
now properly called Auplopus.

I am not in a position to straighten out all the confusion caused
by Haupt’s paper, but I wish to consider seven genera which he
described from the neotropics, all of which can be promptly rele-
gated to synonymy. There are, however, several remarkable new
genera and subgenera of this tribe in South America, wdiich both
Haupt and Banks (1946) were unaware of, and I shall use this
opportunity to describe these taxa and to present a revised key to
neotropical Auplopodini.

I am much indebted to Professor J. O. Hiising, of the Zoolo-
gisches Institut, Halle, for permitting me to borrow some of the
specimens that Haupt studied, including several types.

Pseudageniellci Haupt

Pseudageniella Haupt, 1959, pp. 23, 46 (type-species: Pompilus rusticus

Fabricius, 1804, monotypic and by designation).

^Published with the aid of a grant from the Museum of Comparative
Zoology.

Present address: Dept, of Entomology and Zoology, Colorado State Univ.,
Fort Collins, Colo. 80521.

Manuscript received by the editor > June 7 1 1973 .

212

1973]

Evans — - Neotropical Pompilidae

213

I have not seen the type of rusticus, but the specimens that Haupt
had before him agree perfectly with the type of Priophanes insolens
Banks, 1946, and they are from the same locality. This is a well-
marked and evidently common species, and these specimens agree
well with Fabricius’ description. I therefore do not hesitate to con-
sider insolens Banks a synonym of rustica Fabricius (new syn-
onymy). Townes (1957) places insolens in Ageniella , subgenus
Ameragenia , an assignment with which I concur; thus P seudageniella
Haupt falls as a synonym of Ameragenia Banks, 1945 (new syn-
onymy). In his key, Haupt states that this genus is from the Ne-
arctic region, an obvious error, since he states on p. 46 that his
material is from Brazil.

Allageniella Haupt

Allageniella Haupt, 1959, pp. 23, 46 (type-species: Allageniella obsoleta
Haupt, 1959, monotypic and by designation).

Haupt separated this genus from the preceding on exceedingly
minor characters, and in fact his obsoleta and his specimens of
rustica differ primarily in size and coloration. I have compared the
type specimen of obsoleta Haupt with that of Priophanes plagosa
Banks, 1946, and found them to be conspecific; again, both are from
the same locality. Townes (1957) also places plagosa in Ageniella,
subgenus Ameragenia. Thus Allageniella is to be regarded as a
synonym of Ameragenia Banks, obsoleta a synonym of plagosa Banks
(new synonymies).

Brachyagenia Haupt

Brachyagenia Haupt, 1959, pp. 25, 59 (type-species: Brachyagenia nigra
Haupt, 1959, monotypic and by designation).

By comparison of type specimens, B. nigra is to be regarded as a
synonym of Ameragenia thione Banks, 1946 Brachyagenia thus rep-
resents still another synonym of Ameragenia Banks (both new syn-
onymies) .

P ar ageniella Haupt

Parageniella Haupt, 1959, pp. 26, 61 (type-species: Priocnemis rufofemoratus
Taschenberg, 1869, monotypic and by designation).

The type-species is a well-marked Argentinian form and I have
little doubt that Haupt and Banks identified it correctly. Banks
placed rufofemorata in Priophanes, but like most of Banks’ South

214

Psyche

[September

American Priophanes the species will run to Arneragenia in Townes’
( 1 957 ) key. I would regard Parageniella as still another synonym
of Arneragenia (new synonymy).

Cosmagenia Haupt

Cosmagenia Haupt, 1959, pp. 28, 63 (type-species: Agenia amabilis Tas-
chenberg, 1869, by designation).

I have studied Haupt’s material of amabilis and found it con-
specific with Ageniella amoena Banks, 1946. Since Haupt presum-
ably had access to Taschenberg’s types, it seems safe to place amoena
in the synonymy of amabilis (new synonymy). This species is prop-
erly placed in the genus Priocnemella Banks, 1925, and Cosmagenia
can thus be relegated to the synonymy of that genus (new syn-
onymy) .

Compsagenia Haupt

Compsagenia Haupt, 1959, pp. 29, 66 (type-species: Compsagenia laevipes
Haupt, 1959, monotypic and by designation).

Study of the type specimen of laevipes shows it to be conspecific
with the type of Nannochilus ' obscurus Banks, 1946 (new syn-
onymy). Townes (1957) has correctly placed Nannochilus in the
synonymy of Minagenia Banks, 1934, and Compsagenia Haupt
should now be added to the list (new synonymy). Since levipes
Cresson, 1869, also belongs to this genus, Haupt’s laevipes is both
a homonym and a synonym. Townes assigns Minagenia to the sub-
family Ceropalinae, tribe Minageniini. I would assign it to the
Pepsinae, but I do not pretend to understand how the genera should
be grouped within that subfamily; at any rate Minagenia does not
belong in the Auplopodini.

Anapriocnemis Haupt

Anapriocnemis Haupt, 1959, pp. 25-26, 60-61 (type-species: Pompilus flavipes
Guerin, 1836, by designation).

P. flavipes is a well known Chilean species which has been as-
signed by Townes (1957) to the genus Priocnemis, subgenus Sphic-
tostet fins Kohl, along with the other two species assigned by Haupt
to Anapriocnemis. Since the type of Sphictostethus has a brachypter-
ous female, it might be argued that flavipes is not correctly assigned
to that subgenus (though I would not so argue) ; in any case Ana-
priocnemis may be placed in the synonymy of Priocnemis Schiodte,

1973]

Evans — Neotropical Pompilidae

215

which is placed in the Pepsini in Townes’ classification (new syn-
onymy) .

Mystacagenia, new genus
Type-species. — M. variegata, new species

Generic characters. — Females with the general features of Age-
niella , including wing venation (shown in Fig. 2) ; males unknown.
Length of known species 5. 5-8. 5 mm; variously colored, with banded
wings. Maxillary palpi very long, capable of reaching apex of front
coxa; mentum with a few thin setae; mandibles with a basal swell-
ing just below which there is a group of long, pale setae which overly
and partially conceal the mandible; mandibles slender apically, with
a small tooth on the inner margin that is strongly set off from the
shaft. Labrum partially exposed, with a deep median notch; clypeus
about as wide as lower face ; malar space well developed, at least half
as long as width of mandibles at their base; antennae unusually
slender (Figs. 1, 3). Propodeum sloping evenly, its surface smooth
and devoid of setae; legs devoid of setae except for very minute ones
on the tibiae and tarsi; claws dentate. First abdominal segment hour-
glass shaped at extreme base ; last segment bearing a number of setae,
without a smooth pygidial area.

Key to species ( females )

1. Front of head, including mouthparts, white in color; vertex

bearing several strong, curved, white setae just behind the
ocelli (Fig. 3) ; hind wings with a subapical band

al biceps , new spec:es

Front of head largely ferruginous; vertex without prominent
setae; hind wings hyaline 2

2. Abdomen largely fuscous except basal and apical segments white;

fore wings hyaline, with a strong band below the stigma and a
narrow band over the transverse median vein ; length of fore

wing 5 mm bellula, new species

Abdomen with all segments irregularly blotched with brown and
white; fore wings opaque whitish, crossed by three brown
bands, the outer two connected above; length of fore wing
7.5 mm variegata , new species

Mystacagenia variegata, new species

Holotype , — 9, Nova Teutonia, Santa Catarina, Brazil, 21 Jan.
1956 (Fritz Plaumann) [Coll. H. K. Townes, Ann Arbor, Mich.].

2l6

Psyche

[September

Fig. 1. Anterior view of head of Mystacagenia variegata n. sp., type 2 .
Fig. 2. Wings of same specimen, color pattern not shown. Fig. 3. Anterior
view of head of M. albiceps n. sp., type 2 . Fig. 4. Wings of Dimorpha-
genia naumanni n. sp., allotype 2. Fig. 5. Mandible of same specimen.
Fig 6. Mandible of Ageniella ( Cyrtagenia ) innuba n. sp., type 2. Fig. 7.
Lateral view of head of same specimen. Fig. 8. Subgenital plate of Pi-
morphagenia naumanni n. sp., type $ . Fig. 9. Genitalia of same specimen,
ventral aspect.

1973]

Evans — Neotropical Pompilidae

217

Description. — Length 8.4 mm; fore wing 7.4 mm. Head light
rufous, with a pair of black spots on upper front ; front and temples
with a reticulate pattern of white; malar space, mandibles, and
lab rum mostly white; thoracic dorsum mainly light rufous, sides with
broad streaks of light rufous and white, also some black on posterior
parts of mesopleura and metapleura ; propodeum white, with a pair
of broad, longitudinal black bands ; abdomen mainly whitish or some-
what cream in color, tergite 1 with a small amount of black latero-
basally and medioapically, tergites 2-5 with much black basally, ter-
gite 6 mostly pale; venter mostly pale, but all pale markings of
abdomen irregularly tinged with brown ; antennae stramineous and
partially infuscated on basal 0.3, again at basal vein, this band con-
nected through the first submarginal cell with another band partially
crossing the wing at the second submarginal ; hind wings hyaline.
Body with pale, inconspicuous pubescence; erect setae absent except
on clypeus, mouthparts, venter and apex of abdomen.

Clypeus 3 X as wide as high, its apical margin sinuate, with a
broadly rounded median lobe; front broad, middle interocular dis-
tance .60 X head width ; upper interocular distance .70 X lower
interocular distance; vertex very weakly arched between eye tops,
subcarinate behind ocelli; postocellar line: ocello-ocular line = 4:5;
antennae very slender, third segment 6.6 X as long as its apical
width, 1.2 X upper interocular distance (Fig. 1). Pronotum rather
flat dorsally, its posterior margin broadly angulate; postnotum
widened at the midline; wing venation shown in Fig. 2.

Mystacagenia bellula, new species

H'olotype. — $, Avispas, 30 mi. from Marcapata, 'Cusco, Peru,
1 -1 5 Oct. 1962 (Luis Pena) [Coll H. K. Townes, Ann Arbor,
Mich.].

Description. — Length 6.2 mm; fore wing 4.7 mm. Head testa-
ceous to somewhat orange, with small dark blotches at center of
inner eye margins and a larger dark blotch in front of anterior
ocellus; temples, malar space, and area below antennal sockets more
or less white; mandibles, labrum, and palpi white; thorax and pro-
podeum rufous except propleura and anterior corners of pronotum
white; abdomen dark brown except all of segments 1 and 6 and much
of sternite 5 contrastingly white; scape white, flagellum white on
basal half, testaceous on apical half, streaked with fuscous on lower
surface throughout; legs white, with a complex pattern of dark
brown and a small amount of rufous at base of middle and hind

2 1 8

Psyche

[September

coxae. Wings clear hyaline, fore wing with a brown band over
transverse median vein (weakly extended along basal vein) and a
much broader brown band nearly crossing wing at stigma. Body
pubescence very fine and inconspicuous; body with scarcely any erect
hairs except for those on mandibles, a few on clypeus, and some thin
ones on apex and venter of abdomen.

Clypeus 2.7 X as wide as high, shaped as in the preceding species;
front less broad than in variegata , middle interocular distance .55 X
head width; upper interocular distance .73 X lower interocular dis-
tance; vertex very weakly arched above eye tops; postocellar line:
ocello-ocular line — 6:7; antennae very slender, third segment 6 X
as long as its apical width, 1.2 X upper interocular distance. Pro-
notal disc rather flat, very short, posterior margin broadly angulate;
metanotum angularly projecting backward medially; midline of pro-
podeum weakly impressed; wing venation differing from that of
variegata in no important details.

Mystacagenia albiceps, new species

Flolotypc. — $, Avispas, 30 mi. from Marcapata, Cusco, Peru,

1 -1 5 Oct. 1962 (Luis Pena) [Coll. H. K. Townes, Ann Arbor,
Mich.].

Description. — Length 5.6 mm; fore wing 4.5 mm. Head and
mouthparts entirely white except vertex and occiput blotched with
testaceous; thorax and propodeum predominantly rufotestaceous,
blotched with fuscous across much of pronotal disc, center of meso-
scutum, base and apex of scutellum, along pleural sutures, and over
most of venter; abdomen rufotestaceous, blotched with fuscous on
sides of tergites 1-4, tergites 2 and 3 also with small lateral white
spots; scape mostly white, flagellum brown, darkened toward apex;
legs mostly rufotestaceous, coxae blotched with fuscous, apices of
femora and most of tibiae blotched with fuscous and annulated with
white. Wings hyaline except hind wing with a preapical brown band,
apex clear; fore wing brownish at base, across basal and transverse
median veins, and in a broad band below stigma, the last band ex-
tended along radial vein. Pubescence delicate, inconspicuous; clypeus
with several white setae in addition to the tufts on the mandibles,
ocellar area also with several strong, curved, white setae; scutellum
and apex and venter of abdomen with sparse, weaker setae.

Clypeus 2.4 X as wide as high, apical margin weakly convex;
head subcircular in anterior view; middle interocular distance .60 X
head width ; upper interocular distance .68 X lower interocular dis-

1973]

Evans — Neotropical Pompilidae

219

tance; vertex strongly elevated above eye tops, especially at ocellar
triangle; postocellar line: ocello-ocular line — 2:1; third antennal
segment 7.5 X as long as its apical width, 1.1 X upper interocular
distance (Fig. 3). Pronotum short, disc sloping and with no flat
dorsal surface; postnotum transverse, not projecting backward medi-
ally. Wing venation similar to that of variegata but stigma unus-
ually wide, third submarginal cell smaller, only 1.5 X as wide as
high, removed from wingtip by twice its own width.

Dimorphagenia, new genus

Type-species. — D. naumanni, new species.

Generic characters. — With the general features of the Auplo-
podini, including the wing venation (shown in Fig. 4) and the form
of the first abdominal segment; length 7-10 mm; wings unbanded,
lightly tinged with brown. Female: maxillary palpi of moderate
length ; mentum with a number of strong setae arising near base and
directed forward, much as in Auplopus ; mandibles slender, with
scattered, strong bristles (Fig. 5) ; labrum wholly concealed; clypeus
not extending under lower margins of eyes; malar space about one
third as long as width of mandibles at their base; temples well de-
veloped, not strongly receding, nearly as wide as eyes; vertex ex-
tended well above eye tops ; ocellar triangle located well before
vertex crest; propodeum with smooth contours, slope low and even;
legs relatively smooth, but middle and hind tibiae bearing numerous
spines of moderate length ; claws dentate, tooth arising rather close
to outer ray; apical tergite with a flat pygidial area which is devoid
of setae but is minutely punctate and shagreened. Male (Fig. 10) :
head remarkably enlarged, much wider than thorax, vertex far above
eye tops and ocelli, temples much wider than eyes; malar space about
half as long as width of mandibles at their base; antennae elongate,
capable of reaching middle of abdomen; tarsal claws and spines of
tibiae as in female; first abdominal segment much expanded from the
base, but with no evidence of a lateral seam. Subgenital plate tongue-
shaped, midline only weakly elevated (Fig. 8) ; genitalia with the
basal hooklets absent, parameres elongate, digiti broad and abruptly
truncate apically (Fig. 9).

Remarks. — This genus is most closely related to Auplopus, dif-
fering in the less strongly petiolate abdomen (especially in the male),
the less well defined pygidial area, presence of a short malar space,
broad clypeus with a slightly concave apical margin, and several
other features. The male genitalia differ in no important details
from those of Auplopus.

220

Psyche

[September

Dimorphagenia naumanni, new species

Holotype. — c?, Limoncoche, Prov. Napo, Ecuador (oo° 24'S,
76° 36'W) 7 May 1971 (Martin G. Naumann, nest 2048) [Mus.
Comp. Zool., no. 32105].

Description >of male type. — Length 7 mm; fore wing 6 mm.
Entire body testaceous except center of front and (to a lesser degree)
vertex blotched with medium brown ; legs wholly testaceous ; an-
tennae testaceous darkened to medium brown beyond basal third,
flagellar segments narrowly ringed with fuscous apically. Wings very
lightly tinged with brown; stigma testaceous. Pubescence pale,
inconspicuous. Body largely devoid of erect setae except for strong
bristles on clypeus and mandibles, scattered setae on front, vertex,
and thoracic dorsum (but not propodeum), and numerous short
setae toward apex of abdomen.

Clypeus 2.8 X as wide as high, apical margin weakly concave;
middle interocular distance .68 X head width, 1.4 X eye height;
upper and lower interocular distances subequal; postocellar line:

1973]

Evans — Neotropical Pompilidae

221

ocello-ocular line = 2 :5 ; in lateral view, distance from eye tops to
vertex crest .7 X eye height, temples about 1.5 X eye width; third
antennal segment 4.3 X as long as wide, equal to slightly less than
half upper interocular distance. Maximum width of thorax only .7
that of head ; pronotum weakly expanded dorsally, its midline de-
pressed. Wing venation as in female; terminalia as figured (Figs.
8, 9) ; lateral view of body shown in Fig. 10.

Allotype. — 9) same data as type except dated 3 July 1971 [Mus.
Comp. Zool.].

Description of female allotype. — Length 10 mm; fore wing
8 mm. Head, thorax, and propodeum dark brown, somewhat shin-
ing; abdomen rufous except base of first segment black; antennae
dark brown; coxae, trochanters, and tarsi dark brown, femora and
tibiae rufous. Wings tinged with yellowish brown; stigma light
brown. Pubescence cinereous to light brown, rather conspicuous on
coxae, pleura, and propodeum. Body with fairly numerous pale,
erect hairs, including some on thoracic dorsum and pleura, pro-
podeum, coxae, and especially the abdominal venter; hind femora
with scattered short hairs.

Clypeus 2.5 X as wide as high, its apical margin weakly concave;
middle interocular distance .63 X 'head width, 1.1 X eye height;
upper interocular distance very slightly exceeding lower interocular
distance; vertex broadly rounded off well above eye tops, distance
from posterior ocelli to top of vertex exceeding postocellar line;
postocellar line : ocello-ocular line — 2:5; temples strong, although
roundly contracted from behind the eyes, in lateral view not quite
as wide as eyes ; antennae not especially elongate, third segment only
about half the upper interocular distance. Maximum width of thorax
only slightly less than that of head ; features of pronotum and post-
notum as described for male ; wing venation as in Fig. 4 ; legs and
abdomen as described under generic heading.

Paratypes. — 2 9?> same data as allotype [U.S. Nat. Mus., Brit-
ish Mus.].

Variation. — Both paratypes are slightly smaller than the allotype
(fore wing 7.3, 7.5 mm) but there are no differences worthy of note.

Remarks. — Despite the great difference in head structure in the
two sexes, there is close agreement in all other essential features,
and there can be no question that these are male and female of one
species. This is the only case known to me in the Pompilidae in
which sexual dimorphism involves a major difference in head size.
In this connection the following notes provided by Martin G.

222

Psyche

[September

Naumann may be of interest (his nest no. 2048; see type designation
for locality).

This was a nest of Stelopolybia sp., a social vespid that typically
nests in cavities. In this case the nest occupied several cavities inside
a large carton ant nest ( Azteca sp.) attached to a tree trunk, 2 m
above the ground. On May 7, a wasp was seen walking about on
the surface of the ant nest. It was captured and proved to be a male
pompilid (the type of this species). On June 21 both wasp and ant
nest were heavily damaged by children, but on July 3 both wasps
and ants were still active, and the nest was harvested by chloro-
forming it and catching the contents in a sac. The three female
pompilids were found among the vespids, the ants, and the rubble.

Structure of the females suggests strongly that they build mud
cells: this is the usual function of stiff bristles on the labium and a
smooth pygidial plate. In this instance it seems probable that they
were utilizing a part of one of the cavities inside the ant nest and
being tolerated by the ants and the vespids. I suggest that the large
head of the male may enable it to pass as a worker Azteca ant.
These ants are polymorphic, and the larger workers commonly are
macrocephalic. In this instance the workers were considerably
smaller than the male Dimorphagenia , but they were of a similar pale
color and the larger workers decidedly macrocephalic. Presumably
macrocephaly does not occur in the female sex because it would
render them unable to perform their usual hunting and nest-building
activities. Macrocephaly in the male suggests that the male is more
than a passive inhabitant of the nest; perhaps the presence of several
such males inhibits attacks by ants and social wasps. One can only
hope that the relationships of these insects can some day be worked
out in detail.

Genus Ageniella Banks
Cyrtagenia, new subgenus

Type-species. — Ameragenia fallax Aide.

Subgeneric characters. — Females with the general features of
Ageniella s. str. except as follows (males unknown). Mandibles un-
usually broad, with a small tooth located close to the apex (Fig. 6) ;
clypeus with rather sharp anterolateral corners and with a median,
apical angulation; front, in lateral view, either abruptly subangulate
a short distance above the antennal sockets, then flat to the vertex
crest, or flattened all the way from the antennal sockets to the vertex
(Fig. 7), in either case with a median prominence just above the

1973]

Evans — Neotropical Pompilidae

22 3

sockets. Pronotum short, with a somewhat flattened dorsal part; pro-
podeum with smooth contours, without erect setae or with a very
few setae on each side; legs relatively smooth, but middle and hind
tibiae with several rows of very small spines; brush on inner side of
hind tibia continuous to apex. Third submarginal cell receiving
second recurrent vein .4 the distance from the base; anal vein of
hind wing reaching media well before cubital fork. Known species
with the wings unbanded, the antennae dark but with a white annu-
lus near the middle.

Remarks. — -Arnold {1934) described a genus from Africa in
which the female has the front more or less angulate in profile,
Arpactomorpha. However, in this genus the angulate portion has a
median groove, and below the angulation there is an oblique im-
pression on each side of the face. Furthermore, in Arpactomorpha
the mentum has a beard composed of four or five long bristles arising
from the base, whereas in Cyrtagenia there are only a few weak
setae arising along the length of the mentum, as usual in A geniella.

I doubt if there is any close relationship between these two groups.

Key to species ( Females )

Angulation of front well above antennal sockets; some of the ab-
dominal tergites with lateral white spots; pubescence fine and

relatively inconspicuous fallax (Arle)

Front forming a nearly flat, oblique slope from the antennal sockets
to the vertex (Fig. 7) ; abdomen without white spots; pubescence
unusually coarse and hoary innuba, new species

A geniella (Cyrtagenia) fallax (Arle) new combination

Amerag enia fallax Arle 1947, pp. 426-428, figs. 23-25.

Aide’s description and figures are excellent, and there seems no
need to redescribe the species at this time. Arle had a single female,
from near Rio de Janeiro. The species appears to be widely dis-
tributed, as I have seen females from Teresopolis and Nova Teu-
tonia, Brazil; Oran and Tucuman, Argentina; and Avispas, near
Marcapata, Peru. These females are exceedingly variable in color.
All have a pale annulation on the antennae and at least small spots
on the sides of the abdomen, but the other maculations described by
Arle may be much reduced or even absent. At the other extreme,
the specimen from Peru is exceedingly ornate, having ivory spots
over much of the head and thorax, as well as a median stripe on the

224

Psyche

[September

propodeum and lateral stripes on the first tergite. It is possible that
more than one species is involved, but on the basis of presently avail-
able material I am inclined to think not.

Ageniella (Cyrtagenia) innuba, new species

Plolotype. — ?, Nova Teutonia, Santa Catarina, Brazil, Jan.
1966 (Fritz Plaumann) [Mus. Comp. Zool., no. 32106].

Description of female type. — Length 9 mm; fore wing 8.3 mm.
Head black except marked with white as follows: apical .8 of cly-
peus; narrow inner orbits, with extensions toward antennal sockets;
a median streak before anterior ocellus; small spots at eye tops; lower
outer orbits and malar space; mandibles white basally, then testa-
ceous, with dark tips; palpi testaceous; basal 5 antennal segments
black (except scape with a small white spot), next three segments
mainly white, remainder dark brown above, light brown below.
Thorax and propodeum black; abdomen rufous except first segment
black basally; legs rufotestaceous except coxae, trochanters, and
femora partially infuseated; tarsi in part whitish. Wings hyaline,
veins and stigma dark brown. Pubescence coarse, cinereous, giving
the body a somewhat hoary appearance; propodeum with a. few short
erect hairs on each side and abdominal venter and apical tergite
setose, but body otherwise without erect hairs.

Clypeus 2.6 X as wide as high; malar space .2 X width of mandi-
bles at their base; middle interocular distance .59 X head width;
upper interocular distance .95 X lower interocular distance; posto-
cellar line: ocello-ocular line = 2:3; vertex rather sharp, distance
from posterior ocelli to crest about equal to postocellar line. Front
rather flat from vertex crest to antennae, which arise from a pro-
tuberance, as shown in Fig. 7 ; third antennal segment equal to
.72 X upper interocular distance. Hind tibia with a faint longi-
tudinal impression between the two uppermost rows of small spines.

Paratypes. — BRAZIL : 1 ?, same data as type but collected
January 1965 [U.S. Nat. Mus.]; 1 $, Teresopolis, 11 March 1966
(H. & M. Townes) [Coll. H. K. Townes].

Variation. — Both paratypes have a small white spot on each
posterior pronotal lobe. The abdomen of the Teresopolis specimen
is dusky ferruginous, the legs darker than in the type. The topo-
typic paratype is slightly smaller than the type (fore wing 8 mm)
and has a white spot on the third antennal segment as well as a
large one on the scape.

1973]

Evans — Neotropical Pompilidae

225

Key to Neotropical genera of Auplopodini ( Females )
(Modified from Banks, 1946, and Townes, 1957)

1. Apex of front tibia on outer side with a. strongly differentiated,

curved, hooklike spine; clypeus large, extending well beneath

bottoms of eyes 2

Apex of front tibia without a strong, curved spine that is well
differentiated from the other spines 3

2. Last segment of middle and hind tarsi spined beneath ; lower part

of mesopleurum with a projection; clypeus strongly emarginate

Phanochilus Banks

Last segment of all tarsi smooth beneath ; mesopleurum without a
prominence; clypeus truncate Priocnemella Banks

3. Mandibles with a basal tuft of long, pale setae which cover much

of the mandibles; malar space at least half as long as width of

mandibles at their base (Figs. 1, 3)

Mystacagenia , new genus

Mandibles without such modification, simple and with scattered
setae; malar space less than half as long as width of mandibles
at their base, often nearly absent 4

4. Apical tergite covered with bristles and without a differentiated

pygidial area; mentum with or without a few thin setae scat-
tered along its length Ageniella Banks

Apical tergite with a median area which is devoid of setae and
more or less smooth, often polished ; mentum with a group of
stout setae arising near the base and directed forward 5

5. Malar space about one third as long as width of mandibles at

their base; temples prominent, nearly as wide as eyes; tooth of
claws quite close to outer ray (males macrocephalic, Fig. 10)

D im orp hagenia, new genus

Malar space small or absent, mandibles and lower eye margins
nearly in contact; temples narrow, receding; tooth of claws
well separated from outer ray Auplopus Spinola

Key to Neotropical subgenera of Ageniella ( Females )

1. Propodeum with an abundance of erect hair 2

Propodeum without hair or with a few inconspicuous hairs on

the sides 4

2. Mentum at most with a few short, inconspicuous setae; mostly

small species, under 14 mm Ameragenia Banks

Mentum with a number of rather long setae; larger species,
mostly over 15 mm 3

226

Psyche

[September

3. Hind tibiae serrate in profile, also quite strongly spinose; most

species with a prominence on lower part of mesopleurum

Alasagenia Banks

Hind tibiae smooth, non-serrate, and with only small spines;
mesopleurum without a prominence Lissagenitt Banks

4. Front, in lateral view, somewhat angulate (either at the antennal

sockets or between sockets and ocelli), above the angulation
quite flat; mandibles unusually broad, the tooth small (Fig. 6)

Cyrtcigenia, new subgenus
Front, in lateral view, more or less gently rounded; mandibles
not modified as above 5

5. Hind tibia not at all serrate, smooth or with rows of very small

spines Ageniella Banks

Hind tibia serrate in profile 6

6. Brush on inner side of hind tibia with a subapical interruption;

pronotum somewhat elongate N emagenia Banks

Brush on inner side of hind tibia without a subapical interruption

Priophanes Banks

References

Arle, R.

19+7. Nouvelles especes de Pompilidae du Bresil (Hymenoptera) . Rev.
de Ent., 18: 416-428.

Arnold, G.

1934. Psammocharidae of the Ethiopian region. Part 2. Ann. Trans-
vaal Mus., 15: 283-399.

Banks, N.

1946. Studies of South American Psammocharidae. Part 1. Bull. Mus.
Comp. Zool., 96 : 311-525.

Evans, H. E.

1966. A revision of the Mexican and Central American spider wasps
of the subfamily Pompilinae (Hymenoptera: Pompilidae). Mem.
Amer. Ent. Soc., no. 20, 439 pp.

Haupt, H.

1959. Elemente einer systematischen Aufteilung der Macromerinae m.
(Hymenoptera-Sphecoidea). Nova Acta Leopoldina, (9) 21, no.
141, 74 pp.

Townes, H.

1957. Nearctic wasps of the subfamilies Pepsinae and Ceropalinae.
Bull. U.S. Nat. Mus., no. 209, 286 pp.

NOTES ON I-IETEROONOPS AND TRIAERIS
(ARANEAE; OONOPIDAE)

By Arthur M. Chickering
Museum of Comparative Zoology

In this short note are some additions and corrections to my pre-
viously published revisions.

Eleteroonops spinimanus (Simon)

Figures 1-4

Oonops spinimanus Simon, 1891: 563, fig. 6. The female holotype from St.

Vincent, B. W. I. is in the British Museum (Natural History). Simon,

1892: 445; 1893: 294; Petrunkevitch, 1911: 128; 1929: 67, figs. 53-57;

Gertsch, 1936: 8.

Heteroonops spinimanus, — Dalmas, 1916: 203, 217; Bryant, 1940: 205;

Roewer, 1942: 276; Bonnet, 1957: 2185; Chickering, 1969: 154, figs.

28-32.

I have been much interested in Eleteroonops spinimanus (Simon)
for many years. Simon (1891) described the species from females
collected on St. Vincent, B. W. I. In 1892 he reported the species
from Venezuela. Dr. Petrunkevitch, in his study of Puerto Rican
spiders, (1929), stated that he had males and females for study in
the collection of the American Museum of Natural History. These
were collected in 1915 in San Juan, Cayey, Naranjito and Coamo
Springs. He regarded the males, taken in these localities, as belong-
ing with the females and described a male from San Juan as the
male of Oonops spinimanus Simon. This identification has been
widely accepted up to the present time. I have had the specimens
that were apparently studied by Dr. Petrunkevitch also on loan from
the American Museum of Natural History. It has been very dis-
appointing to find almost all of the specimens, believed to be those
studied by Dr. Petrunkevitch, in a very dismembered and almost
useless condition. I think there is no question about the status of the
females involved but I am obliged to regard the identification of the
males as open to serious doubts. Dr. Petrunkevitch noted a con-
siderable degree of variation among the males in respect to the
appearance of the palpal conductor and embolus, indicating, perhaps,
a mixture of species. I have spent much time in searching through
my extensive collection of O'onops /or males which could be matched
with the well established females but without success. I am of the
opinion that the males identified as Oonops spinimanus Simon by

227

228

Psyche

[September

from above. Fig. 2. Right palpal patella; nearly dorsal view. Fig. 3. Right
palpal femur; retrolateral view. Fig. 4. Epigynal area of female from
St. Vincent; viewed from below.

Dr. Petrunkevitch really belong with Oonops castellus Chickering,
now believed to be rather widely distributed among the West Indies.
I readily concede, however, that there is no certainty at the present
time.

Records. In addition to the records cited above the following
should now be added in order to bring the record up to date: Dr.
Gertsch recognized the species from Florida in 1936, where it is
now known to be fairly common (Chickering, 1969). Miss Bryant
(1940) reported the finding of females in Cuba. I have taken many
females in the following localities during my collecting trips in
1954, 1957-1958, 1963-1964, 1965 and 1966: Jamaica, W. I. where
the species seems to be abundant; St. Thomas, St. John and St. Croix,
U. S. Virgin Islands; Puerto Rico, W. I.; St. Lucia, St. Kitts, Nevis
and St. Vincent, all in the British West Indies; Trinidad, W. I.;
Panama Canal Zone and parts of Panama, particularly in the
mountainous regions; and finally in Costa Rica.

Triaeris pusillus (Bryant), new combination

liytanis pusilla Bryant, 1942: 326, figs. 13-14. The female holotype from
St. Croix, V. I. is in the Museum of Comparative Zoology, examined.
Triaeris reticulatus Chickering, 1968: 354, figs. 6-13. The male holotype
from St. Croix, V. I. is in the Museum of Comparative Zoology, new

SYNONYMY.

1973]

Chickering — Heteroonops and Triaeris

229

The female, regarded by Miss Bryant as representing a new
species of Hytanis, was completely overlooked during my study of
the genus Triaeris (1968). At that time I believed that I had a
new species of the genus represented by a male from St. Croix, V. I.
Because of the close similarity of structure and coloration, I believed
that a female from Nevis, B. W. I., belonged with the male from
St. Croix, V. I. As a result of my examination of the holotype of
Hytanis pusilla Bryant I think it is logical to believe that this
female belongs with my male from the same locality and that it
represents a species of Triaeris. This leaves the status of the female
from Nevis in some doubt. This species may be a new one but I
am not yet certain about this. Further careful collecting among the
numerous West Indian Islands is obviously needed.

References

Bonnet, Pierre

1957. Bibliographia Araneorum. Toulouse. 2(3).

Bryant, Elizabeth

1940. Cuban Spiders in the Museum of Comparative Zoology. Bull.

Mus. Comp. Zool. 86(7) : 249-532, 22 pis.

1942. Notes on the spiders of the Virgin Islands. Bull. Mus. Comp.
Zool., 89: 317-363.

Chickering, A. M.

1968. The Genus Triaeris Simon (Araneae, Oonopidae) in Central
America and the West Indies. Psyche, 75 (1): 351-359.

1969. The Family Oonopidae (Araneae) in Florida. Psyche, 76: 144-
162, 41 figs.

Dalmas, Compte de

1916. Revision du Genre Orchestina E. Simon. Ann. Soc. Entom. France,
85: 203-258, 30 figs.

Gertsch, W. J.

1936. Further Diagnoses of New American Spiders. Amer. Mus. Novi-
tates, No. 852: 1-27, 4 pis.

Petrunkevitch, Alexander

1911. A synonymic index-catalogue of spiders of North, Central, South
America, etc. Bull. Amer. Mus. Natur. Hist., 29: 1-809.

1929. The spiders of Porto Rico. Pt. 1. Trans. Connecticut Acad. Arts
and Sci. 30: 7-158, 150 figs.

Roewer, C. Fr.

1942. Katalog der Araneae. 1 : 1-1040.

Simon, E.

1891. On the spiders of the island of St. Vincent. Pt. 1 Proc. Zool.
Soc. of London, Nov. 17, 1891: 549-575.

1892. Voyage de M. E. Simon au Venezuela. Ann. Soc. Entom. France,
61 : 423-462, 1 pi.

1892-1895. Histoire naturelle des Araignees. Deuxieme Edition. 1
Librairie Encyclopedique de Roret, Paris.

THE UTILIZATION OF VARIOUS ASCLEPIAS SPECIES
BY LARVAE OF THE MONARCH BUTTERFLY,
DANA US PLEXIPPUS

By James M. Erickson*

Dept, of Entomology, Cornell University
Ithaca, New York, 14850

Introduction

Plants of the genus Asclepias are well known to contain high
concentrations of cardiac glycosides (cardenolides) . It is generally
surmised that such cardiac stimulants function to protect plants
containing them against insect and vertebrate herbivores (Euw et ah
1967). In addition, some insects which are adapted to feed on
Asclepias plants store cardiac glycosides apparently as a means of
protection against vertebrate predators which find the compounds
distasteful (Brower and Brower 1964, Brower 1969). It was by
means of such an herbivore-predator interaction, bluejays feeding
on monarch butterflies, Danaus plexippus L. (Brower 1969, Brower
et ah 1968), that a spectrum of palatability was detected among
monarch butterflies reared on various species of milkweed. It is
surmised that this spectrum of toxicity to a vertebrate predator reflects
a spectrum of concentrations of cardiac glycosides in the different
species of the insects’ larval food plants (Brower and Brower 1964,
Brower et al. 1967, Brower 1969). However, quantitative and
qualitative data for cardenolides in Asclepias leaves are at best in-
complete (Reynard and Morton 1942, Kupchan et al. 1964, Duffey
1970, Singh and Rastogi 1970, Feir and Suen 1971, Duffey and
Scudder 1972, Scudder and Duffey 1972, and Eggermann and
Bongers 1972).

The following experiments were undertaken to determine
whether a differential response by D. plexippus larvae to their host
plants could be detected by measuring their efficiency of food utiliza-
tion and whether such a response would support the concept of a
spectrum of toxicity. It seemed conceivable that detoxication or
storage of cardiac glycosides might require expenditure of energy
and could be detected by a lowered feeding efficiency. In addition,
Brower et al. (1972) have speculated that there is a reduction in

^Present address: Dept, of Biological Sciences, California State Univer-
sity, Hayward, California 94542.

M anuscript received by the editor June 18, 1973.

230

1973]

Erickson — Danaus plexippus

231

general viability of monarch adults which have stored higher con-
centrations of cardiac glycosides. To test this hypothesis the fertility
and fecundity of adult monarchs was determined.

Methods and Materials

Groups of intact larvae of D. plexippus 1 were taken from the
second generation of a culture founded from wild insects taken near
Ithaca, New York, and were reared in the laboratory on one of the
following species of Asclepias 1 host plants: A. curassavica , A. syriaca,
A. incarnata, or A. tuberosa. Newly molted 4th-instar larvae were
placed individually in glass petri dishes (Pyrex, 100mm X 15mm)
lined on the bottom with a piece of Whatman No. 1 filter paper.
Mature and uninjured leaves of the native species were gathered in
the field each day from plants growing in open sunlit areas, and
leaves of A. curassavica were collected from plants grown in the
greenhouse. All leaves were sealed in plastic bags and used within
2 hours. These randomly collected leaves were split along the mid-
rib, one half weighed and offered to the larvae and the other half
used to determine the percent dry matter in the leaf material
(Waldbauer i960, 1964). Leaves were replaced and feces collected
every 24 hours.

All the larvae were placed in a controlled temperature room,
except for the period of time each day during which new food was
offered to the larvae and the feces collected. The room was kept
relatively constant with day-night temperatures of 22° and 180,
respectively, and with a relative humidity of approximately 55%.
The photoperiod was regulated at a 16-8 hour light-dark cycle.

The dry weight of food ingested was estimated following the
techniques of Waldbauer (i960, 1964), except that plant material
was lyophilized instead of oven-dried. The dry weight of the food
utilized or assimilated was assumed to be the dry weight of the food
ingested minus the dry weight of feces. An additional group of
larvae was reared along with the experimental larvae, and these
were sacrificed to determine the dry weights, and thus, the per-
centage dry matter of the larvae. Indices of food utilization were
determined following the methods of Waldbauer (i960, 1964,
1968). Many terms have been used both by ecologists and by
physiologists to describe various measures and indices of food

Specimens of the insects used in this research have been deposited in the
Cornell University Insect Collection, Lot 1023, Sublot 14. Specimens of the
plants have been deposited in the Bailey Hortorium, Cornell University.

232

Psyche

[September

utilization and efficiency. Relationships between many of these terms
are discussed by Kozlovsky (1968) and Waldbauer (1968).

As an index of digestibility the ratio of the amount of food
assimilated to the amount of food ingested, referred to as the
‘Assimilation Efficiency’ (Odum, 1971) or the ‘Coefficient of Di-
gestibility’ (Waldbauer 1964, 1968, House 1965), was used. In
practice, this measure is only an approximation since the numerator
(as determined by the standard gravimetric technique) does not
quite represent the amount of food actually assimilated (Waldbauer
1968). This slight error is due to the presence of metabolic wastes
in the feces in addition to the undigested food (Lafon 1951), but
Hiratsuka (1920) and Waldbauer (1964, 1968) point out that this
difference between true and measured assimilation efficiencies is
negligible.

The efficiency with which ingested food is converted to biomass
is calculated by dividing the dry weight of food ingested into the
dry weight gained by the larva. This index, referred to by physiolo-
gists as the ‘Efficiency of Conversion of Ingested Matter’ (Wald-
bauer 1968) and by ecologists as the ‘Ecological Growth Efficiency’
(Odum 1971), is an overall measure of an animal’s ability to utilize
for growth the food ingested.

The efficiency with which digested food is converted to biomass is
calculated by dividing the dry weight of food assimilated into the
dry weight gained by the larva. This index, referred to by Wald-
bauer (1968) as the ‘Efficiency of Conversion of Digested Matter’
and by Odum (1971) as the ‘Tissue Growth Efficiency’, decreases
as the proportion of digested food metabolized for energy and main-
tenance of physiological functions increases (Waldbauer 1968).

The relative growth rate is calculated by dividing the mean dry
weight of the larva times the duration of the instar in days into the
dry weight gained by the larva during the stadium (Waldbauer
1968). This index reflects the rate at which biomass is added by a
larva corrected for any size differential between groups of larvae.

The ‘Respiratory Coefficient’ (Lindeman 1942) is described as
the ratio of respiratory and maintenance loss to the net secondary
production or biomass increase. This coefficient is calculated by
dividing the total calories lost through respiration and maintenance
by the total calories added to the insects’ biomass. This ratio is
what may be termed an ‘Energy Production Cost Ratio’ ; the smaller
the coefficient or ratio, the more efficient the larva is at allocating
energy to biomass, the larger the coefficient, the greater the number

1973]

Erickson — Danaus plexippus

233

of calories lost through respiration and maintenance per calorie
allocated to biomass.

Of general interest to ecologists is the ‘Principle of Allocation’
described by Cody (1966). Organisms have a limited amount of
energy to spend and will be selected to partition this energy in dif-
ferent ways depending upon changing physiological or environmental
conditions. Any activity of an organism, or more precisely, the
energy expenditure for that activity, can be viewed only in relation
to all other demands for energy. To descry any increased ‘cost’ of
detoxifying or incorporating cardenolides by D. plexippus larvae
when reared on Asclepias host plants with known toxicity spectrum,
a larval energy budget based on dry weights was constructed. Calo-
rific values of the larval food plants, feces and larvae were deter-
mined by means of a Phillipson non-adiabatic microbomb calorimeter
(Gentry and Wiegert Inst. Inc., Aiken, C.S.) (Phillipson 1964).
The lyophilized plant material was subjected to five replications
whereas lyophilized larvae and feces were subjected to three replica-
tions for the determination of calorific values.

The nitrogen content of the leaf material was determined by the
Kjeldahl method for total nitrogen (Williams 1964). A minimum
of three replicate samples were obtained for each of the plant species.

For the purposes of food utilization and efficiency determinations,
the experiment was concluded when the larvae molted into the
ultimate instar. The larvae were then reared through to the adult
stage on the same experimental plants that they fed upon in the
utilization experiments. All adult females were hand paired and
placed individually in a 1 meter square screen cage with one A.
curassavica and one A. syriaca plant of approximately the same age
and condition. Records were kept each day of the number of eggs
layed per female and the percentage of these eggs which were fertile.

The data are generally presented as a mean and standard error
for the larvae in any particular treatment group. The various
experimental parameters and indices were subjected to one-way
analysis of variance to determine differences in efficiencies or develop-
mental rates among the various larval food plants.

Results

Various plant parameters differed greatly among the four Asclepias
species offered to the monarch larvae (Table 1). The leaves of
A. tuberosa were significantly higher in dry matter content than the
other three species. No significant difference in caloric content could

Table 1. Dry weight, calorific values and nitrogen content of Asclcpias species offered to the larvae of the

234-

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1973]

Erickson — Danaus plexippus

235

be found among the four Asclepias species. The nitrogen content
of the leaf material did not vary significantly for A. curassavica ,
A. syriaca, and A. incar nata but the leaves of A. tuber osa contained
a little more than one half the total nitrogen content of the other
plants examined.

The proportion of ingested food which was digested and as-
similated (‘assimilation efficiency’) averaged about 50% for all
experimental host plants except for A. syriaca which was signifi-
cantly higher (P<0.05) with a mean of approximately 58%.
(Table 2). The efficiency with which ingested matter was con-
verted to biomass ranged from 29.59% for larvae reared on A-
syriaca to 16.39% for larvae reared on A. tuberosa (P<O.Oi)y
whereas the efficiency of conversion of digested matter ranged from
53-3°% for larvae reared on A. curassavica to 34.04% for larvae
reared on A. tuberosa (P<o.6i). The duration of the 4th stadium
also varied significantly (P<o.oi) with larvae on A. curassavica
having the shortest duration (2.15 days) and larvae on A. syriaca
having the longest duration (2.55 days). The amount of biomass
gained per day per larva also varied significantly (P<0.05) with
larvae reared on A. curassavica gaining the most weight (16.42
mg/day) and larvae on A. incarnata the least (12.70 mg/day).
Of great interest, however, is that the relative growth rate did not
vary significantly for the larvae feeding on the four Asclepias species.

The total allocation of energy by the larvae on the different
host plants can be found in Table 3. The larvae reared on A.
tuberosa ingested and assimilated almost twice as many calories as
larvae reared on A. curassavica or A. syriaca. The total number of
calories allocated to larval biomass did not vary to any extent among
the Asclepias species; ingestion of more calories by larvae reared on
A. incarnata and A. tuberosa was balanced by greater costs for res-
piration and maintenance. The ratio of the total number of calories-
allocated to an increase in larval biomass to the total calories
assimilated ranged from approximately 65% by larvae reared on
A. curassavica to approximately 32% for larvae reared on A. tuber-
osa. The respiratory coefficient also ranged from 0.53 (A. curas-
savica) to 2.16 (A. tuberosa).

There was no difference in the mean length of the larval stage or
the pupal stage among larvae reared on the various Asclepias species
(Table 4). Larval survival was 100% on all host plants except
A. incarnata where 90% of fhe larvae survived to the adult stage.
Fecundity and fertility does not vary significantly; the mean number
of eggs layed per female ranged from 103.7 for females reared on

Table 2. The utilization of food by 4th-instar larvae of the monarch butterfly, D. plexippus,

236

[September

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1973]

Erickson — Danaus plexippus

237

A. curassavica to 117.2 for females reared on A. syriaca. The mean
per cent of eggs hatching ranged from 87.3% (A. incarnata ) to
93.2% (A. tuberosa).

Discussion

One of the major concerns of modern ecology is the description
and explanation of the energetic relationships between and within
various communities. A knowledge of the food utilization efficiency
of insects is thus of particular importance to ecology since insects
exert a substantial influence and impact on almost all terrestrial or
fresh water communities. The ecological significance of such energy
utilization studies have been extensively reviewed (Englemann 1966,
Phillipson 1966, and others).

It seems apparent that adaptive nutritional differences in host
plants must be sought on a quantitative level and that meaningful
comparisons of food utilization and nutrition will not emerge until
quantitative studies are carried out. The determination of absolute
requirements for dietary constituents depends upon the measurement
of food or nutrient intake. Differences in food efficiency can be
demonstrated only be measuring intake and growth. Measurement
of the food intake and the utilization of this food elucidates to a
great degree the physiological processes occurring in an insect since
patterns of utilization may be different although food sources are
similar in their ability to support growth. For instance, low food
intake may be offset by a high utilization of ingested or digested food
and a very high food intake may well lead to a very low efficiency
in the utilization of ingested or digested matter.

In this experiment, the assimilation efficiency of the larvae did not
vary significantly among the various Asclepias host plants except for
larvae reared on A. syriaca which had an efficiency about 8% higher
than larvae on the other host plants. This means that the larvae
reared on the various host plants were digesting and excreting ap-
proximately equal amounts of food. The efficiency with which
ingested food and digested food are utilized varied significantly with
larvae reared on A. curassavica and A. syriaca having the highest
efficiencies and larvae reared on A. tuberosa the lowest (Table 2).
The efficiency with which digested food is utilized for growth will
vary not only with the maintenance and respiration requirements for
energy but also with the balance of nutrients in the food source.
Larvae reared on A. tuberosa ingested almost twice as much food
during the 4th instar as larvae reared on the other three host plants

Table 3. The allocation

238

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973]

Erickson — Danaus plexippus

239

(see Table 3). As the ingestion rate increases, there is a negative
correlation with the efficiency with which ingested or digested matter
are utilized (Waldbauer 1968). This reduction in overall gross and
net efficiency has been shown for Prodenia larvae (Soo Hoo and
Fraenkel 1966) and by House (1965). The larvae reared on A.
tuberosa may be viewed as being extremely ‘wasteful’ of the potential
caloric content of their food, most likely passing great quantities of
food through the digestive tract to secure a supply of some limiting
nutrient or substance. The leaf material does not vary to any sig-
nificant degree among the Asclepias species tested in terms of caloric
content (Table 1), whereas the leaves of A. tuberosa contain a little
more than one half the nitrogen content of the other three species.

It has been suggested that the water content of food material will
greatly affect the utilization of this material. It is generally the case
with the swallowtail butterfly, Papilio polyxenes , that the efficiency
of utilization of food decreases as the dry matter content of the food
plant increases (Erickson and Feeny, in preparation). In this ex-
periment, there is a significant difference in the dry matter content
of the leaves, with A. tuberosa having the highest dry matter con-
tent at approximately 21% and A. incar nata and A. syriaca the
lowest dry matter content at about 17% (Table 1). It has been
shown in this laboratory, that by varying the water content of leaf
material, the utilization of food by the cecropia moth, Hyalophora
cecropia is greatly affected (J. M. Scriber, in preparation). The
effect is shown not in an increased intake rate, as occurs in monarch
larvae, but in lengthened larval development times, in which the
total food intake is increased but the rate of this intake remains
relatively constant. It thus appears that the dry matter content of
Asclepias leaves may have some influence on the utilization efficiencies
but not to the degree found in this experiment.

In keeping with Cody’s (1966) ‘Principle of Allocation’, I at-
tempted to descry any increased ‘cost’ for D. plexippus larvae to
detoxify or to incorporate cardenolides. It is well known that A.
curassavica is highly toxic and contains at least 15 cardenolidies
(Kupchan et al. 1964, Duffey 1970), three of which have been
isolated from distasteful D. plexippus larvae (Parsons 1965, Reich-
stein 1967) incorporated there as a defense against vertebrate preda-
tors (Brower and Brower 1964, Brower et al. 1967, Brower 1969).
Asclepias syriaca has been found to be only slightly toxic to verte-
brates and to contain at least seven cardiac glycosides (Reynard and
Morton 1942, Duffey 1970, and others). Asclepias incarnata and
A. tuberosa are known to contain cardenolides but in much lower

240

Psyche

[September

Table 4. Length of pupation, fecundity and fertility of monarch butterflies
raised on various plants of the genus Asclepias.

Host plant

Number of
animals

Mean length
of pupation
(days)

Mean total
number of eggs
per female
± SE12

Mean percent
of eggs
hatching
± SE13

A. curassavica

12

10.31

105.7 ± 8.7

91.6 ± 4.9

A. syria ca

17

10.67

117.2 ± 10.6

89.1 ± 5.7

A. incarnata

12

11.02

110.3 ± 9.4

87.3 ± 6.3

A. tuberosa

9

10.57

106.0 ± 9.6

93.2 ± 4.8

T.os (df 3,20) = 3.10
F.oi (df 3,20) =4.94
2F = 1.13
3F = 0.73

concentrations ( Duffey 1970, Singh and Rastogi 1970) and only
marginally toxic (Hansen 1924, Heal et al. 1950). It is found,
however, that contrary to expected results, larvae reared on the
highly toxic A. curcissavica gained more biomass per day, spent the
shortest time in the 4th instar, and were the most efficient at utilizing
and converting digested matter into biomass (Table 2). In addition,
larvae reared on A. curassavica utilized approximately 65% of the
assimilated energy to produce biomass whereas only 32% of the
assimilated energy was allocated to biomass for larvae reared on
A. tuberosa (Table 3), and larvae reared on A. curassavica had the
lowest ‘Respiratory Coefficient’ at 0.53 compared to larvae reared on
A. tuberosa which had a value of 2.16. This means that larvae
reared on the A. curassavica host plant were allocating about 2
calories for biomass for every calorie lost through respiration and
maintenance, wJhereas larvae reared on A. tuberosa allocated ap-
proximately 0.5 calorie to biomass for every calorie lost through
respiration and maintenance. It thus appears that there is little
measurable ‘cost’ to detoxify or incorporate caridac glycosides by
monarch larvae since the larvae grew and developed most rapidly
on the most toxic Asclepias food plant tested.

1973]

Erickson — Danaus plexippus

241

Recently, Brower et al. (1972) have shown a general decrease in
cardiac glycoside content of migrating monarch butterflies as the
specimens are collected farther south. They feel selection may be
operating against high cardiac glycoside content (larvae reared on
A. curassavica or A. humistrata ) since, although high concentrations
of cardiac glycosides in the butterfly confer greater protection from
predators, these authors surmise that these high concentrations de-
crease the viability of the insect. In this experiment, there was no
significant difference in either the number of eggs deposited or the
number of these that were fertile among the adult females reared
on the four Asclepias species (Table 4). It does not appear at least
over a couple of generations, that larvae reared on A. curassavica are
any less viable than larvae reared on much less toxic Asclepias species.

The importance of nitrogen for larval growth and development
cannot be overemphasized. House (1961, 1962) and Dadd (1973)
have discussed the qualitative requirements of proteins and amino
acids for larval development. There appears to be an optimal nitro-
gen level, which varies from species to species, that produces maximal
larval growth (Dadd 1961, House 1959, Vanderzant 1958). The
fecundity of Dacus dorsalis Hendel increased with an increase in the
protein content of the diet (Hagen 1958), whereas low protein
levels greatly prolonged the developmental period of Drosophila
melanogaster (Sang 1956). In this experiment, larvae reared on
A. tuberosa had the second fastest developmental time of the 4
groups of larvae, yet this plant species contained a little more than
one half the nitrogen content in the leaves than was contained in
the other leaves of the other host plants. Larvae of the monarch
butterfly appear, therefore, to have an adaptive strategy which al-
lows them to best utilize a resource that is in limited supply. Al-
though ‘wasteful’ in the caloric sense, these larvae are able to secure
the necessary supply of nitrogen needed for the later adult stage by
increasing their total intake of food (Table 3). A similar situation
has been demonstrated in this laboratory involving the utilization of
crucifer plants by Pieris rapae (Slansky and Feeny, in preparation).
This ability of an insect to compensate for decreased nutrient con-
tent of its food is discussed by House (1965) and McGinnis and
Lasting (1966). It thus appears that the low nitrogen content of
A. tuberosa has a somewhat limiting effect on the monarch larvae
but this low nutrient content only limits or regulates the efficiency
with which food is utilized by the larvae and does not limit the
larval growth or consumption rates to a significant degree.

^42

Psyche

[September

Acknowledgments

The author wishes to thank Drs. L. P. Brower, P. P. Feeny and
J. G. Franc! emont for reading the manuscript and offering helpful
suggestions. Many thanks also go to Ms. Sherry Rehr for technical
help and also offering suggestions on the manuscript. The work was
supported in part by Hatch Grant NY (C) 1 3941 3 of Dr. Paul
Feeny.

Literature Cited

Brower, L. P.

1969. Ecological chemistry. Sci. Amer. 220: 22-29.

Brower, L. P., and J. V. Z. Brower.

1964. Birds, butterflies, and plant poisons: A study of ecological chem-
istry. Zoologica 49: 137-159.

Browter, L. P., J. V. Z. Brower, and J. M. Corvino.

1967. Plant poisons in a terrestrial food chain. Proc. Nat. Acad. Sci.
57: 893-898.

Brower, L. P., W. N. Ryerson, L. L. Coppinger, and S. C. Glazier.

1968. Ecological chemistry and the palatability spectrum. Science 161:
1349-1352.

Brower, L. P., P. B. McEvoy, K. L. Williamson, and M. A. Flannery.

1972. Variation in cardiac glycoside content of monarch butterflies from
natural populations in Eastern North America. Science 117:
426-428.

Cody, M. L.

1966. A general theory of clutch size. Evolution 20: 174-184.

Dadd, R. H.

1961. The nutritional requirements of locusts. V. Observations on es-
sential fatty acids, chlorophyll, nutritional salt mixtures and the
protein or amino acid components of synthetic diets. J. Insect
Physiol. 6: 126-145.

1973. Insect nutrition: Current developments and metabolic implica-
tions. Ann. Rev. Entomol. 18: 381-420.

Duffey, S. S.

1970. Cardiac glycosides and distastefulness: some observations on pal-
atability spectrum of butterflies. Science 169: 78-79.

Duffey, S. S., and G. G. E. Scudder.

1972. Cardiac glycosides in North American Asclepiadaceae, a basis
for unpalatability in brightly colored Hemiptera and Coleoptera.
J. Insect Physiol. 18: 63-78.

Eggermann, W., and J. Bongers

1972. Die Wirtswahl von Oncopeltus fasciatus Dali. (Heteroptera :
Lygaeidae) : Bindung an Asciepiadaceen durch wirtsspezifische
Glykoside. Oecologia (Berl.) 9: 363-370.

Englemann, M. D.

1966. Energetics, terrestrial field studies, and animal productivity. IN:
Advances in Ecological Research, ed. J. B. Cragg. Academic
Press, London and New York. Vol. 3, pp. 73-115.

1973]

Erickson — - Danaus plexippus

243

Euw, J. V., L. Fishelson, J. A. Parsons, T. Reichstein, and M. Rothschild.

1967. Cardenolides (heart poisons) in a grasshopper feeding on milk-
weeds. Nature 214: 35-39.

Feir, D., and J. S. Suen.

1971. Cardenolides in the milkweed plant and feeding by the milkweed
bug. Ann. ent. Soc. Amer. 64: 1173-1174.

Hansen, A. A.

1924. The poison plant situation in Indiana. J. Amer. Vet. Med.
Assoc. 66 : 351.

Heal, R. E., E. F. Rogers, R. T. Wallace, and O'. Starnes.

1950. A survey of plants for insecticidal activity. Lloydia 13: 89-162.

Hiratsuka, E.

1920. Researches on the nutrition of the silkworm. Bull. ser. Exp. Sta.
Japan, 1: 257-315.

House, H. L.

1959. Nutrition of the parasitoid Pseudosarcophaga affinis (Fall.) and
of other insects. Ann. N.Y. Acad. Sci. 77: 394-405.

1961. Insect nutrition. Annu. Rev. Entomol. 6: 13-26.

1962. Insect nutrition. Annu. Rev. Biochem. 31: 653-672.

1965. Effects of low levels of the nutrient content of a food and of
nutrient imbalance on the feeding and the nutrition of a phyto-
phagous larva, Celero euphorbiae (L.) (Lepidoptera : Sphingidae).
Can. Ent. 97 : 62-68.

Kozlovsky, D. G.

1968. A critical evaluation of the trophic level concept. I. Ecological
efficiencies. Ecology 49: 48-60.

Kupchan, S. M., J. R. Knox, J. E. Kelsey, and J. A. Saenz Renauld.

1964. Calotropin, a cytotoxic principle isolated from Asclepias curas-
savica L. Science 146: 1685-1686.

Lafon, M.

1951. Quelques documents sur l’appetit et la consommation alimentaire
chez les insectes. Ann. Nutr., Paris, 5: 485-504.

Lindeman, R. L.

1942. The trophic-dynamic aspect of ecology. Ecology 23 : 399-418.

McGinnis, A. J., and R. Kasting.

1966. Comparison of tissues from solid- and hollow-stemmed spring
wheats during growth. IV. Apparent dry matter utilization and
nitrogen balance in the two-striped grasshopper, M clanoplus
bivittatus (Say). J. Insect Physiol. 12: 671-678.

Odum, E. P.

1971. Fundamentals of ecology. W. B. Saunders Company, Philadel-
phia, Pennsylvania. 3rd edition, 574 p.

Phillipson, J.

1964. A miniature bomb calorimeter for small biological samples.
Oikos 15: 130-139.

1966. Ecological energetics. William Clowes and Sons Ltd., London.
57 p.

Reynard, G. B., and J. B. S. Norton.

1942. Poisonous plants of Maryland in relation to livestock. Maryland
Agr. Exp. Sta. Tech. Bull. A 10.

244

Psyche

[September

Scudder, G. G. E., and S. S. Duffey.

1972. Cardiac glycosides in the Lygaeinae (Hemiptera: Lygaeidae).
Can. J. Zool. 50: 35-42.

Singh, B. and R. P. Rastogi.

1970. Cardenolides — Glycosides and Genins. Phytochemistry 9: 315-331.

SooHoo, C. F. and G. Fraenkel.

1966. The consumption, digestion, and utilization of food plants by a
polyphagous insect, Prodenia eridania (Cramer). J. Insect
Physiol. 12: 711-730.

Vanderzant, E. S.

1958. The amino acid requirements of the pink bollworm. J. econ. Ent.
51: 309-311.

Waldbauer, G. P.

1960. Feeding and growth on solanaceous and non-solanaceous plants
by normal and maxillectomized larvae of the tobacco hornworm,
Protoparce sexta (Johan.) (Lepidoptera : Sphingidae). Ph.D.
thesis. University of Illinois, Urbana, Illinois. 134 p.

1964. The consumption, digestion, and utilization of solanaceous and
non-solanaceous plants by larvae of the tobacco hornworm, Proto-
parce sexta (Johan.) (Lepidoptera: Sphingidae). Ent. exp. &
appl. 7: 253-269.

1968. The consumption and utilization of food by insects. Adv. Insect
Physiol. 3 : 229-282.

Williams, P. C.

1964. The colorimetric determination of total nitrogen in feeding stuffs.
Analyst 89: 276-281.

COPULATORY PATTERN SUPPORTS GENERIC
PLACEMENT OF SCHIZOCOSA A VIDA
( WALCKENAER ) (ARANEAE: LYCOS ID AE)1

By Jerome S. Rovner

Department of Zoology, Ohio University, Athens, Ohio 45701

The wolf spider genus Schizooosa was established in 1904 by
Chamberlin. He subsequently (1908) transferred several species
(bilineata, crassipes, and saltatrix ) from their former placement in
the genus Lycosa to the genus Schizocosa. Such a transfer was also
recommended for Lycosa avida Walckenaer by Gertsch and Wallace
(r937)* Their designation was accepted by some workers (e.g.,
Fitch, 1963; Dondale, 1969) but not by all (e.g., Kaston, 1948,
1972).

In a recent paper (Rovner, in press) I noted that the pattern of
palpal insertions during mating in the species of Schizocosa studied
so far is qualitatively distinct from that seen in Lycosa spp. Through-
out most of the copulation in Schizocosa spp., one palp is inserted a
number of times prior to each shift to the opposite palp. At the
beginning of copulation the number of insertions in a series by each
palp is relatively small. The number of insertions per series soon
reaches a maximum and then, for the remainder of the copulation,
gradually declines. In Lycosa spp., on the other hand, alternation of
palps typifies the entire copulation. Re-insertions of the same palp
are uncommon, constituting only a small percentage of the total
number of insertions (Rovner, 1972).

Quantitative differences in copulatory behavior also may aid in
characterizing these two genera. The duration of copulation and the
total number of palpal insertions are much greater in Schizocosa
saltatrix than in Lycosa spp. (Rovner, in press) .

With these parameters in mind, I observed mating behavior in
Schizocosa avida. The data obtained supported Gertsch and Wal-
lace’s (1937) reclassification of this species from Lycosa to Schizo-
cosa.

Methods

Penultimate individuals of S. avida were collected during early
May, 1973, in a field near Amesville (Athens Co.), Ohio, USA.

JThis study was supported in part by grant no. GB 35369 from the
National Science Foundation.

Manuscript received by the editor July 23, 1973

245

246

Psyche

[September

Molting to the adult instar occurred during early June, and ob-
servations were made during late June and early July. Spiders were
housed separately and visually isolated from each other. They were
fed mealworms (larvae of Tenebrio rnolitor ) weekly and had a
constant water supply.

Virgin spiders were paired in arenas for observation under fairly
constant temperature (23-25°C) and humidity (60-65% RH) con-
ditions. After observing one pair in a preliminary trial, I filmed
portions of copulation in a second pair. The palpal insertion patterns
of five other pairs were recorded for later analysis.

Results

Copulatory behavior was similar in all seven pairs of S. avida
observed. Throughout most of copulation, each palp was inserted
repeatedly a number of times prior to a shift to the opposite palp.
The length of such series showed the same pattern of variation in
nearly all cases : few insertions per series at the beginning, increasing
shortly thereafter to a maximum number of insertions per series,
then a gradual decrease, eventually ending with a minimal number
of palpal insertions. (In one male, insertions of the left palp began
at the maximum series length; i.e., several “introductory” series of
relatively short length were not shown by this male’s left palp, al-
though they did occur in the right palp.)

In the five recorded matings, each palp was inserted an average of
158 times, for a mean total of 316 insertions per copulation. These
copulations lasted 2.8 ± 1.63 (SD) hours. Other data are sum-
marized in Table I.

Table I. Palpal insertions during copulation in five pairs of Schizocosa
avida.

Total no.

Total no.

Insertions per series

Pair no.

of series

of insertions

Mean ± SD

Range

1

21

241

11.5 ± 14.7

2-60

2

11

254

23.1 ± 9.4

4-35

3

31

213

6.9 ± 5.0

1-21

4

34

371

10.9 ± 10.6

1-48

5

37

501

13.5 ± 10.4

1-39

Mean

26.8

316.0

13.2 ± 15.1

1973]

Rovner — Schizocosa avida

247

Discussion

The copulatory pattern observed in Schizocosa avida was similar
to that described in S. bilineata and S. crassipes (Montgomery, 1903)
and S. saltatrix (Rovner, in press). The occurrence of a series of
insertions of one palp prior to a shift to the other palp differs from
the rather strict alternation of palps seen in Lycosa chaperi (Bhat-
nagar and Sadana, 1965), L. gulosa (Kaston, 1936), L. helluo
(Kaston, 1936; Nappi, 1965), L. punctulata (Rovner, unpublished
data), and L. rabida (Montgomery, 1903; Kaston, 1936; Rovner,
1972). Furthermore, the pattern of variation in series length, which
seems typical of Schizocosa spp. (Rovner, in press), was also present
in S. avida. Thus, Gertsch and Wallace’s (1937) placement of the
former Lycosa avida into the genus Schizocosa on the basis of
morphology has been strengthened by these behavioral data. The
large number of palpal insertions (about 300) and the relatively
long duration of copulation (about 2-3 hours) in S. avida also indi-
cate a greater affinity to Schizocosa spp. than to Lycosa spp. (Rov-
ner, in press).

Another aspect of behavior may be worth investigating in this
regard. Knost and Rovner (in prep.) found that post-immobilization
tying down of prey occurred in L. punctulata and L. rabida but not
in S. crassipes. Whether this distinction is -genus-specific or not must
await studies of feeding behavior in other members of these two
genera.

Literature Cited

Bhatnagar, R. D. S. and G. L. Sadana

1965. Sexual behaviour in Lycosa chaperi Simon (Arachnida: Ara-
neida). J. Bombay Nat. Hist. Soc. 62: 568-573.

Chamberlin, R. V.

1904. Notes on generic characters in the Lycosidae. Canad. Ent. 36:
173-178.

1908. Revision of North American spiders of the family Lycosidae.
Proc. Acad. Nat. Sci. Philad. 60: 158-318.

Dondale, C. D.

1969. Two new species of the spider genus Schizocosa (Araneida:
Lycosidae) from the Great Lakes region. Canad. J. Zool. 47:
751-758.

Fitch, H. S.

1963. Spiders of the University of Kansas Natural History Reservation
and Rockefeller Experimental Tract. Misc. Publ. Mus. Nat.
Hist. U. Kansas No. 33. 202 pp.

Gertsch, W. J. and H. K. Wallace

1937. New American Lycosidae with notes on other species. Amer.
Mus. Nov. No. 919: 1-22.

248

Psyche

[September

Kaston, B. J.

1936. The senses involved in the courtship of some vagabond spiders.
Entomol. Amer., n. s. 16: 97-167.

1948. Spiders of Connecticut. Bull. State Geol. Nat. Hist. Surv. Hart-
ford No. 70. 874 pp.

1972. How to Know the Spiders. Wm. C. Brown Co., Dubuque, Iowa.
289 pp.

Knost, S. J. and J. S. Rovner

In prep. Feeding behavior in wolf spiders (Araneae: Lycosidae) :
multiple-prey capture and tying-down.

Montgomery, T. H., Jr.

1903. Studies on the habits of spiders, particularly those of the mating
period. Proc. Acad. Nat. Sci. Philad. 55: 59-149.

Nappi, A. J.

1965. Notes on the courtship and mating habits of the wolf spider
Lycosa helluo Walckenaer. Amer. Midi. Nat 74: 368-373.
Rovner, J. S.

1972. Copulation in the lycosid spider Lycosa rahida Walckenaer: a
quantitative study. Anim. Behav. 20: 133-138.

In press. Copulation in the lycosid spider Schizocosa saltatrix (Hentz) :
an analysis of palpal insertion patterns. Anim. Behav.

THE MALE GENITALIA OF BLATTARIA.

X. BLABERIDAE. PYCNOSCELUS , STILPNOBLATTA ,
PROSCRATEA (PYCNOSCELINAE), AND
DIPLOPTERA (DIPLOPTERINAE) *

By Louis M. Roth
Pioneering Research Laboratory
U.S. Army Natick Laboratories
Natick, Massachusetts 01760

McKittrick (1964) grouped the Pycnoscelinae, Diplopterinae,
Panchlorinae, and Oxyhaloinae, in the Panchloroid Complex of
Blaberidae. The male genitalia of the latter two subfamilies have
been described (Roth, 1971a, 1971b). In this paper I shall illustrate
the male genitalia of several species of Pycnoscelinae and two species
of Diplopterinae.

Materials and Methods

The genitalia were treated with 10% KOH and mounted in
Permount. The source of each of the specimens illustrated is given
using the following abbreviations: (ANSP) — Academy of Natural
Sciences, Philadelphia; (BMNH) = British Museum (Natural
History); (L) = Zoological Institute, Lund, Sweden; (MCZ) =
Museum of 'Comparative Zoology, Harvard University; (VM) —
Vienna Museum Natural History, Vienna, Austria. Geographical
collection data and the names of specialists who identified the speci-
mens, if known, follow these abbreviations. The number preceding
the abbreviation refers to the number assigned to the specimen and
its corresponding genitalia (on a slide) which are deposited in their
respective museums.

Results and Discussion
Pycnoscelinae

Pycnoscelus surinamensis (Linn.) is the type species but it is
parthenogenetic and normally only exists as females. P. indicus
(Fab.) is bisexual and apparently the parent stock from which
surinamensis arose. Occasionally parthenogenetic males occur in
cultures of surinamensis but they are non-functional when mated to
parthenogenetic females (Roth, 1967).

* Manuscript received by the editor August 27, 1973

249

250

Psyche

[September

1973]

Roth — Blattaria

251

The arrangement of the male genital phallomeres is shown in
figure 1. In Pycnoscelus indicus all 3 phallomeres are well devel-
oped and sclerotized (Figs. 7-9). The L2d is separated from L2vm;
the obliquely more or less truncate ends of these 2 sclerotized struc-
tures (Figs. 7, 10, 21, 24, 31) have the appearance of having been
broken off and separated from L2vm. The outer lower curved por-
tion of L2d is spicular (Figs. 1, 7, 10-13), and the underlying
prepuce is densely “hairy” but otherwise not unusually shaped
(Fig. 7, P). The curved genital hook (R2) lacks a subapical
incision, is heavily sclerotized, somewhat truncate at the apex, the
inner curved margin with (Figs. 8, 14-18) or without (Figs. 19-20)
small projections. The Li is very well developed with the cleft
turned upward and its margins heavily sclerotized (Figs. 1, 9). The
genitalia of Pycnoscelus surinctmensis (Figs. 21-25), and Pyncoscelus
nigra (Brunner) (Figs. 31-33) are indistinguishable from those of
P. indicus. Habitus photographs of P. indicus and P. nigra are shown
in figures 2 and 4.

The shapes of the L2d and R2 (Figs. 26, 27, 29, 30) of Pycnos-
celus semivitreus Princis (Fig. 3) differ from those structures in
P. indicus ; P. surinamensis , and P. nigra; however, the shape of Li
in all 4 of the above species is similar (cf. Figs. 9, 23, 28, 33 (dis-
torted in preparation)). Two of the genital phallomeres of Pycnos-
celus striata (Kirby) are distinguishable from those of the other
species of the genus. Its I>2d (Fig. 34) differs in shape, lacks the
spicular surface characteristic of the outer lower curved region and
is only slightly separated from L2vm (cf. Fig. 31). The curved
portion of R2 (Fig. 35) of striata is more elongate and slender than
in semivitreus (Figs. 27, 29) and more uniform in width than in
P. indicus (Figs. 14-20) or nigra (Fig. 32).

Princis (1964) included Stilpnohlatta in the Pycnoscelidae and
the genitalia of S. opaca (Walker) (Figs. 37-39) tends to support
this conclusion, though I relegate his family to subfamily rank
(McKittrick, 1964). Especially notable is the marked similarity in
appearance of the Li of Stilpnohlatta (Fig. 39) with those of
Pycnoscelus (Figs. 23, 36). The L2d (Fig. 37) of S. opaca is
greatly reduced and irregular in outline and is widely separated from

Fig. 1. Male genitalia (dorsal view). Top. (106 MCZ). Pycnoscelus
indicus. Zamboanga, Philippine Islands, (det. Roth). Bottom. (70 BMNH).
Proscratea complanata. Sao Gabriel, Rio Negro, Brazil, 27.IX.1927, J. F„
Zikan. (LI — first sclerite of left phallomere; L2d — dorsal sclerite of L2 'r
L2vm — ventromedial sclerite; M — saclike membrane above L2d; R2 —
hooked sclerite of right phallomere).

252

Psyche

[September

Figs. 2-6. Males of Pycnoscelinae. 2. Pycnoscelus indicus. Sakaerat
District, Thailand. From a laboratory colony. 3. (100 MCZ). Pycnoscelus
semivitreus. Manila, Philippine Islands, (det. Princis). This species was
described (Princis, 1967, p. 148) from 2 $ $ rom Java, Depok. 4. (25
BMNH). Pycnoscelus nigra. Southwest China, Yunnan, Ho-an (leg. J. W.
Gregory, 26.Y.1922). 5. (1447 L). Pycnoscelus striata. Kariorang, Borneo,
(det. Princis). 6. (144 ANSP). Stilpnoblatta opaca. Butawa, Modera, S. P.
Ceylon (det. by Hebard as S. bengalensis (Sauss)). (scale — 5 mm).

1973]

Roth — Blattaria

253

Figs. 7-20. Male genital phallomeres of Pycnoscelus indicus. 7-9. L2d,
R2, and Ll from a specimen originating from Hawaii (laboratory culture).
P = prepuce. 10-13. L2d’s. 10. Sakaerat District, Thailand (laboratory
culture). 11. (101 MCZ). Pasay, Philippine Islands. 12-13. Hawaii (lab-
oratory culture). 14-20. R2’s. 14. Sakaerat District, Thailand (laboratory

culture). 15-19. Hawaii (laboratory culture). 20. (101 MCZ). Pasay,

Philippine Islands, (scale — 0.2 mm; figures 10-13 magnified to scale shown
in fig. 7; figures 14-20 magnified to scale shown in fig. 8).

254

Psyche

[September

Figs. 21-30. Male genital phallomeres of Pycnoscelus spp. 21-25. P. surl-
namensis. Two males which occurred in a culture originating from Fraser
Island, Australia. 26-30. P. semivitreus. 26-28. (102 MCZ). Manila, Philip-
pine Islands (det. Princis). 29-30. (100 MCZ). From $ shown in figure 3.
(scale — 0.2 mm; figures 25, 27, 29 magnified to scale shown in fig. 22).

1973]

Roth — Blattaria

255

Figs. 31-39. Male genital phallomeres of Pycnoscelinae. 31-33. (25

BMNH). Pycnoscelus nigra ; LI (Fig. 33) was distorted in preparation,
(from 3 shown in figure 4). 34-36. (1447 L). Pycnoscelus striata (from
$ shown in figure 5). 37-39. (144 ANSP). Stilpnoblatta opaca (from $
shown in figure 6). (scale — 0.2 mm).

256

Psyche

[September

L2vm. The R.2 (Fig. 38) is broad, relatively short and, as in
Pycnoscelus , lacks a subapical incision.

Princis (1964) lists 4 valid species of Proscratea: peruana Sauss.,
inequalis (Walker), funebris Burmeister, and complanata (Perty).
Brunner (1865) synonymized P. peruana with P. complanata with, a
(?) and Kirby (1904) listed them as synonyms. Hebard (1926)
placed funebris as a synonym of complanata , but Rehn (1932) felt
that this was not warranted until additional information became
available. Princis (1963) concluded that the above synonymies were
incorrect, and showed differences in pronotal shapes and color mark-
ings of the 4 species. I have examined the type of P. inequalis and
find that its genitalia are so different from those of P. complanata
(type of genus) that it undoubtedly does not belong to this genus.

I collected P. complanata (det. by Gurney) in Brazil and estab-
lished a colony which was maintained for several years at the Natick
Laboratories. Habitus figures of adults and a nymph (from the cul-
ture) are shown in figures 40-44. The adult pronotal markings may
vary (Figs. 40-42) (see Rehn, 1932, p. 71) and the pattern of the
specimen shown in figure 42 resembles that shown by Princis (1963,
p. 148) for funebris. The specimen provisionally determined by Rehn
as peruana has pronotal markings (Fig. 45) similar to complanata.

Princis (1964) placed Panchlora, Proscratea, and Phortioecoides
in the Panchloridae. Male genitalia clearly place Phortioecoides in
the Zetoborinae (Blaberidae) (Roth, 1970a). The male phallomeres
of the Panchlorinae (5 genera) are notable for their marked reduc-
tion or absence (Roth, 1971b, Gurney and Roth, 1972). The geni-
talia of Proscratea are not characteristic of the Panchlorinae.

Hebard (1926) and Rehn (1932) believed that Proscratea be-
longed to that section of the Perisphaeriinae which included Para-
nauphoeta Brunner and its allies. I have examined the male genitalia
of 6 species of Paranauphoeta and all 3 phallomeres differ markedly
from those of Proscratea.

McKittrick (personal communication) placed Proscratea in the
Diplopterinae. However, the shape of the L2d and R2 in Proscratea
are more like those of Pycnoscelus and I tentatively place Proscratea
in the Pycnoscelinae. McKittrick (1964) shows Diploptera , Leuro-
lestes, and Pycnoscelus , as arising from a common stock. Phoetalia
(= Leurolestes) which she placed with Diploptera in the Diplop-
terinae belongs to the Blaberinae (Roth, 1970b). The shape of the
spermatophore of Proscratea complanata looks like a bowling pin and
strongly resembles the spermatophore of Diploptera suggesting a
relationship between these 2 genera. However, other genera in

1973]

Roth — Blattaria

257

Figs. 40-45. Proscratea spp. 40-44. P. complanata. From laboratory colony
which originated from Serra Tamendaui, Rio Negro, Amazonas. 40. Bra-
chypterous male. 41-42. Macropterous males. 43. Brachypterous female.
44. Female nymph. 45. (129 ANSP). P. “ peruana ” $, Hacienda San Juan,
Colonio Perene, Peru, June 23, 1920; Cornell Univ. Exped. (det. Rehn).
(scale = 5 mm; scale for figures 40-44 shown in fig. 43).

Figs. 46-57. Male genital phallomeres of Proscratea spp. 46-48. (129

ANSP). P. “ peruana ” (from $ shown in figure 45). 49-57. P. complanata.
49-51. Tapurucuara, Rio Negro, Amazonas. 53, 56. Urucurutuba, Rio
Madeira, Amazonas. 54, 57. Serra Tamendaui, Rio Negro, Amazonas,
(scale — 0.2 mm).

1973]

Roth — Blattaria

259

different subfamilies (e.g., Capucina [Zetoborinae,] Nauphoeta and
Gromphadorhina [Oxyhaloinae] ) have spermatophores ( Graves,
1969) somewhat similar in shape to those of Diploptera and Pros -
crated.

The L2d of Proscratea is well developed, somewhat crescent-
shaped (but variable) and widely separated from L2vm (Figs. 46,
49, 52-54). The membrane above L2d is modified to form a sac-like
projection whose surface is covered with microspicules; this structure
is not found in the other Pycnoscelinae. The R2 is short, stout, and
lacks a subapical incision (Figs. 47, 50, 5 5_5 7 ) * The shape of the
cleft in the Li of Proscratea does not curve upwards (Figs. 48, 51)
as it does in Pycnoscelus (Fig. 9) or Stilpnoblatta (Fig. 39). The
genitalia of Proscratea oomplanata (Figs. 46-48) are indistinguishable
from the specimen provisionally determined by Rehn (1932, pp. 71-
72 ) to be Proscratea peruana ( Figs. 49-5 1 ) .

Because of the differences between the Li and the presence of the
modified membrane over the L2d of Proscratea , I suggest 2 tribes in
this subfamily:

1. Pycnoscelini : Pycnoscelus , Stilpnoblatta

2. Proscrateini : Proscratea

Diplopterinae

Princis (1965) placed the genera Diploptera and Diplopterina in
the Diplopteridae. The genitalia of Diplopterina are closer to cer-
tain members of the Perisphaeriinae (unpublished observations) and
I consider it to be a member of this subfamily. McKittrick (1964)
included 2 genera, Diploptera and Phoetalia {— Leurolestes) in
Diplopterinae. As indicated above, the genitalia of Phoetalia place
it in the Blaberinae (Roth, 1970b). Diploptera punctata (Esch-
sdholtz) is the only viviparous cockroach known and, at present, I
consider this genus to be the only member of the Diplopterinae.
Other members of the Blaberidae, whose reproduction has been in-
vestigated, are ovoviviparous. Princis (1965) lists 7 species of
Diploptera and it would be of interest to determine if those, other
than punctata, are viviparous also.

The arrangement of the phallomeres of Diploptera minor Brunner
is shown in figure 78. The male genital phallomeres of D. punctata
(type of genus) are shown in figures 60-62. The curved hook (R2)
lacks a subapical incision, is more slender and elongate, and more
strongly curved (Fig. 61) than in most of the species of Pycnos-
celinae. The inner margin of the curved portion of the hook has

26o

Psyche

[September

Figs. 58-67. Supra-anal and subgenital plates, and genital phallomeres of
male Diploptera spp. 58-62. D. punctata (from laboratory colony originating
in Hawaii). 58. Supra-anal plate (dorsal). 59. Subgenital plate (ventral).
60-62. Genital phallomeres. 63-67. (67 BMNH). D. sp., Honolulu, Hawaii.
63. Supra-anal plate (dorsal). 64. Subgenital plate (ventral). 65-67. Genital
phallomeres. (scale: supra-anal and subgenital plates — 0.5 mm; phallo-
meres — 0.2 mm) .

1973]

Roth — Blattaria

261

Figs. 68-77. Supra-anal and subgenital plates, and genital phallomeres of
Diploptera spp. 68-72. (68 BMNH). Samoa Island. 68. Supra-anal plate

(dorsal). 69. Subgenital plate (ventral). 70-72. Genital phallomeres. 73-
77. (69 BMNH). Henderson Island, South Pacific. 73. Supra-anal plate
(dorsal). 74. Subgenital plate (ventral). 75-77. Genital phallomeres.
(scale: supra-anal and subgenital plates = 0.5 mm; phallomeres — 0.2 mm).

Figs. 78-79. (17 VM). Diploptera minor. Type $, Philippines. 78.

Genitalia and subgenital plate (SP) (ventral). P = prepuce; other abbre-
viations as in figure 1. 79. Supra-anal plate (dorsal). The terminal seg-
ments of the cerci were missing. The paraprocts are visible beneath the
supra-anal plate in the cleared specimen, (scale — 0.5 mm).

Psyche

L2vm

1973]

Roth — Blattaria

263

minute undulating irregularities reminiscent of some specimens of
P. indicus (cf. Fig. 61 with Figs. 14-15, 18). The L2d is apparently
represented by a small sclerotized region of the prepuce widely sep-
arated from the apex of L2vm (Fig. 60). The vertical downward
curvature of the cleft and indentation on the outer margin of the
lower lobe of Li (Fig. 62) is unique for this phallomere in the
Blaberidae; the lower lobe of Li lacks setae.

The supra-anal and subgenital plates of D. punctata are shown in
figures 58-59. The genital phallomeres of 3 other undetermined
specimens of Diploptera (Figs. 65-67, 70-72, 75-77) are indistinguish-
able from D. punctata. The subgenital plates of these specimens
(Figs. 64, 69, 74) are similar to D. punctata (Fig. 59), but the
shapes of the supra-anal plate of 2 of the specimens differ somewhat.
In the specimen shown in figure 63, there is a deep invagination on
the posterior margin ; however because of the slight asymmetry of the
indentation this may be an aberrant punctata. This specimen is from
Hawaii and although D. punctata is widespread in the Pacific it is
the only species of the genus recorded from Hawaii (Princis, 1965).
Except for size, the phallomeres (fig. 78) of D. mimr are almost
indistinguishable from D. punctata; the sclerotization (L2d) of the
prepuce found in D. punctata is absent in minor. The supra-anal
plate of D. minor is slightly more rounded (fig. 79) than that of
punctata (fig. 58).

Acknowledgements

The writer thanks Dr. David Rentz, Academy of Natural Sciences,
Philadelphia; Dr. David Ragge, British Museum (Natural History),
London; Dr. Karl Princis, Zoological Institute, Lund, Sweden; Dr.
Howard Evans, Museum of Comparative Zoology, Harvard Univer-
sity; and Dr. A. Kaltenbach, Vienna Museum of Natural History,
Austria, for the loan of museum specimens. I am grateful to Mr.
Samuel Cohen for taking the photographs. I collected Proscratea
complanata as a member of Phase C of the Alpha Helix expedition
to the Amazon in 1967. I thank the National Science Foundation for
support on the Amazon expedition under Grant NSF-GB-5916.

References

Brunner, von Wattenwyl, C.

1865. Noveau systeme des Blattaire. Vienna, 426 pp.

Graves, P.

1969. Spermatophores of the Blattaria. Ann. Entomol. Soc. Amer., 62:
595-602.

264

Psyche

[September

Gurney, A. B, and L. M. Roth.

1972. A generic review of the cockroaches of the subfamily Panchlori-
nae (Dictyoptera, Blattaria, Blaberidae). Ann. Entomol. Soc.
Amer, 65: 521-532.

Hebard, M.

1926. The Blattidae of French Guiana. Proc. Acad. Nat. Sci. Phil.,
78: 135-244.

Kirby, W. F.

1904. A synonymic catalogue of Orthoptera. Vol. I. London, pp. 61-205.
McKittrick, F. A.

1964. Evolutionary studies of cockroaches. Cornell Univ. Agr. Exp.
Sta. Mem. No. 389, 197 pp.

Princis, K.

1963. Kleine Beitrage zur Kenntnis der Blattarien und ihrer Verbrei-
tung. VII. Opusc. Entomol., 28: 147-155.

1964. Orthopterorum Catalogus. Edit. M. Beier. Pars. 6. Blattariae:
Subordo Blaberoidea. ’s - Gravenhage, pp. 174-281.

1965. Orthopterorum Catalogus. Edit. M. Beier. Pars. 7. Blattariae:
Subordo Blaberoidea. ’s - Gravenhage, pp. 284-400.

1967. Kleine Beitrage zur Kenntnis der Blattarien und ihrer Verbrei-
tung. X. Opusc. Entomol., 32: 141-151.

Rehn, J. A. G.

1932. Wissenschaftliche Ergebnisse der schwedischen entomologischen
Reisen des Herrn Dr. A. Roman in Amazonas 1914-1915 und
1923-1924. Arkiv for Zoologi, 24A: 1-71.

Roth, L. M.

1967. Sexual isolation in parthenogenetic Pyconoscelus surinamcnsis
and application of the name Pycnoscelus indicus to its bisexual
relative (Dictyoptera: Blattaria: Blaberidae: Pycnoscelinae) . Ann.
Entomol. Soc. Amer., 60: 774-779.

1970a. The male genitalia of Blattaria. III. Blaberidae: Zetoborinae.
Psyche, 77: 217-236.

1970b. The male genitalia of Blattaria. IV. Blaberidae: Blaberinae.
Psyche, 77: 308-342.

1971a. The male genitalia of Blattaria. VI. Blaberidae: Oxyhaloinae.
Psyche, 78: 84-106.

1971b. The male genitalia of Blattaria. VIII. Panchlora, Anchohlatta,
Biollcya, Pelloblatta, and A chroblatta (Blaberidae: Panchlorinae) .
Psyche, 78: 296-305.

CAMBRIDGE ENTOMOLOGICAL CLUB

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No, 4

PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol. 80 December, 1973

CONTENTS

The Mating Behavior of Brochymena quadrapustulata (Fabricius)
(Hemiptera, Pentatomidae) . George Gamboa and John Alcock ....

Patterns of Abdominal Fusions in Male Boreus (Mecoptera).

K. JV. Cooper

Speocolpodes, a New Genus of Troglobitic Beetles from Guatemala
(Coleoptera, Carabidae). T. C. Barr, Jr

The Stabilimenta of Nephila clavipes and the Origins of Stabili-
mentum-building in Araneids. M. H. Robinson and B . C. Robinson

The Larva and Pupa of Carpelimus debilis Casey (Coleoptera:
Staphilinidae). Ian Moore and E. F. Legner

Generic Diversity in Phase Rhythm in Formicine Ants.

E. S. McCluskey

The Male Genitalia of Blattaria. XI. Perisphaeriinae. L\. M. Roth

New Species, Records, and Synonyms of Chilean Theridiid Spiders
(Araneae, Theridiidae) . IV . C. Sedgwick

Latitudinal Gradients in Larval Feeding Specialization of the World
Papilionidae (Lepidoptera) . J. M. Scriber

The 100th Anniversary of the Cambridge Entomological Club

Author and Subject Index for Volume 80

265

270

271

277

289

295

305

349

355

374

375

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1973-1974

President .
Vice-President .
Secretary .

Treasurer .

Executive Committee

B. K. Holldobler
W. D. Winter, Jr.
H. E. Nipson
F. M. Carpenter
R. E. SlLBERGLIED
L. P. Lounibos

EDITORIAL BOARD OF PSYCHE

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University

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The Lexington Press. Inc.. Lexington. Massachusetts

PSYCHE

Vol. 80 December, 1973 No. 4

THE MATING BEHAVIOR OF
B ROC HYMEN A QUADRA PUSTULATA (FABRICIUS)*

By George Gamboa and John Alcock
Department of Zoology,

Arizona State University,

Tempe, Arizona 85281

Pentatomid reproductive behavior has been the subject of a number
of papers (e.g. Kullenberg, 1947; Teyrovsky, 1949; Sou thwood and
Hine, 1950; Leston, 1955; Kaufmann, 1966; Mitchell and Mau,
1969; Tostowaryk, 1971; Alcock, 1971; Fish and Alcock, 1973).
These studies have revealed considerable diversity and complexity in
the courtship activities of male pentatomids, raising questions about
the evolution and ecological significance of these behaviors. Answers
to these questions ' will require additional comparative data. We
present information here on Brochymena quadrapustulata ; loose
aggregations of this cryptic species were observed on grapefruit
( Citrus paradisi ) and desert broom ( Baccharis sarothroides ) in
suburban Tempe, Arizona, from 25 February to 17 May 1973. In
addition to written records of field observations of eight courtships,
super-8 films of three separate courtships leading to copulation were
utilized for detailed analysis of the mating behavior of B. quadra-
pustulata.

Results

Eight complete and three incomplete courtships were observed
between 1145 and 1540 hrs. The components of mating are out-
lined chronologically below and illustrated in Fig. 1.

(1) The male approaches the female (she may be moving or im-
mobile at the time) and touches her with his antennae. If
moving, the female may freeze with her abdomen held close to
the branch on which she was walking or escape by running
away.

* Manuscript received by the editor October 9, 1973

265

[December

266 Psyche

Figure 1. A diagram of the courtship of Brochymcna quadrapustulata.

A. The male’s crab-like movements on the anterior dorsum of the female.

B. The male moves to the rear of the female. C. The view from above and
from the side of the male antennating the venter of the female’s abdomen.
D. The view from above and from the side of the male about to achieve
genital linkage. E. Genital linkage in an end-to-end position. F. The male
drumming on the side of the female with its hindlegs.

1973]

Gamboa & Alcock — Brochymena

267

(2) The male seizes the anterior or posterior dorsum of the female
with his legs. Unreceptive females respond by breaking away
from the male or by moving off with their partner clinging to
them.

(3) If the female is immobile, the male moves so as to face his
mate while continuing to grasp the dorsum of her body. He
then begins a rapid crablike walk from side to side over the
anterior female dorsum. Receptive females gradually raise their
abdomen in response to this activity. The male’s back and forth
movements varied from 20-34 in number and lasted from 23-43
seconds in four filmed courtships.

(4) At some point the male moves off the anterior dorsum of the
female and continues out along the side of his mate while con-
stantly antennating her lateral surface.

(5) The male, upon reaching the posterior of the female, sweeps
his head under her elevated abdomen ; his antennae move rapidly
up and down in alternating strokes touching the female’s
abdominal venter. The male may prod and lift the body of
unresponsive females which have not voluntarily raised their
abdomens.

(6) As the male’s head passes under the body of the female he
begins a tight 180° turn that brings him into the end-to-end
precopulatory position. Unreceptive females may dash off down
a branch as the male executes this maneuver.

(7) The male elevates his abdomen while sweeping the aedeagus in
a zig-zag pattern against the venter of the female’s abdomen.
Each sweep brings the male aedeagus progressively closer to
the female genitalia; linkage occurs when mutual genital con-
tact occurs.

(8) Upon linkage, the insects’ bodies jerk violently from side to
side for several minutes. The coupled pair may move a short
distance during this time.

(9) The male rapidly drums the sides of the female’s abdomen with
his hindlegs (also abserved by Ruckes, 1938, for B. sulcata ).
Spells of drumming are interrupted by pauses during which the
male rests its hind legs on the dorsum of the female’s abdomen.
This continues as long as the partners are linked. One pair of
stinkbugs remained in copulo for at least 75 min but had sepa-
rated when observed 45 min later.

Discussion

This report provides additional evidence that among the Penta-

268

Psyche

[December

tomidae it may not be uncommon for males to initiate copulation
while facing directly away from the female. This behavior is rare
among the Heteroptera (Weber, 1930). Below we have summarized
the major methods by which pentatomids achieve genital linkage.

Genital linkage initiated in an end-to-end position

Brochymena (this paper), Euschistus (Alcock, 1971), Nezara
(Mitchell and Mau, 1969), Chlorochroa and Cosmopepla (Fish
and Alcock, 1973), Perillus (Esselbaugh, 1948), Podisus (Olsen,
1910)

Genital linkage initiated with male above female, both facing in same
direction

Brochymena (Ruckes, 1938), Dolycoris (Teyrovsky, 1949),
Calidea (Kaufmann, 1966)

Genital linkage initiated with female above male, both facing in same
direction

Podisus (Tostowaryk, 1971)

We also have records of end-to-end matings in Phyanta pallido-
virens. Males of this species, upon making contact with a female,
begin antennating the surface of her body while moving to the tip
of her abdomen. There they may prod and lift the female’s body
with their head before turning away from her. The female, if
receptive, raises her abdomen slightly. Unlike other species which
initiate copulation in a dismounted position, the male’s body often
forms a right angle with the female instead of a 180° angle. The
male then kicks lightly at the rear of its partner’s wingcovers and
abdomen with its hind legs before inserting the aedeagus into the
female’s genital opening.

As Fish and Alcock (1973) have noted, species which employ -the
end-to-end method have highly similar courtship routines. Common
characteristics include (1) male antennation of the female, (2) at-
tempts by the male to lift the female’s abdomen with its head,
(3) abdominal elevation by receptive females, and (4) tactile stim-
ulation of the venter of the female’s abdomen with the antennae and
aedeagus of the male. The male’s “goal” in courtship appears to be
to induce the female to adopt a position that will make insertion
of the aedeagus relatively easy.

The unusual feature of the courtship of B. quadrapustulata is the
rapid crab-like movement of the male over the head and thorax of
the female, a behavior that may have evolved from efforts of males
in the past to prevent physically the escape of females. Now the
action may serve as a releaser of abdominal elevation by receptive

1973]

Gamboa & Alcock — Brochymena

269

females. Interestingly, very similar behavior has been reported for
the distantly related African pentatomid Calidea dregii, a member
of the Scutellerini (Kaufmann, 1966).

Finally the fact that different members of the same genus (e.g.
Brochymena , Podisus) may exhibit basically different methods of
initiating genital linkage suggests that this component of reproduc-
tive behavior is evolutionarily labile. The reasons for evolutionary
changes of this sort remain obscure.

Acknowledgments

We thank Elinor Lehto for the identification of the plants men-
tioned in the paper. Alan R. Hardy of the California Department
of Agriculture provided the identifications of Brochymena and
Thyanta for which we are grateful. This research was supported
in part by NSF grant GB-35269.

References

Alcock, J.

1971. The behavior of a stinkbug, Euschistus conspersus Uhler (Hemip-
tera, Pentatomidae) . Psyche 78: 215-228.

Esselbaugh, C. O.

1948. Notes on the bionomics of some midwestern Pentatomidae. En-
tomol. Americana 28: 1-73.

Fish, J. and J. Alcock

1973. The behavior of Chlorochroa ligata (Say) and Cosmopepla
bimaculata (Thomas), (Hemiptera, Pentatomidae). In press,
Entomol. News.

Kaufmann, T.

1966. Notes on the life history and morphology of Calidea dregii
(Hemiptera: Pentatomidae: Scutellerini) in Ghana, West Africa.
Ann. Entomol. Soc. Amer. 59: 654-659.

Kullenberg, B.

1947. Uber Morphologie und Funktion des Kopulationsapparats der
Capsiden und Nabiden. Zool. Bidrag Fran Uppsala 24: 217-418.

Leston, D.

1955. The function of the conjunctiva in copulation of a shieldbug,
Peizodorus lituratus (Fabricius) (Hemiptera, Pentatomidae)..
J. Soc. Brit. Entomol. 5: 101-105.

Mitchell, W. C. and R. F. L. Mau

1969. Sexual activity and longevity of the southern green stinkbug
Nezara viridula. Ann. Entomol. Soc. Amer. 62: 1246-1247.

Olsen, C. E.

1910. Notes on breeding Hemiptera. J. N. Y. Entomol. Soc. 18: 39-42.
Ruckes, H.

1938. Courtship and copulation in Brochymena sulcata Van D. Bull.
Brook. Entomol. Soc. 33: 89-90.

270

Psyche

[December

SOUTHWOOD, T. R. E. AND D. J. HlNE

1950. Further notes on the biology of Sehirus bicolor (L.). Entomol.
Mon. Mag. 86: 299-301.

Teyrovsky, V.

1949. Praeconnubia and courtship in terrestrial bugs. Acta Acad. Nat.
Hist. Moravo-Silesiacae, Brno 21: 1-16.

Tostowaryk, W.

1971. Life history and behavior of Podisus modestus (Hemiptera:
Pentatomidae) in a boreal forest in Quebec. Can. Entomol. 103:
662-674.

Weber, H.

1930. Biologie der Hemipteren. Julius Springer, Berlin.

Patterns of Abdominal Fusions in Male Boreus (Mecop-
tera). — Since publication of my comments on the fusion of
terminal tergal and sternal abdominal plates in male Boreus (Psyche,
yg: 277, 1972), I have examined two males of Boreus vlasovi
Martynova (determined by Dr. Martynova) from Ashkhabad,
Turkmeniya, U. S. S. R., through the kindness of Prof. F. M.
Carpenter. Contrary to my listing based on the original description,
these males do not have the 9th tergum fused to its sternum. Both
have their 8th and 9th terga and sterna free, namely (0,0), just as
in the North American Boreus brevicaudus Byers, B. brumalis Fitch,
B. nivoriundus Fitch, and B. noboperates Cooper. Byer’s figure
(Ann. Ent. Soc. Amer. 47 : 491, fig. 15, 1954) shows the North
American B. reductus Carpenter also to be (0,0). — K. W. Cooper,
Dept, of Biology, University of California, Riverside, Calif., 92502.

SPEOCOLPODES , A NEW GENUS OF
TROGLOBITIC BEETLES FROM GUATEMALA
(COLEOPTERA: CARABIDAE)*

By Thomas C. Barr, Jr.

School of Biological Sciences
University of Kentucky
Lexington, Kentucky, U. S. A. 40506

In early January, 1973, Mr. Henry Frania and Mr. Michael
Shawcross visited Seamay Cave, Alta Verapaz, Guatemala, and
discovered two specimens of a remarkable troglobitic carabid beetle.
“One was collected after dusk running over flowstone near pools
of water several hundred yards into the cave. The second was
found the next morning, less than 200 yards from the entrance
under a rock in a dry, silt-covered flowstone pool, but still well
within the dark zone.” (H. Frania, in litt.). Subsequently Mr.
Frania referred the specimens to me for study.

The Seamay Cave beetle is of special interest for two reasons:

1 ) It is the first troglobitic beetle known from Guatemala, and
the cave is the farthest south of any troglobitic beetle type locality
in North America.

2) The beetle clearly belongs to the Agonini and in body form
resembles the species of Rhadine LeConte, but the possession of long
setae on the underside of the tarsi and conspicuous lobes on the fourth
tarsal segments indicates that its affinities lie not with Rhadine but
with the large, heterogeneous group of agonines currently placed in
Colpodes M’Leay.

Only one other group of colpodines is known to regularly inhabit
caves and to include apparent troglobites, — the various species of
Mexisphodrus Barr, from the uplands of central Mexico. These
have recently been described by Barr (1965, 1966) and Hendrichs
and Bolivar (1966, 1973). Although the sphodrine appearance of
some of the first species of Mexisphodrus which came to my attention
led me to consider them primitive sphodrines, more recent study of
the material now available suggests that they are not true sphodrines
at all, but a specialized, rather distinctive-looking line of colpodines.
The status of the name Mexisphodrus will depend upon future
taxonomic revisions of the colpodines, but I believe it will be useful

* Manuscript received by the editor November 4, 1973.

271

272

Psyche

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at least at the subgeneric level because of the distinctive habitus of
its component species.

The Seam ay Cave beetle does not appear to be close to Mex-
isphodrus despite their cavernicolous habitats and despite their in-
clusion within the colpodines. In Mexisphodrus there are well-formed
eyes or eye rudiments, the head is rounded, the pronotum is more
or less cordiform with very broadly reflexed sides, the elytral humeri
are well developed and prominent, and the color is dark violaceous-
ferruginous. In the Seamay Cave beetle the eye rudiments are
minute, the head is twice as long as wide and its sides are subparallel,
the pronotum is barrel-shaped, the elytral humeri are completely
rounded, and the color is rufotestaceous.

Speocolpodes, new genus

Slender, elongate, rufotestaceous cave beetles with minute vestiges
of eyes and wings; front and middle tarsi with penultimate segment
deeply and asymmetrically bilobed ; head twice as long as wide, sides
subparallel ; pronotum widest at middle, hind angles blunt and obtuse.

Integuments generally glabrous; microsculpture on dorsum of head
obsolete, obsoletely transverse on pronotal disc, finely and densely
transverse on elytra.

Eye rudiment visible only as minute, translucent spot beneath
cuticle ; labrum with anterior margin broadly convex, 6-setose ; two
clypeal setae; clypeofrontal groove joining postantennal groove and
terminating in lunate depression between anterior and posterior
supraorbital punctures. All mouthparts elongate and slender; mandi-
bles falcate, right mandible with large tooth ; maxillary palps with
terminal segment fusiform, a little shorter than penultimate segment;
glossa with apex broadly rounded and bisetose; labial palps bisetose;
mentum tooth grooved, truncate; two prebasilar setae each side of
submentum behind suture.

Pronotum barrel-shaped, longer than wide, widest at middle, apex
slightly narrower than base; two pairs of marginal setae; hind angles
obtuse and somewhat rounded, not produced. Metepisterna short,
24 as wide as long.

Elytra elongate-elliptical, humeri completely rounded, subapical
sinuation absent ; scutellar stria very brief, obsolescent ; scutellar
puncture setiferous; two disca.1 punctures in apical half, on third
interval against second stria; umbilicate series 6 + 2 + 6, seventh
and eighth punctures rather widely spaced ; a single puncture near
apex of seventh stria.

1973]

Barr — Speocolpodes

273

Antenna very long and slender, attaining apical fourth of elytra;
pubescence beginning near base of fourth segment. Tarsi with first
four segments densely setose beneath, last segment glabrous beneath,
claws smooth ; fourth segment bilobate, conspicuously so on pro- and
mesotarsi, inner lobe longer than outer, fourth metatarsal segment
feebly bilobate; two or three basal segments of meso- and metatarsi
with feeble lateral ridge.

Male unknown.

Type species: Speocolpodes franiai, new species.

Speocolpodes franiai, new species

Etymology : Patronymic honoring Mr. Henry Frania, codiscoverer
of the species.

Diagnosis: The genus is at present monobasic; the single species
is distinguished from all other known species of North American
Anchomeninae by the combination of bilobed fourth tarsal segments,
vestigial eyes, rufotestaceous color, and slender, elongate body form.

Description : Length 9.5-10.2 mm. Rufotestaceous, shining, pol-
ished ; integuments generally glabrous except for fixed setae ; micro-
sculpture obsolescent on dorsum of head, evanescently transverse on
pronotal disc, finely and densely transverse on elytral disc. Body
form slender and elongate, appendages all slender and elongate.

Head 2.1 times longer than wide (excluding mandibles), sides
subparallel, feebly convergent at neck; eye vestigial, represented only
by minute rudiment seen as translucent spot through cuticle, about
0.09 X 0.15 mm. in holotype; labrum 0.4 as long as wide, anterior
margin evenly convex; clypeofrontal grooves short, divergent behind
level of antennal bases, joining feebly impressed postantennal groove
at anterior supraorbital puncture and ending in lunate depression
between anterior and posterior supraorbitals ; mandibles elongate,
slender, falciform, left mandible 1.40 mm. long in holotype; maxil-
lary palps with terminal segment fusiform, about 0.85 as long as
penultimate segment, total length of palp (holotype) 1.77 mm.,
segments in ratio of 1.0 : 4.0 : 3.6 : 3.0; labial palps with two setae
on basal half of penultimate segment (distal seta possibly irregular:
short or broken on two of four palps examined, absent on other two
palps), total length of palp (holotype) 1.23 mm., segments in ratio
of 1:4:3; mentum tooth prominent, grooved, truncate at apex;
submentum with two prebasilar setae each side behind suture.

Pronotum 1.25 times longer than wide, apex width 0.8 times base
width and 0.67 times maximum width, which occurs near middle;

274

Psyche

[December

Figure 1. Speocolpodes franiai, new genus and species. Holotype female,
length 9.5 mm,, Seamay Cave, Alta Verapaz, Guatemala. [Second antennal
segment inadvertently omitted on left side of figure; this segment is shown
correctly on right side.]

1973]

Barr — Speocolpodes

275

disc convex, a little deplanate toward sides ; anterior angles depressed,
sides rounded but shallowly sinuate before middle and before hind
angles, which are obtuse, moderately rounded and reflexed ; base
unmargined ; marginal setae placed in margin, anterior pair at mid-
dle, posterior pair before hind angles. Prosternum slightly flattened
along mid-line; posterior process truncate, but not very sharply so.

Elytra 1.8 times longer than wide, humeri completely rounded,
sides slightly sinuate in basal fourth, subapical sinuation obsolete;
disc convex, striae deep and uninterrupted, intervals convex; scutellar
stria very short, obsolescent; first stria either truncating or joining
second stria at base, fourth and fifth striae joining in apical fourth
and not attaining apex; scutellar puncture setose; two discal punc-
tures on third interval touching second stria, anterior puncture be-
hind middle, posterior puncture in apical fifth; umbilicate series
6 + 2 + 6, spacing between punctures 6 through 9 wide and a little
irregular; whips apparently (some broken?) in punctures 1, 9, and
13, but not excessively long; seventh stria with a single preapical
puncture.

Metathoracic wings vestigial, about 0.7 mm. long (holotype).
Abdominal tergites pale, unpigmented, translucent (as in troglobitic
Trechinae). Metepisterna short, anterior margin 0.75 as long as
lateral margin.

Appendages all slender and elongate. Antenna 0.85 times total
body length, first three segments glabrous, pubescence beginning at
basal sixth of fourth segment; fourth segment one-third longer than
third segment; beginning with scape, segments in length ratio as
follows: 1 .00 : 0.44 : 1.10 : 1.52 : 1.27 : 1.22 : 1.14 : 1.10 :
1. OO : 0.87 : 0.79 (holotype). Tarsus with fourth segment bilobate,
inner lobe longer than outer, lobes prominent on pro- and mesotarsus
but inconspicuous on metatarsus; first four segments with long setae
beneath, fifth segment glabrous beneath; meso- and metatarsi with
feeble ridge on outer side of basal two or three segments ; total length
of protarsus 1.20 mm. in holotype, segments in length ratio of 1.00 :
O.55 10.48 : 0.29 (excluding lobes) : 1.55.

Male unknown.

Holotype: Female (American Museum of Natural History, New
York), Seamay Cave, near Sena.hu, Alta Verapaz, Guatemala,
January 9, 1973, collected by Henry Frania. Approximate elevation
3000 fett; approximate coordinates of type locality as furnished by
Mr. Frania, 89° 50' X 150 23'.

Paratype: Female, same locality, January 8, 1973, collected by
Michael Shawcross. Deposited in T. C. Barr collection, University
of Kentucky.

276

Psyche

[December

Measurements of Holotype: total length 9.5 mm., head (excluding
mandibles) 2.32 mm. long X 1.08 mm. wide, pronotum 1.95 mm.
long X 1.55 mm. wide, elytra 5.53 mm. long X 2.99 mm. wide,
antenna 8.13 mm. long.

Discussion

Speocolpodes was presumably derived from a colpodine stock with
eyes and wings, possibly from a troglophile ancestor which frequented
moist, dark microhabitats. Its exact position among the hundreds of
species of colpodines inhabiting Mexico and Central America is
difficult if not impossible to determine until the taxonomy and
phylogeny of the group is better understood.

Seamay Cave, located on the Finca Seamay near Senahu, is one
of three caves in which the catopine leiodid beetle Ptomaphagus
giaquintoi Jeannel (1949) is known to occur (Peck, 1970). P.
giaquintoi has small though probably functional eyes and apparently
functional metathoracic wings, and is at most a troglophile, even
though it shows certain cave adaptations such as long antennae.
Although Peck visited Seamay Cave and collected P. giaquintoi there
(in June and August), he did not find Speocolpodes in that cave or
in nearby Cueva Sepacuite, the largest of three caves on Finca
Sepacuite. It is conceivable that the carabids appear seasonally in
the cave, and that concentrated search during January or February
would yield more specimens.

Literature Cited

Barr, Thomas C., Jr.

1965. A new cavernicolous sphodrine from Veracruz, Mexico (Coleop-
tera: Carabidae). Coleopterists’ Bulletin, 19: 65-72.

1966. New species of Mexisphodrus from Mexican caves (Coleoptera:
Carabidae). Psyche, 73: 112-115.

Hendrichs S., Jorge, and C. Bolivar Y Pieltain

1966. Hallazgo de un nuevo Mexisphodrus cavernicola en el Estado de
Hidalgo (Mexico): M. gertschi nov. sp. Ciencia, 25: 7-10, pi. 1.
1973. Un nuevo esfodrino ciego del Sotano de San Agustin, Oaxaca,
Mexico (Coleopt., Carab.). Ciencia, 28: 37-41.

Jeannel, R.

1936. Monographic des Catopidae. Mem. Mus. Nat. Hist. Nat., n. s., 1:
1-473.

Peck, Stewart B.

1970. A systematic revision and the evolutionary biology of the
Ptomaphagus (Adelops) Beetles of North America (Coleoptera:
Leiodidae: Catopinae). Harvard University, unpublished doc-

toral dissertation.

THE STABILIMENTA OF NEPHILA CLA PIPES
AND THE ORIGINS OF
STABILIMENTUM-BUILDING IN ARANEIDS*

By Michael H. Robinson and Barbara C. Robinson
Smithsonian Tropical Research Institute,

P.O. Box 2072, Balboa, Canal Zone

Structures of multi-strand (ribbon) silk are built into the webs
of certain araneid and uloborid spiders. These devices are widely
known as stabilimenta (following Simon 1895). Marples (1969)
has objected to the functional connotations of the term and has
called such structures decorations. However, we feel that there is
no point in abandoning a term that has acquired a. designatory value
largely independent of functional implications.

The stabilimenta built by araneids differ, from species to species,
in form, complexity and disposition within the web. There can also
be differences between the stabilimenta built by different develop-
mental stages within the same species. Some species build disc
stabilimenta at one stage in the life cycle and linear stabilimenta at
another (almost always later) stage. Despite these differences (ex-
amples in Robinson & Robinson 1970, Ewer 1972, general discussion
with references in Kaston 1964) araneid stabilimenta have construc-
tional features in common. With the exception noted below, all the
stabilimenta that have been described consist purely of silk that is
laid down between structural elements of the web in a zig-zag
manner. In linear stabilimenta, the subject of this paper, the zig-
zags bridge the gap between adjacent radii. An exception to this
common constructional feature occurs in the stabilimenta built by
some species of Cyclosa in which the devices incorporate discarded
remnants of prey and other debris, and may also contain egg sacs.
These stabilimenta are perhaps best regarded as a special case and
could be described as ‘composite.’

As far as we are aware, there has been no record of stabilimentum
building by spiders of the genus Nephila prior to its discovery in
Nephila maculata (Fabricius) by Robinson & Robinson (1973).
These authors found that N. maculata builds a perpendicular linear
stabilimentum in very rare instances. The structure is built only by
immature females and shares the constructional features described

* Manuscript received by the editor November 4, 1973.

277

278

Psyche

[December

Table 1. Results of web census of Nephila maculata (July 1973) at
localities in southwest Canal Zone, between Rodman and
Arraijan.

Adults

Immatures*

Totals

Number with stabilimenta

Perfect

0

Skeleton

0

Perfect

0

Skeleton

12

12

Number with ribbon silk

at hub

52

0

152

18

222

Total number of webs

in the census

65

0

188

22

275

% of all webs with hub silk 80.7%

% of all webs with stabilimenta 4.+%

% of skeleton webs with stabilimenta 54.5
^Females above 10mm in size.

above. It is of multi-strand silk laid down between adjacent radii
(see Robinson & Robinson 1973, Figure 5).

In this paper we describe examples of stabilimenta built by imma-
ture females of Nephila clavipes (L.) and discuss the functional and
evolutionary implications of the rare phenomenon of stabilimentum
building by Nephila species.

The stabilimenta of Nephila clavipes
Nephila clavipes is widely distributed in the New World tropics
and extends into subtropical regions to the north and south (see
Bonnet 1958). Notes on the web structure and ecology of this
species appear in studies by Peters (1953, 1954, 1955). In Panama
the species is relatively common in most lowland areas where trees
and bushes persist. Our preliminary studies of seasonal abundance
suggest that it becomes rare towards the end of the dry season
(April-May). Populations build up again during the wet season
with adults becoming numerous by July-August. We first discovered
a stabilimentum in the web of N. clavipes in a garden at Rio Indio,
Arraijan, Republic of Panama in July 1973. After this discovery
we censussed 275 webs of adults and immatures (Table 1) in forest-
edge areas in the southwestern part of the Canal Zone around the
Interamerican Highway.

As a result of the census we discovered 12 stabilimenta, all of
which were in skeleton webs (see below) of immature females.
Thus stabilimenta were present in only 4.4% of the webs examined.
The census included 65 adult webs and 210 webs of immature fe-
males. Of the latter 22 were without viscid elements and are
referred to as skeleton webs. Fifty-four percent of the skeleton webs

1973]

Robinson & Robinson — Nephila

279

Fig. 1. Stabilimentum in perfect web of juvenile female Nephila clavipes.
Dimensions in text.

28o

Psyche

[December

contained stabilimenta. The original stabilimentum discovered at
Arraijan was in a functional web and differed in detail from those
found in skeleton webs. No stabilimenta of this type were encoun-
tered during the census. We photographed the first-found stabili-
mentum and one of the type found in skeleton webs and also col-
lected several of the latter on black paper to analyse in detail. The
accompanying photographs were made without coating the web to
enhance its visibility since this could have obscured the detail of the
stabilimenta by rendering structural members differentially more
conspicuous than in the natural state. Web photography was ac-
complished by using two flash units, one placed at right angles to
the web and one discharging along the plane of the web from below.

Figure i shows the stabilimentum in the complete web. The
immature female was ca. 17mm in length and occupied a web ap-
proximately 29cm in height (between upper and lower foundation
threads). The web was in a shaded location surrounded by bushes
and had a dense and complex barrier web dorsal to the spider (below
the sloping main sheet) and a second less dense barrier web above
the web, ventral to the spider. The zig-zags of stabilimentum silk
were laid down across three inter-radial gaps and in the lower part
also followed a branched radius, but were mainly concentrated
between two radii. The whole structure, of golden ribbon silk,
formed a somewhat imperfect perpendicular stabilimentum, quite
dense over 22mm of its length, and about 33mm in total length.
As can be seen from the figure, the stabilimentum extended from the
hub region into the viscid spiral zone.

The stabilimenta found in skeleton webs all consisted of much
longer structures in which silk was deposited between much more
widely spaced radii with the attachment points more widely dis-
persed on the structural elements. Figure 2 shows a section of one
such stabilimentum in which stabilimentum silk was deposited en-
tirely on a single radius for part of the length of the structure. All
these stabilimenta were relatively inconspicuous and could have been
overlooked in casual examination of the webs. One skeleton web
stabilimentum that we collected had zig-zags of multi-strand silk
extending over seven inter-radial spaces and was 30mm wide (maxi-
mum) and over 75mm long.

The structure and function of skeleton webs

Several species of araneid spiders that we have studied cease to
build complete webs at some stage and rest for one or more days on
skeleton webs. We noted this phenomenon, without explanation, in

1973]

Robinson £sf Robinson — Nephila

28

Fig. 2. Part of stabilimentum (ca. 8.5 mm long) from skeleton web of
N. clavipes.

282

Psyche

[December

the case of Argiope argentata (Fabricius) during a study involving
the examination of 2614 adult webs of this species (Robinson &
Robinson 1970: 644). Examination of our unpublished data on the
nature of skeleton webs shows that they were not simply old webs
but constructions with a small number of radii, no viscid spiral and
no well-defined structural spiral. Stabilimenta in such webs were
extremely long, had widely spaced zig-zags and frequently had areas
where the stabilimentum silk was deposited on top of a single radius.
Thus the constructional details of these stabilimenta were strikingly
parallel to those seen in the stabilimenta found in skeleton webs of
N. clavipes. This can be seen by comparing Figure 2 herein with
Figure 3 of Robinson & Robinson (1970: 646). In our study of
Nephila maculata we found adult females ceased web renewal several
days before egg-laying but remained on old, or skeleton, webs (Rob-
inson & Robinson 1973). We now suspect that the examples of
skeleton webs that we found in Argiope argentata were constructed
by females that were about to lay eggs and that in both cases these
webs function as resting platforms at a stage when food capture has
become unnecessary. In the case of Nephila clavipes the skeleton
webs of immature females almost certainly function as moulting
platforms. (We were fortunate to see one female ecdyse whilst on
a skeleton web and found two others with cast cuticles still present.)
The spider moults below the web, hanging on a silk line attached
to its hub. When moulting is complete, the spider eventually swings
back onto the skeleton web and assumes a normal predatory stance.

The skeleton webs of Nephila clavipes that we examined (22)
were characterized by small area (compared with the perfect webs
of the same developmental stage) and the small number of radii that
were present. The hub regions appeared to be normal and barrier
webs were present in all cases. Strong (thick) bracing threads
connected the hub region to one or other of the barrier webs, or
sometimes to both of these structures. Such threads are present in
most of the webs of Nephila clavipes and produce a conical distortion
of the hub above the spider’s resting area. In this region there are
often deposits of sheet silk laid down irregularly over the hub ele-
ments and base of the bracing line(s). Of the 275 webs that we
censussed, 80.7% had conspicuous silk deposits on this region. The
significance of this aspect of web structure is discussed below.

Discussion

Three central (and related) questions result from our study of
the stabilimenta of Nephila clavipes and N. maculata:

1973]

Robinson & Robinson — Nephila

283

1 . What is the function of these devices ?

2. Why are they built so rarely?

3. Why are they found only in the webs of juveniles?

There have been numerous functional interpretations of the ribbon
stabilimenta of araneids. These are briefly summarized in Table 2
and expounded in more detail by Robinson & Robinson (1970) and
Ewer (1972). The assumption that a unitary explanation is neces-
sary for all the structures that are semantically united by being
described as stabilimenta is invalid on logical grounds alone, as
Robinson & Robinson (ibid, 654) point out. (It is also important
to stress that spiders at different stages in development may be sub-
jected to attacks by different spectra of predators, because of size
differences or different web-site preferenda. For this reason juvenile
and adult stabilimenta could differ in both mode of operation and
overall function.)

In exploring the possible function (s) of Nephila stabilimenta we
will treat the two basic forms described above separately. The
stabilimenta built in skeleton webs differ strikingly from the single
ribbon stabilimentum found in a perfect web. The latter and the
rare stabilimenta of Nephila maculata are essentially similar.

We believe that most of the forms of defensive function listed in
Table 2 can be securely rejected in the case of the stabilimenta found
in skeleton webs, for the reasons set out below :

1. Direct concealment can be rejected because the device does not
cover the region where the spider rests.

Table 2. Hypotheses of stabilimentum
see Robinson & Robinson 1970)
Antipredator function

1. Direct concealment of spider 1.

(requires that the device

covers the spider)

2. Disguise (requires that the
device appears continuous with
the body or legs of the spider)

3. Deflection (requires that the 1

predator attacks the device

rather than the spider) 2.

4. Advertisement (requires that the
predator seeks to avoid the web
that is advertised by the device)

5. Shielding (requires that the

device strengthens the hub 3.

against penetration by predators)

function (for unattributed sources

Mechanical function
The device in some way allows
the spider to adjust the state of
the completed web after it has
applied its own weight to the hub
region.

Other functions
The device acts as a visual
attraction to insects (Ewer 1972)
The device protects the web
against damage by large insects
(which the spider could not
subdue) by making it conspicuous
(Ewer 1972)

The device acts as a sunshield
thereby eliminating the need for
costly postural thermoregulation.

284

Psyche

[December

2. Disguise can be rejected because the structure is by no means
conspicuous and, in any case, is separated from the spider by a
wide gap.

3. Deflection can be rejected because of this same lack of con-
spicuousness (resulting from the wide dispersal of the opaque silk
deposits). The same argument applies to advertisement.

All these forms of defense operate visually and it is necessary to be
cautious about conclusions concerning the visual acuity of predators.
One of the lessons of ethological studies is that in many situations
animals actually attend to relatively simple stimuli even when they
are quite capable of discriminating more complex ones. There is an
excellent treatment of this subject in Hinde (1970: 57-80). Despite
this qualification, we feel that a visually-operating function for the
skeleton-web stabilimenta of N. clavipes is highly improbable.

The defensive function of reinforcing the hub against penetration
by predators that strike through the web can be rejected because the
silk is simply not deposited in the hub region. The structure could
conceivably form a barrier to spider-hunting insects that might other-
wise fly through the skeleton web and attack the spider from below.
This function is improbable in view of the limited area covered by
the device.

A function as an insect attractant is highly improbable, not only
because the device is inconspicuous but also because the web has no
viscid spiral and cannot, therefore, trap prey. (In addition, the
spiders do not feed in the period immediately prior to moulting.)
Web protection is probably achieved by the barrier webs and this
function of the stabilimentum can be rejected for this reason. The
structure is not sufficiently dense to act as an effective sunshield and,
furthermore, is in the wrong place. (Note that some araneids adopt
complex postures, at times, in order to minimize heat absorption,
Krakauer 1972, Robinson & Robinson 1973.)

The fact that the stabilimentum occurs most commonly in skeleton
webs could be correlated with a mechanical function. The laying
down of zig-zags on those radii that are immediately below the point
at which the moulting spider attaches itself to the hub may stabilize
this attachment. This would ensure a secure moulting base within
an area protected by the barrier webs. From this base the spider
could safely go through the fairly vigorous process of withdrawing
from the old cuticle. The moulting process involves much pulling
and jerking and if the spider loses connection with the web, and
drops to the ground, the result may be fatal. The reinforcement of

1973]

Robinson & Robinson — Nephila

285

the skeleton web could also strengthen the platform on which the
spider assumes its normal stance as the new cuticle hardens. If these
are the mechanical functions of stabilimentum building at this stage,
it is necessary to explain why this solution to the problem of building
a strong moulting platform has been adopted. The omission of the
viscid spiral is explicable on the grounds that if insects were trapped
in a moulting platform, their struggles could dislodge the moulting
spider at a critical moment. In addition, feeding seems to be in-
hibited at this stage and expenditure on prey-capture systems would
be wasteful. (It is also possible that silk production is inhibited prior
to moulting, perhaps because of the urgent biosynthetic priority of
moulting itself.) If the viscid spiral is omitted, the platform could
still be stabilized by the deposition of structural silk in the form of a
temporary spiral. Constructing a spiral is probably a more expensive
process than simply reinforcing the relevant radii by the act of ‘over-
stitching’ them with zig-zags. The latter involves only a small num-
ber of excursions from the hub region. Studies of the building of
skeleton webs could provide a more detailed basis for further specu-
lation about this problem. A function related to strengthening a
moulting platform would account for the virtual restriction (see
Table 1) of stabilimentum building to the skeleton webs of immature
Nephila clavipes.

The dense perpendicular stabilimenta found in perfect webs of
Nephila maculata (and the single example from N. clavipes shown in
Figure 1.) could function defensively by disguise or deflection. In
this case it is only necessary to assume that some predators are in-
hibited from attacking large spiders to explain the restriction of the
devices to the webs of immatures. Thus if the stabilimentum in-
creased the apparent size of the juvenile spider (disguise) or was
itself mistaken for a small spider (deflection) it could inhibit or
deflect attacks on the otherwise attack-eliciting juvenile. This ex-
planation is plausible but unsatisfactory in view of the extreme rarity
of the stabilimenta.

A mechanical function for such infrequently-built structures could
only be justified if webs required additional strengthening (tension-
ing or bracing) on rare occasions. This is possible but we have no
evidence that it is probable.

It is possible to assume that the rare phenomenon of stabilimentum
building in perfect webs is an aberrant expression of a behavior that
is functional in the context of the skeleton web. (We do not know
whether Nephila maculata builds stabilimenta in such webs.) If this
were so, the greater density of the structure could result from the

286

Psyche

[December

fact that radii are separated by much shorter distances. This possi-
bility could have interesting implications from the evolutionary stand-
point (see below) .

Rarity of a functional structure (and behavior) could be explained
by assuming that the present state of the two species represents an
early stage in the evolution of stabilimentum-building. Conversely,
if the structures are regarded as presently non-functional, they could
be looked on as vestigial. Although no definite conclusion is possible
about the function of the stabilimenta in perfect webs, it is worth
exploring the implications of these two assumptions.

If it is assumed that the behavior is vestigial, it must be concluded
that it has lost its original function (s). If it is assumed that the
original function was defensive, then either it is no longer an effective
defense against predators or the spider has evolved more effective
defenses. The first possibility could result from a change in the
spectrum of predators exerting selection pressure on defensive de-
vices or from an advance in the behavior of the predators that
rendered the device ineffective. Nothing useful can be said about
these two possibilities. If the spider has evolved more effective de-
fensive adaptations these should be detectable at present. The most
obvious present-day defensive devices of Nephila species are the
spider’s escape behavior and its barrier webs. Escape behavior is,
we think, unlikely to be a recent specialization. Kaston (1964)
regards the barrier webs as primitive on grounds that seem reasonably
secure. We conclude that the probability that the structure is a
vestigial defensive device is extremely low.

Similar arguments suggest a rejection of the hypothesis that the
stabilimenta are vestigial bracing devices. Presentday Nephila spp.
brace the hub regions of their webs with strong lines attached to the
barrier webs. These structures are probably a primitive feature of
web construction and would seem to make a bracing stabilimentum
redundant.

The contrary assumption that these Nephila species are at a stage
close to the origins of stabilimentum-building behavior may have
greater heuristic value. As far as we know, there has been no at-
tempt to explain the origins of ribbon stabilimenta. Since the struc-
tures are added on to structural members and are not simply addi-
tions of structural silk, any theories of derivation must take this into
account.

Most (if not all) araneids use ribbon silk for a. variety of pur-
poses (see Kaston 1964). Given this faculty, the first stage in the
evolution of stabilimenta could be the use of this silk to reinforce

1973]

Robinson CSf Robinson — Nephila

287

moulting platforms (to minimize constructional effort as suggested
above). Once this behavior has evolved, lines of further variation
(and selection) are possible. The step(s) to the production of a
dense perpendicular band in a functional web could follow several
evolutionary paths. Nephila species often renew the radii and viscid
elements of one side of the web while leaving the other side un-
touched (see Peters 1955). This process frequently results in the
formation of a perpendicular renewal line. If this renewal line were
reinforced with zig-zag silk (an extension of skeleton web rein-
forcement behavior) a rudimentary stabilimentum could result (in a
functional web). Such an origin for the perpendicular stabilimentum
is at least possible. Assuming that the mechanical function of over-
stitching could be exploitable in further situations (other than the
partial renewal of a web) the behavior would be a potential starting
point for further selection.

In addition, behavior such as this, with a function originally re-
lated to the mechanisms of web renewal, might incidentally have an
anti-predator effect. A conspicuous junction line might be margin-
ally concealing, deflecting, advertising or disguising. Selection ex-
erted by predators on further variations in this behavior might then
result in transformation of the structure and function of the device.
This means that stabilimenta could have evolved under different
selection pressures to fulfill different functions in different species of
spiders. This view seems to us to be at least as probable as the
assumption that all stabilimenta have a common function.

Summary

1. The stabilimenta built by Nephila clavipes (and N. maculata )
are characterized by being laid down as zig-zags of ribbon silk
between adjacent radii.

2. Such devices are rare in complete webs but relatively common in
the skeleton webs of immature N. clavipes that are built as moult-
ing platforms. The situation with regard to skeleton webs built
by N. maculata is not known.

3. A functional explanation of the stabilimenta built into such
skeleton webs is that they reinforce an otherwise reduced, simpli-
fied and possibly unstable structure. A secure base for moulting
is regarded as essential to the development of the spider.

4. Defensive or other non-mechanical functions for the stabilimenta
in skeleton webs are, we think, highly improbable.

5. Perpendicular stabilimenta in complete Nephila webs are so rare
as to make functional explanation difficult.

288

Psyche

[December

6. The hypothesis that Nephila stabilimenta are vestigial seems to
be largely untenable.

7. We suggest that the devices built into skeleton webs could repre-
sent a primitive stage of stabilimentum construction. Hypothetical
steps leading from this stage to the evolution of the perpendicular
stabilimentum could include one at which zig-zag ‘overstitching’
was used to reinforce a line of junction between old and new
sections of a partially renewed web. The further evolution of
stabilimenta from this stage could have followed different func-
tional pathways in different species.

References

Bonnet, P.

1958. Bibliographia Araneorum Vol. II. N-S. Douladore, Toulouse.
Ewer, R. F.

1972. The devices in the web of the West African spider Argiope
jlavipalpis. J. nat. Hist., 6 : 159-167.

Hinde. R. A.

1970. Animal Behaviour. McGraw Hill Book Co. N.Y. 2nd ed.
Kaston, B. J.

1964. The evolution of spider webs. Amer. Zool., 4: 191-207.
Krakauer, T.

1972. Thermal responses of the orb-weaving spider Nephila clavipes
(Araneae: Argiopidae). Am. Midi. Nat., 88: 245-250.

Marples, B. J.

1969. Observations on decorated webs. Bull. Br. Arachnological Soc.,
l: 13-18.

Peters, H. M.

1953. Beitrage zur Vergleichenden Ethologie und Okologie Tropischer
Webenspinnen. Z. Morph. Okol. Tiere., 42: 278-306.

1954. Estudios adicionales sobre la estructura de la red concentrica de
las aranas. Comun. Inst. Trop. El Salvador, 1: 1-18.

1955. Contribuciones sobre la etologia y ecologia comparada de las
aranas tejedoras tropicales. Comun. Inst. Trop. El Salvador,
1-2: 37-46.

Robinson, M. H. & B. Robinson.

1970. The stabilimentum of the orb web spider, Argiope argentata:
an improbable defence against predators. Can. Ent., 102: 641-655.

1973. The ecology and behavior of the giant wood spider Nephila
maculata (Fabricius) in New Guinea. Smith, cont. Zool., 149: 1-76.

Simon, E.

1892-1895. Histoire naturelle des Araignees. Roret, Paris.

THE LARVA AND PUPA OF
CARPELIMUS DEBILIS CASEY
(COLEOPTERA: STAPHYLINIDAE)*

By Ian Moore and E. F. Legner
Division of Biological Control, Department of Entomology
University of California
Riverside, California 92502

On a number of occasions Carpelimus debilis Casey has been
found to be numerous at a marine salt marsh at La Salina, near
La Mision de San Miguel, Baja California Norte, Mexico. This
salt marsh was described in some detail by Moore (1964) and its
Coleoptera discussed, a key was given by Moore for the separation
of larvae of some of the Staphylinidae found there but no detailed
descriptions or illustrations were presented, nor were pupae men-
tioned. We are now taking this opportunity to describe and illustrate
the larva and pupa of one of those insects.

In July and August, 1971, a very large colony of C. debilis was
found in an area about two meters square in Moore’s Zone One.
The substrate was a mixture of sand and mud of a consistency
which would support the weight of a person with slight sinking.
Present with the Carpelimus but in much lesser numbers were Tachys
vittiger Le Conte, Thinobius frizzelli Hatch and Ochthebius sp. and
large numbers of larvae, almost entirely Carpelimus. Thousands of
adult C. debilis , hundreds of larvae and three pupae were collected.
The pupae were staphylinid pupae of about the size of Carpelimus so
it seems very likely that we have properly assigned them. A few
meters away in Zone One was another small area of similar material
but too wet to support the weight of a person. That area contained
large numbers of the same species of Ochthebius but few, if any,
other insects. Insects were collected in these areas by the water
flotation method in which a shovel of substrate is placed in a pail of
water and the insects removed as they surface (Moore, 1954). This
method brings adults and some active larvae to the surface but the
inactive pupae probably do not surface readily, so few were en-
countered.

Carpelimus is a large genus of wo rid- wide distribution with sev-
eral species often found together. Over 352 species were described

* Manuscript received by the editor September 10, 1973

289

290

Psyche

[December

through 1969. Because of their small size and generally uniform
appearance, members of the genus are difficult to identify. Larvae of
Staphylinidae are notoriously difficult to associate with adults except
in rare circumstances such as the present one. They are not easy to
rear, the larvae often being canabalistic when confined in close
quarters. Only twice before have larvae of this genus been made
known. Paulian (1941) described and illustrated the larva of the
Holarctic species C. bilineatus Stephens and Kasule (1968) illus-
trated parts of a British species (as Trogophloeus sp.). No pupa of
the genus has been described.

Larva of Carpelimus debilis Casey

Length 3.25 mm. (largest spms.). Body elongate, somewhat con-
vex, parallel, pale cream-colored with the sclerites pale piceus and
the extremities somewhat darker. Head oval, about as wide as long;
with three ocelli in an uneven row on each side ; epicranial suture
about one-half the length of head. Labrum longer than wide, nar-
rowed and truncate in front. Antennal fossae located at sides of
head above bases of mandibles. Antennae three-segmented ; first seg-
ment about as long as wide; second segment a little wider than first
and about twice as long as wide with the modified acorn seta near
the apex nearly as large as third segment, born at an obtuse angle ;
third segment less than half as wide as second, about as long as wide,
with a small modified acorn-like seta at apex. Mandibles arcuate,
with three small equal teeth arranged in a triangle at apex. Maxil-
lary palpi three-segmented ; first segment about twice as long as wide ;
second segment a little narrower than first, narrowed from base to
pointed apex. Lacinia triangular, widest at base, spinose or inner
edge. Labial palpi two-segmented ; first segment longer than wide ;
second segment narrower than first, about as long as wide. Pro-
notum, mesonotum and metanotum wider than long, the last two
each wider than preceding. Abdominal tergites each wider than
metanotum, more than twice as wide as long, fifth tergite widest,
pseudopod subcylindrincal, slightly longer than wide, slightly nar-
rowed from base to apex. Urogomphus one-segmented, pointed,
slightly longer than pseudopod. Many specimens, La Salina, Baja
California Norte, Mexico, Salt Marsh, August 3, 1971, Ian Moore
collector.

The only noticeable difference between this larva and Paulian’s
description of that of C. biliniatus is that in debilis the apex of the
mandible is formed of three small teeth arranged in a small triangle
whereas in biliniatus Paulian’s illustration shows two small subequal

1973]

Moore & Legner — Carpelimus

29

Figures 1-4. Carpelimus debilis. Fig. 1, larva; Fig. 2, venter of pupa
Fig. 3, side of pupa; Fig. 4, dorsum of pupa.

292

Psyche

[December

Figures 5-8. Carpelimus debilis. Fig. 5, antenna of larva; Fig. 6, max-
illa of larva; Fig. 7, urogomphus and pseudopod of larva; Fig. 8, mandible
of larva.

1973]

Moore & Legner — Carpelimus

293

teeth, one before the other, with a minute denticle between, and his
description states that the apex of the mandible is bidentate.

The pupae of Coleoptera are so poorly known that it is difficult
to choose with certainty characters which will be of diagnostic value.
However, studies by Jerome Rozen (1959, 1963a, 1963b) indicate
that location of tuberculate setae on various parts of the body, length
and shape of elytra, wings and urogomphus are probably most useful
so those characters have been given the most attention in the follow-
ing description.

1

Pupa of Carpelimus debilis Casey

Pupa exarate, elongate, pale, not chitinized in any part; various
parts of the body with long slender setae, each arising from a sur-
face so slightly elevated that it can hardly be said to be tuberculate.
Head venterally reflexed so that only a small part is visible from
above; with three ocelli in a row on each side; on each side with
two setae at front margin, one discal seta and three setae at lateral
margin. Pronotum irregularly ovoid, on each side with two setae
at front margin, one at lateral margin and two at posterior margin.
Elytra very little longer than wide, standing at right-angles to body,
posterior margin, when viewed from above, with an obtusely angu-
late projection in the middle, without setae. Wings narrower than
and about twice as long as elytra, held away from the body at an
angle of about 45 degrees, without setae. Mesothorax and meta-
thorax without setae. First abdominal segment on each side with
one discal and one lateral seta. Abdominal segments two through
six on each side with a single lateral seta. Urogomphus minute, with
a tuft of hair at apex.

Three specimens from La Salina, Baja California Norte, Mexico,
Salt Marsh, August 3, 1971, Ian Moore collector.

Acknowledgments

We are grateful particularly to Robert E. Orth of the University
of California at Riverside for laboratory help with this and other
studies.

Literature Cited

Kasule, F. K.

1968. The larval characters of some subfamilies of British Staphylini-
dae (Coleoptera) with keys to the known genera. Trans. Royal
Entomol. Soc. London 120: 115-138; 116 figs.

294

Psyche

[December

Moore, Ian

1954. An efficient method of collecting dung beetles. Pan-Pac. Entomol.
30: 208.

1964. The Staphylinidae of the marine mudi flats of southern California
and northern Baja California (Coleoptera). Trans. San Diego
Soc. Natur. Hist. 13 : 269-284, 4 figs.

Paulian, Renaud

1941. Le premier etats des staphylinoidea. fitude de morphologie com-
paree. Mem, Mus. Hist. Natur. Paris, N. ser., 15: 1-361, 1365
figs., 3 pt.

Rozen, Jerome G.

1959. Systematic study of the pupae of the Oedermeridae (Coleoptera).
Ann. Entomol. Soc. Amer. 52: 299-303, 28 figs.

1963a. Two pupae of the primative suborder Archostemata (Coleop-
tera). Proc. Entomol. Soc. Wash. 65 : 307-310, 7 figs.

1963b. Preliminary systematic study of the pupae of Nitidulidae (Co-
leoptera). Amer. Mus. Novitates. 2124: 1-13, 30 figs.

GENERIC DIVERSITY IN PHASE OF RHYTHM
IN FORMICINE ANTS

By E. S. McCluskey
Departments of Physiology and of Biology
Loma Linda University, Loma Linda, California 92354

Ants are abroad through most of the day and night. But the
species composition of this 24-hour patrol changes from one part of
the day to the next (Talbot 1953; Wilson 1971). For example, in
Michigan the maximum foraging activity of Lasius neoniger is at
night, of Myrmica americana in the early morning and late after-
noon, and of Formica pallidefulva nitidiventris in the middle of the
day (Talbot 1946, 1953).

Likewise the mating flights of ants occur at different hours for
different species (Kannowski 1963; Talbot 1945). The flight times
may be similar for closely-related species (Kannowski 1959a).

If one were to look at many species of one genus, would he find
them to be similar as to time of day of foraging or of mating flight?
Or does each genus span the 24 hours in terms of its various species?
The aim of this report is to quantitatively compare species diversity
with generic diversity of such phase relationships in one tribe of ants,
the Formicini. The biosystem atics of much of this group, particu-
larly of the genera Lasius, Acanthomyops, and Formica, is relatively
well known on morphological and zoogeographic grounds (Wilson
and Regnier 1971).

The comparisons are based on a compilation of literature records
for as many species and genera as possible in this tribe (Figs. 1 and 2).
A genus was included if there were records for three or more species.
About a third of the species of Acanthomyops, of Cataglyphis, of
Lasius, and of Myrmecocystus are represented in the records cited
here, but a smaller fraction of the large genus Formica. These
genera are all from North Temperate latitudes. (For a preliminary
report see McCluskey, 1972.)

The workers could be classified as nocturnal, diurnal, etc. But in
the absence of single or definite beginning points or midpoints of
activity in most of the literature records, another method was used
to reduce each rhythm to one point for comparison with other species:
If the ants do not normally come above ground at all (e.g., Acan-
thomyops species), the species is plotted as an X at the extreme left
(Fig. 1) ; if nocturnal only, one position farther to the right; if out

295

296

Psyche

[December

as late as sunrise, another position to the right; etc. As far as pos-
sible, only summer records were used so as to make directly com-
parable.

It can be seen that the species in Acanthornyops , in Formica , and
in Cataglyphis are closely grouped within each genus. The species
in Lasius appear less so, but they barely overlap those of Formica or
Cataglyphis.

(none) night sunrise morning midday

X

X

x Acanthomyops

X

X

X

Lasius

X X

X

(X)

X X

X

X

Myrmecocystus

(X)

X (X) X

(X)

X X

(X)

X X

X X

X X

Formica * 5

XXX

X

Cataglyphis

X

x

X

(X)

X

Figure I

1973]

McCluskey — Generic Diversity

297

Scoring an X in the leftmost column of Fig. i as “i”, next to the
left (“night”) as “2”, and finally the rightmost column as “5”,
permits an analysis-of-variance comparison of the variation within
a genus with the variation between the genera :

ss

df

ms

F

P<

among genera

66.1

4

16.5

16.5

.001

within genera

36.9

37

1.0

Thus the likeness within genera is greater than that between genera.

A different type of analysis confirms this conclusion. The “none”
aboveground activity was now omitted, since that is not really a time
character (thus eliminating Acanthomyops and some Lasius species) ;
the “night” species were arbitrarily considered as (out until) 5 AM,
“sunrise” 7 AM, “morning” 10 AM and “midday” 1 PM. These
hours were treated as angles of a circular distribution, and the mean
angles of the different samples (genera) were compared by Watson

Fig. 1. Worker surface activity (limited mainly to summer records, for
most direct comparison of species). Each X represents one species and
shows its nearest approach to midday; those based on the most limited
cited records are enclosed in (). The leftmost column “NONE” indicates
that the species does not usually come above ground at all. Following are
species and literature sources represented: FORMICA : bradleyi (Wheeler
and Wheeler 1963), dakotensis (Talbot 1971), exsectoides (Andrews 1927,
1929; McCook 1877), fusca (Morisita 1939), fusca lemani (Brian 1955),
lasioides (Wheeler and Wheeler 1970), neogagates (Talbot 1953), ob-
scuripes (Weber 1935), pallidefulva nitidwentris (Talbot 1946, 1953, 1965),
polyctena (Bruns 1954; Chauvin 1965a, b), pratensis (Stebaev 1971; Stebaev
and Reznikova 1972), Sibylla (Wheeler 1917), subintegra (Talbot and
Kennedy 1940), subnitens (Ayre 1958, 1959), subpilosa (Stebaev 1971;
Stebaev and Reznikova 1972), subpolita (Mallis 1941), ulkei (Holmquist
1928). LASIUS: emarginatus (Tohme 1969), flavus (Bernard 1968; Odum
and Pontin 1961; Talbot 1965; Wilson 1955), fuliginosus (Wilson 1955),
minutus (Kannowski 1959b; Talbot 1965), neoniger (Talbot 1946, 1953,

1965) , niger (Eidmann 1926; Morisita 1939), sitiens (Wilson 1955),

sitkacnsis (now pallitarsis ) (Talbot 1965; Wilson 1955), speculiventris
(Talbot 1965), umbratus (Starcke 1937; Talbot 1965; Wilson 1955).
ACANTHOMYOPS : claviger, interjectus , latipes, and murphyi (Talbot

1963; Wheeler and Wheeler 1963). MYRMECOCYSTUS: lugubris (Cole

1966) , mclliger orbiceps (now placodops) (Wheeler 1908b), mexicanus

(Cole 1966; LaRivers 1968), mexicanus hortideorum (McCook 1882),

mimicus (Cazier and Statham 1962; Leonard 1911), mojave (Cole 1966;
LaRivers 1968; Leonard 1911), pyramicus (Smith 1951), wheeleri (Snelling
1971). CATAGLYPHIS: albicans (Delye 1968), albicans viaticoides

(Tohme 1969), altisquamis (Tohme 1969), bicolor (Delye 1968; Pickles
1944; Tohme 1969; Wehner and Duelli 1971), bicolor setipes (Gupta 1970),
bombycina (Delye 1968), frigida (Tohme 1969), lucasi (Baroni Urbani 1969;
Delye 1964). An annotated table giving the details of support for Figs. 1
and 2 is available from the author.

298

Psyche

[December

AM

HOURS

PM

6

8

10 12 14 16

18 20

F

(f)

F

F

[ Formica

(f)

F

F F

F

F

F F F F

(F)

(f)

F

Lasius f

F

(f)

F

F

>F

F

F

F

F (F)

(F)

Acanthomyops f

F

F

F

Fig. 2. Flight hours. Each F represents for one species the halfway
point between the earliest and latest literature records of flight; () indicate
the most fragmentary records. The following are represented: FORMICA :
dakotensis (Talbot 1971), fusca (Kannowski 1959a; Talbot 1965), montana
(Kannowski 1963; Kannowski and Johnson 1969), neogagates (Talbot
1966), obscuripes (Clark and Comanor 1972; Talbot 1971), obscuri'ventris
(Talbot 1964), opaciventris (Scherba 1961), pallidefulva nitidiv entris (Tal-
bot 1948), pergandei (Kannowski and Johnson 1969), pratensis (Eidmann
1928; Wheeler 1908a), rufa (Donisthorpe 1927; Standen 1909), rufibarbis
(Forel 1874a), sanguinea (Forel 1874b), subintegra (Kannowski 1963;
Talbot and Kennedy 1940), subnitens (Ayre 1957), ulkci (Scherba 1958;
Talbot 1959). LASIUS: alienus (Gosswald 1932; Hall 1887), brunneus

(Forel 1874b; Schenck 1852), carniolicus (Kutter 1946), cmarginatus (Forel
1874b, 1928), flavus (Donisthorpe 1927; Forel 1874b; Talbot 1965; Wilson
1955), fuliginosus (Wilson 1955), ininutus (Kannowski 1959a; Talbot 1965),
nearcticus (Wilson 1955), neoniger (Kannowski 1963; Talbot 1945, 1965;
Wilson and Hunt 1966), nevadensis (Cole 1956), nlger (Donisthorpe 1927;
Forel 1874b), pallitarsis (Medler 1958; Talbot 1965), speculiventfis (Kan-
nowski 1959a; Talbot 1965), subumbratus (Kannowski 1971), umbratus
(Crawley 1913; Forel 1874a, 1875; Kannowski 1963; Rau 1934)*.

ACANTHOMYOPS: clavigcr (Talbot 1963, 1973), inter jectus (Talbot

1963), latipes (Gregg 1963; Talbot 1963, 1973), murphyi (Talbot 1963),
subglaber (Talbot 1973).

*The morning record (Rau 1934) is most unusual for this species (Kan-
nowski 1963) and I omitted it in plotting the midpoint in the graph. Craw-
ley (1913) and Forel (1874a, 1875) may possibly refer to a sibling species
of umbratus (cf. Wilson 1955).

1973]

McCluskey — Generic Diversity

299

and Williams’ (1956; cf. Batschelet 1965, but only for a two-
sample case) test: if the samples are considered random, Fql N_q —
[ (N-q) (2R-R) ] / [ (q— 1 ) (N-2R.) ] = 3.58 and P ’<f.05
[N = 33 (species), q — 4 (genera), R. refers to the combination
vector for all the species in each genus, and R refers to the com-
bination vector of all genera] .

Figure 2 shows that for the three genera of this tribe with enough
species records, the flight hours of the species within a genus are
strikingly similar, exceptions being the one morning species of Lasius
and the two afternoon species of Formica. Comparison of the flight
hours by the Watson and Williams test again shows the likeness
within genera to be greater than that between genera (P < .001).

It seems noteworthy that a single rhythm character would char-
acterize a genus this well. A preliminary test for generality of the
within-genus likeness of flight hour was made on eleven genera from
five subfamilies of ants (all of those from which literature records
of at least three species per genus were at hand). The Watson and
Williams test gives a value of P < .001 ; this is true also if the three
formicine genera are omitted and only the other eight considered.

By including all species records no matter what tribe, it is pos-
sible in a few cases to rank workers from early to late within each
given locality (McCluskey, unpublished). Again species appear
grouped generically, giving further suggestive evidence of a taxo-
nomic or historical explanation of the phase differences (as opposed
to a strictly ecological or geographical explanation).

An example of a physiological character earlier shown to correlate
well with a previously-established classification is that by Priesner
(1968). Male moths throughout the family Saturniidae were shown
to react to pheromone from the females in a manner roughly cor-
responding to taxonomic grouping.

It should be added that some of the phase relationships considered
here might in fact be explained by response to temperature, since
these are records from the field rather than a constant temperature
laboratory. But this consideration does not change the fundamental
nature of the conclusion drawn. Further, certain ants which have
been studied in constant temperature exhibit a phase relationship to
the light cycle similar to that in the field (McCluskey 1963, 1965,
1969; McCluskey and Carter 1969).

Summary

Literature records of field mating flight and worker rhythms in-
dicate the phase to be much more alike from species to species within

300

Psyche

[December

a genus than between genera. The analysis is based on about a third
of the world species in each of several genera of the tribe Formicini.
Comparison in more genera, from several subfamilies, again indicates
this prominent within-genus likeness of behavior.

Acknowledgements

I thank P. Y. Yahiku for help with statistics, and L. R. Brand,
D. G. Kissinger, and R. E. Ryckman for reading the manuscript.

Literature Cited

Andrews, E. A.

1927. Ant-mounds as to temperature and sunshine. J. Morphol. 44: 1-20.

1929. The mound-building ant, Formica exsectoides F., associated with
tree-hoppers. Ann. Entomol. Soc. Amer. 22: 369-391.

Ayre, G. L.

1957. Ecological notes on Formica suhnitens Creighton (Hymenoptera:
Formicidae). Insectes Soc. 4: 173-176.

1958. Some meteorological factors affecting the foraging of Formica
subnitens (Hymenoptera: Formicidae). Insectes Soc. 5: 147-157.

1959. Food habits of Formica subnitens Creighton (Hymenoptera: For-
micidae) at Westbank, British Columbia. Insectes Soc. 6: 105-114.

Baroni Urbani, C.

1969. Una nuova Cataglyphis dei monti delPAnatolia. Fragmenta En-
tomologica 6: 213-222.

Batschelet, E.

1965. Statistical methods for the analysis of problems in animal ori-
entation and certain biological rhythms. American Institute of
Biological Sciences, Washington, D.C. 57 p.

Bernard, F.

1968. Les fourmis (Hymenoptera Formicidae) d’Europe occidentale et
septentrionale, 360. Masson et Cie, Paris.

Brian, M. V.

1955. Food collection by a Scottish ant community. J. Anirn. Ecol. 24:
336-351.

Bruns, H.

1954. Z. Tierpsychol. 11: 151-154 (cited by Sudd 1967).

Cazier, M. A., and M. Statham.

1962. The behavior and habits of the myrmecophilous scarab Cre-
mastocheilus stathamae Cazier with notes on other species (Co-
leoptera: Scarabaeidae) . J. N. Y. Entomol. Soc. 70: 125-149.

Chauvin, R.

1965a. Enregistrement de l’activite dans les conditions naturelles sur
les pistes de Formica polyctena. Insectes Soc. 12: 131-144.

1965b. L’activite sur les pistes et les fourmilieres de Formica polyctena,
p. 47-53. In R. Chauvin [ed.], La distribution temporelle des
activites animales et humaines, Masson et Cie, Paris.

Clark, W. H., and P. L. Comanor.

1972. Flights of the western thatching ant, Formica obscuripes Forel,
in Nevada (Hymenoptera: Formicidae). Great Basin Naturalist
32: 202-207.

1973]

McCluskey — Generic Diversity

301

Cole, A. C.

1956. Studies of Nevada ants. II. A new species of Lasius (Chthono-
lasius) (Hymenoptera: Formicidae). J. Tennessee Academy

Science 31: 26-27.

1966. Ants of the Nevada Test Site. Brigham Young University Science
Bull., Biological Series 7(3) : 1-27.

Crawley, W. C.

1913. Further experiments on the temporary social parasitism in ants
of the genus Lasius, Fab., with a note on Antennophorus uhlmanni.
Entomologist’s Record Journal Variation 25 : 135-138.

Dei.ye, G.

1964. Sur le peuplement myrmecologique de quelques ergs du Sahara
nord-occidental. Travaux Institut Recherches Sahariennes Univ.
Alger 23: 165-169.

1968. Recherches sur 1’ecologie, la physiologie et l’ethologie des fourmis
du Sahara. Thesis, Marseille. 155 p.

Donisthorpe, FI.

1927. British ants, their life-history and classification, 2nd ed. Rout-
ledge, London. 436 p.

Eidmann, H.

1926. Die koloniegriinding der einheimischen ameisen. Z. Vergl. Phy-
siol. 3: 776-826 (cited by Wilson 1955).

1928. Weitere beobachtungen uber die koloniegrundung einheimischer
ameisen. Z Vergl. Physiol. 7: 39-55 (cited by Kannowski 1963).

Forel, A.

1874a. Denkschr. Schweiz. Ges. Naturw. 26: 407-408 (cited by Donis-
thorpe 1927).

1874b. Les fourmis de la Suisse. Nouv. Mem. Soc. Helv. Sc. Nat.
Zurich 26: 447 p. (cited by Kannowski 1963).

1875. Bull. Soc. Vaudoise Sci. Natur. 14: 69 (cited by Donisthorpe
1927).

1928. The social world of the ants, vol. 1, p. 416. Putnam’s, New York.

Gosswald, K.

1932. Okologische studien fiber die ameisenfauna des mittleren mainge-
bietes. Z. Wiss. Zool. 142: 1-156.

Gregg, R. E.

1963. The ants of Colorado, p. 488. University of Colorado, Boulder.

Gupta, C. S.

1970. Ecological studies on the ant Myrmecocystus setipes Forel. Guru-
kul Kangri Vishwavidyalaya Journal Scientific Research 2:
134-148.

Hall, C. G.

1887. Swarms of Lasius niger, L., var. alienus, Forst., near Dover.
Entomologist’s Monthly Magazine 24: 91.

Holmquist, A. M.

1928. Notes on the life history and habits of the mound-building ant,
Formica ulkei Emery. Ecology 9: 70-87.

Kannowski, P. B.

1959a. The flight activities and colony-founding behavior of bog ants
in southeastern Michigan. Insectes Soc. 6: 115-162.

1959b. The use of radioactive phosphorus in the study of colony dis-
tribution of the ant Lasius minutus. Ecology 40: 162-165.

302

Psyche

[December

1963. The flight activities of formicine ants Symp. Genet. Biol. Ital.
12: 74-102.

1971. Unusual occurrence of winged ants in beach drift. Prairie Natu-
ralist 3 : 61-64.

Kannowski, P. B., and R. L. Johnson.

1969. Male patrolling behavior and sex attraction in ants of the genus
Formica. Anim. Behav. 17: 425-429.

Kutter, H.

1946. Lasius ( Chthonolasius ) carniolictis Mayr, eine neue Schweizera-
meise. Mitt. Schweiz. Entomol. Ges. 19: 698-699 (cited by Wilson
1955).

LaRivers, I.

1968. A first listing of the ants of Nevada (Hymenoptera : Formici-
dae). Biological Society Nevada Occasional Papers 17: 1-12.

Leonard, P.

1911. The honey ants of Point Loma, California. Trans. San Diego
Society Natural History 1 (3): 85-97.

Mallis, A.

1941. A list of the ants of California with notes on their habits and
distribution. Southern California Acad. Sci. Bull. 40: 61-100.

McCluskey, E. S.

1963. Rhythms and clocks in harvester and Argentine ants. Physiol.
Zool. 36: 273-92.

1965. Circadian rhythms in male ants of five diverse species. Science
150: 1037-1039.

1969. Field and laboratory timing of the flights of a harvester ant.
American Zoologist 9: 566 (abstract).

1972. Rhythm characters distinguish certain genera of ants. American
Zoologist 12(3):xxviii (abstract).

McCluskey, E. S., and C. E. Carter.

1969. Loss of rhythmic activity in female ants caused by mating. Comp.
Biochem. Physiol. 31: 217-226.

McCook, H. C.

1877. Mound-making ants of the Alleghenies, their architecture and
habits. Trans. Amer. Entomol. Soc. 6: 253-296.

1882. The honey ants of the Garden of the Gods, and the Occident ants
of the American plains, p. 29-30. Lippincott, Philadelphia.

Medler, J. T.

1958. A note on swarming and emergence of ants (Hymenoptera:
Formicidae). Proc. Entomol. Soc. Wash. 60: 258.

Morisita, M.

1939. On the daily rhythm of activity of ants. II, Activity of Lasius
niger L. Ecol. Rev. (Sendai) 5: 179-184 (Japanese, with English
summary) .

Odum, E. P., and A. J. Pontin.

1961. Population density of the underground ant, Lasius flavus, as
determined by tagging with P32. Ecology 42: 186-188.

Pickles, W.

1944. The territory and activities of the Algerian ant Cataglyphus
bicolor F. s-sp. mcgalocola Foist. Entomologist’s Monthly Maga-
zine 80 : 61-63.

1973]

McCluskey — Generic Diversity

303

Priesner, E.

1968. Die interspezifischen wirkungen der sexuallockstoffe der Satur-
niidae (Lepidoptera) . Z. Vergl. Physiol. 61: 263-297.

Rau, P.

1934. Notes on the behavior of certain ants of St. Louis Co., Mo. Trans.
Acad. Sci. St. Louis 28: 207-215.

Schenck, A.

1852. Beschreibung nassauischer ameisenarten. Jahrb. Ver. Naturk.
Nassau 8(1) : 3-149 (cited by Kannowski 1963).

Scherba, G. M.

1958. Reproduction, nest orientation and population structure of an
aggregation of mound nests of Formica ulkei Emery (Formici-
dae). Insectes Soc. 5: 201-213.

1961. Nest structure and reproduction in the mound-building ant For-
mica opaciventris Emery in Wyoming. J. N. Y. Entomol. Soc. 69:
71-78.

Smith, M. R.

1951. Two new ants from western Nevada (Hymenoptera : Formici-
dae). Great Basin Naturalist 11 : 91-96.

Snelling, R. R.

1971. Studies on California ants. 6. Three new species of Myrmeco-
cystus (Hymenoptera: Formicidae). Los Angeles County Museum
Contributions Science 214: 1-16.

Standen, R.

1909. Gull and ants. Lancashire Nat. 2: 190 (cited by Kannowski
1963).

Starcke, A.

1937. Retouches sur quelques fourmis d’Europe. II. Lasius groupe
umbratus Nylander. Tijdschr. Ent. 80: 38-72 (cited by Wilson
1955).

Stebaev, I. V.

1971. Structure of the protected territory of Formica pratensis and
interaction of ants from the same and different species. Zool.
Zh. 50: 1504-1519 (Russian, with English summary).

Stebaev, I. V., and J. I. Reznikova.

1972. Two interaction types of ants living in steppe ecosystem in
South Siberia, USSR. Ekologia Polska 20: 103-109.

Sudd, J. H.

1967. An introduction to the behaviour of ants. Arnold, London. 200 p.

Talbot, M.

1945. A comparison of flights of four species of ants. Amer. Midland
Naturalist 34: 504-510.

1946. Daily fluctuations in aboveground activity of three species of
ants. Ecology 27: 65-70.

1948. A comparison of two ants of the genus Formica. Ecology 29:
316-325.

1953. Ants of an old-field community on the Edwin S. George Reserve,
Livingston County, Michigan. Contributions Laboratory Vetebrate
Biology University of Michigan 63 : 1-13.

304

Psyche

[December

1959. Flight activities of two species of ants of the genus Formica.
Amer. Midland Naturalist 61: 124-132.

1963. Local distribution and flight activities of four species of ants of
the genus Acanthomyops Mayr. Ecology 44: 549-557.

1964. Nest structure and flights of the ant Formica obscuriventris
Mayr. Anim. Behav. 12: 154-158.

1965. Populations of ants in a low field. Insectes Soc. 12: 19-48.

1966. Flights of the ant Aphaenogaster treatae. J. Kans. Entomol. Soc.
39: 67-77.

1971. Flights of the ant Formica dakoiensis Emery. Psyche 78: 169-179.
1973. Five species of the ant genus Acanthomyops (Hymenoptera:
Formicidae) at the Edwin S. George Reserve in southern Michi-
gan. Great Lakes Entomologist 6: 19-22.

Talbot, M., and C. H. Kennedy.

1940. The slave-making ant, Formica sanguinea subintegra Emery, its
raids, nuptual flights and nest structure. Ann. Entomol. Soc.
Amer. 33: 560-577.

Tohme, G.

1969. Repartition geographique des fourmis du Liban (Hymenoptera:
Formicoidea) . Thesis, Toulouse. 77 p.

Watson, G. S., and E. J. Williams.

1956. On the construction of significance tests on the circle and the
sphere. Biometrika 43 : 344-352.

Weber, N. A.

1935. The biology of the thatching ant, Formica rnfa obscuripes Forel,
in North Dakota. Ecol. Monogr. 5: 165-206.

Wehner, R., and P. Duelli.

1971. The spatial orientation of desert ants, Cataglyphis bicolor before
sunrise and after sunset. Experientia 27: 1364-1366.

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

1963. The ants of North Dakota. University of North Dakota, Grand
Forks. 207 p.

1970. The natural history of Manica (Hymenoptera: Formicidae).

J. Kans. Entomol. Soc. 43 : 129-162.

Wheeler, W. M.

1908a. Comparative ethology of the European and North American
ants. J. Psychol. Neurol. 13: 404-435 (cited by Kannowski 1963).
1908b. Honey ants, with a revision of the American Myrmecocysti.

Bull. Amer. Mus. Natur. Hist. 25: 345-397.

1917. The mountain ants of western North America. Amer. Acad.
Arts Sci., Proc. 52: 457-569.

Wilson, E. O.

1955. A monographic revision of the ant genus Lasius. Mus. Comp.
Zool. Bull. 113 : 1-201.

1971. The insect societies, p. 214. Harvard University, Cambridge.
Wilson, E. O., and G. L. Hunt.

1966. Habitat selection by queens of two field-dwelling species of ants.
Ecology 47: 485-487.

Wilson, E. O., and F. E. Regnier, Jr.

1971. The evolution of the alarm-defense system in the formicine ants.
Amer. Natur. 105: 279-289.

THE MALE GENITALIA OF BLATTARIA.

XI. PERISPHAERIINAE*

By Louis M. Roth
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01760

In this paper I shall illustrate and characterize the male genitalia
of 11 genera (indicated with an * in Table 1) of cockroaches which
belong to the Perisphaeriinae. Three recent classifications which
include these genera are shown in the table. I follow McKittrick
(1964) in placing all of those discussed here under the Blaberidae:
Perisphaeriinae, and Beier (1961) in using tribes to separate them.
Rehn (1922) discussed the characteristics of Pilema, Cyrtotria , Dero-
calymma , Zuluia , and Hostilia, and Princis (1963) devised keys to
some South African species of the above genera, as well as those of
Bantua , P erisphaeria, and Blepharodera.

Materials and Methods

All genitalia were treated with 10% KOH, dehydrated, cleared,
and mounted in Permount. The source of each of the specimens
examined is given using the following abbreviations: (ANSP) =

Academy of Natural Sciences, Philadelphia; (BMNH) = British
Museum (Natural History), London; (CUZM) = Zoological
Museum, Copenhagen; (L) = Zoological Institute, Lund, Sweden;
(HO) = Hope Department of Entomology, Oxford, England;
(M'CZ) = Museum of Comparative Zoology, Harvard University,
Cambridge, Mass. Locality data and names of specialists who identi-
fied the specimens, if known, follow these abbreviations; most of the
species were determined by Princis or Rehn. The number preceding
the abbreviations refers to the number assigned the specimen and its
corrsponding genitalia (on a slide) Which are deposited in their re-
spective museums.

Results and Discussion

The normal arrangement of the phallomeres of male genitalia of
the Perisphaeriinae, as seen in dorsal view, is illustrated in figure 1 .
Based on male genitalia, I suggest the erection of 4 tribes for the

* Manuscript received by the editor October 10, 1973

305

306

Psyche

[December

Table i. Three recent classifications of some of the species of
Blattaria discussed in this paper (marked with an •)

Beier(i96i) McKittrick ( 1964)

Princis (i960)
Derocalymmidae
Apterinae

Apt era Sauss.

Elliptica
Sauss. & Zehn.

Elliptohlatta
Sauss. & Zehn.

Gymnonyx
Sauss. & Zehn.

Thoracopygia
Sauss. & Zehn.

Derocalymminae

* Bantua Shelf.
Compsagis
Chop.

*Cyrtotria

Stal

* Derocalymma
Burm.

Glomerihlatta

Bey-Bienko

*Hostilia

Stal

*P erisphaerus
Serv.

*Pilema Sauss.
Platysilpha
Shelf.

Pronaonota
Sauss. & Zehn.

*Pseudoglo?neris

Brunner

*Zuluia Rehn

Blaberidae

Perisphaeriinae

Apterini

Aptera

Elliptica

Elliptohlatta

Gymnonyx

Thoracopygia

Derocalymmini
Peris phaerus
Glomerihlatta
Pseud ogho m eris
Pronaonota
Cyrtotria
Host ilia
Compsagis
Derocalymma
Platysilpha
and others

Perisphaeriini
Perisphaeria
Blepharodera
Poecilohlatta
Paras phaeria
Gyna

Paranauphoeta
Paraplecta
and others

Brachycolinic
Brachycola Serv.
Hormetica Burm.
P arahor m etica
Brunner
and others

Blaberidae
Perisphaeriinae
Perisphaeria
(— II yposphaeria )
Derocalymma
Eustegasta Gerst.a
Gyna
Pilema
Perisphaerus
Isoniscus Borg.a

1973]

Roth — Blattaria

307

Princis (i960) Beier (1961) McKittrick (1964)

Perisphaeriidae
Gyninae
Evea Shelf.

Gyna Brunnerb
Paranauphoeta
Brunner

Paraplecta Shelf.

Progonogamia

Rehn

Thliptoblatta
Sauss. & Zehn.

Perisphaeriinae

*Blepharodera

Burm.

Parasphaeria

Brunnerd

*Perisphaeria

Burm.

Poeciloblatta
Sauss. & Zehn.

*Trichoblatta
Sauss. & Zehn.

Zetoborinaee
Lanxoblatta Hebard
Phortioeca Sauss.

Schistopeltis Rehn
Schizopilia Burm.

Tribonoidea Shelf.

T ribonium Sauss.

Zetobora Burm.

Zetoborella Hebard

aPrincis (1960) placed these 2 genera in Blaberidae: Laxtinae. Beier
(1961) placed Eustcgasta in the Panchlorinae : Laxtini. The $ genitalia
of Panchlorinae are characterized by a marked reduction or absence of one
or more of the phallomeres (Roth, 1971; Gurney and Roth, 1972). The
genitalia of Eustegasta are quite different from members of the Panchlorinae
(unpublished observations).

bThe genitalia of Gyna spp. have been described (Roth, 1972).

°I (1970a) followed McKittrick (1964) and placed this tribe in Blaberi-
nae.

dThe genitalia of Parasphaeria ovata (Blanchard) (South America) is
more typical of members of the Epilamprinae and I placed it in this sub-
family (Roth, 1973).

eI (1970) followed McKittrick (1964) in placing this subfamily closer to
the Blaberinae (Blaberoid Complex) than to the Perisphaeriinae (Epilam-
proid Complex).

308

Psyche

[December

Figure 1. (62 BMNH). Male genitalia (dorsal) of Cyrtotria latipennis

(Kirby), British East Africa (from $ shown in Fig. 37). Left (L) and
right (R) phallomeres numbered according to McKittrick (1964); C =: cleft
of LI; UL and LL — upper and lower lobes of LI, respectively.

genera studied here; additional tribes will undoubtedly be required
for other genera of this subfamily (unpublished observations).

Perisphaeriini. — The upper lobe of Li is usually about the same
height or slightly larger than the lower lobe and it has a snoutlike
extension (accidentally lost in some of the illustrated preparations) ;
setae are absent from the lower lobe. The L2d is variable in shape*
its surface distinctly spicular (at least in part), or reticulate (at
least in part) and usually with a smooth lateral extension on the
right side. R2 with or without a subapical incision. In this tribe
I include the African genera Peris phaeria, Blepharodera , Pilema ,.
Hostilia, Bantua, and Cyrtotria , which can be further subdivided
into groups.

Tribal Group I. The entire or at least part of the surface of L2d
is spicular (not reticulate) : Perisphaeria, Blepharodera, Pilema, and
Hostilia.

1973]

Roth — Blattaria

309

Figures 2-9. Adult males of Perisphaeria spp. 2. (1424 L). P. virescens
(Brunner), (det. Princis). 3. (40 CUZM). P. scabra (Thunb.). Cap
bono Spei, Dec. 1816. 4. (1420 L). P. scabra. Holub, South Africa (det.
Princis). 5. (1419 L). P. saxicola (Esch.) Capetown, Table Mountain,
2400' (det. Princis). 6. (1422 L). P. ruficornis (Sauss. and Zehn.). Swart-
bergpas, Platberg, Cape Province, South Africa, ca. 5000' (det. Princis).
7. (1421 L). P. stylifera Burm. Manhica (det. Princis). 8. (161 ANSP).

P. stylifera. Durban, (det. by Rehn as Hyposphaeria giebeliana (Sauss.
and Zehn.). 9. (1423 L). P. aspera (Walker). Baberton, Levetzow (det.
Princis). (scale = 5 mm).

3io

Psyche

[December

Figures 10-18. Male genital phallomeres of Perisphaeria spp. 10-12.
(1424 L). P. vires cens (from $ shown in Fig. 2). 13-15. (40 CUZM).
P. scabra (from $ shown in Fig. 3). 16-18. (1420 L). P. scabra (from $
shown in Fig. 4). (scale = 0.2 mm).

Figures 19-27. Male genital phallomeres of Perisphaeria and Cyrtotria
spp. 19-21. (1419 L). P. saxicola (from $ shown in Fig. 5). 22-24. (1422
L). P. ruficornis (from $ shown in Fig. 6). 25-27. (15 HO). Cyrtotria
sp. (from $ shown in Fig. 105). (scale = 0.2 mm).

1973]

Roth — Blattaria

312

Psyche

[December

Figures 28-36. Male genital phallomeres of P erisphaeria spp. 28-30.
(1421 L). P. stylifera (from $ shown in Fig. 7). 31-33. (161 ANSP).
P. stylifera (from $ shown in Fig. 8). 34-36. (1423 L). P. aspera (from
$ shown in Fig. 9). (scale = 0.2 mm).

1973]

Roth — Blattaria

313

Figures 37-42. Adult males of Cyrtotria, Pilema, and Hostilia spp. 37.
(62 BMNH). C. latipenms (holotype of Thysanoblatta latipennis Kirby)
British East Africa. 38. (53 BMNH). P. dentata Sauss. and Zehn. Pre-
toria, South Africa (det. Princis). 39. (989 L). P. cribrosa (Sauss.).
Pretoria, South Africa (det. Princis). 40. (990 L). P. thoracica (Walker).
Mamathes, Basutoland (det. Princis). 41. (991 L). P. reflexa (Walker).

Zululand, Mtubatuba Dist., Dukuduku, South Africa (det. Princis). 42.
(1406 L). H. clavigera (Kirby). Blouberg, Transvaal (det. Princis).
(scale = Figs. 37, 41, 42 = 5 mm; Figs. 38-40 = 10 mm).

[December

314

Psyche

48

49

50

1973]

Roth — Blattaria

315

Figures 43-50. Adult males of Hostilia , Blepharodera, and Bantua spp.
43. (170 ANSP). H. proterva (Stal). Pinetown, Natal, South Africa (det.
Rehn). 44. (1405 L). H. proterva. Stanford Hill, Durban, Natal, South
Africa (det. Princis). 45. (1426 L). Blepharodera discoid'a'is (Brunner).
Cap. 1872 (det. Princis). 46. (1425 L). B. ciliata Burm. Hopefield, C.P., X.
1883 (det. Princis). 47. (34 CUZM). B. ciliata. Cap. bono Spei, June 1917
(det. Princis). 48. (1404 L). Bantua scabra (Walker). Okahandja, S.W.
Africa (det. Princis). 49. (999 L). B. repentina Rehn. Kruger National Park,
South African Transvaal (det. Princis). 50. (1403 L). B. robusta. (Shelf.)
Sandspruit, 15 miles northeast of Petersburg, South African Transvaal (det.
Princis). (scaled 5 mm).

3i6

Psyche

[December

Figures 51-58. Male genital phallomeres of Hostilia spp. 51-53. (1405
L). H. proterva (from $ shown in Fig. 44). 54-56. (170 ANSP). H.
proterva (from $ shown in Fig. 43). 57-58. (1406 L). H. clcwigera (from
$ shown in Fig. 42). (scale = 0.2 mm) .

1973]

Roth — Blattaria

317

Figures 59-70. Male genital phallomeres of Pilema spp. 59-61. (989 L).
P. crihrosa (from $ shown in Fig. 39). 62-64. (53 BMNH). P. dentata
(from $ shown in Fig. 38). 65-67. (900 L). P. thoracica (from $ shown
in Fig. 40). 68-70. (991 L). P. reflexa (from $ shown in Fig. 41).

(scale = 0.2 mm) .

3i8

Psyche

[December

Figures 71-79. Male genital phallomeres of Blepharodera spp. 71-73.
(1426 L). B. discoidalis (from $ shown in Fig. 45). 74-76. (34 CUZM).
B. discoidalis (from $ shown in Fig. 47). 77-79. (1425 L). B. ciliata
(from $ shown in Fig. 46) . (scale = 0.2 mm) .

1973]

Roth — Blattaria

319

Figures 80 88. Male genital phallomeres of Bantua spp. 80-82. (1403 L).
B. robusta (from $ shown in Fig. 50). 83-85. (148 ANSP). B. robusta
(Shelford), paratype of Bantua armigera Rehn. 86-88. (54 BMNH). B.
dispar. Distant Coll, (labeled Derocalymma pallipcs Kirby), (scale =
0.2 mm).

320

Psyche

[December

Figures 89-97. Male genital phallomeres of Bantaa and Cyriotria spp.
89-91. (999 L). B. repentina (from $ shown in Fig. 49). 92-94. (1404 L).
B. scabra (from $ shown in Fig. 48). 95-97. (993 L). C. graniger (from
$ shown in Fig. 99). (scale — 0.2 mm) .

1973]

Roth — Blattaria

3 2

In Perisphaeria 2 species groups can be distinguished based on the

R2 :

Species Group i. R2 with a distinct, deep, subapical incision
(Fig. 23). P. stylifera (Figs. 7-8, 28-33), P- aspera (Figs. 9, 34-
36); these 2 species also have very similar shaped L2d’s (Figs. 28,
34) ; P. ruficornis (Figs. 6, 22-24).

Species Group 2. R2 without a subapical incision, or, if present,
weakly represented. P. virescens (Figs. 2, 10-12) has a slight in-
cision in the middle of the hook (Fig. 11) and its L2d lacks any
extension (Fig. 10). P. scahra (Figs. 3, 13-18) and P. saxicola
(Figs. 9, 19-21) lack incisions in their R2’s (Figs. 14, 17, 20) ; both
species have smooth outgrowths arising from the right side of the
spicular L2d (Figs. 13, 16, 19).

In Hostilia (Figs. 42-44, 51-58) the lower margin and left side
of L2d is spicular and a smooth hook-like extension arises from the
upper right corner (Figs. 51, 54, 57). R2 has a shallow subapical
incision (Figs. 52, 55).

In Pilema (Figs. 39-41, 59-70), the outer margin of R2 is rela-
tively straight. Three species, cribrosa , dentata, and thoracica have
deep subapical incisions (Figs. 60, 63, 66), whereas reflexa (Fig. 69)
has a relatively shallow one. Thoracica and reflexa have smooth
outgrowths from the right side of L2d (Figs. 65, 68), which appear
to be lacking or weakly indicated in cribrosa and dentata (Figs. 59,
62).

In Blepharodera (Figs. 45-47, 71-79) the spicules on L2d are
very minute; the L2d outgrowth is broad in P. discoidalis (Fig. 71)
and much narrower in B. ciliata (Figs. 74, 77). R2 of discoidalis
has a minute lobe at about the middle of the hook (Fig. 72), whereas
in ciliata there is no subapical incision (Figs. 75, 78).

Tribal Group II. Part of the L2d surface on the left is reticulate
(not spicular) (e.g. Figs. 80, 121), but otherwise very similar to
Group I having a lateral extension on the right: Bantua (Figs. 48-
50, 80-94) and Cyrtotria (Figs. 19-21, 94-144).

All of the 5 species of Bantua examined have R2’s with deep sub-
apical incisions that extend almost to the middle of the curved part
of the hook (Figs. 81, 84, 87, 90, 93). Their L2d’s are also basi-
cally similar in having a smooth extension on the right side arising
from the reticulate portion of the sclerite (Figs. 80, 83, 86, 89, 92).

Cyrtotria is very close to Bantua as evidenced by the similarity of
the L2d’s of some of the species. There are 2 groups of Cyrtotria
based on the presence or absence of a subapical incision:

Species Group 1. Subapical incision on R2 absent. C. pallicornis
(Figs. 108, 109-m), C. inarshalli (Figs. 106, 107, 121-126), C.
latipennis (Figs. 1, 3, 7, 101, 136-144).

322

Psyche

[December

1973]

Roth — Blattaria

323

Figures 98-108. Adult males of Cyrtotria spp. 98. (63 BMNH). C.
gibhicollis (Stal). Natal, (labeled Perisphaeria linearis Walker, selected
as a lectotype by K. Princis). 99. (993 L). C. graniger (Sauss. and Zehn.).
Mudugh Prov., Somalia (det. Princis). 100. (997 L). C. tuherculata Shel-
ford. Lukuledi, German East Africa (det. Princis). 101. (151 ANSP). C.
latipennis (Kirby). N.E. Africa (det. Princis; Rehn det. the specimen as
C. capucina, but the genitalia are similar to the type of latipennis (cf. Figs.
142-144 and 136-138) ; the L2d of capucina (Fig. 130) differs from that of
latipennis (Fig. 136). 102. (998 L). C. orienialis (Chopard). Kasenyi, East
Congo (idet. Princis). 103. (56 CUZM). C. orientalis Kasenyi, East Congo.
104. (996 L). C. capucina (Gerst.). Kakamego (det. Princis). 105. (15 HO).
C. sp. Damaraland, South Africa. Shelford selected this specimen as the
$ holotype of C. poduriformis (Walker) but Princis (personal communi-
cation) claimed that he had no authority to select an additional type to
the $ holotype in the British Museum; the L2d surface (Fig. 25) and out-
growth differ somewhat from the L2d of the specimen which Princis believes
to be poduriformis (Fig. 133). 106. (995 L). C. marshalli Shelford. Es-
pungabera (det. Princis). 107. (14 HO). C. marshalli, holotype, Salisbury,
Mashonaland, 5000'. 108. (992 L). C. pallicornis (Kirby). Pretoria, South
African Transvaal (det. Princis). (scale = 5 mm).

324

Psyche

[December

Figures 109-117. Male genital phallomeres of Cyrtotria spp. 109-111.
(992 L). C. pallicornis (from $ shown in Fig. 108). 112-114. (55 CUZM).
C. tuberculata. Fsiolo, Kenya (det. Princis). 115-117. (56 CUZM). C.
orientalis (from $ shown in Fig. 103). (scale — 0.1 mm).

1973]

Roth — Blattaria

325

Figures 118-126. Male genital phallomeres of Cyrtotria spp. 118-120.
(63 BMNH). C. gibbicollis (from $ shown in Fig. 98). 121-123. (14 HO)..
C. marshalli (from $ shown in Fig. 107). 124-126. (995 L). C. marshallt
from $ shown in Fig. 106). (scale =0.2 mm).

Psyche

[December

Figures 127-135. Male genital phallomeres of Cyrtotria spp. 127-129.
(998 L). C. orientalis (from $ shown in Fig. 102). 130-132. (996 L).
C. capucina (from $ shown in Fig. 104). 133-135. (55 BMNH). C. podu-
riformis. Natal, South Africa (det. Princis). (scale = 0.1 mm).

1973]

Roth — Blattaria

327

Figures 136-144. Male genital phallomeres of Cyrtotria latipennis. 136-
138. (62 BMNH) . (from $ shown in Fig. 37). 139-141. (995 L). Kitui

(det. Princis). 142-144. (151 ANSP). (from $ shown in Fig. 101). (scale
=: 0.1 mm) .

328

Psyche

[December

Figures 145-148. Adult males of Zuluia spp. 145. (1408 L). Z. litho-
strota Rehn. Letsletle Valley, Gravelotte Dist., South African Transvaal
(det. Princis). 146. (154 ANSP). Z. pubescens (Sauss.). Durban, paratype
of Z. mitella Rehn. 147. (145 ANSP). Z. abscissa (Walker). Durban
(det. Rehn; Princis [1960, p. 141] states that Rehn’s identification of ab-
scissa was an error and that the specimen was Z. lithostrota Rehn. The
L2d of this specimen (Fig. 158) is more like that of abscissa (Fig. 155)
than lithostrota (Fig. 149)). 148. (1407 L). Z. abscissa (det. Princis).

(scale = 5 mm).

1973]

Roth — Blattaria

329

Figures 149-160. Male genital phallomeres of Zuluia spp. 149-151.
(1408 L). Z. lithostrota (from $ shown in Fig. 145). 152-154. (154 ANSP).
Z. pubescens (from $ shown in Fig. 146). 155-157. (1407 L). Z. abscissa
(from $ shown in Fig. 148). 158-160. (145 ANSP). Z. abscissa (from $
shown in Fig. 147). (scale = 0.1 mm).

330

Psyche

[December

Species Group 2. Subapical incision on R.2 present. C. graniger
(Figs. 95-97, 99), C. orient alis (Figs. 102, 103, 127-129), C. capu-
cina (Figs. 104, 130- 132), C. poduriformis (Figs. 133-135), C.
tuberculata (Figs. 100, 112-114), C. gibbicollis (Figs. 98, 118-120),
C. sp. (Figs. 25-27, 105).

Zuluiinii — L2d is an unevenly outlined plate, the left side of
which is scale-like or smooth; R2 with a subapical incision. Li sim-
ilar to Bantua. I place Zuluia (Figs. 145-160), an African genus,
in this tribe. Z. abscissa (Figs. 155, 158) has a lobe-like extension
on the left side of L2d which is absent in Z. lithostrota (Fig. 149)
and Z. pubescens (Fig. 152).

Rehn (1932) stated that Zuluia is a relative of Derocalymma and
H ostilia, but in general, the relationship appeared closer to Hostilia.
The cf genital phallomeres do not support this suggestion.

Derocalymmini. — Li, which is very similar in all the species
examined, has a deep cleft which may curve upwards slightly;
normally the margins of the 2 lobes are close together (e.g., Figs.
187, 199), but in some preparations the phallomere was distorted by
pressure of the coverslip so that the sclerotized margins were widely
separated (e.g., Figs. 178, 196). The upper lobe is usually somewhat
narrower and more elongate than the lower lobe, and the latter lacks
setae. L2d is a broad flat sclerotized plate of variable shape, slightly
separated from L2vm, and whose surface appears weakly wrinkled
or scale-like. The R2, generally similar in all species examined, is
unmodified, lacking an apical notch or subapical incision. I place
Derocalymma (Figs. 161-227), an African genus, in this tribe.

Based on the shape of L2d, I separate 2 species groups:

Species Group 1. The L2d variable in shape, but lacking a
finger-like projection on the right side: D. lampyrina (Figs. 161,
1 73-1 78 ) , D. kalahari (Figs. 163, 182-184), D. granulata (Figs.
164, 185-190), D. porcellio (Figs. 165, 191-196), D. silphoides
(Figs. 166, 197-199), D. rajiformis (Figs. 167, 200-205), D. cru-
ralis (Figs. 168, 206-208), D. versicolor (Figs. 169, 209-2 14), D.
sp. (Figs. 162, 179-181 ).

Species Group 2. Distinct finger-like projection extending from
the right side of the L2d: D. costata (Figs. 170, 21 5-21 7), D.
pluteus (Figs. 1 71, 218-223), D. scruposa scruposa (Figs. 172,
224-226).

Rehn (1932) in a. study of Derocalymma , using female characters,
arranged the species in what seemed to him to be a natural phylo-
genetic order. He considered D. pluteus as the least specialized of
the genus (the $ shows no traces of caudo-lateral diverging “gut-

33i

1973]

Roth — Blattaria

Figures 161-166. Adult males of Derocalymma spp. 161. (37 CUZM).
D. lampyrina Gerstaecker. (det. Princis). 162. (141 ANSP). D. sp. Ger-
man East Africa (det. by Rehn as lampyrina and by Princis as versicolor ;
the L2d (Fig. 141) differs from both these species (see Figs. 173, 176 and
209, 212). 163. (1414 L). D. Kalahari Rehn. Khomashochland, S.W. Africa
(det. Princis). 164. (1413 L). D. granulata Sauss. and Zehn. S.W. Africa
(det. Princis). 165. (1416 L). D. porcellio Gerstaecker. Morogoro, German
East Africa (det. Princis). 166. (1411 L). D. silphoides Bolivar. Abachaus,
Darmarland (det. Princis). (scale = 5 mm).

332

[December

Psyche

Figures 167-172. Adult males of D erocalymma spp. 167. (38 CUZM).
D. rajiformis Rehn. Kenya (det. Princis). 168. (1410 L). D. cruralis
(Stal). Bipindi, South Cameroon (det. Princis). 169. (1412 L). D. versi-
color Burmeister. Southern Rhodesia (det. Princis). 170. (1417 L). D.
costata Rehn. Cameroon (det. Princis). 171. (1409 L). D. pluteus Rehn.
Haut-Luapula, Kansenia (det. Princis). 172. (1418 L). D. scruposa scru-
posa Rehn. Togo, Misahohi-Mitte (det. Princis). (scale = 5 mm).

1973]

Roth — Blattaria

333

Figures 173-181. Male genital phallomeres of Derocalymma spp. 173-
178. (1415 L). D. lampyrina. Okavango, Runtu, S.W. Africa (det. Princis).
176-178. (37 CUZM) . (from $ shown in Fig. 161). 179-181. (141 ANSP).
D. sp. German East Africa (from $ shown in Fig. 162). (scale = 0.1 mm).

334

Psyche

[December

Figures 182-190. Male genital phallomeres of Derocalymma spp. 182-184.
(1414 L). D. kalahari. (from $ shown in Fig. 163). 185-190. D. granulata.
185-187. (1413 L). (from $ shown in Fig. 164). 188-190. (143 ANSP).

Damara (det. Rehn). (scale = 0.1 mm) .

Figures 191-199. Male genital phallomeres of D erocalymma spp. 191-
196. D. porcellio. 191-193. (1416 L). (from $ shown in Fig. 165). 194-
196. (167 ANSP). Tendaguru, Lindi, German East Africa (det. Hebard).
197-199. (1411 L). D. silphoides (from $ shown in Fig. 166). (scale =
0.1 mm).

1973]

Roth — Blattaria

335

336

Psyche

[December

Figures 200-208. Male genital phallomeres of Derocalymma spp. 200-
205. D. rajiformis. 200-202. (38 CUZM). (from $ shown in Fig. 167).
203-205. (169 ANSP). Bukama (Musee du Congo) (det. Rehn). 206-208.
(1410 L). D. cruralis (from $ shown in fig. 168). (scale = 0.1 mm).

1973]

Roth — Blattaria

337

Figures 209-217. Male genital phallomeres of Derocalymma spp. 209-
214 D versicolor. 209-211. (142 ANSP). Gaberones, Bechuanaland (det.
Rehn). 212-214. (1412 L). (from $ shown in Fig. 169). 215-217. (1417 L).
D. costata (from $ shown in Fig. 170). (scale = 0.1 mm).

Psyche

[December

Figures 218-226. Male genital phallomeres of D erocalymma spp. 218-
223. D. pluteus. 218-220. (157 ANSP). Paratype, Mayumbe, Congo. 221-
223. (1409 L). (from $ shown in Fig. 171). 224-226. (1418 L). D. scru-
posa scruposa (from $ shown in Fig. 172). (scale = 0.1 mm).

1973]

Roth — Blattaria

339

ters”, the mediocephalic portion of the disk virtually without a me-
dian carinula, and the ventro-lateral faces of the pronotum are not
inflated or bullate, their surface deplanate). The L2d of pluteus
with its finger-like projection falls into group 2 along with costata
and scruposa, two species which are almost at the opposite extremes
of Rehn’s listing. The finger-like extension could represent a “primi-
tive” character which was lost in the species placed in group i.
However, it is also possible that the L2d of pluteus represents a more
highly evolved form with the extension developing from species which
lacked it.

Perisphaerini. — Unfortunately, the spelling of this tribe differs
only slightly from the Perisphaeriini, but Rehn (1951) used Peri-
sphaerus to erect the tribe based on wing venation.

Li of this tribe is distinctive in that the upper lobe is considerably
larger than the lower lobe, with a snoutlike projection over the cleft
opening and a patch of numerous, moderately spaced setae on the
lower lobe (e.g., Fig. 44). The preputial membrane is almost
entirely modified to form the L2d which is weakly separated from
the sclerotized L2vm ; a small part of the preputial membrane on the
right side may not be sclerotized, and minute microtrichia can be
seen in this region. The hook (R2) with (Figs. 240, 281, arrows)
or without an apical notch ; no other genera, here examined, have an
apical notch: Perisphaerus (Figs. 227-259), Pseudo gloineris (Figs.
260-262), and Trichoblatta (Figs. 263-289). Three of 16 known
species of Trichoblatta are African, the others, as well as all the
species of Perisphaerus and Pseudo gloineris are Asian and adjoining
islands.

The genera and species can be grouped according to the degree of
development of the apical notch on R2 :

a. Apical notch absent: Pseudo gloineris gloineris (Figs. 260-
262), Trichoblatta sericea (Figs. 264, 288-289).

b. Apical notch poorly defined: Perisphaerus flavipes (Figs.

257-259) (the shape of L2d of this species (Fig. 257) differs from
all the others of the tribe), P. armadillo (Figs. 233-235, 248-251),
and P. glomeriformis (Figs. 231-232, 254-256).

c. Apical notch well developed: Perisphaerus contiguus (Figs.
227, 236-238), P. flavicornis (Figs. 228, 239-241), P. semilunatus
(Figs. 230, 242-244), P. aterrimus (Figs. 229, 245-247), Tricho-
blatta magnifica (Figs. 268, 271-273), T. oniscina (Figs. 267, 274-
276), T. tarsalis (Figs. 264, 277-279), T. nigra (Figs. 269, 280
282), T. guerini (Figs. 266, 283-285).

340

Psyche

[December

Figures 227-235. Adult males of Perisphaerus spp. 227. (176 ANSP).
P. contiguus (Sauss.). Mt. Lamington, N.E. Papua (det. Hebard). 228. (39
CUZM). P. flavicornis Burm. Java. 229. (983 L). P. aterrimus (Herbst).
Kangean Island, Oengkokop (det. Princis). 230. P. semilunatus (Hanitsch).
Sakaerat, Khorat Province, Thailand (det. Princis). 231-232. (104 MCZ).
P. glomcriformis Lucas. Samar, Catarman, Philippine Islands (det. Princis).
232. Tooth (arrow) on underside of pronotum. 233-235. P. armadillo Ser-
ville. 233. (989 L). Prai Jawang, East Sumba (det. Princis). 234-235.
(109 MCZ). Zamboanga, Philippine Islands (det. Princis). 235. Tooth
(arrow) on underside of pronotum. (scale: Figs. 232, 235 — 1 mm; all

others — 5 mm).

1973]

Figures 236-244. Male genital phallomeres of Perisphaerus spp. 236-
238. (176 ANSP). P. contiguus (from $ shown in Fig. 227). 239-241. (39
CUZM). P. flavicornis (from $ shown in Fig. 228). 242-244. P. semi-
lunatus (from $ shown in Fig. 230). (scale = 0.1 mm).

Roth — Blattaria

342

Psyche

[December

Figures 245-253. Male genital phallomeres of Perisphaerus spp. 245-
247. (983 L). P. aterrimus (from $ shown in Fig. 229). 248-251. P. arma-
dillo. 248-249. (109 MCZ). (from $ shown in Fig. 234). 250-251. (982 L).
(from $ shown in Fig. 233). 252-253. (180 ANSP). P. aeneus (Brunner).
Sibuyan Island (det. Hebard). (scale = 0.1 mm).

Figures 254-262. Male genital phallomeres of Perisphaerus and Pseudo-
glomeris spp. 254-256. (104 MCZ). Perisphaerus glomeriformis (from $
shown in Fig. 231). 257-259. (51 BMNH). Perisphaerus flavipes (Sauss.).
Kinta Valley, Perak (det. Princis; this specimen was det. as Perisphaeria
lucasiana Sauss. and Zehn. (a synonym) byHanitsch). 260-262. (46 BMNH) .
Pseudoglomeris glomeris (Sauss.). Matale, Ceylon (det. Princis).

344

Psyche

[December

Figures 263-270. Adult males of Trichoblatta spp. 263. (987 L). T.
pygmaea (Karny). Ryukyu (det. Princis). 264. (985 L). T. tarsalis

(Walker). Kambaiti, N.E. Burma, 7000' (det. Princis). 265. (984 L). T.
sericea (Sauss.). Madras, India (det. Princis). 266. (8 BMNH). T. gucrini
Sauss. and Zehn. “Givalior” (det. Princis). 267. (988 L). T. oniscina
(Gerstaecker) . Urwald, Ukaika (det. Princis). 268. (986 L). T. magnified
(Shelford). Tonkin, Chapa, Indo-China (det. Princis). 269-270. (103 MCZ).
269. T. nigra (Shiraki). Formosa (det. Princis). 270. Tooth (arrow) on
underside of pronotum. (scale: Figs. 263-269 5 mm; Fig. 270= 1 mm).

1973]

Roth — Blattaria

345

p ppli

Figues 271-279. Male genital phallomeres of Trichoblatta spp. 271-273.
(986 L). T. magnified (from $ shown in Fig. 268). 274-276. (988 L). T.
oniscina (from $ shown in Fig. 267). 277-279. (985 L). T. tarsalis (from
$ shown in Fig. 264). (scale — 0.1 mm) .

346

Psyche

[December

Figures 280-289. Male genital phallomeres of Trichoblatta spp. 280-282.
(103 MCZ) . T. nigra (from $ shown in Fig. 269). 283-285. (8 BMNH) .
T. guerini (from $ shown in Fig. 266). 286-287. (987 L). T. pygmaea
(from $ shown in Fig. 263). 288-289. (984 L). T. sericea (from $ shown
in Fig. 265). (scale =0.1 mm).

1973]

Roth — Blattaria

347

Some genera of Perisphaeriincie have a characteristic tooth (Figs.
232, 235, 270) on each side underneath the pronotum into which
the apex of the tegmina fit. Princis (i960) and Beier (1961) used
this structure as a characteristic of Derocalymminae and Derocalym-
mini, respectively. The shape of the tooth may have value as a
species character (cf. Figs. 232 and 235).

Shelford (1908) summarized Saussure and Zehntner’s (1895)
analysis of the evolution of the form of the pronotum in the Peri-
sphaeriinae. The complex type of the pronotum of genera such as
Pilema and Cyrtotria appear to have evolved from a more simple
form. Pilema thoracica has been found in family groups of a female
and nymphs in round holes in the ground about 6 inches in depth.
Shelford suggests that the shovel-like pronotum of Pilema is prob-
ably used for digging and evolved as a result of a change to this
habitat. Derocalymma spp. live under heavy stones and their flat-
tened form would enable them to crawl through narrow crevices for
security from predators. Some species of Cyrtotria also have the
pronotum adapted for digging, but others haven’t; nothing is known
of the habits of this genus. The Bantua pronotum is intermediate
in structure between Cyrtotria and Derocalymma , but nothing is
known of the genus other than it has been collected under a log.
Shelford suggests that the Derocalymma form originated from a form
like Pilema evolving through a Bantua- like stage.

Summary

Based on male genitalia, 1 1 genera of Perisphaeriinae are placed
in 4 tribes as follows :

1. Perisphaeriini : Perisphaeria , Blepharodera , Pilema , Hostilia,
Bantua , and Cyrtotria .

2. Zuluiini : Zuluia.

3. Derocalymmini : Derocalymma.

4. Perisphaerini : Perisphaerus, Pseudo glomeris, and Trichoblatta.

Acknowledgements

I thank the following for the loan of museum specimens : Dr.
David Rentz, Academy of Natural Sciences of Philadelphia; Dr.
David Ragge, British Museum (Natural History) ; Dr. Karl Prin-
cis, Zoological Institute, Lund, Sweden; Dr. M. W. R. de V.
Graham, Hope Department Entomology, Oxford, England ; Dr.
Howard Evans, formerly at the Museum of Comparative Zoology,
Harvard University; Dr. S. L. Tuxen, Zoological Museum, Copen-

348

Psyche

[December

hagen, Denmark. I am grateful to Mr. Samuel H. Cohen for taking
the photographs.

References

Beier, M.

1961. H. G. Bronns Klassen und Ordnungen des Tierreichs. Arthro-
poda, Insecta, Blattopteroidea, Blattodea. 5, Div. Ill, Book 6.
Part 4, pp. 587-848. Leipzig.

Gurney, A. B. and L. M. Roth

1972. A generic review of the cockroaches of the subfamily Panchlori-
nae (Dictyoptera, Blattaria, Blaberidae). Ann. Entomol. Soc.
Amer. 65: 521-532.

McKittrick, F. A.

1964. Evolutionary studies of cockroaches. Cornell Univ. Agric. Exp.
Sta. Memoir 3 89, 197 pp.

Princis, K.

1960. Zur systematik der Blattarien. Eos 36: 427-449.

1963. Blattariae. In South African Animal Life. Stockholm, pp. 1-318.
Rehn, J. A. G.

1922. Contributions to our knowledge of the Dermaptera and Ortho-
ptera of the Transvaal and Natal. Part 1. Dermaptera and
Blattidae. Ann. Transvaal Mus. 9: 1-99.

1932. African and Malagasy Blattidae (Orthoptera) . Part II. Proc.
Acad. Nat. Sci. Philadelphia 84: 405-511.

1933. Dermaptera and Orthoptera of the De Schauensee South African
Expedition, — Part 1. Proc. Acad. Nat. Sci. of Philadelphia 85:
39-66.

Rehn, J. W. H.

1951. Classification of the Blattaria as indicated by their wings (Orthop-
tera). Mem. Amer. Entomol. Soc. No. 14: 1-134.

Roth, L. M.

1970. The male genitalia of Blattaria. III. Blaberidae: Zetoborinae.
Psyche 77: 217-236.

1970a. The male genitalia of Blattaria. IV. Blaberidae: Blaberinae.
Psyche 77: 308-342.

1971. The male genitalia of Blattaria. VIII. Panchlora, Anchoblatta,
Biolleya, Pelloblatta, and Achroblatta (Blaberidae: Panchlorinae) .
Psyche 78: 296-305.

1972. The male genitalia of Blattaria. IX. Blaberidae. Gyna spp.
(Perisphaeriinae) , Phoraspis , Thorax, and Phlebonotus (Epilam-
prinae). Psyche 98: 185-217.

1973. Paramuzoa (Nyctiborinae) , a new cockroach genus previously
confused with Parasphaeria (Epilamprinae) . Psyche 80: 249-264.

Saussure, H. de and L. Zehntner.

1895. Revision de la Tribu des Perisphaeriens. Insectes Orthopteres de
la famille des Blattides. Rev. suisse Zool. 3 : 1-59.

Shelford, R.

1908. Some new genera and species of Blattidae, with notes on the
form of the pronotum in the subfamily Perisphaeriinae. Ann.
and Mag. Hist. Ser. 8, 1: 157-177.

NEW SPECIES, RECORDS, AND SYNONYMS OF
CHILEAN THERIDIID SPIDERS
(ARANEAE, THERIDIIDAE)*

By Walter C. Sedgwick
Bar M Ranch, Boston, Ga. 31626

This paper is a supplement to Levi (1967), without which it
would not have been possible. Since the northern provinces of Chile
are so arid, most previous collecting has been done in the southern
part. My collecting was mostly in the provinces of Santiago and
Valparaiso, as well as those few enclaves and rivers in the northern
desert which support vegetation. As my trip to Chile occurred during
the dry Chilean summer, I concentrated on collecting under stones
and using a head-light at night. This yielded an abundance of speci-
mens of Theridion. One Dipoena and two species of Theridion are
new, another is a new record for Chile. There is one new synonym
of a Chilean theridiid and two new synonyms as a result of exam-
ining type specimens of Nicolet’s Araneidae which had been placed
in the wrong family.

The new Dipoena cartagena is unusual because its mouth parts
are specialized (Fig. 14), apparently for holding onto legs of ants,
Dipoends prey. I do not know of other instances of similar adapta-
tion of mouth parts; but they may have been overlooked in the
minute specimens. Mouth parts of spiders are conservative and
hardly differ in different genera.

I am very grateful to Dr. Willard Whitcomb for allowing me
to accompany him on his trip, Dr. and Mrs. Luciano Campos for
their hospitality, Drs. Cortez and Charlin for their patience and
kindness during the trip, and Luis Cartagena and Luis Alvarez for
their collecting enthusiasm. I am especially and most sincerely grate-
ful to Dr. Herbert Levi of Harvard for the drawings and the con-
tinuing advice and help he has given me throughout the preparation
of this paper and for examining the Nicolet types. Dr. M. Hubert
of the Museum National d’Histoire Naturelle, Paris, found the
Nicolet types and made them available to me for study.

Achaearanea tepidariorum (C. L. Koch) (Levi, 1955: 32, figs.
69-70, 83-84; 1963b: 215, map 1; 1967: 5, 7-8.) Additional rec-
ord. Coquimbo Province: Pampilla, 14. II. 73, one 9 (L. Alvarez).

Achaearanea acoreense (Berland) ( Achaearanea geochares Levi,
1955: 20, figs. 39-40, 46. Achaearanea teja Levi, 1967: 8, figs. 8-10.

* Manuscript received by the editor December 5, 1973.

349

350

Psyche

[December

new synonymy.) Additional record. Valparaiso Province: La
Cruz, 20. I. 73, three cT , five $.

Steatoda grossa (C. L. Koch) (Levi, 1962: 26; 1967: 7.) Addi-
tional record. Tarapaca Province: Agricultural Experiment Station,
Azapa, 27. I. 73, under the outer layer of a banana tree, two $, six
immatures.

Steatoda sabulosa (Tullgren) (Levi, 1962: 30, figs. 25-27; 1967:
7.) Additional record. Coquimbo Province: Portilla, 20. I. 73,
2600 m elev. under a rock, one 9-

Theridion amarga Levi ( Theridion foliaceum, — Levi, 1963c:
536, figs. 44-46. Theridion amarga Levi, 1967: 7-8.) Additional
record. Valparaiso Province: La Cruz, 20. I. 73, two <$ , three ?.

Theridion volubile Keyserling (Levi, 1963c: 527, figs. 10-13.)
Previously known from Venezuela to Peru. Additional record. Tar-
apaca Province: Agricultural Experiment Station, Azapa, under the
outer layer of a banana tree, two 9; and Taltape, Camarones Valley
in Cartaderid sp. (Foxtail), 29. I. 73, one cf.

Theridion whitcombi n.sp.

Figures 1-4

Holotype. Female from Agricultural Experiment Station, Azapa,
Tarapaca Province, Chile 27. I. 1973 (W. Sedgwick) in the Mu-
seum of Comparative Zoology. It is named after Dr. Whitcomb for
the encouragement he has given me over many years. It was found
under the outer layer of a banana tree.

Description. Carapace yellow white, with a triangular dark band
beginning one-third of the distance from the posterior median eyes to
the cervical groove and which narrows from the anterior to form a
blunt point at the cervical groove. It is indented anteriorly (Fig. 1).
There is a black band around the carapace ending at the first coxa,
continuing in the form of a very thin line with a black spot the
width of the anterior median eyes in the center of the clypeus. The
eye region is light. The legs have black spots, and the distal half of
the tarsi are a very light brown. The size of the two specimens is
different. The abdomen has two white spots slightly less than their
diameter apart between the genital groove and spinnerets, with a
pair of smaller white spots anterior to the spinnerets (Fig. 4). The
dorsal abdominal pattern is similar to Theridion elli having a wavy
white band which narrows posteriorly. It is bordered on both sides
with a dark brown band (Fig. 1). Total length 1.84 mm. Carapace
0.69 mm long, 0.59 mm wide. First femur, 1.22 mm; patella and

1973] Sedgwick — Theridiid Spiders 351

Figs. 1-4. Theridion ivhitcombi n. sp. 1. Carapace and abdomen of
female. 2. Dorsal view of epigynum. 3. Epigynum. 4. Venter of abdomen.

Figs. 5-10. Theridion elli n. sp. 5. Dorsal view of epigynum. 6. Epi-
gynum. 7. Female. 8. Venter of abdomen. 9, 10. Left palpus. 9. Mesal.
10. Ventral.

Figs. 11-14. Dipoena cartagena n. sp. 11. Carapace and abdomen of
male. 12. Left palpus, ventral view. 13. Carapace, chelicera, endite. 14.
Endites and labium.

Scale lines, 0.1 mm; except 1, 7, 11, 13, 1.0 mm.

352

Psyche

[December

tibia, 1.25 mm; metatarsus, 0.91 mm; tarsus, 0.47 mm. Second
patella and tibia., 0.75 mm; third, 0.53 mm; fourth, 0.84 mm.

Diagnosis. This species differs from Theridion volubile by the
shape of the internal genitalia, the ducts of which loop less than those
of Theridion volubile (Fig. 2).

Record. One female paratype collected with holotype deposited in
the Museum of Comparative Zoology.

Theridion elli n.sp.

Figures 5-10

Holotype. Female from Antumapu, Santiago Province, Chile 18.
I. 1973 (L. Alvarez) in the Museum of Comparative Zoology. The
specific name is a noun in apposition, after a friend who has en-
couraged me greatly in arachnology.

Description. Carapace light brown, with a wide margin around
the sides, turning into a thin black line at the clypeus. The eye
region is dark (Fig. 7). The legs are generally spotted in mid and
distal portions, the distal half of the tarsi being a light brown. The
abdomen is variable in size and shape, but always has a white spot
at each end of the genital groove and two smaller spots located be-
tween the genital groove and the spinnerets (Fig. 8). The dorsal
pattern of the abdomen consists of seven white spots stretching from
near the pedicel to near the spinnerets. The first two are small, the
third larger, and the subsequent spots become smaller towards the
spinnerets. The spots are bordered in black which is sometimes a
continuous border, sometimes in spots (Fig. 7). The male is similar.
Both male and female were found under rocks and papers. Total
length of female 2.13 mm. Carapace 0.88 mm long, 0.75 mm wide.
First femur, 1.25 mm; patella and tibia, 1.41 mm; metatarsus, 1.00
mm; tarsus, 0.53 mm. Second patella and tibia, 0.88 mm; third,
O.59 mm; fourth, 0.91 mm. Total length of male 1.47 mm. Cara-
pace 0.69 mm long, 0.63 mm wide. First femur, 1.31 mm; patella
and tibia, 1.34 mm; metatarsus, 1.06 mm; tarsus, 0.56 mm. Second
patella and tibia, 0.84 mm; third, 0.59 mm; fourth, 0.84 mm.

The female type has a short, blind extra duct originating from the
left seminal receptacle (Fig. 5).

Diagnosis. This species can be differentiated from Theridion volu-
bile and Theridion whitcombi by the epigynum (Fig. 6) in which
the ducts are not visible (as in Theridion volubile ) and which dif-
fers in shape from that of Theridion whitcombi. The ducts open in
black spots of the depression (Figs. 5, 6).

1973]

Sedgwick — Theridiid Spiders

353

Records. Santiago Prov. : Santiago, Dept, of Agriculture Building
at University of Chile, three cf, $ paratypes (W. Sedgwick).
Antumapu two 9 paratypes (L. Alvarez). (9, <$ paratypes placed
in American Museum of Natural History, New York.)

Dipoena cartagena n.sp.

Figures 11-14

Holotype. Male from Bosques al Sur de Quintero, Valparaiso
Province, Chile, 20. II. 1973 (Luis Cartagena) in the Museum of
Comparative Zoology. It was found under a leaf. It is named for
the collector. The specific name is a noun in apposition.

Description. Carapace, sternum, legs olive-brown. Tarsi of legs
light colored. Dorsum of abdomen with a black pattern and white
pigment spots in areas without black (Fig. 11). Sides and venter
black. Venter with a pair of white spots side by side. Posterior
median eyes 0.6 diameters of anterior medians. Laterals the radius
of anterior medians. Anterior median eyes slightly more than their
diameter apart, almost touching laterals. Posterior median eyes
slightly less than one diameter apart, about their diameter from
laterals. The labium and endites are modified, the endites with a
distal brush and the labium with a transparent lobe on each side
(Fig. 14). Total length 2.0 mm. Carapace 0.98 mm long, 0.90 mm
wide, 0.76 mm high. First femur, 1.52 mm. Second patella and
tibia, 1. 00 mm; third, 0.69 mm; fourth, 1.18 mm.

Dipoena species feed on ants. This specimen is unusual because
unlike others of the genus, the endites seem to be specially adapted
probably for holding the legs of the ant while the spider pierces it
with its fangs.

Diagnosis. This species is close to Dipoena pumicata (Keyserling)
but differs from this and related species by the twisted embolus and
plain projecting conductor of the palpus (Fig. 12).

Phoroncidia pennata (Nicolet), new combination.

Gasteracantha pennata Nicolet, 1849, in Gay, Historia Fisica y Politica
de Chile, 10(3): 474, pi. 5, fig. 5. Female type in poor condition from
Chile. No. 5524 in the Museum National d’Histoire Naturelle, Paris,
examined July 1973.

Ulesanis pennata , — Simon, 1894, Histoire Naturelle des Araignees 1: 556.

Glyptogona pennata, — Simon, 1895, Histoire Naturelle des Araignees 1:
867. Roewer, 1942, Katalog der Araneae, 1 : 891.

Phoroncidia coquimbo Levi, 1964, Bull. Mus. Comp. Zool., 131: 77, figs.
38-41, $, $. Female holotype from Hacienda Illapel, Prov. Coquimbo,

Chile, in the Institut Royal des Sciences Naturelles de Belgique, Brussels.
NEW SYNONYMY.

354

Psyche

[December

Nicolet’s type is in very poor condition and has the lateral pro-
jections of the abdomen relatively longer than in previous specimens.

Phoroncidia variabilis (Nicolet), new combination
Gasteracantha variabilis Nicolet 1849 in Gay, Historia Fisica y Politica
de Chile, 10(3): 476. Female holotype from Chile, no. 5523 in the Museum
National d’Histoire Naturelle, Paris, examined, July 1973.

Glyptogona pennata, — -Simon, 1895, Histoire Naturelle des Araignees 1:
1 : 867. Roewer, 1942, Katalog der Araneae, 1 : 891.

Phoroncidia margamarga Levi, 1964, American spiders of the genus
Phoroncidia (Araneae: Theridiidae). Bull. Mus. Comp. Zool. 131: 77,

figs. 30-34, 2. Female holotype from Marga Marga, Prov. Valparaiso,

Chile, in the Institut Royal des Sciences Naturelles de Belgique, Brussels.
Levi, 1967, Bull. Mus. Comp. Zool. 136: 18, figs. 16-18, $. new synonymy.

References Cited

Levi, H. W.

1955. The spider genera Coressa and Achaearanea in America north
of Mexico (Araneae: Theridiidae). Am. Mus. Novit., no. 1718:
1-33.

1962. The spider genera Steatoda and Enoplognatha in America (Ara-
neae, Theridiidae). Psyche, 69: 11-36.

1963a. American spiders of the genus Achaearanea and the new genus
Echinotheridion (Araneae, Theridiidae). Bull. Mus. Comp. Zool.,
129: 187-240.

1963b. American spiders of the genus Theridion (Araneae: Theridi-
idae). Bull. Mus. Comp. Zool., 129: 481-589.

1967. The theridiid spider fauna of Chile, Bull. Mus. Comp. Zool., 136:
1-20.

LATITUDINAL GRADIENTS IN
LARVAL FEEDING SPECIALIZATION
OF THE WORLD PAPILIONIDAE ( LEPIDOPTERA) *

By J. Mark Scriber
Department of Entomology
Cornell University, Ithaca, N. Y. 14850

Introduction

A major question arising from ecological theory is whether or not
there are relatively more specialized species in the lower or tropical
latitudes than in the higher or more temperate latitudes. There
exists a great deal of speculation concerning this question, but little
actual data. This article presents an analysis of such data for one
family of Lepidoptera ; the Papilionidae.

The subject of niche breadth and degree of competitive overlap
has generated tremendous interest in the field of ecology in the last
two decades. In terms of the spatial model of the niche first pro-
posed by Hutchinson (1957) and later expanded by Slobodkin
(1962), Levins (1968), and MacArthur (1968), niche breadth is
the “distance through” a niche hypervolume along a particular
resource gradient within a community (Colwell and Futuyma, 1971 ;
Whittaker, et al., 1973).

Although there is presumed to be an apparent limit to the simi-
larity and hence the number of competing species that can coexist
(MacArthur and Levins, 1967), the packing of species (MacArthur,
1969a) appears to be roughly proportional to the environmental
variance, with less overlap, or looser packing, occurring with in-
creased environmental fluctuation or perturbation. May and Mac-
Arthur (1972) imply that niche overlap may be relatively inde-
pendent of small to moderate environmental fluctuations, but it
remains true that greater packing could still be attained by special-
ization, and the question concerning what ultimately limits niche
width remains unsettled. Also, in a more stable environment of the
tropical latitudes, with larger productivity one might expect that
marginally scarce resources would become adequate to support new
species at the extremes of the resource gradient (Odum, et al., i960;
Connell and Orias, 1964). In a more stable low-latitude tropical
environment (MacArthur, 1969a), old lakes (Kozhov, 1963), the

* Manuscript received by the editor October 18, 1973

355

356

Psyche

[December

benthos (Sanders, 1968, 1969) or coral reefs (Kohn, 1968) we
expect closer packing and greater numbers of competitors per com-
munity (Levins, 1968; MacArthur and Levins, 1967; Vandermeer,
1970) with narrower realized niches and increased ecological dif-
ferentiation and niche specificity (Whittaker, et ah, 1954; Klopfer,
1959; Selander, 1965; Millar, 1969).

Although species diversity varies from one community to the next,
it is often greatest in the lower latitudes. Sanders (1969) incorpo-
rates many of the earlier proposed explanations of differences in
species diversity (Pianka, 1966; Paine, 1966; Connell and Orias,
1964; MacArthur, 1965, 1969b) in his stability-time hypothesis.
According to this hypothesis, the greater number of species in the
tropics implies a greater number of biotic interactions than in the
temperate regions. The results of competition would be most obvious
in the form of niche divergence and increased specializations over a
resource gradient. The corresponding decrease in predictability and
stability of environmental conditions with an increase in latitude
may mean that species in temperate regions are influenced more by
the physical environment than by competitive interactions (Slobodkin
and Sanders, 1969; Janzen, 1970). Realized niches may be smaller
in the tropical or low latitudes than in higher latitudes because of
the expected difficulties of specializing in an unpredictable environ-
ment and of generalizing in an extremely diverse and competitive
environment. The following investigation was undertaken in an
attempt to determine the correctness of the premise that realized
feeding niches are relatively more specialized in the tropical and sub-
tropical latitudes than in the higher latitudes.

Methods

My approach was to choose a single taxonomic group that is cos-
mopolitan, whose habits are relatively well known and which all
nicely fall into a single ‘guild’ (Root, 1967). The 538 species of
Papilionidae (Lepidoptera) of the world, based on Munroe’s classi-
fication, met these criteria adequately.

In order to analyze a niche breadth it was necessary to choose a
criterion that could be compared over a range of latitudes and not
one that varied between communities. Therfore, I investigated larval
host plant relationships arbitrarily designating those species feeding
on more than one taxonomic family of plants as “generalists,” or
wide-niched, species. By using food-plant range as such, latitudinal
comparisons of food-plant specialization, or trophic niche breadth

1973]

Scriber — Papilionidae

357

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[December

Fig. 1. Latitudinal gradients in species richness for the New World
Fapilionidae (Modified from Slansky, 1972)

were simplified considerably. As explained in the discussion, it was
not desired to use degree of morphological correlation with feeding
specialization as an index of niche breadth.

Because larvae generally remain upon the host plant throughout
most of their existence, the food-plant is especially likely to be the
key factor in the niche of most butterfly species since, to a large
extent, it is also the shelter, substrate, or habitat upon which allelo-
chemical co-evolution (Ehrlich and Raven, 1965; Whittaker and
Feeny, 1971) of the entire ‘component community’ (Root, 1973)
takes place. In this sense it is essentially difficult to differentiate
between ‘niche’ and ‘habitat’ (Whittaker, Levin, and Root, 1973),
and their term ‘ecotope’ might be more appropriate since it entails
the inter-community as well as the intra-community variables.

I have compiled range maps of each of the 538 species of Papil-
ionidae of the world (Munroe, i960) using the techniques of
Slansky (1972). I used Goode’s Atlas (Espenshade, i960) and
locality data recorded in the literature, primarily in Rothschild
(1895), Rothschild and Jordan (1906), Aurivillus (1908-1910),

1973]

Scriber — Papilionidae

359

Bollow (1929), Fruhstorfer (1908), Jordan (1907, 1908-1909),
von Rosen (1929), Seitz (1906), Stichel (1906), Shirozu (i960),
D’Almeida (1966), and Common and Waterhouse (1972).

The number of species known to occur in each ten degree latitude
belt was recorded. The worldwide data were divided into the three
geographical sections in figures 1, 2, and 3 to simplify the analysis.
The components of each of these figures are shown in tabular form
(Table 1).

The species of known larval food-plants were compiled from the
literature, although care was used in interpreting recorded host plant
records (see Ehrlich and Raven, 1965; and Shields, Emmel, and
Breedlove, 1969). The plant classification follows Willis (1973).
A supplementary table with each species of Papilionidae with its
latitudinal range, food-plant families fed upon, and citations for
these food plant records is available from the author and at the
following libraries: Cornell University (Entomology), Harvard
(M.C.Z.), and the U.S.N.M.1

With those species feeding on more than one taxonomic family of
plants as my “generalists” or wide-niche species, a species was counted
in all parts of its range as being a wide-niche species, or generalist,
if it feeds on more than one plant family anywhere in its range.

Latitudinal gradients in niche breadth were prepared by enumer-
ating the species for each ten degree belt of latitude and calculating
the percentage of species exhibiting a “general” feeding strategy.

Although the foodplants of the Papilionidae are probably as well
known as those of any other taxon of similar magnitude, there are
still gaps in the record. In the case of a species of unknown food-
plant preference I have assumed that the host plant family range
utilized is, in general, similar to those of closely related species of
the group. There are, for example, many tropical Troidini for which
the precise species of Aristolochia utilized are not known.

Longitudinal and altitudinal variations in species richness are an
important and interesting part of distributional patterns, but were:
not considered in this study.

Results

Species richness is greatest in the tropical latitudes for each of the-
three geographical regions (Figures 1, 2, 3). The significance of
this trend for Papilionidae in the New World is discussed by Slansky

The Grace Griswold Fund of the Department of Entomology of Cornell
University assisted with the expense of having the supplementary tables of
data copied.

36o

Psyche

[December

Fig. 2. Latitudinal gradients in species richness for Eurasian and Afri-
can Papilionidae.

(x973)- The smaller number of species of Papilionidae occurring
in the 20-30 degree North latitude belt of figure 2 is presumably due
to the Sahara and Arabian deserts. The great number of species in
the 20-40 degree North latitude belts in figure 3 is correlated to a
large degree with the strong radiation especially of the Parnassiinae
and also of the Papilioninae subfamilies in the mountainous Hima-
laya region and the Tibetan Plateau. (See Table 1.) There is a
relatively sudden drop in species richness south of 10 degrees South
latitude in figure 3, the biogeographical relevance of which is un-
certain.

Figure 4 shows the overall pattern of latitudinal gradients in the
number of species of Papilionidae, corrected for distributional over-
lap of certain species that occurred in more than one of the earlier
three figures. Also the number of species for each tribe of Papilioni-
dae feeding on two or more plant families and the overall percentage
of generalized feeders for each latitudinal belt are shown in relation

1973]

Scriber — Papiliomdae

36i

to this pattern. Note that there is a considerable decrease in the per-
centage of generalists toward the lower tropical latitudes (Figure 5).

Discussion

Although the absolute number of generalists is not strikingly dif-
ferent, it is readily apparent that the percentages are considerably
less in the lower latitudes. This fact appears to support the premise
that there are relatively fewer generalists in the tropics and relatively
more in the higher latitudes. There is, however, no sharp division,
but instead a gradient of increasing relative degree of specialization
from the extremes of latitude to the equatorial regions. It is not
known whether increased competition could be the cause or the result
of the increasing diversity gradients toward the lower latitudes, nor
is it certain whether the observed gradients in feeding niche breadth
are due to ecological release from competition in higher latitudes or
just the inability to specialize in their unpredictable and unstable
environmental conditions.

It was assumed throughout this study that the niche breadth as
measured here is that of the ‘realized’ niche as opposed to the ‘funda-
mental’ (Odum, 1971). This study of feeding niche breadths in-
volves a cross section of both ‘evolutionary’ and ‘ecological’ time
(Slobodkin, 1962) and the differentiation at both intra- and inter-
population levels. That is, there are probably areas of active compe-
tition where we are not viewing only the neat results of evolutionary
processes that have already and permanently narrowed the realized
hostplant ranges from those plant families which are physiologically
exploitable, but there are also the currently unfolding consequences
of ecological interactions that someday may or may not result in more
specialized herbivores. It should be noted that the relative merits of
either specializing or generalizing may involve ‘qualitative’ and
‘quantitative’ chemical defense of the host plants (Feeny, 1974) as
well as other ecological factors besides competition alone.

Emmel and Emmel (1969) have shown that for Papilio indra
(Reakirt) and Papilio rudkini (Comstock) direct competition of
sympatric populations is avoided most of the time by temporal isola-
tion and utilization of mutually exclusive food-plant families. In
one instance however they found that food did become the limiting
resource and the separate host preferences broke down when the
favored food became scarce. In this situation both Umbelliferae and
Rutaceae were then utilized by the insects of these populations.
Further evidence that competitive interactions leading to adaptations

362

Psyche

[December

Fig. 3. Latitudinal gradients in species richness for Indo-Australian and
Eastern Asian Papilionidae.

that permit the coexistence of several species of butterflies should
probably be sought in the larval rather than the adult stages has
been obtained by Young (1972). The critical dimension of the
niche (or ecotope) may in fact be something other than the range of
larval hostplants utilized ; however, as well as being one that is rela-
tively easy to measure, it is assumed here to be a crucial parameter
in most cases.

In a relatively recently speciated group of swallowtails in the U.S.,
the polyphagous Papilio glaucus (L.) is apparently entirely excluded
geographically by three species which are sympatric and which have
subdivided the range of plant families utilized by P. glaucus
(Brower, 1958a; 1959). Besides their specialized feeding habits,
isolating mechanisms such as seasonality, coloration, and mating
preferences appear to have been involved in the Pleistocene specia-
tion. There is no clear cut distinction between ecological and evolu-
tionary time, and the importance of trophic specialization is uncertain

1973]

Scriber — P apilionidae

363

in this and similar cases such as the four species of the P. demodocus
(Esper) group of swallowtails that have evolved on Madagascar,
or the three species of the P. nireus (L.) group in West Africa that
are presumed to have speciated during the Tertiary when the climate
and vegetation were rapidly changing (Owen, 1971). Furthermore,
because the author is in this study to a large degree uncertain of
regional food plant preferences that might distinguish a polymorphic
population of specialized individuals from a monomorphic population
of generalized individuals, no attempt has been made to differentiate
any components of niche breadth as described by Roughgarden

(1972).

It is probably fair to assume that neither the latitudinal gradients
in diversity nor those for trophic niche breadth remain constant for
any long periods of time. The phylogenetic histories of the Papilioni-
dae are certainly intertwined with the zoogeographical movement and
changes in the flora of major continents (Hovanitz, 1958; Carcas-
son, 1964; and Kostrowicki, 1969), and this paper is merely an
attempt to look at the present state of these continually changing
phylogenies.

Besides regional preferences there can be temporal changes in food-
plant preferences and these are frequently accentuated by human agri-
culture. Owen (1971) points out that several species of African
papilionids have recently expanded their ranges and seasonal abun-
dance by switching from their wild Rutaceous foodplants onto culti-
vated varieties of Citrus which, unlike the wild plants, maintain
their leaves throughout the dry season and are spreading rapidly with
agricultural development. The extent to which members of the
Papilionidae have become secondarily polyphagous on completely new
plant families or else by revitalized usage upon re-exposure to ances-
tral hostplant families is uncertain. It is very likely that besides the
general ecological pressures such as competition with more specialized
herbivores, there may well be both ‘fixed’ and ‘variable’ metabolic
‘cost’ associated with any extension of the normal range of food
plants utilized by a species (Feeny, 1974). In Papilvo demoleus for
example the principal natural foodplants for Papilio demoleus
stheneles (Macleay) in Australia are Psoralea tenax and P. patens
which are members of the Leguminosae family (D’Abrera, 1971 ;
McCubbin, 1971; and Common and Waterhouse, 1972), while in
Ceylon and India the presumed ancestral P. demoleus demoleus (L.)
feed primarily upon various Rutaceae, such as Aegle , Clausenia, Gly-
oosmis, and Citrus (Woodhouse, 1950; and Igarashi, 1966). Al-
though the females of P. d. stheneles oviposit upon Citrus in Queens-

364

Psyche

[December

Number of Generalized
Feeders *

latituc

Number of Species

e

A

B

c

Percentaae
. Generalized
i Feeders

e World Papiltonida

D

tota

60-70

4

2

1

3 !

42.86 %

50-60

^19

2

7

9

47.37 %

40-50

2

10

2

14

26.42 %

30-40

ifiM-

1

4

17

4

26

22.61 %

20-30

125

5

13

5

23

18.40 %

10-20

A 160

7

9

5

21

13.13 %

0-10

iiifif

- J224

8

9

6

23

10.27 %

0-10

iiifiP

256

8

11

4

23

9.02 %

10-20

155

10

10

3

23

14.84 %

20-30

j 89

1

10

7

2

19

21.35 %

30-40

>

3

6

1

10

38.46 %

40-50

f2

1

1

1

50.00 %

1

50 ' 150 250

Fig. 4. Latitudinal gradients in species richness and percentage of gen-
eralized species of Papilionidae of the world. A — Parnassini ; B = Lep-
tocircini; C = Papilionini ; D = Troidini.

land (Australia), the larvae are unsuccessful and eventually die when
left to feed upon leaves of this relatively recent introduction (Ed-
wards, 1948; and Straatman, 1962). It appears thus that in at least
some populations metabolic specializations or adjustments may have
coevolved along with the regional restriction in food-plant choice such
that Citrus is perhaps in some way no longer physiologically suitable
for those individuals. It should be pointed out that individuals fiom
some Australian populations can and do successfully utilize the native
rutaceous Alicrocitrus australis (D abrera, I970> and those in New
South Wales can be reared successfully upon Citrus (see Stiaatman,
1962) .

It is interesting that in Africa Papilio demodocus (Esper), a spe-
cies which is very closely related to and once consideied a race of
P. demoleus, is primarily a Rutaceae-feeder over most of its range.
It is however polymorphic (Clarke, et al. 1963) and also utilizes the
Umbelliferae in South Africa w^here the richness of this plant family
is relatively high (Dethier, 1941). Similarly, the P. machaon group
of Umbellifer feeding species in Sardinia, East Asia, and North
America are also close to P. demoleus in ancestry (Munroe, i960).

1973]

Scriber — Papilionidae

365

Further speciation has apparently taken place in those regions where
Umbelliferae are most diverse. Several of these species, however,
are still able to utilize rutaceous food plants.

Feeding patterns of the Papilionidae and phytophagous insects in
general, range from strict monophagy (stenophagy) , in which a single
species of foodplant is utilized, to wide polyphagy (euryphagy) in
which many species, genera, or families may be utilized (Brues, 1920;
Dethier, 1954). The exploitation of a particular hostplant involves
many adjustments on the part of the insect to the plant’s micro-
community. This includes host-specific predators (Brower, 1958b)
and parasites (Read et al., 1970). Facultative polyphagy in some
Papilio larvae, conveying the ability to take advantage of oviposi-
tional mistakes of the adult, may allow some escape from the intense
mortality rates that can result from host-specific parasitism (Stride
and Straatman, 1962). Polymorphisms in larval color patterns can
also be involved (Clarke et al., 1963), but perhaps the most im-
portant factor in this coevolving community is the metabolic adjust-
ments on the part of the insect to the secondary chemistry of the
plant (Fraenkel, 1959; Jermy, 1966; and Thorsteinson, i960). As
pointed out by Ehrlich and Raven (1965), the plasticity of the
chemoreceptive response and the potential for physiological adjust-
ments to secondary plant chemicals may be very important factors in
determining the potential for evolutionary radiation in a phytopha-
gous species. Since the possibility exists that taxonomically general-
ized feeders may in fact be allelochemical specialists that cue in on
one key set of chemicals in the plants, feeding niche breadth may well
take on new meaning.

Increased specialization by a species is generally presumed to in-
crease the competitive ability of a species largely because the species
utilizes its resources more efficiently (Emlin, 1966; MacArthur and
Levins, 1964; MacArthur and Pianka, 1966; Morse, 1971; Rosen-
zweig, 1966; and Schoener, 1971). Presumably, the less efficient
species should be completely eliminated (MacArthur, 1972) or
crowded to the periphery to become a specialist on another niche
dimension (McNaughton and Wolf, 1970). In this sense, the rela-
tive efficiencies of exploiting a critical limiting resource may in large
part determine the niche breadth.

Previous attempts to measure niche breadths have had varying de-
grees of success. Various works in the literature attempt to relate
morphological characters, behavioral repertoires or habitat selection
response with niche breadth or overlap (Klopfer, 1962, 19675
Schoener, 1969; Van Valen, 1965; and Wellington, 1968). Upon

366 Psyche [December

Fig. 5. Latitudinal gradients in feeding specialization for the Papilioni-
dae of the world.

ecological release from competition upon islands (MacArthur and
Wilson, 1967), various workers have found some evidence that spe-
cies niches do appear to broaden in the sense that there is increased
morphological variance, behavioral plasticity, and dominance (Grant,
1967; Lack and Southern, 1949; Patrick, 1967; Vaurie, 1957; Wil-
liams, 1969; and Wilson, 1961). There is, however, some question
whether these measures actually reflect niche breadth of the species
(Soule and Stewart, 1970 ; Willson, 1969). I have employed a
somewhat different measure here, and by altering the number of
families or genera used as a criterion for ‘generalist’, the amplitude
of the observed latitudinal gradient would vary but probably not the
overall pattern. Other studies have described a ‘generalist’ in some-
what different terms as those species able to maintain populations
over a broad range of environmental types or substrates (McNaugh-
ton and Wolf, 1972), or those exhibiting a greater variance or
breadth of morphology or feeding behavior (Levins, 1968; Rough-
garten, 1972; Schoener, 1971). These also may or may not reflect
actual niche breadth as such, but latitudinal comparisons within a
taxocene would probably be possible to prepare in these terms as well.

It is interesting that many species of Papilionidae are relatively
specialized on certain tropical and sub-tropical plant families (Sian-

1973]

Scriber — Papilionidae

367

sky, 1972). In the higher latitudes where fewer species of most
major families of host plants occur there has probably been some
breakdown of the phylogenetic larval host plant preferences that tie
some species to these major evolutionary host plant groups, perhaps
favoring polyphagy at the plant family level. The Papilio troilus
L. and P. glaucus L. groups, for instance, have expanded their diets
considerably from the Lauraceae and Magnoliaceae they are thought
to have originally fed upon. Thus, besides ‘ecological release’ and
the inability to specialize in unpredictable and unstable environments
as possible explanations of the observed gradients in feeding special-
ization, a third possibility exists. Perhaps there was a necessary
alteration of those allelochemical specializations developed over a
long period of evolutionary time, either as species of Papilionidae
extended their ranges into the higher latitudes where their usual host
plant families never existed, or as they remained in higher latitudes
where their ancestral hosts had since been eliminated, perhaps by
climatic changes.

The precise reason for Papilionidae being more specialized in their
feeding habits in tropical latitudes is uncertain and will probably
remain largely as such. One contributing factor for this is that no
one knows whether polyphagy in general is the ancestral condition
from which more specialized feeding habits arose or not. The fact
exists, however, that there are greater percentages of feeding special-
ists in the tropical latitudes, and that this percentage decreases as one
moves toward the higher latitudes. These data do not suggest that
species are more specialized in the tropical than temperate latitudes,
but that more species are specialized.

Summary

Latitudinal gradients in food plant specialization are examined for
the swallowtail butterflies of the world (Papilionidae). It is found
that the absolute number of generalized feeders existing in each ten
degree belt of latitude is fairly constant, but the relative percentage
of generalized species is significantly higher in the higher latitudes.
This fact is discussed in terms of trophic niche breadth and species
diversity of the Papilionidae in tropical and temperate regions.

Acknowledgements

This work was supported by N.S.F. Grant #GB 33398 (P.P.F.).

I wish to thank Drs. Paul Feeny, J. G. Franclemont, Robert
Poole, and Frank Slansky, Jr. for reading the manuscript or supple-

368

Psyche

[December

ment and offering instructive criticisms. Special thanks are extended
to Frank Slansky, Jr. for helpful data concerning certain Papilioni-
dae, to J. G. Franclemont for generous amounts of his time and as-
sistance, and to Kathleen Scriber for assistance in the preparation of
parts of the manuscript and supplement.

Literature Cited

Aurivillius, C.

1908-10. Papilio. In (A. Seitz ed.) The Macrolepidoptera of the
world. The African Rhopalocera. Alfred Kernen. Stuttgart. 13:
11-28.

Bollow, C.

1929. Parnassius. In (A. Seitz edi.) The Macrolepidoptera of the
World. The Palearctic Butterflies. Alfred Kernan. Stuttgart.
Suppl. 1 : 20-83.

Brower, L. P.

1958a. Larval foodplant specificity in butterflies of the Papilio glaucus
group. Lepid. News 12: 103-114.

1958b. Bird predation and foodplant specificity in closely related Pro-
cryptic insects. Amer. Nat. 92: 183-187.

1959. Speciation in butterflies of the Papilio glaucus group. Parts I
and II. Evol. 13 : 40-63; 212-228.

Brues, C. T.

1920. The selection of foodplants by insects with special reference to
Lepidopterous larvae. Amer. Nat. 54: 313-332.

Carcasson, R. H.

1964. A preliminary survey of the zoogeography of African butterflies.
E. Afr. Wildlife J. 2: 122-157.

Clarke, C. A., C. G. C. Dickenson, and P. M. Sheppard.

1963. Larval color pattern in Papilio demodocus. Evol. 17: 130-137.
Colwell, R. K. and D. J. Futuyma.

1971. On the measurement of niche breadth and overlap. Ecology 52:
567-576.

Common, I. S. B. and D. F. Waterhouse.

1972. Butterflies of Australia. Angus and1 Robertson Ltd., Brisbane.
498 p.

Connell, J. H. and E. Orias.

1964. The ecological regulation of species diversity. Amer. Nat. 98:
399-414.

D’Abrera, B.

1971. Butterflies of the Australian region. Lansdown Press. Melbourne.
415 p.

D’Almeida, R. F.

1966. Catalogo dos Papilionidae Americanos. Sociedade Brazileiro
Entomologica. Sao Paulo, Brazil. 366 p.

Dethier, V. G.

1940. Life histories of Cuban Lepidoptera. Psyche 47 : 14-26.

1941. Chemical factors in determining the choice of foodplants by
Papilio larvae. Amer. Nat. 75: 61-73.

1973] Scriber — ■ P apilionidae 369

1954. Evolution of feeding preferences in phytophagous insects. Evol.
8: 33-34.

Ehrlich, P. R. and P. H. Raven.

1965. Butterflies and plants: a study in coevolution. Evol. 18: 586-608.
Emlin, J. M.

1966. The role of time and energy in food preferences. Amer. Nat.
100: 611-617.

Emmel, T. C. and J. F. Emmel.

1969. Selection and host plant overlap in two desert Papilio butterflies.
Ecology 50: 158-159.

Feeny, P. P.

1974. Biochemical coevokition between plants and their insect herbi-
vores. In Coevolution of Animals and Plants (L. E. Gilbert and
P. H. Raven eds.) Symp. 1st Int. Congr. Syst. & Evol. Biol.
Boulder, Colorado (August, 1973). Univ. Texas Press, Austin
(in press).

Fraenkel, G.

1959. The raison d’etre of secondary plant substances. Science 129:
1466-1470.

Fruhstorter, H.

1908. Parnassius. In (A. Seitz ed.) The Macrolepidoptera of the
World. The Indo-Australian Rhopalocera. Alfred Kernen. Stutt-
gart. 9: 109-112.

Goode, J. P.

1960. World Atlas. Espenshade, E. B. (ed.). Rand McNally Co., Chi-
cago. 288 p.

Grant, P. R.

1967. Bill length variability in birds of the Tres Marias Islands,
Mexico. Can. J. Zool. 45 : 805-815.

Hovanitz, W.

1958. Distribution of butterflies in the new world. In (C. L. Hubbs
ed.) Zoogeography, (p. 321-368). Amer. Assoc. Adv. Sci. Publ.
51.

Igarashi, S.

1966. Butterflies of Nepal ; Immature Stages. Special Bull. Lep. Soc.
Japan (no. 2) 1-74.

Janzen, D. H.

1970. Herbivores and the number of tree species in tropical forests.
Amer. Nat. 104: 501-528.

Jermy, T.

1966. Feeding inhibitors and food preferences in chewing phytophagous
insects. Ent. Exp. et Appl. 9: 1-12.

Jordan, K.

1907. Papilio. In (A. Seitz ed.) The Macrolepidoptera of the World.

The American Rhopalocera. Alfred Kernen. Stuttgart. 5: 11-45.
1908-09. P apilionidae. In (A. Seitz ed.) The Macrolepidoptera of the
World. The Indo-Australian Rhopalocera. Alfred Kernan.
Stuttgart. 9: 11-109.

Klopfer, P. H.

1959. Environmental determinants of faunal diversity. Amer. Nat.
93: 337-342.

370

Psyche

[December

1962. Behavioral aspects of Ecology. Prentice Hall. Englewood Cliffs,
New Jersey. 161 p.

1967. Behavioral stereotypes in birds. Wilson Bull. 79: 290-300.
Klopfer, P. H. and R. H. MacArthur.

1960. Niche size and faunal diversity. Amer. Nat. 94: 293-300.

1961. On the causes of tropical species diversity: niche overlap. Amer.
Nat. 95: 223-226.

Kohn, A. J.

1968. Microhabitats, abundance and food of Conus on atoll reefs in
the Maidive and Chagos Islands. Ecolgy 49: 1046-1061.

Kostrowicki, A. S.

1969. Geography of the Palearctic Papilionidae (Lepidoptera) . Pan-
stwowe Wydawnictwo Naukowe, Poland. 380 p.

Kozhov, M.

1963. Lake Baikal and its life. W. Junk Publ. The Hague. 344 p.
Lack, D. and H. N. Southern.

1949. Birds on Tenerife. Ibis 91 : 607-626.

Levins, R.

1968. Evolution in changing environments. Princeton Univ. Press.
Princeton, N.J. 120 p.

MacArthur, R. H.

1965. Patterns in species diversity. Biol. Rev. 40: 510-533.

1968. The theory of the niche. In R. C. Lewontin (ed.). Population
Biology and Evolution. Syracuse Univ. Press, Syracuse, N.y.
1969a. Species packing, or what competition minimizes. Proc. Nat’n.
Acad. Sci., U.S. 64: 1369-1371.

1969b. Patterns of communities in the tropics. Biol. J. Linn. Soc. 1 :
19-30.

MacArthur, R. H. and R. Levins.

1964. Competition, habitat selection, and character displacement in a
patchy environment. Proc. Nat. Acad. Sci. U.S. 51: 1207-1210.
1967. The limiting similarity convergence, and divergence of coex-
isting species. Amer. Nat. 101: 377-385.

MacArthur, R. H. and E. R. Pianka.

1966. On optimal use of a patchy environment. Amer. Nat. 100:
603-604.

MacArthur, R. H. and E. O. Wilson.

1967. The theory of island biogeography. Princeton Univ. Press.
Princeton, N.J. 203 p.

May, R. M. and R. H. MacArthur.

1972. Niche overlap as a function of environmental variability. Proc.
Nat. Acad. Sci. U.S. 69 : 1109-1113.

McNaughton, S. J. and L.L. Wolf.

1970. Dominance and the niche in ecological systems. Science 67:
131-139.

Millar, R. S.

1969. Competition and species diversity. In Diversity and stability in
ecological systems. Brookhaven Symp. Biol. 22: 63-70.

Morse, D. H.

1971. The insectivorous bird as an adaptive strategy. Ann. Rev. Ecol.
and Syst. 2 : 177-200.

1973]

Scriber — P apilionidae

371

Munroe, E.

1960. The classification of the Papilionidae (Lepidoptera). Can. Ent.
suppl. 17: 1-51.

Odum, E. P.

1971. Fundamentals of Ecology. 3rd Edition. W. B. Saunders Co.
Philadelphia. 574 p.

Odum, H. T., J. Cantlon, and L. S. Kornicker.

1960. An organizational hierarchy postulate. Ecol. 41: 395-399.

Owen, D. F.

1971. Tropical butterflies. Clarendon Press, Oxford. 214 p.

Paine, R. T.

1966. Food web complexity and species diversity. Amer. Nat. 100:
65-75.

Patrick, R.

1967. The effect of invasion rate, species pool, and size of area on
the structure of the diatom flora. Proc. Nat. Acad. Sci. U.S. 58;
1335-1342.

PlANKA, E. C.

1966. Latitudinal gradients in species diversity: a review of the con-
cepts. Amer. Nat. 100: 33-46.

Read, D. P., P. P. Feeny and R. B. Root.

1970. Habitat selection by the aphid parasite Diaeretiella rapae (Hy-
menoptera: Braconidae) and hyperparasite Charips brassicae

(Hymenoptera : Cynipidae). Canad. Ent. 102: 1567-1578.

Root, R. B.

1967. The niche exploitation pattern of the blue-gray gnat catcher.
Ecol. Monogr. 37: 317-350.

1973. Organization of a plant-arthropod association in simple and
diverse habitats: the fauna of collards ( Brassica oleracea ).

Ecol. Monogr. 43 : 95-124,
von Rosen, K.

1929. Papilla. In (A. Seitz ed.) The Macrolepidoptera of the World.
The Palearctic Butterflies. Alfred Kernen. Stuttgart. Suppl. 1 :
7-20.

Rosenzweig, M. L.

1966. Community structure in sympatric carnivora. J. Mammal. 47:
602-612.

Rothschild, W.

1895. A revision of the Papilios of the Eastern hemisphere, exclusive
of Africa. Novitates Zoologicae Vol. II. Hazell, Watson and
Viney. London. 167-463.

Rothschild, W. and K. Jordan.

1906. A revision of the American Papilios. Novitates Zoologicae 8.
Hazell, Watson and Viney. London. 411-753.

Roughgarden, J.

1972. Evolution of niche width. Amer. Nat. 106: 683-718.

Sanders, H. L.

1968. Marine benthic diversity: a comparative study. Amer. Nat. 102:
243-282.

1969. Benthic marine diversity and the stability-time hypothesis pp. 71-
81. In Diversity and stability in ecological systems. Brookhaven
Symposium in Biology 22.

372

Psyche

[December

Schoener, T. W.

1969. Models of optimal size for solitary predators. Amer. Nat. 103:
277-313.

1971. Theory of feeding strategies. Ann. Rev. Ecol. and Syst. 2:
369-404.

SCRIBER, J. M.

1972. Confirmation of a disputed foodplant of Papilio glaucus. J.
Lepid. Soc. 26: 235-236.

Scudder, S. H.

1889. The butterflies of the eastern United States and Canada. S. H.
Scudder. Cambridge. Vol. 2.

Seitz, A.

1906. Papilio. In (A. Seitz ed.) The macrolepidoptera of the world.
The palearctic butterflies. Alfred Kernen. Stuttgart. 1: 7-18.
Selander, R. K.

1965. Sexual dimorphism and differential niche utilization in birds.
Condor 68: 113-151.

Shields, O., J. F. Emmel and D. E. Breedlove.

1969. Butterfly larval foodplant records and a procedure for reporting
foodplants. J. Res. Lepid. 8: 21-36.

Shirozu, T.

1960. Butterflies of Formosa in Colour. Hoikusha, Osaka. 481 p.
Slansky, F.

1972. Latitudinal gradients in species diversity of the New World swal-
lowtail butterflies. J. Res. Lepid. 11(4): 201-218.

Slobodkin, L. B.

1962. Growth and regulation of animal populations. Holt, Rinehart,
and Winston. New York. 184 p.

Slobodkin, L. B. and A. L. Sanders.

1969. On the contribution of environmental predictability to species
diversity, p. 82-95. In Diversity and stability in ecological sys-
tems. Brookhaven Symp. Biol. 22.

SoulE, M. and B. R. Stewart.

1970. The “niche-variation” hypothesis: a test and alternatives. Amer.
Nat. 104: 85-97.

Stichel, H.

1906. Parnassius. In (A. Seitz ed.) The Macrolepidoptera of the
World. The Palearctic Butterflies. Alfred Kernen. Stuttgart. 1 r
18-36.

1907. Parnassius. In (A. Seitz ed.) The Macrolepidoptera of the
World. The American Rhopalocera. Alfred Kernen. Stuttgart.
5: 45-48.

Straatman, R.

1962. Notes on certain Lepidoptera ovipositing on plants which are
toxic to their larvae. J. Lepid. Soc. 16: 99-103.

Stride, G. O. and R. Straatman.

1962. The host plant relationship of an Australian swallowtail, Papilio'
aegeus, and its significance in the evolution of host selection.
Linn. Soc. New South Wales 87: 69-78.

Thorsteinson, A. J.

1960. Host selection in phytophagous insects. A. Rev. Ent. 5 : 193-218.

1973]

Scriber — Papilionidae

373

Trimen, R. and J. H. Bowker.

1889. South African Butterflies. Trubner and Co. London. 3: 191-254.
VanValen, L.

1965. Morphological variation and width of ecological niche. Amer.
Nat. 99: 377-390.

Vaurie, C.

1957. Systematic notes on palearctic bird. No. 25. Motacillidae : the
genus Motacilla. Amer. Mus. Novitates 1832: 1-16.

Wellington, W. G.

1968. Qualitative changes in populations in unstable environments.
Canad. Ent. 96: 436-451.

Whittaker, R. H. and P. P. Feeny.

1971. Allelochemics : Chemical interactions between species. Science

171: 757-770.

Whittaker, R. H., S. A. Levin and R. B. Root.

1973. Niche, habitat and ecotope. Amer, Nat. 107: 321-338.
Whittaker, R. H., R. B. Walker and A. P. Kruckeberg.

1954. The ecology of serpentine soils. Ecol. 35: 258-288.

Williams, E. E.

1969. The ecology of colonization as seen in the zoogeography of
Analine lizards on small islands. Quant. Rev. Biol. 44: 345-384.

Willis, J. C.

1973. A dictionary of the flowering plants and ferns. 8th ed. Cam-
bridge Univ. Press. Cambridge. 1245 p.

Willson, M. F.

1969. Avian niche size and morphological variation. Amer. Nat. 103:
531-542.

Wilson, E. O.

1961. The nature of the taxon cycle in the Melanesian ant fauna.
Amer. Natur. 95: 169-193.

Woodhouse, L. G. O.

1950. The butterfly fauna of Ceylon. 2nd ed. The Columbo Apothe-
caries Co. Ltd. Columbo. 230 p.

Wynter-Blyth, M. A.

1957. Butterflies of the Indian region. Bombay Nat. Hist. Soc., Bom-
bay. 523 p.

Young, A. M.

1972. Community ecology of some tropical rain forest butterflies. Am.
Midi. Nat. 87(1): 146-157.

374

Psyche

[December

THE HUNDREDTH ANNIVERSARY OF THE
CAMBRIDGE ENTOMOLOGICAL CLUB

The Cambridge Entomological Club, founded on January 9, 1874,
will celebrate its 100th anniversary at a Centennial Meeting, Tues-
day, April 9, 1974. As part of the celebration, Professor Thomas
Eisner of Cornell University will deliver the Centennial Lecture on
Monday, April 8, at 5 p.m., in the main lecture room, Biological
Laboratories (Divinity Ave.), Cambridge. On Tuesday at 4 p.m.
the entomological section (4th floor) of the Museum of Comparative
Zoology and the new M.C.Z. Laboratories will hold open house for
Club members and visiting entomologists. The Centennial Meeting
on Tuesday will be held at the Harvard Faculty Club, Quincy
Street, Cambridge, at 6 p.m. Visiting entomologists are cordially
invited to attend both the Lecture and the Meeting, provided that
arrangements are made in advance with the secretary of the Entomo-
logical Club.

The March number of the Club’s journal, Psyche, which began
publication in May, 1874, has been designated the Centennial Issue
and will contain, among other articles, a history of the Club.

— F. M. Carpenter, editor

PSYCHE

INDEX TO VOLUME 80, 1973

INDEX TO AUTHORS

Barr., T. C., Jr. Speocolpodes, a New Genus of Troglobitic Beetles from
Guatemala (Coleoptera, Carabidae). 271

Benforado, J. and K. H. Kistler. Growth of the Orb Weaver, Araneus
diadematus, and Correlation with Web Measurements. 90

Carmichael, L. D. Correlation Between Segment Length and Spine Counts
in Two Spider Species of Araneus (Araneae: Araneidae). 62

Chickering , A. M. Notes on Heteroonops and Triaeris (Araneae; Oonopi-
dae). 227

Cooper, K. IV. Patterns of Abdominal Fusions in Male Boreus (Mecop-
tera). 270

Erickson, J. M. The Utilization of Various Asclepias Species by Larvae of
the Monarch Butterfly, Danaus plexippus. 230

Evans, H. E. Cretaceous Aculeate Wasps from Taimyr, Siberia (Hymen-
optera). 166

Evans, H. E. Studies on Neotropical Pompilidae (Hymenoptera). IX. The
Genera of Auplopodini. 212

Gamboa, G. and J. Alcock. The Mating Behavior of Brochymena quad-
rapustulata (Fabricius) (Hemiptera, Pentatomidae) . 265

McCluskey, E. S. Generic Diversity in Phase Rhythm in Formicine Ants.
295

Moore, /. and E. F. Legner. The Larva and Pupa of Carpelimus debilis
Casey (Coleoptera: Staphilinidae) . 289

Porter, C. C. A New Species of Anacis from Northwest Argentina (Hy-
menoptera, Ichneumonidae). 83

Ramousse, R. Body, Web-building and Feeding Characteristics of Males
of the Spider Araneus diadematus (Araneae: Araneidae). 22

Robinson, M. H. The Evolution of Cryptic Postures in Insects, with Special
Reference to Some New Guinea Tettigoniids (Orthoptera) . 159

Robinson, M. H. and B. C. Robinson. The Stabilimenta of Nephila clavipes
and the Origins of Stabilimentum-building in Araneids. 277

Roth, L. M. Paramuzoa (Nyctiborinae) , a New Cockroach Genus Previously
Confused with Parasphaeia (Epilamprinae) . 179

3 75

Roth, L. M. The Male Genitalia of Blattaria. X. Blaberidae. Pycnoscelus,
Stilpnoblatta, Proscratea (Pycnoscelinae), and Diploptera (Diplopteri-
nae). 249

Roth, L. M. The Male Qenitalia of Blattaria. XI. Perisphaeriinae. 305

Roth, L. M. and K. Princis. The Cockroach Genus Calolampra of Aus-
tralia with Descritions of New Species (Blaberidae). 101

Rovner, J. S. Copulatory Pattern Supports Generic Placement of Schizo-
cosa avida (Walckenaer) (Araneae: Lycosidae). 245

Scriber, J. M. Latitudinal Gradients in Larval Feeding Specialization of
the World Papilionidae (Lepidoptera). 355

Sedgwick, IV. C. New Species, Records and Synonyms of Chilean Theridiid
Spiders (Araneae, Theridiidae) . 349

Shear, IV. A. The Milliped Family Rhiscosmididae (Diplopoda: Chor-
deumida: Striarioidea) . 189

van Helsdingen, P. J. Annotations on Two Species of Linyphiid Spiders
Described by the Late Wilton Ivie. 48

Wheeler, G. C. and J. Wheeler. Ant Larvae of Four Tribes: Second Sup-
plement (Hymenoptera : Formicidae: Myrmicinae). 70

Wheeler, G. C. and J. Wheeler. Supplementary Studies on Ant Larvae:
Cerapachyinae, Pseudomyrmecinae and Myrmecinae. 204

Young, A. M. Notes on the Life Cycle and Natural History of Parides
areas mylotes (Papilionidae) in Costa Rican Premontane Wet Forest. 1

376

INDEX TO SUBJECTS

All new genera, new species and new names are printed in capital type.

A New Species of Anacis from
Northwest Argentina (Hymenop-
tera, Ichneumonidae) , 83

Ageniella (Cyrtagenia) fallax, 223
Ageniella (Cyrtagenia) innub a, 224
Anacis tucumana, 85

Annotations on Two Species of Liny-
phiid Spiders Described by the
Late Wilton Ivie, 48

Ant Larvae of Four Tribes: Second
Supplement (Hymenoptera : Formi-
cidae: Myrmicinae), 70

Araneus, 22, 62, 90
Araneus diadematus, 22, 90
Archaepyris minutus, 174

Body, Web-building and Feeding
Characteristics of Males of the
Spider Araneus diadematus (Ara-
neus: Araneidae), 22

Boreus, 354

Brochymena quadrapustulata, 265

Calolampra CONFUSA, 121
Calolampra darlingtoni, 105
Calolampra elegans, 103
Calolampra fenestrata, 123
Calolampra ignota, 109
Calolampra insularis, 153
Calolampra mjoebergi, 138
Calolampra pernotabilis, 153
Calolampra propinqua, 140
Calolampra queenslandica, 146
Calolampra solida, 134
Calolampra subgracilis, 119
Cambridge Entomological Club, 374
Carpelimus debilis, 289

Celonophamia taimyria, 175

Chilean theridiid spiders, 349

Cockroach Genus Calolampra of
Australia with Descriptions of
New Species (Blaberidae), 101

Copulatory Pattern Supports Generic
Placement of Schizocosa avida
(Walckenaer) (Araneae; Lycosi-
dae), 245

Correlation Between Segment Length
and Spine Counts in Two Spider
Species of Araneus (Araneae:
Araneidae), 62

Cretabythus sibiricus, 172

Cretaceous Aculeate Wasps from
Taimyr, Siberia (Hymenoptera),
166

Cryptic Postures in Insects, 159

Cryptocerus, 79

Dimorphagenia naumanni, 219

Diploptera, 249

Dipoena cartagena, 353

Evolution of Cryptic Postures in In-
sects, with Special Reference to
Some New Guinea Tettigoniids
(Orthoptera) , 159

Formicine ants, 295

Generic Diversity in Phase Rhythm
in Formicine Ants, 295

Growth of the Orb Weaver, Ara-
neus diadematus, and Correlation
with Web Measurements, 90

Heteroonops s pinimanus , 227

Hypocleptes rasnitsyni, 176

Hundredth Anniversary of Cam-
bridge Entomological Club, 374

3 77

Latitudinal Gradients in Larval
Feeding Specialization of the
World Papilionidae (Lepidoptera) ,
355

Larva and Pupa of Carpelimus de-
bilis Casey (Coleoptera: Staphi-
linidae), 289

Leptothorax , 70
Linyphiid Spiders, 48

Macromischa, 70

Male Genitalia of Blattaria. X.
Blaberidae. Pycnoscelus , Stilpno-
blatta, Proscratea (Pycnoscelinae)
and Diploptera (Diplopterinae) ,
249

Male Genitalia of Blattaria. XI.
Perisphaeriinae, 305

Mating Behavior of Brochymena
quadrapustulata (Fabricius)

(Hemiptera, Pentatatomidae) , 265

Milliped Family Rhiscosmididae
(Diplopoda: Chordeumida: Stria-
roidea), 189

Monarch Butterfly, 230
Mystacagenia albiceps, 218
Mystacagenia bellula, 217
Mystacagenia variegata, 215

Nephila clavipes, 2 77

New Species, Records, and Syno-
nyms of Chilean Theridiid Spiders
(Araneae, Theridiidae) , 349

Notes on HeteroonoPs and Triaeris
(Araneae: Oonopidae), 227

Notes on the Life Cycle and Natural
History of Parides areas mylotes
(Papilionidae) in Costa Rican
Premontane Wet Forest, 1

Ocymyrmex, 75

Orb Weaver, 23, 62, 90

Or eon elides recurvatus, 55

Papilionidae, 355
Paras pacria, 183

Paramuzoa (Nyctiborinae), a New

Cockroach Genus Previously Con-
fused with Parasphaeia (Epilam-
prinae) , 179

Parides areas mylotes, 1

Patterns of Abdominal Fusions in
Male Borens (Mecoptera), 354

Phoroneidia pennata, 353
PlTTOECUS PAUPER, 171
Pompilidae, 212
Procryptocerus, 79
Proscratea, 249
Protamisega khatanga, 177
Pycnoscelus , 249

R h i s c oso m id es, 191
Rhiscosomidcs trinitarium, 203
Rogeria, 74

Schizocosa avida, 245

Speocolpodes, a New Genus of Trog-
lobitic Beetles from Guatemala
(Coleoptera, Carabidae), 271

Speocolpodes franiai, 272

Stabilimenta of Nephila clavipes and
the Origins of Stabilimentum-
building in Araneids, 277
Stilpnoblatta, 249

Studies on Neotropical Pompilidae
(Hymenoptera) . IX. The Genera
of Auplopodini, 212

Supplementary Studies on Ant
Larvae: Cerapachyinae, Pseudo-

myrmecinae, and Myrmecinae, 204

Taimyrsphex pristinus, 168
T aranuenus ornithes, 48
T etramorium, 76
Theridion elli, 352
Theridion whitcombi, 350
Triaeris pusillus, 228
Triglyphothrix, 78

Utilization of Various Asclepias
Species by Larvae of the Monarch
Butterfly, Danaus plexippus, 230

378

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 is of Dr. Hermann A.
Hagen, Professor of Entomology and Curator in the Museum of Compara-
tive Zoology at Harvard University from 1870-1893. Professor Hagen,
along with S. H. Scudder, provided the initiative for the founding in 1874
of the Cambridge Entomological Club and its journal Psyche, now in their
100th year. [The illustration is a reproduction of an engraving in the
Archives of the Harvard College Library.]

BACK VOLUMES OF PSYCHE

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

F. M. Carpenter

Editorial Office, Psyche
16 Divinity Avenue

Cambridge, Mass. 02138

FOR SALE

Classification of Insects, by C. T. Brues, A. L. Melander and
F. M. Carpenter, Published in March, 1954, as volume 108 of the
Bulletin of the Museum of Comparative Zoology, with 917 pages
and 1219 figures. It consists of keys to the living and extinct families
of insects, and to the living families of other terrestrial arthropods;
and includes 270 pages of bibliographic references and an index of
76 pages. Price $14.00 (cloth bound and postpaid). Send orders
to Museum of Comparative Zoology, Harvard College, Cambridge,
Mass. 02138.

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