PSYCHE

A Journal of Entomology

Volume 68

1961

Editorial Board

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

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

E. O. Wilson H. E. Evans

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

Cambridge, Mass., U. S. A.

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

Vol. 67, no. 4, Dec., 1960: July 7, 1961

Vol. 68, no. 1, March, 1961 : August 2, 1961

Vol. 68, nos. 2-3, June-Sept., 1961: December 29, 1961

PSYCHE

A JOURNAL OF ENTOMOLOGY

Established in 1874

Vol. 68

March, 1961

No. 1

CONTENTS

Australian Carabid Beetles V. Transition of Wet Forest Faunas from
New Guinea to Tasmania, P. J. Darlington , Jr. 1

A Reconsideration of the Genus Epipompilus ( Hymenoptera : Pompilidae).

H. E. Evans 25

The Rediscovery and Probable Phylogenetic Position of Psilopsocus
(Psocoptera) . E. L. Mockford

38

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1961-62

President ...J. J. T. Evans, Harvard University

Vice-President C. Walcott, Harvard University

Secretary A. R. Brady, Harvard University

Treasurer F. M. Carpenter, Harvard University

Executive Committee R. W. Taylor, Harvard University

S. K. Harris, Boston University

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

P. J. Darlington, Jr., Head Curator of Insects, Museum of Com-
parative Zoology

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

E. 0. Wilson, Associate Professor of Zoology Harvard University
H. W. Levi, Associate Curator of Arachnology, Museum of Com-
parative Zoology

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

PSYCHE is published quarterly by the Cambridge Entomological Club, the
issues appearing in March, June, September and December. Subscription
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Checks and remittances should be addressed to Treasurer, Cambridge Ento-
mological Club, Biological Laboratories, Harvard University, Cambridge, Mass.

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

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

Authors contributing articles over 8 printed pages in length may be required
to bear a part of the extra expense, for additional pages. This expense will
be. that of typesetting only, which is about $6.00 per page. The actual cost of
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AUTHOR’S SEPARATES

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

The December i960 Psyche (Vol. 67, no. 4) was mailed July 7,
1961. The present issue of Psyche (Volume 68, no. 1) is pub-
lished with the aid of a National Science Foundation Grant
(G 15817) to the Cambridge Entomological Club.

The Lexington Press. Inc., Lexington, Massachusetts

PSYCHE

Vol. 68 March, 1961 No. 1

AUSTRALIAN CARABID BEETLES V.1 TRANSITION OF
WET FOREST FAUNAS
FROM NEW GUINEA TO TASMANIA

By P. J. Darlington, Jr.

Museum of Comparative Zoology, Cambridge, Mass.

Introduction

Beetles of the family Carabidae (predaceous ground beetles) are
numerous in tropical rain forest in New Guinea and numerous also
(but less diverse) in cool south temperate rain forest in Tasmania,
but no species and hardly any genus is common to the two faunas, and
even the dominant tribes are different. However there is no single
boundary between the New Guinean and Tasmanian faunas, but a
broad and complex transition, which I shall try to describe.

My interest in this part of the world began with the Harvard
Australian Expedition of 1931-1932, when I collected Carabidae in
eastern Australia north to part of the Cape York Peninsula, as well
as in southwestern Australia. In 1943-1944 I spent eleven months in
New Guinea as an army entomologist, and was able to collect
Carabidae especially in lowland rain forest at Dobodura, Papua, while
hospitalized there, and in mountain forest on the Bismarck Range,
Northeast New Guinea, in lieu of leave. I have sorted and arranged
my own and much borrowed material and am now more than half way
through writing “The Carabid Beetles of New Guinea” (see Darling-
ton 1952), so that I have a good knowledge of New Guinean Carabi-
dae. Recently, from December 1956 to June 1958, I have been again
in eastern Australia, traveling and living in a small truck with my
wife and fourteen-year-old son, and collecting Carabidae in practically
every important piece of wet forest from the northern tip of Cape

Tarlier parts of this series are listed in the reference list at the end of
this paper.

]

2

Psyche

[March

York to the southern tip of Tasmania.2 A brief itinerary with maps
and list of localities has been published (1961). Information and
collections obtained during this trip have enabled me to correlate other
information and write the present paper. New genera and species re-
ferred to now (but not by name) will be described in forthcoming
numbers of Psyche and Breviora.

The Forests

My “wet forests” are rain forests as classified in “The Australian
Environment” (CSIRO 1950, 77-96). That is, they are dense, ever-
green (non-deciduous) forests with closed canopies, often (in tropical
rain forest) with many woody vines, but with comparatively little
low vegetation, the ground being covered with dead leaves and leaf
mold rather than grass or herbs.

Two main types of rain forest exist in the Australian Region:
tropical (including subtropical) (Figs. 1, 2) and south temperate
(Figs. 3, 4). Tropical rain forest is widely distributed in New
Guinea at low and middle altitudes, although in the drier country of
southern New Guinea it is replaced by op n savannah woodland like
that of much of northern Australia. Tropical rain forest occurs also
on the eastern edge of Australia in separate tracts spaced irregularly
from parts of Cape York south through Queensland and northern
New South Wales (map, Fig. 6). The best of this forest in tropical
and subtropical Australia as well as in New Guinea is real, Malaysian-
type rain forest, although some tracts in Australia are lighter and
seasonally drier, and light rain forest sometimes grades into semi-
deciduous monsoon forest.

The northernmost rain forest in Australia is the tip-of-peninsular
(Fockerbie or Somerset) tract on the tip of Cape York. It is lowland
rain forest, but somewhat depauperate (see p. 17) .

2This trip was supported in part by a fellowship of the John Simon Guggen-
heim Memorial Foundation. I am especially indebted to Dr. L. J. Webb, of
the Commonwealth Scientific and Industrial Research Organization, for in-
formation on the distribution of rain forest in Queensland, to many members
of the Queensland Department of Forestry who aided or guided us in the field,
and to Mr. P. J. Killoran, of the Queensland Department of Native Affairs,
who arranged our visit to Bamaga and the tip of Cape York. I very much
regret that I do not have space to acknowledge other assistance in detail here.

Explanation of Plate 1

Fig. 1. Tropical rain forest, Lake Barrine, Atherton Tableland, North
Queensland (P. J. D. 1932).

Fig. 2. Interior of tip-of-peninsular (tropical) rain forest, from edge of
new clearing, Lockerbie, Cape York, Queensland (P. J. D. 1958). This is
the habitat of Mccynognathus.

Psyche, 1961 Vol. 68, Plate

Psyche, 1961

Vol. 68, Plate 2

4

1961]

Darlington — Australian Carabid Beetles

5

Next in order southward is a gap more than ioo miles wide of
drier, open savannah woodland (Fig. 5) in which may be an isolated
piece of rain forest near the head of the Jardine River, unknown
biologically (Brass 1953, pp. I54> 161).

Next is the mid-peninsular rain forest system. It extends irregularly
and with perhaps slight interruptions from near Iron Range and Mt.
Tozer south to the “Rocky Scrub” east of Coen. Altitudinally it ex-
tends from near sea level ( e.g . at Iron Range) to about 2,000 ft. on
the higher summits of the Mcllwraith Range. It includes fairly heavy
rain forest, although its quality varies locally.

Fig. 5. Rather dry savannah woodland northeast of Coen, Cape York
peninsula. (P. J. D. 1932). Such woodland is an effective barrier to rain
forest Carabidae in the tropics.

Next, after another gap more than 150 miles wide of drier, open
woodland, is the base-of-peninsular or main tropical rain forest system
of North Queensland. Outlying pieces of semi-rain forest of this
system are within sight of Cooktown, and heavier rain forest begins on
the coastal mountains (Mt. Amos, Mt. Finnigan) about 20 miles to

Explanation of Plate 2

Fig. 3. South temperate rain forest, Lake St. Clair, Tasmania (P, J, D.
1957). On left is transitional wet forest with overstory of big eucalypts;
center, heavy rain forest including N othofagus.

Fig. 4. Interior of old south temperate rain forest, Cradle Valley, northern
Tasmania (courtesy Mr. H. J. King, Honorary Photographer, and Mr. Frank
Ellis, Director, Queen Victoria Museum, Launceston).

6

Psyche

[March

the south. From here an irregular system of good rain forests extends
somewhat discontinuously but with no very wide breaks south to and
across the Atherton Tableland and farther south along a series of
plateaus and ranges to the Mt. Spec plateau (Paluma Range) almost
within sight of Townsville. Much of this forest system lies between
1,000 and 5,000 ft. altitude, but areas of good rain forest belonging
to it occur (or occurred before being cleared) also on the coastal plain
east of the Atherton Tableland and in the Mossman-Daintree region.

From the southern end of the main tropical rain forest system to
below Rockhampton is a gap of nearly 500 miles of dry, open wood-
land broken only (so far as I know) by two noteworthy islands of
rain forest. One is at about 3,000-4,000 ft. on the crest of the Elliot
Range, within sight of (southeast of) Townsville but separated from
the northern rain forests by a low, comparatively dry valley. The
other, more important island of rain forest is on the Eungella Range
about 40 miles inland from Mackay, at about 2,000-4,000 ft. altitude.
Scattered fragments of semi-rain forest, for example near Proserpine
(Repulse Bay) and Yepoon (Byfield), are relatively unimportant so
far as carabid distribution is concerned.

South of Rockhampton, in the edge of the south temperate zone,
begins what I call the subtropical rain forest system. The first piece
of (rather poor) rain forest of this system is on Mt. Jacob east of
Many Peaks. Other tracts are widely scattered in southeastern
Queensland at low altitudes as well as on mountains (Blackall Range,
Bunya Mts., Mt. Tamborine, McPherson Range on the New South
Wales border, etc.). The different forest tracts vary in quality, but
the best of them approximate tropical rain forest. This system of
rain forests extends into northeastern New South Wales at rather
low altitudes, although much of it has now been cleared. The more
important pieces that still remain are listed and briefly described in
my published locality list (1961). The most southern good tract that
seemed to me to be tropical-type rain forest is on “Mt. Dorrigo”, on
the lower (eastern) edge of the Dorrigo Plateau, at about 30° 20' S.,
but small pockets of more or less similar forest occur still farther
south, even south of Sydney, especially in wet ravines.

South temperate rain forest (see again Figs. 3, 4) is different in
aspect from tropical rain forest (fewer vines, etc.) and different
botanically, often dominated by southern beeches (Nothofagus) .
Such forest is widespread in southwestern Tasmania and occurs in
isolated tracts elsewhere in Tasmania (see paper referred to above for
details). Isolated tracts of similar forest occur on plateaus and moun-
tains in southern Victoria including the Otway Ranges southwest of

1961]

Darlington — Australian Carabid Beetles

7

Melbourne and some of the southern “Victorian Alps” east of Mel-
bourne. This kind of forest occurs also, at wide intervals, on isolated
plateaus in eastern New South Wales, notably on the plateau of the
Mt. Royal Range (Barrington Tops and Tomalla Tops) at about
310 50' S. and on the higher part of the Dorrigo-Ebor Plateau
(especially at Point Lookout in New England National Park) at
about 30° S. Both these plateaus reach about 5000 ft. above sea level.
The northernmost Nothofagus in Australia is still farther north, on
the southern border of Queensland, where small tracts of old trees
exist on the highest points of the McPherson Range, at about 28° 20'
S. and 4,000 ft. altitude. Nothofagus does not occur on the mountains
of tropical North Queensland but is dominant in New Guinea in
mountain forests between about 6,500 and 10,000 ft. (Womersley
and McAdam 1957, p. 25). However, south temperate groups of
Carabidae do not occur in the New Guinean Nothofagus forests.

The distribution of tropical (including subtropical) and south
temperate rain forest is shown, rather diagramatically, on the accom-
panying map (Fig. 6). The map is based partly on the vegetation
map in “The Australian Environment” (CSIRO 1950, pp. 88-89)
and on Brass’s (1953, p. 152) map of Cape York rain forests, but
many details are modified according to my own observations. In most
cases rain forest is not continuous within the boundaries shown, but
occurs as irregular, sometimes discontinuous tracts and strips inter-
spersed with savannah woodland (in the north) and/or sclerophyll
forest (in the south). The two kinds of rain forest overlap widely in
New South Wales. Within the area of overlap south temperate rain
forest is usually above (at higher altitude than) tropical rain forest,
but there is some mixing.

The Carabidae

The wet-forest Carabidae of New Guinea and Australia, including
Tasmania, are numerous, diverse, and complex in ecology and distri-
bution. They form three general ecological groups. Those that live on
the ground without being specially associated with surface water are
mesophiles or geophiles. Those that live on the ground beside streams
or ponds or in swamps are hydrophiles. And those that live on tree
trunks or in foliage above the ground are arboreal. According to my
(i943, P- 41) rough analysis of the Australian carabid fauna, at least
half the species are geophiles, not quite a quarter hydrophiles, and not
quite a quarter arboreal. The carabid fauna of New Guinea divides
in something like the same way, although I cannot yet give exact
figures.

Psyche, 1961

Vol. 68, Plate 3

NEW GUINEA

TASMANIA

1961]

Darlington — Australian Carabid Beetles

9

State of wings of Carabidae is correlated with ecology and distri-
bution. Most Carabidae in most parts of the world have fully devel-
oped inner wings and can fly, but some have lost their wings (except
for vestiges) and become flightless. The Australian carabid fauna
includes an unusually large proportion of flightless species : according
to my rough analysis (loc. cit.), nearly 45% of all Australian Carabi-
dae have atrophied wings, and many genera and even some tribes are
wholly flightless. Most hydrophiles and arboreal forms have retained
their wings and can fly, but about 75% of Australian geophile Carabi-
dae are flightless, and flightless groups are common everywhere in
Australia, at low and high altitudes and in wet and dry climates, and
some are well represented in the tropical as well as the temperate parts
of the continent. In New Guinea flightnessess is rare among lowland
Carabidae. This accords with the general rule that most Carabidae in
most wholly tropical lowland areas are winged. On mountains in
New Guinea, however, as on many tropical mountains elsewhere,
flightless geophile Carabidae are numerous.

New Guinea-Tropical Australian Relationships
Probably the first fact that strikes entomologists collecting in the
rain forests of tropical Australia is that some of the insects are species
that occur in New Guinea. This is expected. The Australian rain
forests themselves are predominantly New Guinean (or Malaysian)
both in aspect and in botanical relationships (CSIRO 1950, pp. 95-
96; Brass 1953? P* 154) ; many mammals in the North Queensland
rain forests belong to New Guinean genera or even species; and so
do many birds. Some Carabidae are common to New Guinean and
Australian rain forests. For example Syleter papua Dari, extends to
the tip of Cape York, living on the ground in shaded swamps. Morion
longipenne Putz. of New Guinea extends to the main North Queens-
land rain forests, on and in fallen logs. And Violagonum violaceum
(Chd.) is common in rain forest in New Guinea and eastern Australia
south at least to near Rockhampton, in accumulations of dead leaves
on the ground and in thick foliage. Besides shared species like these
(there are many others among Carabidae) the New Guinean and
Australian rain forests share some geographically restricted genera,
for example Platycoelus ( Chlaenioidius) , Loxandrus , and Stricklandia,

Explanation of Plate 3

Fig. 6. Distribution of rain forests in eastern Australia. Solid lines enclose
principal areas of tropical (including subtropical) rain forest; broken lines,
of south temperate rain forest. In most cases rain forest is not continuous
within the boundaries shown but occurs in discontinuous or scattered tracts.
See text for further details.

10

Psyche

[March

as well as many more-widely distributed genera. Up to a point, there-
fore, the Carabidae agree with the forest trees, mammals, and birds
in showing a considerable number of species and genera common to
the rain forests of New Guinea and tropical Australia.

When I was collecting on the Atherton Tableland in northeastern
Australia in 1932, I found not only many Carabidae of obviously New
Guinean groups but also, in rain forest, many species of Australian
groups not known to occur in New Guinea. Included were striking
endemic species of Notonomus , Trichosternus, Leiradira , Pamborus,
and Mystropomus. Knowing, as I did, that the rain forests of
Australia and New Guinea had much in common, and knowing that
the Carabidae of New Guinea were poorly collected, I imagined in
New Guinea a rich fauna of the genera just named, perhaps in rain
forest at middle altitudes, but wholly unknown. It was a sort of El
Dorado for the future, to a young and enthusiastic carabid student.
But now that I have collected in New Guinea and seen thousands of
Carabidae collected there by other persons, I know that this El Dorado
does not exist, and I know why. All the Carabidae common to the
New Guinean and Australian rain forests are winged and probably
fly. All the genera mentioned above as represented in rain forest on
the Atherton Tableland are wholly flightless, and I know now that
there is no direct relationship between any flightless Carabidae of the
New Guinean and Australian rain forests.3

The difference between the flightless Carabidae of Australia and
New Guinea goes far beyond mere differences of species and genera.
The composition and origins of the two faunas are fundamentally
different. Flightless Carabidae are numerous everywhere in Australia,
even at low altitudes in the tropical part of the continent including
Cape York. Many of the species belong to wholly flightless genera or
even flightless tribes that have evidently been in Australia a long time.
Derivatives of old Australian flightless groups dominate the flightless
ground-living carabid fauna of tropical rain forest in Australia. In
New Guinea, in contrast, no primarily flightless groups of Carabidae
occur at low altitudes. A very few species of the primarily winged

3If tiger beetles are considered Carabidae, Tricondyla aptera 01. is an
exception to this rule. The genus Tricondyla is primarily Oriental and is
wholly flightless. Nevertheless T. aptera has reached New Guinea, probably
rather recently (it is only slightly differentiated there), and has got beyond
New Guinea to the mid-peninsular rain forests of Cape York. (It has reached
the Solomon Islands and New Hebrides too.) It is a good sized (nearly an
inch long), big-eyed, ant-like, active insect, which lives on tree trunks in rain
forest. It has probably dispersed on floating trees, which ground-living
Carabidae are not likely to do.

1961]

Darlington — Australian Carabid Beetles

1 1

genera Clivina, Tachys, Lesticus , Tlatycoelus , and Loxandrus have
undergone wing atrophy at low altitudes in New Guinea (Darlington
in press) , but they have evidently done it recently, in situ. Some of the
species are still dimorphic, with fully winged individuals occurring
with the short winged ones, and all the short winged lowland species
are closely related to long winged ones that still exist in New Guinea.
It is only above about 5000 ft. in the mountains that flightless Carabi-
dae become numerous in New Guinea, and they too have apparently
undergone wing atrophy in situ. That is, they have been derived on
the mountains of New Guinea from winged ancestors, and do not
represent flightless stocks of other regions. This is my conclusion after
making formal studies of the New Guinean representatives of the two
principal tribes concerned, the Agonini (Darlington 1952, especially
table p. 108) and Pterostichini (in press).

Besides the change of specific flightless stocks from New Guinea to
Australia there is a change of dominance of tribes. In New Guinea,
Agonini are much more numerous than Pterostichini, and most flight-
less Carabidae of the island are agonines. But in Australia, even in
the tropical rain forest, Pterostichini are overwhelmingly dominant
and include most of the flightless forms. This striking shift of domi-
nance is further discussed on page 22.

The first important finding of the present study, then, is that, al-
though the rain forests of New’ Guinea and tropical Australia are
similar and share many species of plants, mammals, birds, and winged
insects including many winged Carabidae, they have wholly different
faunas of flightless Carabidae, which differ not only in taxonomic
details but also in general ecology (in relation to altitude), in origin
of the flightless stocks, and in relative dominance of tribes.

Transition in Australia: South from the Tropics

Now to be considered is the transition of wet forest carabid faunas
within the limits of Australia and Tasmania.

Five important genera of flightless geophile Carabidae are mentioned
above as occurring in rain forest on the Atherton Tableland. Of
these five genera, Notonomus is most dominant. It is a genus of about
100 species, confined to eastern and southeastern Australia and Tas-
mania except for one species isolated in southwestern Australia. The
genus’ northern limit is between Daintree and Cooktown. It is repre-
sented by several species (some very localized) in the main tropical
rain forest system of North Queensland, where it seems to be confined
to rain forest. It is well represented in the subtropical rain forests of
South Queensland and northern New South Wales and south through

12

Psyche

[March

eastern New South Wales and southern Victoria; in these areas some
species occur not only in rain forest (including south temperate rain
forest) but also in wet sclerophyll forest and good savannah woodland.
However only two groups of the genus reach Tasmania and only one
group (two related, primarily allopatric species) occurs in rain forest
there.

Trichosternus is a genus of 25 or more species confined to eastern
Australia, except that one species is isolated in southwestern Australia
(Darlington 1953, p. 94). The genus’ northern limit is between
Daintree and Cooktown. It occurs (several species, some very local-
ized) throughout the main rain forest system of North Queensland,
where it is apparently confined to rain forest. It is well represented
also in the subtropical rain forest system of South Queensland and
northern New South Wales, and in this area some species occur in
savannah woodland as well as in tropical-type rain forest, and some
have entered south temperate rain forest on the Dorrigo-Ebor plateau
and the Mt. Royal Range. The southern limit of the genus is some-
where in east-central New South Wales, probably not far north of
Sydney.

The northern limit of Leiradira (or of the group of genera that
includes Leiradira) is between Daintree and Cooktown. This genus
too occurs in much of the main tropical rain forest system of North
Queensland, being represented there by several distinct species each
more or less localized, but the genus may be absent in the southern
extension of the main tropical rain forest system south of the Atherton
Tableland. It is represented also by several species in the subtropical
rain forests of South Queensland etc. Its southern limit is apparently
on the lower, eastern edge of the Dorrigo plateau. It is confined to
eastern Australia. It is wholly or chiefly a rain forest genus in all
parts of its range.

The three preceding genera are all Pterostichini. All their species
are flightless geophiles. Additional flightless geophile pterostichines
are localized in all the different rain forest areas of Australia from
Cape York to Tasmania. Examples are Mecynognathus in the tip-of-
peninsular forests; Paranurus in the mid-peninsular forests; Loxo-
genius and undescribed genera in the main tropical rain forest system ;
Nursus s. s.j Liopasa, Ceratoferonia Zeodera, and Notolestes in the
subtropical rain forest system ; Loxodactylus in the wet forests of
southern Victoria; and Rhabdotus ia those of Tasmania. (It should
be added that Australia possesses many winged pterostichines as well
as these and other flightless genera.)

1961] Darlington — Australian Carabid Beetles 13

Of non-pterostichines, Pamborus is noteworthy. It is confined to
eastern Australia and is one of the two known genera of the tribe
Pamborini. (The other is monotypic Maoripamborus in New Zea-
land— Brookes 1944.) The northern limit of Pamborus is probably
near Cooktown. Four species of the genus occur in the main tropical
rain forest system of North Queensland, chiefly or wholly in rain
forest. Six other species occur in South Queensland and New South
Wales. Some of them occur mainly in (sub) tropical rain forest, but
viridis inhabits savannah woodland and some other species occur in
open woods as well as rain forest, and some enter south temperate
rain forest on the high plateaus of north-central New South Wales.
The southern limit of the genus is near the Shoalhaven River about 70
miles south of Sydney. (Old records for Victoria are probably errors.)

The genus Mystropomus is the only Australian representative of the
pantropical tribe Ozaenini. The genus is confined to eastern Australia.
Its northern limit is between Daintree and Cooktown. A single species
(two subspecies) occurs throughout the main tropical rain forest sys-
tem of North Queensland, and is apparently confined to rain forest.
Another, variable species (two subspecies) occurs in the subtropical
rain forest system, and extends into more open woodland. The south-
ern limit of the genus is apparently near Sydney.

These five genera dominate the flightless geophile carabid faunas
of the main tropical and subtropical rain forest systems of eastern
Australia. Their distribution is notable in several ways. All five
genera reach an approximately common northern limit, north of Dain-
tree and south of or near Cooktown. All five genera are widely dis-
tributed both in the main tropical and in the subtropical rain forest
systems. These two forest systems are separated by a wide barrier of
comparatively dry, open forest in which is one important “island” of
rain forest, on the Eungella Range west of Mackay, and all five of
the genera in question are represented there.4 In the tropics, these
genera occur only or chiefly in rain forest,5 although most of them
enter opener forest too in the south temperate zone.

4Of the 5 genera in question on the Eungella Range, the one Pamborus has
close relatives in both North and South Queensland. The one Mystropomus is
a South Queensland species. Of 2 Trichosternus, one probably belongs to a
South Queensland group and the other is doubtful. The one Notonomus be-
longs to a North Queensland group. And the one Leiradira belongs to a
South Queensland subgenus. These genera in the Eungella rain forest there-
fore show 2 close ties with North Queensland (in Pamborus and Notonomus)
and 4 with South Queensland.

5 Trichosternus cordatus Chd. occurs outside rain forest in the southern
edge of the tropics.

14

Psyche

[March

Southward, through New South Wales, rain forest of (sub) tropical
type diminishes in area and quality, and the Carabidae associated with
it diminish too. Of the five genera just discussed, Leiradira may not
extend south of the Dorrigo. T richosternus , Mystropomus , and Pam-
borus go a little farther south, reaching different limits probably in
this order, but do not reach Victoria. And N otonomus reaches Vic-
toria (in numbers) and Tasmania (only one stock in rain forest).
Toward their southern limits, all these genera, except Leiradira , occur
not only in tropical-type rain forest but also in opener forest, and all,
except again Leiradira , have entered or even evolved endemic species
in south temperate rain forest on the Dorrigo-Ebor plateau and the
Mt. Royal Range.

Transition in Australia: North from Tasmania

The ground-living Carabidae of the south temperate rain forest of
Tasmania are dominated by or include flightless genera of four special
tribes in addition to the more widely distributed Pterostichini, Licinini,
etc.

The tribe Broscini is well represented in both the north and the
south temperate zones of the world (Ball 1956) but is absent in the
tropics or nearly so. Some northern broscines have well developed
wings, but I think that all those of the southern hemisphere have atro-
phied wings and are flightless. Four genera occur in Tasmania.
Promecoderus is represented there by several rain forest species and by
other species that live in drier, opener woodland. The genus is widely
distributed across southern Australia, but chiefly in dry forest and
arid country, although one or two species occur in rain forest in
Victoria. Of the other Tasmanian genera, Chylnus is confined to
Tasmania, in wet forest. Per cosoma occurs in Tasmania and the
mountains of southeastern Victoria, in wet forest. And Eurylychnus
occurs in Tasmania, southern Victoria etc. including the Otway
Ranges, and east and north into southern New South Wales, and two
separate stocks of the genus have species isolated (chiefly in south
temperate rain forest) on the Mt. Royal Range and the Dorrigo-Ebor
plateau. The latter is the northern limit of wet-forest broscines in
Australia.

The tribe Trechini (subfamily Trechinae of Jeannel 1926-1928)

Explanation of Plate 4

Fig. 7. Diagram of transition of selected flightless geophile Carabidae in
rain forests of eastern Australia. The 5 genera at bottom of the diagram
are primarily tropical and subtropical ; the other genera and tribes, pri-
marily south temperate. See text for further details.

Psyche, 1961

Vol. 68, Plate 4

Tasmania

i6

Psyche

[March

is world-wide in distribution. It includes both flying and flightless
genera, but the flying genera usually live beside standing or running
water and are not forest-living geophiles. Flightless geophile Trechini
are numerous both north and south of the tropics. In Tasmania they
are numerous in south temperate rain forest and hardly enter other
habitats at low altitudes, although some occur in open country above
timber line, on cold mountain tops. Flightless Trechini are less numer-
ous but still widely scattered in wet forests and on mountain tops in
southern Victoria, including the Otway Ranges (Moore i960), east
nearly to the New South Wales border and north to Mt. Kosciusko
in southern New South Wales, and endemic species perhaps represent-
ing one original flightless stock of spotted " Trechus ” are isolated on
the Mount Royal Range, the Dorrigo-Ebor plateau, and the Mc-
Pherson Range on the Queensland border.6

The tribe Migadopini (Jeannel 1938; Darlington i960, p. 663)
is confined to the southern hemisphere, with different genera localized
in Tasmania and southeastern Australia, New Zealand, and the south-
ern tip of South America, etc. Two flightless genera of the tribe occur
in Tasmanian rain forest: Calyptogonia is confined to Tasmania;
Stichonotus extends to the mainland, but only to the Otway Ranges.
A third Australian genus of the tribe is known from a single specimen
collected long ago near Kiama south of Sydney, and a fourth genus
occurs still farther north, in subtropical forest on the low (c. 2,000
ft.) Comboyne plateau at about 3 1 0 35' S. This last genus, Decogmus,
differs from all other Migadopini in being winged.

Finally, the flightless tribe Agonicini is confined to Tasmania and
southeastern Australia (Moore i960). There are two genera. One
is widely distributed in Tasmania and occurs also in the mountains
of southern Victoria east of Melbourne (B. P. Moore, in letter).
The other is confined to the mainland, including the Otway Ranges
and the “Victorian Alps,” north to Mt. Kosciusko. Agonicines live
on the ground in rain forest, and sometimes in open snow gum woods
on mountains.

Although there are other Carabidae in Tasmanian rain forests
(especially various Pterostichini and Licinini) the four tribes just
discussed make up a large part, and zoogeographically the most im-
portant part, of the flightless wet forest Tasmanian carabid fauna. It
will be seen from details given above that all four tribes occur both in

6 A second “Trechus”, diemensis Bates, extends from Tasmania and south-
eastern Australia north to the McPherson Range, but this species is winged
or dimorphic.

1961]

Darlington — Australian Carabid Beetles

7

Tasmania and on the adjacent mainland of Australia, but that they
all diminish rapidly northward.

The transition of selected elements of the flightless geophile carabid
faunas of tropical and south temperate rain forests is diagrammed in
Fig. 7-

Isolated Australian Faunules

To return to the five carabid genera discussed above as characteristic
of the main tropical and subtropical rain forests, these genera have dis-
tributions that are alike in many details. Within the main (base-of-
peninsular) tropical rain forest system, they all have almost the same
northern limits and (excepting perhaps Leiradira ) the same southern
limits. All are represented on the Eungella Range. In South Queens-
land, all apparently find their northern limit on Mt. Jacob (except
that Trichosternus cordatus extends farther north in drier woodland),
and all extend well into New South Wales, although they reach
different limits there. They illustrate a general fact, that the carabid
faunas of the main tropical and subtropical rain forest systems of east-
ern Australia, although separated by several hundred miles of com-
paratively dry country, are fundamentally similar, dominated by the
same tribes, and share many genera some of which coincide remark-
ably in details of distribution, although some other genera and most
species are different. However three isolated pieces of Australian rain
forest have carabid faunules that do not flt into this main pattern.
They are the tip-of-peninsular and mid-peninsular rain forests of
Cape York and the rain forest on the Elliot Range south of Towns-
ville.

The tip-of-peninsular tract is light rain forest and is limited both
botanically and zoologically. For example, st’nging trees (Laportea) ,
which occur in other Australian rain forests and in New Guinea, are
apparently absent in the tip-of-peninsular forest, and land leeches and
itch mites, which are pests in rain forest elsewhere, are apparently
absent in the tip-of-peninsular tract. The winged Carabidae of this
tract are not remarkable, except that they include New Guinean
species. But the flightless Carabidae form a faunule wholly different
from that of any other rain forest, consisting (so far as I could find)
of only two flightless species. One is Mecynognathus dameli Mach,
an enormous carabid, the largest males 2V2 inches long with mandibles
like stag beetles. The genus occurs nowhere else on earth, although
it may be rather closely related to Paranurus (see below). The other
is a large flightless Clivina (probably kershawi SI.), which is fairly
common both in the rain forest and in adjacent savannah woodland.
The nature of this forest and of its flightless Carabidae suggests that

i8

Psyche

[March

the tip-of-peninsular tract is not a remnant of a larger, continuous
rain forest but has been constituted or reconstituted separately, by
gradual accumulation of a limited variety of plants and animals.

The mid-peninsular rain forests of Cape York are heavier and
more extensive than the tip-of-peninsular tract, more like the base-of-
peninsular forests at least superficially, but their flightless Carabidae
form a second independent faunule. None of the flightless genera
characteristic of the other rain forests is represented in the mid-
peninsular system. In their place is a single large species of Parcimirus.
This is a genus of probably only one, geographically variable species,
which occurs from the tip of Cape York (and islands off the tip) south
to below Cairns mainly in good savannah woodland. In most parts
of its range it apparently does not enter rain forest, but it has done so
in the mid-peninsular system, where it is now widely distributed. It
seems to have invaded this system recently. An earlier invasion of the
tip-of-peninsular rain forest by the ancestral stock of Paranurus may
have produced Mecynognathus . There is also in the mid-peninsular
rain forest a flightless Coptocarpus, but it is small and rare and I am
not sure of its habitat or relationships. And also in this forest is a
large form of Lesticus chloronotus Chd. It is winged, but its distri-
bution and behavior suggest that it may eventually become flightless,
as several stocks of the same genus have done in New Guinea. The
Carabidae, then, suggest that the mid-peninsular rain forest has not
been connected with the main base-of-peninsular system but, like the
tip-of-peninsular tract, has derived or is deriving its flightless Carabi-
dae independently.

The rain forest on the Elliot Range is poorly known. The only
insect collecting ever done in it, so far as I know, was done March 2,
1958, when my son and I climbed from Double Creek to near the
peak of Sharp Elliot and worked for three or four hours in the forest
there. It seemed to be real but rather light rain forest. We found
there series of two conspicuous flightless Carabidae: a very big Nurus
and a Notonomus, both endemic. No trace of the four other genera
(other than N otonomus) discussed above as characteristic of the main
tropical and subtropical rain forests of Australia was found. Judging
from my experience elsewhere, we would probably have found speci-
mens or fragments of other species if the carabid fauna were diverse.
I think, therefore, that the rain forest of the Elliot Range probably
has a limited, endemic faunule of flightless Carabidae presumably
received across a barrier and not by way of continuous rain forest.
The valley that separates the Elliot Range from the main mountain

1961]

Darlington — Australian Carabid Beetles

19

system of North Queensland is not much more than ten miles wide,
but it seems to have been a more effective barrier than the much wider
gaps of dry hilly country between the North Queensland, Eungella,
and South Queensland rain forest areas.

Summary of Transition from New Guinea to Tasmania

The transition of wet forest carabid faunas from New Guinea to
Tasmania involves two main changes. First, between the rain forests
of New Guinea and those of tropical Australia is a complete change
of flightless stocks of Carabidae and also a change from Agonini to
Pterostichini as dominant tribes, although the change is overlain and
superficially concealed by many winged species and genera of Carabi-
dae that are common to New Guinea and Australia and that form a
broad and complex transition, not fully described here. Between the
tropical rain forests of North Queensland and the subtropical ones
of South Queensland etc. are very many changes of species and genera
but no fundamental change in the nature of faunas or in dominant
groups. The second main change is farther south, and is a complex
transition from tropical to south temperate groups. The area of transi-
tion (of overlapping and mixing of faunal elements) is from the
southern edge of Queensland to Tasmania. And the transition in-
volves not only changes of species and genera but a second partial
change of dominant tribes, from Pterostichini as principal dominants
to (in Tasmania) dominance shared by Broscini and Trechini (and
Licinini) as well as some Pterostichini. This change has been de-
scribed as it occurs among selected flightless geophile Carabidae, but
it is reinforced and made more complex by changes of winged Carabi-
dae too.

The whole transition of wet forest carabid faunas from New
Guinea to Tasmania might be described as a very irregular stepcline
of flightless groups overlain by a more regular transition (or cline
of many smaller steps) of winged groups. The flightless Carabidae
of the isolated rain forests of Cape York and the Elliot Range are
outside the main pattern and complicate it, and of course the situa-
tion as a whole is much more complex in detail than I can describe
here.

Historical Duplications : Two Land Bridges

It is a good working principle of zoogeography that situations should
be analyzed first by study of the best known and most significant
groups of animals, especially mammals, but that other groups may add
important details to what the mammals show. In the present case,
two former land bridges are involved: from New Guinea to Australia

20

Psyche

[March

and from Australia to Tasmania. Mammals show, by occurrence of
many identical or closely related species on opposite sides of the exist-
ing water gaps, that both bridges did exist recently and that some
forest-living animals crossed both of them. Carabidae show additional,
different things about the two bridges. In the case of the Australian-
Tasmanian bridge, the Carabidae agree with the mammals. Many
wet forest Carabidae including many flightless ones evidently crossed
this bridge without meeting important ecological barriers, although
cold climate stopped some other animals, especially some reptiles
(Darlington i960, p. 659). In the case of the New Guinea-Austral-
ian bridge, however, the flightless rain forest Carabidae show that
there was an ecological barrier upon the land, and that the barrier
existed for a long time. New Guinea and Australia cannot have
been connected by a continuously rain-forested ridge within the time
of existing carabid faunas. The recent connection was evidently low
and rain forest was probably not continuous across it, although it was
nearly enough continuous to allow certain forest trees, mammals,
birds, and winged insects to get across. These organisms probably
crossed the bridge by way of more or less separate forest “stepping-
stones” and strips of gallery forest that did not allow continuous
passage of flightless rain forest Carabidae, which do not disperse
easily across even narrow gaps of unsuitable land. Rain forest is
discontinuous on Cape York now. The Carabidae suggest that it has
been so for a long time in the past, and that conditions on Cape York
now are like the conditions that existed on the land bridge when New
Guinea and Australia were connected.

Historical Implications : Climatic Fluctuations
The present distribution of wet forest Carabidae shows that many
of them have been able to move up or down the eastern edge of
Australia between North and South Queensland, across what are now
wide gaps of comparatively dry country. The degree of relationships
of different Carabidae in the tropical and subtropical rain forest
systems varies. In some cases ( e . g. Pamborus of the tropicus group)
the North and South Queensland representatives of single original
stocks are only slightly differentiated, but in other cases ( e . g. Leira-
dira and its allies) they have diverged as subgenera or genera. This
suggests either several periods of dispersal and isolation, accompanying
fluctuations of rainfall and rain forest, or occasional trickling of
dominant wet forest Carabidae across the drier gaps of central Queens-
land. In either case wet forest Carabidae seem to have followed a
rather narrow path along the continental divide, and have usually

1961]

Darlington — Australian Carabid Beetles

21

not been able to reach such slightly isolated places as the rain forest
on the Elliot Range. The whole pattern, of occasional or limited
exchange between North and South Queensland and of isolation of
endemic faunules on the Elliot Range and in the Cape York rain
forests, is consistent with climates and forests fluctuating only within
moderate limits, not profoundly.

Ecological Correlations

It is a fact not sufficiently understood by some zoogeographers that
the climatic zones, the differences between tropical and cool temperate
climates, are very important to Carabidae and other insects. In
eastern Australia, where climate is the only permanent barrier to dis-
persal, many old groups of Carabidae are confined to either the tropical
(including subtropical) or the cooler south temperate areas. Evident-
ly whole tribes may persist for long periods in small areas protected
only by climatic barriers, and even dominant tribes do not always
easily cross from one climatic zone to another,

Carabid distribution is correlated with climate and ecology in sev-
eral more specific ways. For example some rain forest Carabidae,
including five genera specially considered above, seem to be more strict-
ly limited to rain forest in the tropics than in the south temperate zone.
This suggests that ecological factors are more intense in the tropics,
as they may well be if temperature and evaporation rates are involved.
That ecological factors are intense in the tropics is suggested also by
groups of Carabidae that occur in diverse habitats in the temperate
zones but enter or cross the tropics only when associated with surface
water, which probably tempers the intensity of tropical climate. I
have discussed this elsewhere (1959, especially pp. 332, 342). In
Australia, for example, the only Trechini that occur in the tropics
are winged hydrophiles: Perileptus and Trechodes by running water
and Trechobembix (which extends north to Cairns) in deep swamps.
Mecyclothorax occurs in many habitats in temperate southern Aus-
tralia, but I found only one species (apparently cordicollis SI.) in the
tropics, in thick vegetation over deep, cool water on the Atherton
Tableland. And Notagonum (“ Agonum ”) submetallicum (White),
which, though always associated with water, occurs in a variety of
waterside habitats in both humid and arid parts of south temperate
Australia, I found in the tropics (Atherton Tableland) only in thick
vegetation over cool, moving water.

There is also a notable correlation of wings and flight of Carabi-
dae with climate and altitude. Carabidae (mostly geophiles) often
become flightless at low altitudes in temperate climates, and on moun-

22

Psyche

[March

tains everywhere, but rarely at low altitudes in the tropics. The few-
ness of flightless Carabidae at low altitudes in New Guinea is an
example. I have discussed this subject, with other examples, else-
where (1943).

Finally there is a partial correlation between size of Carabidae and
climate. Very large Carabidae (over 1 to 2^/2 inches long) are numer-
ous in the forests of warm temperate to tropical eastern Australia but
relatively few or absent in both cool temperate Tasmania and wholly
tropical New Guinea. If Catadromus tenebrioid.es (ol.) is introduced,
as I think it is, the largest carabids in New Guinea are hardly an inch
long and few are that large. I suspect that this correlation has a
complex ecological basis which may include direct action of physiologi-
cal factors, correlation of size with state of wings and flight, and
competition with other insects. Of insects that might compete with
carabids, ants are most obvious. I have suggested (1943, p. 42, Fig. 4)
that ants may take the place of most flightless geophile Carabidae
especially in the lowland tropics.

Geographical History of Carabidae

Carabidae, like other old, complex groups of animals (mammals
etc.), have presumably had complex geographical histories, with suc-
cessive dominant groups evolving, spreading over the world, and
replacing older groups. The present distribution of Carabidae in
the Australian Region may reflect this. Some localized tribes that
are now confined to the cool south temperate zone may be remnants
of an ancient fauna (see Darlington i960 for further discussion of
some of these groups). Pterostichini, now dominant in most of Aus-
tralia, may be more recent and may be replacing more ancient Carabi-
dae. And Agonini may be still more recent, now dominant in New
Guinea (and in the whole tropical Asiatic- Australian area), and
spreading to Australia.

Pterostichini and Agonini tend, as dominant tribes, to be com-
plementary over the world as a whole. I have discussed this before
(1956, pp. 1-3), but what I said then is worth repeating briefly now,
with counts of species brought up to date. Both tribes are cosmopoli-
tan, but unevenly so. In some parts of the world they occur in nearly
equal numbers, in others, one tribe or the other is overwhelmingly
dominant. The tribes tend to be complementary within the Australian
Region, as already indicated. In Australia itself (with Tasmania)
Pterostichini are dominant, with more than 350 known species against
probably less than 20 species of Agonini, a ratio of nearly 40/1. But

1961]

Darlington — Australian Carabid Beetles

23

in New Guinea Agonini are dominant, with considerably more than
ioo known full species (some discovered since my 1952 paper)
against about 40 species of Pterostichini (manuscript in press), a
reversed ratio of about 3/1.

One reason for the number of Agonini in New Guinea is that
species of this tribe have multiplied on the mountains there. In
Australia, however, Pterostichini, not Agonini, have multiplied in
what seem to be comparable habitats on the mountains. This
difference can hardly be accounted for in simple ecological terms but
is probably due to a complex combination of ecological, historical, and
geographical factors. Over the world as a whole, there is a tendency
for Agonini to be better represented in the tropics; Pterostichini, in
the temperate zones. Also it is probable that Agonini, which are
phylogenetically less diverse, are more recent in origin than Pterosti-
chini and that they have dispersed more recently. It is therefore
likely that Pterostichini are dominant in Australia partly because
Australia is more temperate than tropical in climate and partly because
Pterostichini reached Australia before Agonini did, and it is likely
that Agonini are dominant in New Guinea partly because the climate
there is fully tropical and partly because the carabid fauna of New
Guinea is more recent in its origins than that of Australia, as I think
it is. Add to this that the mountain carabid faunas of Australia and
New Guinea have been derived independently, each from the lowland
fauna adjacent to it, and not by dispersal along a connecting mountain
chain, and we have an adequate and probably correct explanation of
the great difference in composition of the carabid faunas on the
mountains of Australia and New Guinea.

As to direction of recent movements of Carabidae, movements of
(winged) species have evidently occurred in both directions between
Australia and New Guinea, although I cannot take space to give
details now. Movements have apparently occurred also in both direc-
tions between the tropical and subtropical forests of Australia. This
is indicated by the relationship of the species now on the Eungella
Range (p. 13), although I am not ready to give further details now.
South of the tropics, patterns of distribution (Fig. 7) suggest
withdrawal of cool temperate groups and southward spreading of
tropical or subtropical groups. This is probably primarily an adjust-
ment to recent warming of climate rather than an invasion of south
temperate habitats by tropical Carabidae, although Pamborus , Tri-
chosternus , and Notonomus have invaded N othofagus forest on high
plateaus in New South Wales.

24

Psyche

[March

References

Ball, G. E. 1956. ... on the classification of the tribe Broscini. . . Coleop-

terists’ Bull., 10: 33-52.

Brass, L. J. 1953. Results of the Archbold Expeditions. No. 68. Summary
of the 19+8 Cape York (Australia) Expedition. Bull. American Mus. Nat.
Hist., 102: 135-206.

Brookes, A. E. 1944. \J\iaoripamborus.~\ Trans. Proc. R. Soc. New Zealand,
73 : 262.

CSIRO (Commonwealth Scientific and Industrial Research Organization)
1950. The Australian environment (2nd ed.). Melbourne. 183 pp.

Darlington, P. J., Jr. 1943. Carabidae of mountains and islands. . .
Ecological Monographs, 13: 37-61.

1952. The carabid beetles of New Guinea. Part 2.
The Agonini. Bull. Mus. Comp. Zool., 107: 87-252.

1953. Australian carabid beetles II. Some new
Pterostichini. Psyche, 60: 90-101.

1956. Australian carabid beetles III. Notes on the
Agonini. Psyche, 63 : 1-10.

1959. The Bembidion and Trechus of the Malay
Archipelago. Pacific Insects, 1: 331-345.

1960. The zoogeography of the southern cold tem-
perate zone. Proc. R. Soc. (London) (B), 152: 659-668.

1961. Australian carabid beetles IV. List of locali-
ties, 1956-1958. Psyche, 67: 111-126.

Jeannel, R. 1926-1930. Monographic des Trechinae, L’Abeille, Vol. 32,
No. 3; Vol. 33; Vol. 35; supplement, Vol. 34, No. 2.

1938. Les migadopides . . . une lignee sub-antarctic. Rev.
francaise d’Ent., 5: 1-55.

Moore, B. P. 1960. Studies on Australian Carabidae (Coleoptera) — 1.
New species of the tribes Agonicini, Trechini, and Pterostichini. Proc. R.
Ent. Soc. London (B), 29: 165-169.

Womersley, J. S., & J. B. McAdam. 1957. The forests and forest condi-
tions in the Territories of Papua and New Guinea. British Commonwealth
Forestry Conference in Australia, 1957. 62 pp.

A RECONSIDERATION OF THE GENUS EPIPOMPILUS
(HYMENOPTERA: POMPILIDAE)1

By Howard E. Evans
Museum of Comparative Zoology

The genus Epipompilus was described by Kohl in 1884, with
maximiliani Kohl, from Mexico, as type. It was next treated by
Ashmead in 1902, who at the same time described a related genus,
Aulocostetkus with bifasciatus Ashmead, from “Peru”, as type,
Haupt, in 1930, erected the tribe Epipompilini for these two genera
and several others; the others were shortly thereafter removed to
another tribe. In 1944 Bradley presented a revision of the American
species of Epipompilus and Aulocostetkus. Ashmead, Haupt, and
Bradley all separated the two genera by whether or not the eyes are
hairy. Since Ashmead said that Epipompilus has glabrous eyes, it is
clear that he was unfamiliar with the genus; and both Haupt and
Bradley admit they had never seen the genus. Thus we have the
curious phenomenon of a genus being treated by three persons, none
of whom had ever seen any specimens belonging to the genus as he
conceived it. As a matter of fact the eyes of maximiliani are hairy, and
Epipompilus as conceived by these three workers is a nonexistent
genus: in actuality the name Epipompilus is a senior synonym of
Aulocostetkus.

This is only one of several sources of confusion in the genus. Ash-
mead described Aulocostetkus by merely placing it in a key and listing
bifasciatus n. sp. as type. His description of bifasciatus can be and has
been considered valid, but he gives no information other than the
generic characters and the type locality (“Peru”), not even as to
color pattern, which is of much value in separating species in this
genus. Haupt used Ashmead’s name for a specimen from Costa Rica,
while Bradley, unable to find Ashmead’s type, followed Haupt while
expressing doubt that he had correctly identified Ashmead’s species.
However, there is a specimen in the U. S. National Museum labeled
as Aulocostetkus bifasciatus Ashmead in Ashmead’s handwriting and
marked as type of that species. But to add to the confusion this speci-
men bears the locality Bahia, Brazil, not “Peru” as it should. Now
Costa Rica (Haupt’s specimen) is actually closer to Peru than is
Bahia, Brazil, but I find it hard to reason away the identification label
in Ashmead’s handwriting. Specimens of this genus are so rare that one
is unlikely to make an error in labeling; in fact I doubt if Ashmead

Published with the aid of a grant from the Museum of Comparative
Zoology at Harvard College.

25

26

Psyche

[March

ever saw any other specimens of the genus. On the other hand, Ash-
mead was a sufficiently careless person so that it is quite believable
that he may have jotted down “Peru” when he meant “Brazil”. At
any rate, I accept this as the type of hifasciatus and have presented a
description of it below, along with a new name for Haupt’s specimen
from Costa Rica.

Still further problems remain. Was Ashmead correct in placing
maximiliani in the synonymy of Cresson’s aztecus, or was Bradley
correct in resurrecting it? If Epipojnpilus and Aulocostethus are
synonyms, what is the status of Banks’ Epicostethus, said to share some
of the characters of both genera? What is the correct generic place-
ment of Epipompilus msularis Kohl, from New Zealand? Finally,
what is the male sex of Epipompilus ?

On the following pages I have presented a brief synopsis of
Epipompilus in which answers to all of these questions are proposed.
I do not mean to imply that all problems in the genus are solved : my
synopsis is based on a mere 18 specimens of these exceedingly rare
insects. There are doubtless undiscovered species, and the males of
most of the species have yet to be found. But at least I hope that I
have supplied a sounder framework for future studies than has
previously been available.

Genus Epipompilus Kohl

Epipompilus Kohl, 1884. Verh. K. K. Zool.-Bot. Gesell. Wien, 34: 57. [Type
species: Epipompilus maximiliani Kohl, 1884 ( — aztecus Cresson 1869)
(designated by Ashmead, 1900)].

Aulocostethus Ashmead, 1902, Canad. Ent., 34: 132. [Type species: Auloco-
stethus hifasciatus Ashmead, 1902 (monobasic and original designation)].
New synonymy.

Epicostethus Banks, 1947, Bull. Mus. Comp. Zool., 99: 445. [Type species:
Epicostethus will'.amsi Banks, 1947 (monobasic)]. New synonymy.

Generic characters. — Maxillary palpi unusually elongate, antepe-
nultimate segment the longest and distinctly longer than third antennal
segment ; labial palpi with the penultimate segment broadly ovate, the
ultimate segment attached to one side of it; mandibles stout, rather
smooth, with a few setae but without a lamina on the inferior margin
which subtends a fimbriate groove, inner margin with a single strong
tooth well back from apex; labrum partially exserted, broad and
short, apical margin of clypeus broadly truncate or arcuately concave ;
flagellum with rather coarse, bristling, semi-erect pubescence, particu-
larly on the inner side of the basal segments of the female and on the
outer side of the entire flagellum of the male; head broader than
high, front with distinct small punctures; eyes densely covered with
short hairs (reduced and scarcely noticeable in some females and in

1961]

Evans — Genus Epipompilus

27

the known males) ; front rather swollen above, between the antennal
bases abruptly declivous to the much lower plane of the area frontalis
and clypeus. Pronotum short or fairly long, sides of disc rather
prominent, streptaulus absent or ill-defined; mesoscutellum and met-
anotum prominent medially, latter with distinct lateral foveae;
postnotum of variable length, front and hind margins subparallel;
propodeum with smooth contours, slope low and even, almost flat
behind ; front femora of female slightly to quite noticeably incrassate ;
front tibiae and tarsi without spines, middle and hind tibiae with or
without scattered short spines; segments of front tarsus of female
unusually short; claws slender, with a strong, subapical tooth which
is nearly parallel to the apical tooth ; ultimate tarsal segments without
spines beneath, pulvillar pads small but giving rise to some strong
setulae. Hind wing with anal lobe small, about .3-. 5 as long as sub-
median cell, anal vein extending very slightly or not at all beyond
junction of transverse median vein, latter vein leaving it at an angle,
oblique, meeting media much before origin of cubitus ; fore wing with
venation extending relatively close to outer wing margin, marginal
cell acute, removed from wing-tip much less than its own length;
three submarginal cells present, second and third receiving recurrent
veins near middle, third much wider at apex than at base. Abdomen
fusiform, in the female somewhat depressed apically, apical sternite
rather flat, even obscurely grooved medially; male with or without
conspicuous brushes of hair on sternites four and five, subgenital plate
of remarkable structure, forming a very slender, hairy process apically,
its basal plate (morphological sternite 8) unusually broad; male
genitalia with short, simple parameres, volsellae short-setose, not
expanded apically, basal hooklets double, aedoeagus small and of
simple structure.

Remarks. — Epipompilus possesses a remarkable array of unusual
structural features ; if one follows the practice of Bradley and Arnold
of splitting the Pompilinae into numerous tribes, there can be no
question that the genus deserves a tribe of its own. Personally, I am
much impressed with certain similarities with A porus and related
genera: the pronotum is similar, the front legs of the female some-
what incrassate, and the head shape and hairiness of the eyes suggestive
of certain Aporini. The male genitalia suggests Allaporus , as does the
venation of the hind wing. Any division of the Pompilinae into tribes
can be no more than tentative until such time as the classification of
the family from a world point of view is more satisfactorily worked
out. In the meantime, I prefer to place Epipo?npilus in the Aporini.

As here construed, the genus Epipompilus is strictly Neotropical

28

Psyche

[March

is distribution. However, Kohl included a New Zealand species,
insularis Kohl, in his conception of the genus, and various workers
since have listed Epipompilus from the Australian region. I have
studied two females determined by Banks as insularis and agreeing
well with Kohl’s description of this species. There can be no question
that this species is closely related to the several Neotropical species
of Epipompilus. Indeed, it agrees well with the above diagnosis except
in the following characters (the males are unknown) : maxillary palpi
not greatly lengthened, about as usual in the family; labial palpi
unmodified; eyes with only minute, scarcely noticeable hairs; trans-
verse median vein of hind wing reaching media a short distance before
origin of cubitus. The generic name Epipompiloides is here proposed
for insularis Kohl, 1884. I know of no other species assignable to
this genus, but the pompilid fauna of the Australian region is, of
course, very inadequately known. This genus is related to Epipo?npi-
lus and should be placed in the Aporini next to that genus.

Key to known species of the genus Epipompilus
Males

Antennae moderately long, crenulate in profile; claws of front tarsus
nearly alike; thorax in considerable part rufous; parameres of
genitalia with extremely long hairs, abdominal venter with hair-

tufts 9. innub us n. sp.

Antennae very short, with coarse, dark pubescence but only very
weakly crenulate in profile; outer claws of front tarsus much more
strongly curved than inner claws; thorax black; parameres and
abdominal venter with only short hairs 6. excelsus (Bradley)

Females

1. Wings wholly fuliginous; abdomen wholly rufous; pronotum

patterned with red and black (Florida and Bahamas)

I. pulcherrimus (Evans)

Wings hyaline, fore wing with two prominent brownish bands;
abdomen not wholly rufous, more or less patterned with black,

rufous, and/or whitish, pronotum all black or all rufous .... 2

2. Abdomen black, with a pattern of whitish spots 3

Abdomen in part rufous, with or without whitish spots 5

3. Body wholly black except for a pair of whitish spots on second
abdominal tergite; hind tibiae with a few short spines; prono-
tum very short, subangulate behind (Ecuador)

2. williamsi ( Banks)

Thorax largely rufous ; hind tibiae without spines 4

1961]

Evans — Genus Epipompilus

29

4. Abdomen with whitish markings on tergites 2 and 5, none on

sternites; posterior lobes of pronotum rufous (Mexico)

3. aztecus ( Cresson )

Abdomen with whitish markings on tergites 2, 3, 4, and 5, also on
sternites 2 and 3; posterior lobes of pronotum whitish (Brazil)
4. bifasciatus (Ashmead)

5. Size larger (8-1 1 mm.) ; abdomen with whitish markings on ter-

gite five, this tergite otherwise black (Brazil) 6

Size small (6-7 mm.) ; abdomen not marked with whitish on
tergite five, tergites five and six brownish-ferruginous (Central
America) 7

6. Antennae black; propodeum black except for limited yellowish

markings; hind tibiae unarmed 5* haupti (Aide)

Antennae rufous except darker apically; propodeum wholly rufo-
castaneous; hind tibiae weakly spinose . 6. excelsus (Bradley)

7. Second abdominal segment with a pair of whitish spots; hind

tibiae unarmed (Panama) 7- delicatus Turner

Second abdominal tergite without whitish spots; hind tibiae with
scattered, short spines (Costa Rica) 8. insolitus n. name

1. Epipompilus pulcherrimus (Evans) new combination
Aulocostethus pulcherrimus Evans, 1955, Ent. News, 66: 150. [Type: $,
Everglades Nat. Park, Florida, 30 December 1953 (U. S. Nat. Mus.)].

Remarks. — Since describing this species from the unique type, 1
have seen one additional specimen, from Mangrove Cay, Andros Is-
land, May-June 1917 (W. M. Mann) [Amer. Mus. Nat. Hist.].
This specimen agrees well with the type except that it is smaller
(about 5 mm. long, fore wing 4.3 mm.) and the pronotum has a
broad median streak of pale rufous as well as being rufous anteriorly
and posteriorly. This species is colored quite differently from any
other. The pronotum is much shorter than in aztecus , nearly as
short as in williamsi. The claws are the same as in aztecus and other
species of the genus, my statement to the contrary in the original
description being in error.

2. Epipompilus williamsi (Banks) new combination
Epicostethus williamsi Banks, 1947, Bull. Mus. Comp. Zool., 99: 446. [Type:
$. Banos, Oriente, Ecuador, 6000 feet, 30 Oct. (Mus. Comp. Zool.)].

Remarks. — Bank’s description is detailed and there is no reason to
repeat it here. The pronotum is short and subangulate behind, the
front femora are barely incrassate, and the hind tibiae have several
spines. The first two of these characters are shared (more or less)

30

Psyche

[March

with pulcherrimus and delicatus, the third with insolitus , excelsus, and
innubus. Thus the species is not as unique as Banks supposed, and
his generic name must be added to the synonymy of Epipompilus. I
have seen no specimens of this species other than Banks’ type.

3. Epipompilus aztecus (Cresson) new combination
Ferreola azteca Cresson, 1869, Proc. Boston Soc. Nat. Hist., 12: 376 [Type: $,
Veracruz, Mexico (Acad. Nat. Sci. Phila.)].

Epipompilus maximillian • Bradley, 1944, Trans. Amer. Ent. Soc., 70: 146
34: 57 [Type: 9, Cuernavaca, Mexico, 1871 (Bilimek) (Vienna Mus.)].
(Placed in synonymy with azteca by Ashmead, 1902). — Haupt, 1930,
Mitt. Zool. Mus. Berlin, 16: 762.

Epipompilus maximilliani Bradley, 1944, Trans. Amer. Ent. Soc., 70: 146
(Misspelling of maximiliani Kohl).

Aulocostethus aztecus Bradley, 1944, ibid., p. 142.

Remarks. — Bradley has recently provided a detailed description
of this species, drawn from Cresson’s type of azteca. He states that
this specimen “does not at all agree with Kohl’s description of maxi-
miliani\ Since I found myself unable to agree with this statement, I
asked to borrow the type of maximiliani from the Vienna Museum
for comparison. At first the type could not be located, but later Dr.
R. M. Bohart visited the museum and at my request searched for
and found it; Dr. Max Fischer then sent it to me by mail, and I took
it to Philadelphia and compared it directly with the type of azteca.
I am very much indebted to Drs. Bohart and Fischer for their assis-
tance with this problem.

The two type specimens differ considerably in size, that of aztecus
being much larger, 13.5 mm. long, fore wing 10 mm.; maximiliani
measures 7.5 mm. long, fore wing 6 mm. The front femora are slight-
ly more incrassate in aztecus (2.6 X as long as maximum width as
compared to 2.75 X in maximiliani) . The whitish maculations are
exactly the same in the two specimens, but they type of aztecus has
the posterior third of the propodeum blackish and the middle and hind
legs blackish except for the white spurs and white streaks on the hind
tibiae; in the type of maximiliani the propodeum is wholly rufous,
the middle and hind coxae rufous above, the middle and hind femora
rufous, and the tibiae partially suffused with rufous. Besides these
two specimens, I have seen one other, a female from Cuernavaca,
taken by my wife inside the window of a house on March 24, 1959.
This specimen is about the same size as the type of aztecus , from
Veracruz, and the front femora are incrassate to the same degree.
However, the propodeum is wholly rufous (as in the type of maximil-
iani, also from Cuernavaca) and the leg coloration intermediate be-
tween that of the two types (middle femora rufous, hind femora

1961]

Evans — Genus Epipompilus

3

black, middle tibiae slightly suffused with rufous but hind tibiae
black and whitish, middle and hind coxae with a small amount of
rufous above). There is no question at all in my mind that these
three specimens are conspecific.

4. Epipompilus bifasciatus (Ashmead) new combination
Aulocostethus bifasciatus Ashmead, 1902, Canad. Ent., 34: 132 [Type: $,
Bahia, Brazil, 19 March 1883 (but stated by Ashmead to be “Peru”)
(U. S. Nat. Mus., no. 58858)]. (Not Aulocostethus bifasciatus of Haupt,
1930; see no. 8. insolitus n. name).

Description of type female. — Length 11 mm., fore wing 8.7 mm.
Head black ; inner orbits narrowly pale yellow up to emargination of
eyes; antennae wholly brownish, darker apically; apical half of
clypeus and labrum pale yellowish; mandibles dull rufous; palpi light
reddish-brown. Thorax rufo-castaneous, except mesonotum with
black streaks over wing bases, and the following pale yellow : posterior
lobes of pronotum, extreme lower posterior corner of mesopleurum,
posterior rim of propodeum (interrupted medially), and apical pos-
terior parts of middle and hind coxae; legs otherwise reddish like
thorax, middle and hind tibiae weakly infuscated, spurs all whitish.
Abdomen black, spotted with pale yellow (almost white) as follows:
two large lateral spots on tergite two, two much smaller spots on
tergite three, two spots on tergite four slightly larger than those on
three, two large spots on tergite five broadly connected by a basal
band; also sternites two and three with small lateral spots. Wings
bifasciate, hyaline with a strong band over the basal and transverse
median veins and a broader band filling the marginal cell and extend-
ing to the posterior wing margin. Body and legs clothed with short,
white hair; eyes short-haired.

Clypeus broadly truncate; labrum small, exserted. First four
antennal segments in a ratio of about 13:5:11:12, segment three .55
X upper interocular line. Head 1.2 X as broad as high; middle inter-
ocular line .57 X width of head; upper interocular line .8 X lower
interocular line. Ocelli in a broad, flat triangle, front angle greater
than a right angle; postocellar line much greater than ocello-ocular
line. Pronotum of moderate length, posterior margin subangulate.
Propodeum with smooth contours except posterior slope finely trans-
versely striolate and with rather long pubescence; median line not
impressed. Posterior tibiae without spines. Fore wing with basal and
transverse median veins interstitial; radial vein somewhat angulate at
junction of second intercubital vein, marginal cell removed from wing-
tip by about .7 its own length.

Remarks. — - This is a rather typical member of the genus, in fact

32

Psyche

[March

rather close to aztecus. As mentioned in the introduction, Ashmead
provided no real description of the species, and Bradley did not see
the type and therefore followed Haupt, who had misidentified the
species. Presumably Ashmead was merely in error when he gave
“Peru” as the type locality of the species, as the type is labeled in
Ashmead’s handwriting.

5. Epipompilus haupti (Aide) new combination
Aulocostethus haupti Arle, 1936, Festschr. fur Embrik Strand, 1: 514 [Type:
9, Serra do Realengo, Rio de Janeiro, Brazil, 7 Oct. 1934]. — Bradley,
1944, Trans. Amer. Ent. Soc., 70: 144. — Banks, 1947, Bull. Mus. Comp.
Zool., 99: 445.

Remarks. — Bradley has provided a translation of the description
of this species. I have studied the specimen mentioned by Banks
(Campinas, Brazil). The legs of this specimen are more extensively
rufous than described for the type, but there is agreement in most
other details; the eyes of this species are more weakly hairy than is
usual in the genus.

6. Epipojnpilus excelsus (Bradley) new combination
Figs. 3 and 4

Aulocostethus excelsus Bradley, 1944, Trans. Amer. Ent. Soc., 70: 143 [Type:
9, Nova Teutonia, Santa Catarina, Brazil, 25 January 1939 (Mus. Comp.
Zool.)]. — Wahis, 1957, Bull, Ann. Soc. R. Ent. Belg., 9 3: 47-49 (Remarks
on color variation).

Remarks. — I have seen several additional females of this species
from the type locality, as well as a female from Rio de Janeiro,
October 1938 (R. C. Shannon) [U. S. Nat. Mus.]. The type, as
well as all the other specimens I have seen, has a pair of connected
whitish spots on the fifth tergite, Bradley’s description being in error
on this point. Wahis has discussed this matter and also pointed out
that some specimens have whitish markings on the second and sixth
tergites.

A male Epipo?npilus in the Canadian National Collections, Ottawa,
is almost certainly that of excelsus , even though it is colored very
differently from the female. Like the type female excelms , it was
taken at Nova Teutonia, Brazil, by Fritz Plaumann, in this case
on 19 June 1946. The spinose hind tibiae, as well as the locality,
suggest that this male belongs here rather than with haupti or bifas-
ciatus.

Description of male . - — Length 5.8 mm., fore wing 4.8 mm. Body
wholly shining black, with a weak bluish luster; face with a pair of
small whitish spots beside and below the antennal sockets, next to the
eyes; apical two thirds of mandibles whitish, the teeth rufous; palpi

1961]

Evans — Genus Epipo?npilus

33

light brown ; antennae dark brown ; tegulae dark brown ; front and
middle legs with the coxae black, suffused with brown apically, re-
mainder of these legs brown, middle femora with a tinge of rufous,
front tibiae yellowish-brown ; hind legs wholly black except tibia
with a sub-basal whitish spot which nearly encircles them ; fore wing
weakly tinged with brownish, especially along the basal vein and on
the apical third, setulae dark, veins and stigma brown.

Maxillary palpi with segments 3-6 in a ratio of about 15:19:15:13.
Mandibles with a single large tooth well back from apex. Clypeus
arcuately emarginate apically, exposing the small labrum. Eyes
strongly convergent below, lower interocular line about .75 X upper
interocular line; middle interocular line. .59 X width of head; head
about 1. 1 5 X as wide as high; ocelli in a broad triangle, postocellar
line 1.3 X ocello-ocular line. Front with distinct punctures which
are separated by about their own diameters. Eyes with minute hairs
except near the tops, where they are somewhat longer. First four
antennal segments in a ratio of about 15:5:8:9, segment three about
1.6 X as long as thick; flagellum short, very weakly crenulate in
profile, with coarse, dark pubescence which is especially long and
suberect on the upper and outer sides of the basal segments.

Pronotum of moderate length, its posterior margin subangulate.
Mesonotum wholly and uniformly covered with small punctures.
Postnotum nearly as long as metanotum, polished, with a median
impression and some weak basal striations. Propodeum with the slope
low and even; median line not impressed. Femora slender; middle
tibiae with a few spines, hind tibiae with many fairly strong spines
above; all tarsi weakly spinose; longer spur of hind tibiae nearly as
long as basitarsus. Claws with the inner tooth of all claws strong,
sloping so that the claws appear bifid; outer claws of front tarsus
much more strongly curved than inner claws. Fore wing with basal
vein arising a very short distance beyond transverse median vein,
basal part of basal vein distinctly arched; marginal cell large, acute
apically, radial vein distinctly angulate at its junction with the
second transverse cubital vein.

Abdomen fusiform, covered with short setae but without distinct
ventral hair-brushes. Subgenital plate (fig. 3) consisting of a long,
hairy apical process arising from a basal section which also bears some
long hairs. Genitalia (fig. 4) with the parameres weakly setose;
volsellae simple, weakly setose; basal hooklets large, double; para-
penial lobes somewhat knobbed apically, very slightly exceeding the
volsellae.

34

Psyche

[March

Fig. 1 Subgenital plate of Epipompilus innubus new species. Fig. 2 Gen-
italia of E. innubus. Fig. 3 Subgenital plate of E. excelsus (Bradley). Fig.
4 Genitalia of E. excelsus. All figures show the ventral aspect.

1961]

Evans — Genus Epipompilus

35

7. Epipompilus delicatus Turner

Epipompilus delicatus Turner, 1917. Ann. Mag. Nat. Hist., (8) 20: 359 [Type:

$, Bugaba, Panama (Champion) (British Mus.)]. — Bradley, 1944, Trans.

Amer. Ent. Soc., 70: 146.

Remarks. — I have not seen the type of this species, nor had Brad-
ley. It is a small species, comparable in size to pulcherrimus and
insolitus. It is reported to have a short pronotum and unarmed hind
tibiae, as well as a color pattern distinct from that of other species.

8. Epipompilus insolitus new name
Aulocostcthus bifasciatus Haupt, 1930. Mitt. Zool. Mus. Berlin, 16: 763. —

Bradley, 1944, Trans. Amer. Ent. Soc., 70: 145 ( Nec Ashmead, 1902,

Canad. Ent., 34: 132; misidentification) .

Type. — <j>, Turrialba, Costa Rica (Coll. H. Haupt, Halle/Saale,
Germany) .

Description (from Haupt, 1930).- — Length 7 mm. Yellowish-
brown, the following black: head, apical half of antennae, tergites 1,
3, and 4, apex of hind femora, and outer base of hind tibia. Fore wing
with two dark brown bands. Hind tibiae with a whitish area behind
the black base, tips of front and hind coxae also whitish. Whole body
with short, thick whitish hair ; eyes and wings hairy.

Wings (Haupt’s fig. 64) yellowish-hyaline, a brown transverse
band before the middle and one before the apex. Fore wing with three
cubital cells, the second somewhat pentagonal, the third trapezoidal,
the latter removed from tip by its own length. Radial cell longer
than second and third cubital cells together, also somewhat higher
than these. Pterostigma cell-like, translucent, somewhat attenuate.
Lower section of basal vein about twice as long as upper and weakly
arched; transverse median vein interstitial. Hind wing with trans-
verse median vein short, oblique, reaching media more than its own
length before origin of cubitus.

Head thick, flattened and weakly concave immediately behind the
eyes, temples barely developed. Ocelli large, forming a weakly acute
angle in front, postocellar and ocello-ocular lines equal. Front
strongly swollen, eyes thick, their inner margins subparallel, the eyes
together about equal to width of front. Antennae filiform, relatively
thick, third segment somewhat shorter than scape. Clypeus short,
trapezoidal, its entire anterior margin weakly arcuately concave. Seg-
ments of maxillary palpi very long, third segment about ten times as
long as thick (Haupt’s fig. 65). Pronotum short, hind margin obtuse-
ly angled, sides parallel, with distinct longitudinal swellings. Meso-
scutum twice as long as pronotum medially; scutellum and metanotum

3 6

Psyche

[March

elevated. Postnotum distinct, half as long as metanotum, lightly
impressed medially. Propodeum somewhat longer than broad, nar-
rowed and less steeply sloping behind, without discernible sculpturing
except indication of a median groove. Fore tarsi without a comb,
second to fourth segments short, the second as long as broad, the
following shorter. Hind tibiae with scattered, short spines. Claws
slender, with a sharp tooth before the apex, also with a distinct fan
of bristles (Haupt’s fig. 84 J) ; claw-comb with a very short plate,
its barbules sparse, surpassing the pulvillus.

Re/narks . — I have not seen this species, but since it has been
described and figured by Haupt in considerable detail, it seems de-
sirable to provide a name for it.

9. Epipompilus innubus new species
Figs. 1 and 2

Type. — cf , Cucharas, 750 m., Valley of Huallaga, Dpt. Huanuco,
Peru, June 1954 (F. Woj'tkowski) [Coll. H. K. Townes].

Description. — Length 6 mm., fore wing 5.7 mm. Head black
except as follows: inner orbits pale yellow up to middle of eyes;
clypeus, labrum, and mandibles pale yellow, almost white ; palpi very
light brown ; antennal sockets connected by a light yellow band ; first
five antennal segments yellowish-brown below, dark brown above,
rest of antenna nearly black. Thorax rufo-ferruginous except shining
blue-black as follows: propleura and extreme anterior parts of prono-
tum, mesosternum and anterior half of mesopleurum, sides of metano-
tum, all of postnotum, all of metapleurum except upper anterior
margin, all of propodeum except for sides of posterior rim, which are
pale yellow. Coxae blackish except middle and hind coxae tipped with
white; middle and hind trochanters blackish; front and middle legs
otherwise light reddish-brown, hind legs nearly black except tarsi
paler and tibiae with a white basal annulus; tibial spurs whitish
except middle and hind spurs suffused with black basally. Abdomen
shining blue-black except apical tergite ivory-white. Wings hyaline,
with dark setulae, veins and stigma brown, fore wing weakly clouded
in and about third submarginal cell.

Maxillary palpi very long, segments in a ratio of about
2:4:8:10:8:7. Mandibles rather smooth, with a few setae, inner
margin with a strong tooth well back from apex. Labrum broad and
short, truncate, exserted well beyond truncate apical margin of
clypeus, latter about twice as broad as high. Front prominent above
antennal orbits, narrow, middle interocular line .56 times width of
head; head nearly 1.2 X as wide as high; ocelli in a broad, flat tri-

1961]

Evans — Genus Epipompilus

37

angle, postocellar line twice the ocello-ocular line. Front with distinct
punctures which are separated by less than their own diameters. Eyes
with very short, barely noticeable hairs. First four antennal segments
in a ratio of about 15:5:13:12, segment three about twice as long as
thick; each flagellar segment, but more particularly the middle ones,
with a distinct swelling below and toward the base, giving the
antennae a somewhat crenulate profile below.

Pronotum very short, its posterior margin subangulate. Mesonotum
with distinct small punctures like the front ; postnotum smooth, trans-
versely striate, about half as long as metanotum. Propodeum with
even contours, median line weakly impressed, surface of declivity very
finely transversely striolate. Femora not notably swollen; middle
and hind tibiae with short spines scattered amongst the pubescence;
longer spur of hind tibia nearly as long as basitarsus; claws of front
and middle legs strongly dentate, those of hind legs obscurely dentate.
Fore wing with basal vein arising well beyond junction of transverse
median vein, basal part of basal vein strongly arched ; marginal cell
large, acute, removed from wing-tip by only about half its length;
other features of wing about as in other species of the genus.

Abdomen fusiform, covered with short setae ; sternites four and five
each with a transverse brush of longer setae, longer on the sides than
medially, the setae curved at their tips; genitalia also giving rise to
some long setae which protrude from sides of subgenital plate. Sub-
genital plate (fig. 1 ) of unusual form, consisting of a long, slender,
hairy process arising from complex basal plates (the modified ultimate
and penultimate sternites). Genitalia (fig. 2) with parameres short,
bearing some very strong setae; volsellae weakly setose, narrow in
ventral view but mesal surface wide and concave; basal hooklets
double, unusually well separated ; parapenial lobes slightly shorter
than volsellae; aedoeagus very small.

Remarks. — The spinose hind tibiae and short pronotum suggest
williamsi as the possible female of this species, and the type localities
of these two are not too far distant (Ecuador and Central Peru).
However, the difference in coloration is great, and it seems to me best
to consider the two distinct for the present.

THE REDISCOVERY AND PROBABLE
PHYLOGENETIC POSITION OF
PSILOPSOCUS (PSOCOPTERA)

By Edward L. Mockford
Illinois State Normal University
Normal, Illinois

The genus Psilopsocus has posed a puzzle to students of the Psocop-
tera since the time of its discovery. The original description by
Enderlein (1903:305) was based on a single specimen. Although
adequate for identification, this description does not permit the genus
to be placed beyond suborder in recent classifications. Enderlein
placed Psilopsocus in the Mesopsocidae, but gave no reasons for this.
Roesler (1944), apparently without re-examining the type, erected
a new family for this genus and placed it in the group Epipsocetae on
the basis of similarity of the lacinia in the Epipsocidae and Psilopso-
cidae.

It is the purpose of this paper to describe a new species of Psilopsocus
from the Philippine Islands, to add to the knowledge of the morph-
ology of the genus, and to re-interpret its phylogenetic position.

Genus Psilopsocus Enderlein, 1903.

In addition to the characters included by Enderlein in the original
description, the following characters are probably important in
delimiting this genus:

1. Male phallic sclerotizations in the form of a simple frame with
no indication of external parameres (fig. 2.).

2. Ovipositor valvulae complete, i.e. three pairs.

3. Female subgenital plate with a slender central process (fig. 1.).

4. Female paraproct with a decided elongation of the posterior
margin (fig. 3.).

5. Male paraproct with a pointed apical process on its posterior
margin (fig. 8.).

6. Labrum not of the Epipsocus type, lacking a pair of diagonal
strap-like sclerites.

The character mentioned by Enderlein of lack of junction of the

Explanation of Plate 5

Psilopsocus nebulosus n. sp. Fig. 1, $, subgenital plate. Fig.2, $, hypandri-
um and phallic frame. Fig. 3, $, left paraproct. Fig. 4, 9, sclerites of 9th
abdominal sternite (dorsal view). Fig. 5, 9, ovipositor valvulae. Fig. 6, $,
lacinial tip. Fig. 7, $, tarsal claw. Fig. 8, $, right paraproct. Fig. 9, $,
tip of pedicel (Ped.) and base of first flagellar segment (fl).

38

Psyche, 1961

Vol. 68, Plate 5

Mockford — Psilopsocus

40

Psyche

[March

bases of Radius and Media + Cubitus in the hindwing is not found
in the species described below, although a fold in the wing membrane
running from the Radius just above this junction, to the wing base
produces the appearance of two closely parallel veins.

In the species described below, the coxal organ is well developed,
with both rasp and tympanum. The rasp is composed of tiny in-
dentations with raised edges.

Psilopsocus nebulosus, new species
Plate 5 and Text Fig. i

Diagnosis: differs from the generotype, P. nigricornis End., pos-
sibly in number of teeth on lacinial tip (6 indicated for nigricornis ,
1 1 in nebulosus) , number of ctenidia on basal segment of hind tarsus
(15 in nigricornis , 16 to 19 in nebulosus) , shape of the pterostigma
(less deep in nebulosus ), and details of the forewing markings
(text figure 1), colorless areas being more numerous in nebulosus , but
there being no colorless areas bordering the wing apex in this species.

Holotype $. Measurements (see table I.).

Morphology: IO/D (distance between eyes divided by greatest
dorsal diameter of eye, method of Badonnel) = 0.78, PO (lateral
diameter of eye from above divided by greatest dorsal diameter of
eye) = 0.94. Apex of lacinia (fig. 6.) broad, with a distinct lateral
and median cusp, the lateral broadest and bearing 10 denticles.
Coeloconic sensilla of first flagellar segment two in number, both
situated at the extreme base of the segment (fig. 9.). Distinct tarsal
ctenidia present only on posterior tarsi, with a row of 17 on Ti and
one each on T2 and T3 (Ti = basal tarsal segment). Hypandrium
weakly sclerotized, its margin rounded except slightly flattened at
the apex; this flattened apex slightly more heavily sclerotized than

Text figure 1. Psilopsocus nebulosus n. sp., $, photomicrograph of right
forewing.

1961]

Mockford — Psilopsocus

4 I

the rest of the hypandrium and bearing a row of tiny cilia (fig. 2.).
Phallic frame (fig. 2.) a simple ring, slightly oblong, rather uniform
in width except somewhat thickened apically and for a short distance
along each side. Within the frame are a pair of membranous lobes.
Paraproct (fig. 8.) with the usual field of trichobothria extending
obliquely downward from its antero-dorsal angle. Posterior margin
of the paraproct extended as a lobe tipped with a pointed process.

Color (in alcohol) : compound eyes black. General body color dull
ivory marked with medium to pale brown. Labrum, antennae, ocellar
interval, and terminal two segments of maxillary palpi medium brown.
Clypeal striations, cloudy spots around compound eyes and antennal
bases, and a band bordering epicranial suture pale brown. Thorax
irregularly mottled with large pale brown areas including most of
the pleura and large areas of the tergal lobes. Brown areas of meso-
thoracic tergal lobes darkest at their borders, pale in their centers.
Legs dull ivory except medium brown on each coxa, distal end of
each femur, distal end of each tibia, and all of each tarsus. Forewing
hyaline, marked with extensive pale brown cloudy bands and spots
as in text figure i. Abdomen ringed with irregular pale brown cloudy
bands. Terminal segments medium brown.

Allotype ?. Measurements (see table I).

Morphology: IO/D = 1.56, PO = 0.72. Tarsal ctenidia present
only on posterior tarsi with a row of 18 on Ti and one each on T2
and T3. Subgenital plate (fig. 1.) with basal pigmented area in
the form of two widely diverging arms. Median process of subgenital
plate broad basally, abruptly narrowing to a slender tongue about
half-way toward its tip; the broad basal portion bearing two large
setae and the slender apical portion bearing many setae of various
sizes. Sclerites of the 9th abdominal sternite (fig. 4.) in the form

Table I. Length (in mm.) of various characters of
Psilopsocus nebulosus

Character

$

$

5

$

$

Entire body

2.87

2.82

2.61

3.03

3-67

Forewing

4.44

4.40

4.44

4.59

4.65

Hind tibia

1.39

1.43

1.43

1.57

1.43

Hind tarsus, Ti*

0.465

0.440

0.476

0.476

0.405

Hind tarsus, T2

O.071

0.071

0.059

0.071

0.059

Hind tarsus, T3

0.1 19

0.1 19

0.131

0.131

0.1 19

First posterior tarsal segment.

42

Psyche

[March

of a transverse ring with three straps radiating from it and a trans-
verse strap basal to it. Ovipositor valvulae (fig. 5.) with first valvula
slender; second valvula broad basally, terminating in a long, slender
process; third valvula a broad flap bearing many setae. Paraproct
(fig- 3-) with field of trichobothria in its antero-dorsal angle. Poste-
rior margin of paraproct markedly protruding; ventral and posterior
margins bearing many setae.

Variation: aside from variations noted in the descriptions and
measurements (table I), four male paratypes have wing and body
markings somewhat paler than the holotype male, but this may be
due to tenerality. IO/D ratios for three male paratypes are 0.69,

0.70, and 0.70; PO ratios for these are 0.88, 0.85, and 0.9 1 ; numbers
of ctenidia on posterior basal tarsal segment are 16, 19, and 18.

Nymph: one nymph taken with adults of P. nebulosus is with
little doubt this species. The association is made on the basis of
similarity in size, color, (except the distal two-thirds of the abdomen
is dark brown in the nymph), and general body shape, also on the
structure of the lacinia and tarsal claws. The lacinia is broad apically
with several indistinct denticles on the outer cusp. The tarsal claw
bears a preapical tooth and a pulvillus of medium width bent at
a decided angle near its base, as in the adult (fig. 7.).

Type locality: Philippine Islands: Mindanao, east slope of Mt.
McKinley, Davao Province, August and September, 1946, in mossy
forest, elevation 6400 feet, H. Hoogstraal collector; holotype 3,
allotype $ , 4 $ paratypes and one nymph, all in collection of Chicago
Natural History Museum.

Discussion : Psilopsocus is apparently very close to the Myopsocidae.

The following points of similarity were noted:

1. Tarsal structure

a. Number of segments same.

b. Distribution of ctenidia same.

c. Both with preapical tooth on tarsal claw.

d. Pulvillus in both of medium width with a decided bend
near its point of attachment.

2. Lacinial structure. The lacinial tip of Lichenomima sparsa has
a broad lateral cusp bearing 13 stubby denticles, and a short median
cusp, hence it shows marked similarity to the lacinial tip of Psilopsocus.

3. Male genitalia.

a. Hypandrial margin. The rounded hypandrial margin of
Psilopsocus is similar to that of several species of Rhaptoneura ,
Phlotodes, and Lichenomima.

1961]

Mockford — Psilopsocus

43

b. Phallic frame. The simple, rounded phallic frame of
Psilopsocus bearing a pair of lateral thickenings and enclosing a pair
of membranous lobes is reminiscent of this structure in Lichenomima
pauliani Bad. (Badonnel, 1955, fig. 529) and Rhaptoneura eatoni
McL. (Badonnel, 1943, fig. 143) . It differs little from this structure
in Lichenomima maxima Sm. (Smithers, 1957* fig- 6).

c. Paraproct. The paraproct of Psilopscus is similar to those of
most Myopsocids in bearing a pointed process on its posterior margin.
Although several species of Myopsocids have two such processes,
there is only one in Phlotodes angolensis Bad. (Badonnel, 1955? fig-
514). The male paraproct of the latter species resembles that of
Psilopsocus in several other respects, including shape of the field of
trichobothria and presence of a roughened area antero-dorsad of this
field.

4. Female genitalia.

a. Subgenital plate. The subgenital plate of Psilopsocus is sim-
ilar in structure to that of Rhaptoneura. The resemblance is espe-
cially marked in the case of R. africana Bad. (Badonnel, 1 955 > fig-
508). In both forms the pigmented basal area consists of a pair of
widely diverging arms; the central process is broad basally, narrowing
abruptly to a slender tongue which bears setae on or near its apex.

b. Ovipositor valvulae. Both Psilopsocus and the Myopsocids
have three pairs of ovipositor valvulae. The second valvula terminates
as a long, slender process in both groups, although this is generally
much longer in the Myopsocidae than in Psilopsocus. The third
valvula is a simple flap bearing many setae in both groups.

c. Sclerites of the ninth abdominal sternite. These sclerites in
several Lichenomina species (Badonnel, 1955, figs. 522-525; Smithers,
1957, fig- 1 1 ) are composed of three main sclerotized areas, and are
thus similar in appearance to the same group of sclerites in Psilopsocus
nebulosus with their three radiating sclerotized straps.

d. Paraproct. The female paraproct is similar in shape, cilia-
tion, and position of the field of trichobothria in the two groups.

The differences between the Myopsocidae and the Psilopsocidae
are not great. The only ones which I have found are ( 1 ) presence in
Myopsocidae of a connection between areola postica and medial stem
in the forewing and absence of this in Psilopsocidae, (2) presence in
Psilopsocidae of a spur vein from the pterostigma and its absence in
Myopsocidae, and (3) much more complex markings of the forewing
in Myopsocidae than in Psilopsocidae, with characteristic alternating
dark and light areas on veins in the former group.

44

Psyche

[March

Pearman (1936) has designated the families Myopsocidae, Psocidae,
and Thyrsophoridae as constituting the group Psocetae. To this group
should be added the Psilopsocidae. It appears that this group repre-
sents an ancient phylogenetic line within the suborder Psocomorpha,
in which the most primitive forms share a broad, multidenticulate
lacinial tip with the Group Epipsocetae. It seems likely that this type
of lacinial tip, found also in the Amphientomidae (Suborder Trocto-
morpha) was present in the earliest forms of the Suborder Psoco-
morpha.

Acknowledgements

I wish to thank the officers of the Chicago Natural History Mu-
seum, Mr. Henry Dybas in particular, for arranging the loan of
material discussed in this paper. The accompanying photomicrograph
(text figure 1) was made by Dr. Robert D. Weigel of the Depart-
ment of Biological Sciences, Illinois State Normal University.

Literature Cited

Badonnel, A. 1943. Faune de France. Psocopteres. Paris, P. Lechevalier
et Fils, 164 pp.

Badonnel, A. 1955. Psocopteres de l’Angola. Comp. Diamant. Angola
Pub. Cult. 26: 1-267.

Enderlein, G. 1903. Die Copeognathen des Indo-australischen Faunenge-
bietes. Ann. Mus. Hung. 1:179-344.

Pearman, J. V. 1936. The taxonomy of the Psocoptera: preliminary sketch.
Proc. R. Ent. Soc. Lond. (B) 5: 58-62.

Roesler, R. 1944. Die Gattungen der Copeognathen. Stett. Ent. Zeit. 105:
117-166.

Smithers, C. N. 1957. Three new species of Myopsocidae (Psocoptera)
from Natal. Proc. R. Ent. Soc. Lond. (B) 26:11-16.

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A JOURNAL OF ENTOMOLOGY

Established in 1874

Vol. 68 June-September, 1961 Nos. 2-3

CONTENTS

A New Earwig in the Genus Vostox (Dermaptera: Labiidae) from the
Southwestern United States and Mexico. W. L. Nutting and A. B.
Gurney 45

Some Comments on Walckenaer’s Names of American Spiders, Based on
Abbot’s Drawings. H.fV. Levi and L. R. Levi 53

The Neotropical Species of the Ant Genus Strumigenys Fr. Smith: Miscel-
laneous Concluding Studies. W . L. Brown , Jr. 58

A Note on the An tAAnamptogenys hartmani Wheeler. W . L. Brown, Jr. 69

Anthicus tobias Marseul, Another Tramp Species (Coleoptera: Anthi-
cidae). F. G. Werner 70

Chemical and Biological Characterization of Venom of the Ant Solenopsis
xyloni McCook. M. S. Blum , J. E. Roberts, Jr., and A. F. Novak 73

Mass Insect Control Programs: Four Case Histories. W. L. Brown, Jr 75

CAMBRIDGE ENTOMOLOGICAL CLUB

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PSYCHE

Vol. 68 June-September, 1961 Nos. 2-3

A NEW EARWIG IN THE GENUS VOSTOX
(DERMAPTERA: LABIIDAE)

FROM THE SOUTHWESTERN
UNITED STATES AND MEXICO1

By W. L. Nutting2 and Ashley B. Gurney3

During the summer of 1958 a single male earwig was taken from a
light trap in southwestern New Mexico and sent to the U. S. National
Museum for identification. Apparently a new species of Vostox, it
was put aside with the hope that more specimens might be collected.
In the fall of 1959, during a study of the Dermaptera in the Univer-
sity of Arizona collection, six adults and three nymphs of this same
earwig were discovered among some undetermined specimens. A fur-
ther search finally resulted in the completion of the series of six males,
seven females, and three nymphs upon which the following description
is based. This new earwig brings the total number of Dermaptera
in the United States, both native and adventive, to 19 species and 1
subspecies.4 Probably not more than six or seven of them are repre-
sentatives of our endemic fauna.

There are about seven previously described species of Vostox , of
which only brunneipennis (Serville) occurs in the United States; the
others are all Neotropical. V. brunneipennis ranges from Virginia.
Indiana, and Illinois south to Florida and westward to eastern Texas,
with a few records from Panama and the states of Vera Cruz and
Sinaloa in Mexico. So far as the available material demonstrates, the
new species ranges from southern New Mexico and Arizona into the
Mexican states of Sinaloa and Baja California. Neither of the species

Arizona Agricultural Experiment Station Technical Paper No. 642.

department of Entomology, University of Arizona, Tucson.

3Entomology Research Division, Agricultural Research Service, U. S. De-
partment of Agriculture, Washington, D. C.

“Several changes, which have occurred in the list of United States Dermap-
tera since the paper by Gurney (1950) appeared, may be noted. Prolabia has
been found to be a synonym of Marava, and M. wallacei (Dohrn) a synonym
of M. arachidis (Yersin), the latter current combination replacing Prolabia
arachidis (see Hincks, 1954). Pyragropsis buscki (Caudell), a recent addition
to the list, occurs in Florida (Gurney, 1959). Prolabia pulchella (Serville)
has been transferred to Laprobia, a genus described as new by Hincks (1960).

45

46

Psyche

[June-September

of V ostox found in the United States is apparently very common over
most of its range, though fairly numerous specimens of hrunneipennis
have been seen from Florida and other southeastern states, where it
occurs beneath loose bark of trees. Spongovostox apicedentatus (Cau-
dell) is similar superficially to the new species and is one of the
commonest earwigs native to the southwestern United States and
northwestern Mexico. The following keys serve to distinguish the
latter and the two United States species of V ostox, in spite of their
general similarity in habitus, size, and coloration.

Keys to Species of V ostox and Spongovostox Found in
Continental United States
(Males)

1. Forceps armed with a conspicuous subapical tooth

Spongovostox apicedentatus (Caudell)

Forceps armed with at least one conspicuous tooth at or consider-
ably anterior to middle (if tooth is absent, forceps are definitely
concave internally on basal third) 2

2. Forceps sparsely tuberculate beneath, not concave internally,

typically bearing a prominent, rounded tooth considerably an-
terior to middle (if two prominent teeth occur, the smaller,
secondary tooth is at the middle) ; pygidium as in Figs, io or
ii ; parameres with conspicuous preapical curvature, Fig. 9.

V ostox hrunneipennis (Serville)

Forceps smooth beneath, generally conspicuously concave internal-
ly on basal third, larger specimens with tooth near middle;
pygidium as in Figs. 2, 3 or 7 ; parameres less conspicuously
curved, Fig. 8. V ostox excavatus , new species

(Females)

1. Forceps armed with a basal, quadrate tooth, projecting but little

beyond dorso-internal margin; abdominal sterna moderately
clothed with fine yellow-brown setae and bearing many long,
brown setae on posterior margins (males and nymphs as well) ;
suggestions of lateral folds on segments four, five, and some-
times six (sometimes subtle but, when prominent, each fold
bearing a long, light brown seta) ; pygidium much like Fig. 5.

Spongovostox apicedentatus (Caudell)

Forceps armed with a large, basal, quadrate tooth, projecting well
beyond dorso-internal margin (Fig. 1) 2

2. Dorsal surface of anal segment with a scattering of prominent

tubercles over posterior third (Fig. 13) ; ventro-internal margin
of forceps prominent and crenulate, dorso-internal margin

1961]

Nutting and Gurney — Genus Vostox

47

broadly rounded and beset with a few widely spaced tubercles,
inner face thus scarcely concave for more than half its length ;
pygidium as in Fig. 12, but scarcely diagnostic.

Vostox bru n n eipen n is (Serville)

Dorsal surface of anal segment comparatively smooth ; dorso- and
ventro-internal margins of forceps prominent and closely set
with small tubercles (almost crenulate), inner face thus dis-
tinctly concave as a longitudinal groove nearly to tip ; pygidium
as in Fig. 4. Vostox excavatus , new species

Vostox excavatus, new species
Figures 1-8

Description. Male (holotype) : Size medium, form usual for genus;
body depressed with sides of abdomen (except for slightly narrower
segments 1 and 10) subparallel and as wide as elytra; abdomen
minutely punctulate above and below, less so on segments 1-3, in-
creasingly so posteriorly, body practically smooth elsewhere; fine,
short setae rather densely covering labrum, antennae and limbs, but
sparse on remainder of body including forceps, elytra and wing scales ;
a few longer setae on posterior margin of head, anterior margin of
pronotum, cephalic faces of femora, near bases of coxae, and on the
posterior margins of all abdominal sterna except the last.

Head cordate in dorsal outline, with greatest width through the
eyes equal to the median length ; occipital margin broadly and obtusely
emarginate ; caudal angle of genae broadly rounded ; eyes not especially
prominent, slightly shorter in length than the postocular portions of
genae ; eedysial cleavage lines very faintly impressed ; antennae broken,
one with 1 1, the other with 12 segments, the first segment equal to the
sixth in length, considerably shorter than the fourth and fifth together.

Pronotum subquadrate, with greatest width at caudal third nearly
equal to its median length, cephalic margin produced mesad to form
a narrow cervical flange, laterocephalic angles obtuse and narrowly
rounded, lateral margins straight and diverging slightly to the broadly
rounded caudal margin, anterior two-thirds of disc convex with
lateral margins flaring upward, thus forming shallow furrows which
broaden and become confluent with the flattened posterior third of
disc; median longitudinal sulcus moderately impressed on convex
portion of disc, but becoming obsolete in posterior third.

Elytra with median length 2.1 times the greatest width of a single
elytron ; lateral margins nearly straight and subparallel, humeral
angles broadly rounded, distal margin subtruncate. Exposed portions

Psyche

[June-September

of wings projecting posteriorly almost one-half the median elytral
length with external margins converging gradually to the truncated
distal extremities.

Abdomen broadened slightly in the middle, with basal segment
notably narrower than anal segment ; lateral folds moderately promi-
nent on second and third terga ; posterior margin of terga four, five,
and six bordered with small tubercles, becoming obsolete laterad ; anal
segment transversely rectangulate with sides subparallel, posterior
margin truncated, but with a small lobe laterad above dorso-internal
margin of each forceps.

Forceps, as in figure 7, about three-fourths as long as the normally
exposed portion of abdomen, relatively smooth, slender, and straight
except for gentle incurving of the apical third, a prominent, slightly
rounded tooth just anterior to middle on dorso-internal margin; inner
faces rather strongly excavate anteriorly, this becoming obsolete
proximad from tooth; ventro-internal margin bearing a few irregular-
ly spaced tubercles anterior to tooth ; pygidium, as in figure 7, with
sides parallel at base, converging acutely to the narrowly rounded
apex; subgenital plate slightly less exposed than the last tergum, its
lateral margins oblique and broadly rounded into the somewhat con-
cave distal margin; concealed genitalia as in figure 8.

Femora moderately inflated, anterior pair most strongly so, and
subequal in length to anterior tibiae; tarsi long, slender, their ventral
margins (particularly of metatarsi) bearing numerous, stiff setae;
posterior metatarsus subequal to the combined length of the remaining
two tarsal segments, the ventral surface with 2 rows of setae along
the outer (lateral) margin, inner (mesal) margin with 2 longitudinal
rows and numerous shorter marginal setae which are arranged in
about 12 to 15 short, oblique, comblike rows to give a “stepped” or
“staircase” effect. (The combs are best seen on clean specimens, in a
mesal view, with magnification of 50 or more times, in a strong light.)

Coloration: Similar to brunneipennis ; head, pronotum, median

third of wing scales and abdomen dark chestnut brown, paler on
antennae, elytra, anal segment and forceps; outer two-thirds of wing
scales yellowish-white; limbs honey yellow; eyes black.

Measurements (in millimeters) : Body length (exclusive of forceps
and pygidium), 9; median length of head, 1.5; length of prono-
tum, 1.5; median length of elytron, 2.5; internal length of exposed
wing scale, 1.1; length of forceps, 4.3.

Female (allotype) : General form as in male, but somewhat more
robust and differing as follows: head broader and longer; eyes larger

1961]

Nutting and Gurney — Genus Vostox

49

and slightly longer than cheeks; antennae broken, one with 9, the other
with 12 segments; abdomen notably wider, but with anal segment
considerably narrower than the basal segment; marginal tubercles
absent from terga four, five, and six ; forceps typically shorter, stouter,
and shaped as in figure 1, with a large quadrate tooth on dorso-internal
margin at base, both dorso- and ventro-internal margins prominent
and irregularly but closely set with small tubercles, inner faces thus
distinctly concave nearly to tips ; pygidium shaped as in figure 4.

Coloration: Differs from male in no important respect except for
being a shade darker over-all, especially on the anal segment and
forceps.

Measurements (in millimeters) : Body length (exclusive of for-
and pvgidium.), 9; median length of head, 1.5; length of pronotum,
1.5; median length of elytron, 2.5; internal length of exposed wing
scale, 1.1 ; length of forceps, 4.3.

Variation: There are five male paratypes, three of which do not
vary significantly in size from the type; the length (in mm.) of
various parts of the smallest specimen (Tucson) follow: body 7.2,
head 1.5, pronotum 1.2, elytron 2.1, wing scale 1.1, forceps 2.8. The
eyes of all but the smallest agree with the type in being shorter than
the genae, whereas in the smallest spec men they are slightly longer.
The complete number of antennal segments varies from 12 to 16.
The Tucson specimen also lacks the marginal tubercles on terga four,
five, and six. Although the forceps of all are distinctively excavated,
the large tooth is absent in the two smaller specimens( Fig. 6), and
its position marked only by a tubercle in the third. The shape of the
pygidium apparently varies considerably as in brunneipennis ; in two
specimens it is unlike the type in that it is truncated at the tip (Figs. 2
and 3 ) . All genitalia are preserved in glycerol and show close agree-
ment with those of the type in the shape of the parameres, details of
the sclerotizcd armature of the basal vesicle, and the bend of the
ejaculatory duct.

The six female paratypes show considerably less variation in size
and configuration of characters; the lengths (in mm.) of various
parts of the smallest specimen (“Vcnodio”) follow: body 8.6, head
1.6, pronotum 1.4, elytron 2.3, wing scale 1.2, forceps 2.2. The length
of the eye of two agrees with the allotype in being longer than the
cheeks, whereas in three it is shorter, and in the remaining specimen
these measurements are equal. The number of antennal segments
ranges from 13 to 16. Most of the paratypes vary but little in the
shades of brown described above; however, the two females from

1961

Vol. 68, Plate 6

Nutting and Gurney — Vostox

V

1961]

Nutting and Gurney - — Genus Vostox

51

Baja California are a dark, smoky brown over-all, nearly black on the
head, and diminishing posteriorly to a dark chestnut brown on the
forceps. The appendages and outer portions of the wing scales are a
lighter smoky brown.

Nymphs: Three nymphs, presumably collected with one of the
adult males, are included with the paratypes. These specimens are
probably more than half-grown, for they range in length from 6.5 to
7 mm., and each bears moderately developed, fused wing pads. The
antennae are 10- to 12-segmented. Each of the first six abdominal
sterna bears two long setae, which are conspicuously arranged in con-
tralateral rows, one-third of the width of the abdomen from each
margin. (This pattern may exist in the adult stage but is not evident
in any of our specimens.) The smooth forceps range from 1.8 to
2.2 mm. in length but show none of the specializations of either sex
beyond the minute tubercles along the dorso- and ventro-internal
margins. Figure 5 shows the configuration of the pygidium which
suggests that all three may be females. Their coloration is similar
to the holotype, except that the outer two-thirds of both pairs of wing
pads are dark brown and the inner third is a lighter, yellow-brown.
Holotype: U.S.N.M. No. 65696

Type locality: Santa Catalina Mts. (2000-3000 ft.), Pima Co.,
Ariz.

The holotype male was collected by Andrew A. Nichol on Au-
gust 15, 1924. In reply to a recent inquiry as to the exact locality, Dr.
Nichol has recalled that it was in the lower parts of either Sabino
Canyon (south slope of the range) or Canada del Oro (north and
west slopes), probably the former. The allotype (U.S.N.M.) was
taken under lights on the bridge over the Salt River (dry), Tempe,
Maricopa Co., Ariz., on July 18, 1947, by Floyd G. Werner.

Paratypes: U. S. National Museum (1 cT, 1$, 2 nymphs) ; Depart-
ment of Entomology, College of Agriculture, University of Arizona,

Explanation of Plate 6.

Figs. 1-8, Vostox excavatus , new species. 1. Forceps of female allotype,
dorsal view. 2. Male pygidium (Tucson), dorsal view. 3. Male pygidium
(Virden), dorsal view. 4. Pygidium of female allotype, ventral view. 5.
Nymphal pygidium, ventral view. 6. Male forceps (Tucson), dorsal view. 7.
Forceps and pygidium of male holotype, dorsal view. 8. Concealed genitalia
of male holotype: Pm, paramere; Pn, penis; BV, basal vesicle; EjD, ejacula-
tory duct. Figs. 9-14, Vostox brunneipennis (Serv.). 9. Left penis and para-
mere (Gainesville, Fla.), dorsal view. 10. Male pygidium (Dallas, Tex.),
dorsal view. 11. Male pygidium (Gainesville), dorsal view. 12. Female
pygidium (Paris, Tex.), ventral view. 13. Female forceps (Mobile, Ala.),
dorsal view. 14. Male forceps (Gainesville), dorsal view. Figs. 1-7, 10-14,
xl5; Figs. 8 and 9, x34. (Drawings by senior author).

52

Psyche

[June-September

I ucson (id", 29, i nymph); Arizona State University, Tempe
( 1 cT ) ; Museum of Comparative Zoology, Cambridge. Mass. ( 1 d1 ,
i?) ; California Academy of Sciences, San Francisco, Calif, (id);
Academy of Natural Sciences of Philadelphia (1$) ; British Museum
(N. H.) , London, England ( i?) .

1 he paratypes are from the following localities :

NEW MEXICO: Virden, Hidalgo Co., one male, Aug. 27, 1958.
light trap, G. L. Nielsen.

ARIZONA: Tempe, one male, Nov. 22, 1955, Jones; Tucson,
Pima Co., one male and three nymphs, Dec. 4, 1924, C. T. Vorhies;
one female, Nov. 7, 1939, Wayne Enloe; two females, Dec. 29,
1939, Tom Embleton ; Sabino Canyon, Sta. Catalina Mts., Pima
Co., one male, July 25, 1955, at light, G. D. Butler and F. G.
Werner; 2 mi. sw. Patagonia (4050 ft., Sonoita Creek bottom,
willow-cottonwood), Sta. Cruz Co., one male, Aug. 21, 1949,
F. H. Parker.

MEXICO: SINALOA, “Venodio”, one female, 1918, Kusche;
BAJA CALIFORNIA, 25 mi. w. La Paz (ca. 500-foot plateau,
relatively rich shrubby vegetation), one female, light trap, Aug. 30,
1959, K. W. Radford and F. G. Werner; 10 mi. sw. San Jose
del Cabo (100 yd. from ocean in sandy wash, sparse shrubs), one
female, light trap, Sept. 1, 1959, K. W. Radford and F. G. Werner.
Aside from the few notes appended to the above localities, there is
no information of any sort available on this apparently rare earwig.
Morgan Hebard (1923, and other papers) described many Orthop-
tera collected by J. A. Kusche in Sinaloa, at “Venvidio”, which
probably is our “Venodio. ” Workers have been unable to locate
either locality since, and Irving J. Cantrall, of the University of
Michigan, has written us of having prepared a manuscript dealing
with the Kusche locality. Thanks to Dr. Cantrall’s cooperation, we
are able to report that Venadillo apparently is the correct name. This
small town is 5 miles northeast of Mazatlan on Mexican Highway
15, which goes to Culiacan.

Literature Cited

Gurney, A. B.

1950. An African earwig new to the United States, and a corrected list of
the Nearctic Dermaptera. Proc. Ent. Soc. Washington 52: 200-203.

1959. New records of Orthoptera and Dermaptera from the United
States. Fla. Ent. 42: 75-80.

Hebard, M.

1922. Dermaptera and Orthoptera from the State of Sinaloa, Mexico,
Part I. Trans. Amer. Ent. Soc. 48: 157-196.

Hincks, W. D.

1954. Notes on Dermaptera, I. Proc. R. Ent. Soc. Lond. (B), 23: 159-163.

1960. Notes on Dermaptera, IV. Ibid. 29: 155-159.

SOME COMMENTS ON WALCKENAER’S NAMES OF
AMERICAN SPIDERS,

BASED ON ABBOT’S DRAWINGS1

By Herbert W. Levi and Lorna R. Levi
Museum of Comparative Zoology, Harvard University

In 1887 McCook rediscovered the Abbot drawings, basis of many
of Walckenaer’s spider descriptions, and initiated a controversy in
spider nomenclature by synonymizing spider names then in use.
Emerton replied that the drawings represent the spiders in so gen-
eral and indefinite a way that identification would only increase the
uncertainty of nomenclature. Banks’ comments about the Walck-
enaer descriptions were blunt: “They rank with ‘hearsay evidence.’
I shall not use them nor list them; I shall ignore them.” Later
Gertsch (1933) expressed the fear that these names would be re-
vived and cause permanent instability: “The problem at hand is not
the question of validity, which should be unchallenged, but one of
recognition.” In 1944 Chamberlin and Ivie made a serious attempt
to establish the Walckenaer names en masse. Their synonymies were
accepted by Archer (1946, 1950), Levi (1954), and Levi and Field
(1954), but not by Gertsch (1953). We were at first inclined to
follow Chamberlin and Ivie in using the Walckenaer names, but
during the course of the theridiid studies, had an opportunity to
examine the Abbot drawings. We are convinced that in the Theri-
diidae at least, Chamberlin and Ivie were ill-advised to attempt the
synonymies, and, indeed, that establishment of such synonymies would
be a disservice to araneology.

English-born John Abbot immigrated in 1776, as a young man, to
Screven County, Georgia, and lived in Georgia for 65 years as a
schoolmaster and naturalist. He painted birds, butterflies and other
animals, and his drawings were sold by John Francillan, a London
silversmith (Dow, 1914). Sixteen volumes of Abbot’s drawings are in
the British Museum (Natural History) in London; many volumes

'We wish to thank the following for reading a draft of this manuscript and
for making suggestions (without implying that they necessarily agree with
the conclusions): Mrs. D. L. Frizzell (Dr. Harriet Exline), Dr. R. Crabill,
Dr. C. Dondale, Prof. E. Mayr, and Dr. W. J. Gertsch. Dr. Gertsch kindly
sent us a manuscript, prepared ten or twelve years ago but never published,
in which he discussed the problem of the Walckenaer names. Although he
believed the names were correctly synonymized by Chamberlin and Ivie
(1944), he pleaded their rejection in the interests of nomenclatural stability.
Dr. Gertsch and Dr. Dondale called our attention to pertinent literature. A
National Science Foundation grant made possible our trip to Europe in 1958,
at which opportunity we examined the Abbot manuscript drawings.

53

54

Psyche

[June- September

are in other institutions, several at the Houghton Library of Harvard
University. According to Chamberlin and Ivie (1944), Walckenaer
purchased drawings from the entomologist Mackay. In the eighteen
forties Walckenaer named and described some of the drawings of
the 1792 Abbot volume (now in the British Museum), in Histoire
Naturelles des Insects Apteres. There is some doubt about the date
of publication of Walckenaer’s second volume. Our personal volume
has two inscriptions, one of Walckenaer, addressed to Mr. Adam
White and dated 4 June 1841, and another presumably in White’s
handwriting: “Adam White Villeneuve, St. Germ. Walckenaer’s
study June 7, 1841.” Thus the publication date is undoubtedly 1841,
not 1842 as stated by Chamberlin and Ivie and Bonnet (1945).

Dr. McCook visited the British Museum in 1887 and his atten-
tion was called to the Abbot drawings of American spiders. In a
report to the Philadelphia Academy of Sciences (1888a), he discussed
some of the questions raised by this discovery. McCook was much
concerned about the changing of names in use, but he also wanted
to credit the earliest author: “the laws of priority must be con-
sidered, and honesty and justice can give no room for considerations
of convenience and sentiment.” Several argiopid names of Hentz
were identified with those of Walckenaer.

McCook’s paper was reviewed by Emerton (1888). Emerton had
looked over the Abbot drawings at the time of his visit to the British
Museum in 1875, “and like Mr. McCook made hasty identifications
of such few of them as I could. ... A comparison of the numbers
shows that only five of these identifications agree with those of Mc-
Cook showing the uncertainty of off-hand identifications of these
drawings by two persons both familiar with the common spiders of
the northern states. The greater number of Abbot’s drawings repre-
sent the spiders only in the most general and indefinite way and it
seems to me improbable that any large number of them can ever be
identified.”

Included in McCook’s self defense (1888b) were excerpts from
a congratulatory letter from Thorell. Banks followed: “The de-
scriptions of new species in Walckenaer’s Insectes Apteres fall into
two classes: descriptions based on specimens, and descriptions based
on figures. The former class are undoubtedly valid and I intend to
accept them wherever I can apply them. Descriptions of figures,
however, I hold, have no claim on the naturalist. Not only are they
based on figures, but the figures have never been published. Many
of the descriptions are sufficient for identification, but most are not.

1961]

Levi and Levi — American Spiders

55

But no matter how complete, they are not descriptions of spiders;
but of figures of spiders. They rank with ‘hearsay evidence’. I shall
not use them nor list them ; I shall ignore them.”

Chamberlin and Ivie (1944) undertook “initially to determine, as
far as possible from available evidence, the proper application of the
names based by Walckenaer upon Abbot’s drawings of the spiders
of Georgia.” Chamberlin made color photographs of Abbot’s drawings
at the time of a London visit, and Ivie spent a month in April 1943
collecting spiders in Georgia. Some other collections were obtained
during brief stops in Georgia in August 1933 and June 1935. Cham-
berlin and Ivie listed the collections (including many determined
juveniles), and synonymized many well established spider names of
many families with names of Walckenaer.

However, in our own examination of the Abbot manuscript draw-
ings, we found that the majority do not show diagnostic characters;
interpretation must be subjective, and authors may differ. For in-
stance, McCook synonymized the name T etragnatha lacerta Walck-
enaer with T etragnatha caudata Emerton ; Chamberlin and Ivie
synonymized the same name with Rhomphaea fictilium (Hentz), of
a different family. Most of McCook’s synonymies concerned argiopid
spiders that have a characteristic dorsal abdominal pattern. However
a modern author has to consider the possibility of sympatric sibling
species.

Further, and to be expected, students working with groups never
revised make errors in identification. Thus Chamberlin and Ivie
synonymized Argyrodes trigonum (Hentz) with Linyphia rufa
Walckenaer. However, the specimens so labelled were not Hentz’s
species, but were Argyrodes furcatus (O.P.-Cambridge) , a species
more common in Georgia. Tidarren fordum (Keyserling) was syn-
onymized with Theridion sisyphoides Walckenaer, but specimens so
labelled were not Tidarren fordum. Female specimens of Theridion
alahamense Gertsch and Archer were misidentified as Theridion amer-
icanum Walckenaer, and the male was described as new. Walckenaer’s
description of T. americanum does not fit T. alabamense. Figure 43,
Walckenaer’s Theridion ansatum , was not recognized as the species
otherwise called Tidarren sisyphoides (Walckenaer), easily recog-
nized by the white line on the posterior part of the abdomen. Probably
half the examined theridiids of the Chamberlin and Ivie Georgia col-
lection had incorrect identifications, and the same may be true of
specimens of other families. While these errors can easily be under-
stood, they invalidate many of the synonymies of Walckenaer’s names.

56

Psyche

[June- September

Chamberlin and Ivie apparently were unaware of the earlier ex-
changes about the synonymies, for they neither referred to them nor
listed them in their bibliography. Also they apparently did not con-
sider the possibility of additional plates, letters and notes concerning
the Abbot spiders, in other libraries. Such sources of evidence might
have relevance in interpreting Abbot’s drawings, which must be con-
sidered the types for the Walckenaer names.

As recently as twenty years ago, some considered the discovery of
an “older name” a matter to* be admired, and priority was frequently
invoked to rationalize changing a name in widespread use. Today
the attitude toward stability of names has changed, partly due to
criticism of zoologists in fields other than taxonomy. Perplexed by
the constant change of names, these other biologists leveled the charge
that some taxonomists were spending more time in historical than in
biological research.

Two approaches to stability of names present themselves: Some
taxonomists think that through priority, the supply of older names
will eventually become exhausted; at the same time, strict priority
leads nomenclature back to the oldest and most uncertain names.
Other taxonomists favor established usage of the name as the basis
of stability, though usage may be hard to define.

The new (1961) Zoological code of Nomenclature combines the
criteria of priority and usage through a statute of limitations. Also,
and more important, the new code emphasizes in its Preamble the
reason for its existence — to keep names stable — a reflection of the
present needs of zoologists. The establishment of sweeping synonymies
of the Walckenaer names based on Abbot’s drawings, so inconsistent
with stability, should be questioned.

A more acceptable treatment of the Walckenaer names was demon-
strated by Bishop (1924), in his revision of the North American
Pisauridae. By restricting himself to one family he was able to treat
each nomenclatural problem individually and authoritatively, rather
than all in one arbitrary sweep. The largest number of specimens,
the greatest grasp of the literature, and the keenest understanding
of the particular spider group were brought into each judgement.

Our purpose, then, is to urge that spider students adopt the Walck-
enaer synonymies proposed by Chamberlin and Ivie only after thor-
ough study of the spider genera in question, including, in addition
to a study of the Abbot drawings, investigation of usage of names,
species problems, and distributions, giving due consideration to the
basic principles of nomenclature : to stability and universality of names.

1961]

Levi and Levi - — ■ American Spiders

57

References

Archer, A. F.

1946. The Theridiidae or comb-footed spiders of Alabama. Paper Ala-
bama Mus. Nat. Hist., no. 22: 1-67.

1950. A study of theridiid and mimetid spiders. Ibid., no. 30: 1-40.
Banks, N.

1901. Notes on some species of Walckenaer, Koch and others. Jour. New
York Ent. Soc., 9: 182-189.

Bishop, S. C.

1924. A revision of the Pisauridae of the United States. Bull. New York
State Mus., no. 252: 1-140.

Bonnet, P.

1945. Bibliographia Araneorum, Toulouse, 1 : 832.

Chamberlin, R. V. and W. Ivie.

1944. Spiders of the Georgia region of North America. Bull. Univ. Utah,
biol. ser., 8 (5): 1-267.

Dow, R. P.

1914. John Abbot of Georgia. Jour. New York Ent. Soc., 22: 65-72.-
Emerton, J. E.

1888. Walckenaer’s names of American spiders. Psyche, 5: 113-114.
Gertsch, W. J.

1933. Notes on American Lycosidae. Amer. Mus. Novitates, no. 693:1-2.

1953. The spider genera Xysticus, Coriarachne and Oxyptila in North
America. Bull. Amer. Mus. Nat. Hist., 102: 415-482.

Levi, H. W.

1954. Spiders of the genus Euryopis. Amer. Mus. Novitates, no. 1666:
1-48.

Levi, H. W. and H. M. Field.

1954. The spiders of Wisconsin. Amer. Midland Nat., 51: 440-467.
McCook, H. C.

1888a. Necessity for revising the nomenclature of American spiders. Proc.

Acad. Nat. Sci. Philadelphia, 1888: 74-79.

1888b. The value of Abbot’s manuscript drawings of American spiders.
Ibid.-. 428-431.

Walckenaer, C. A.

1841. Histoire Naturelle des Insectes Apteres, 2, Paris.

THE NEOTROPICAL SPECIES OF THE ANT GENUS
STRUMIGENYS FR. SMITH:
MISCELLANEOUS CONCLUDING STUDIES

By William L. Brown, Jr.

Department of Entomology, Cornell University

This paper is a continuation of my series on the New World fauna
of the dacetine ant genus Strumigenys Fr. Smith. Earlier parts, con-
taining keys to the abbreviations for measurements and proportions,
may be found in Jour. New York Ent. Soc. 61: 53-59, ioi-iio
(I953)- In addition to these, other parts have been published in the
same journal, in Psyche, and in Studia Entomologica, Petropolis, Bra-
zil. This section is a final one so far as currently available material
in the genus indicates ; only one more section, which will be composed
chiefly of an illustrated key to the New World members of the genus,
is planned at present.

S. micretes and S. lacacoca

Following the descriptions of what I called the emeryi group
(Brown, 1959) the species were discussed as follows:

“The four species emeryi, never marnii , micretes and lacacoca are
very close, and seem, from the limited material available, to replace
one another in a chain extending from Mexico to Panama, and per-
haps beyond. So far as I can see now, the differences are complex
enough and strong enough to indicate that each form is a distinct
species; perhaps together [they constitute] one superspecies. How-
ever, it is not beyond possibility that one or more of these forms inter-
grades with a neighbor. More material is needed.”

Since that writing, material has turned up which, though small in
amount, tends to bridge the gap between S. micretes Brown and S. la-
cacoca Brown, indicating perhaps that they belong to a single variable
species. Nevertheless, the new material poses certain problems itself,
and the discussion next offered is intended to give details that should
help in eventually straightening this complex out.

A sample consisting of parts of four nest series from Boquete, Chiri-
qui Province, Panama (F. M. Gaige leg., see below) contains 25
workers with highly variable preapical mandibular dentition, the den-
ticles varying in number from 1 to 4 in the two mandibles taken to-
gether, and also varying markedly in size, acuteness and position, so
as to bridge virtually completely the chief diagnostic character-gap
between micretes (each mandible with a small but acute preapical
tooth, and a little farther up a minute denticle) and lacacoca (man-

58

1961]

Brown — Strumigenys

59

dibles completely without teeth or denticles basad of the apical fork
in the preapical region). This sample indicates that the preapical
dentition, usually so constant in species of Strumigenys , may some-
times be unreliable. The Boquete sample would clearly indicate syn-
onymy between micretes and lacacoca were it not for one disturbing
fact: the Boquete sample differs from the available samples of both
species in its larger size and in having the promesonotum very distinct-
ly and closely longitudinally striate throughout (arched striate along
the anterior pronotal margin). This sculpture is somewhat shining,
especially on the sides, where a large section becomes smooth or nearly
so. The rest of the sides of the alitrunk are also smooth and shining
for the most part. Postpetiole predominantly smooth and shining,
crossed by a few longitudinal costulae. Propodeal teeth also longer,
more slender and more nearly horizontal than in the micretes or la-
cacoca type series. Anterior coxae smooth or nearly so, shining.

Some specimens of micretes and lacacoca have feebly indicated longi-
tudinal rugulae or costulae on the pronotum (in addition to the medi-
an carinula), but in these the predominant sculpture is the usual
opaque reticulo-punctulation over at least the discal portion. The
size, head wddth and sculptural traits of the Boquete sample could
well be diagnostic of still another species in this close-knit complex, or
they could merely mark a local population of a single variable species
that would also include the types of micretes and lacacoca. For the
present, it seems wise to avoid introducing new species names for mem-
bers of this complex and also to hold off from synonymizing micretes
and lacacoca until the distribution and variation of the complex are
better known. For the convenience of future workers, I list here the
material of the complex that I have studied, with such measurements,
proportions and other observations as I have obtained from them
(n = number of workers measured for each sample) :

Colombiana Farm, Santa Clara, Costa Rica (W. M. Mann leg.),
TL 2.9-3. i, ML 0.70-0.74, ML 0.45-0.47, WL 0.72-0.76 mm; Cl
71-74, MI 62-65 (n = 12), type series of S. micretes. Progreso,
Chiriqui Prov., Panama, (F. M. Gaige leg., no. 332), TL 3. 1-3.3,
HL 0.75-0.76, ML 0.50-0.51 mm; Cl “about as in the type series”
of micretes, MI 66-68 (n = 6), series placed with micretes in the
original description of that species. Boquete, Chiriqui Prov., Panama
(F. M. Gaige leg., nos. 208, 497, 504 and one series with no number)
TL 3.7-4.0, HL 0.86-0.90, HW 0.66-0.68, ML 0.58-0.61, WL 0.92-
O.99 mm; Cl 75-76, MI 67-68 (n = 25), Strumigenys near micretes,
discussed above. Cerro Campana, west of Chorrera, Panama Prov.,
Panama, at about 950 m altitude in montane rain forest (cloud for-

6o

Psyche

[June-September

est), in rotten wood (G. B. Fairchild and W. L. Brown leg., no.
B-86), TL 3.3, HL 0.76, HW 0.52, ML 0.53, WL 0.82 mm; Cl
68, MI 70 (n — 1), specimen here placed as S. lacacoca; this worker
has a single extremely minute denticle in the preapical concavity of
the left mandible, perhaps representing a vestige of a distal preapical
tooth. Rio Chinillo, Panama Canal Zone (T. E. Snyder leg.), TL
2. 8-3. 2, HL 0.70-0.75, ML 0.45-0.48, WL 0.72-0.75 mm; Cl 67-71,
MI 60-66 (n 7), type series of S. lacacoca.

Though the known variation of the nvcretcs-lacdcoca is consider-
ably expanded by the new material reported above, this variation is
mainly away from the direction of S. nevermanni Brown, from Costa
Rica, a species that is generally smaller, with a relatively shorter head
and mandibles and narrow infradental lamellae.

A new species of the T ococae group

Strumigenys fairchildi new species

Holotype worker: TL 4.1, HL 0.96, HW 0.72, ML 0.59, WL
1. 01, scape L 0.68 mm; Cl 75, MI 61.

Aside from its larger size and relatively longer mandibles, this
species closely resembles S. tococae Wheeler from the Amazon Basin.
The upper propodeal teeth are somewhat longer (about as long as
the distance between the centers of their bases, and about twice as
long as the lower teeth), and the lamella between the upper and lower
teeth is lower and more cariniform. The eyes are about the same size
(greatest diameter about 0.14 mm), and the petiole and postpetiole
are similar, but with slightly less voluminous spongiform appendages.
The best distinction lies in the sculpture and pilosity of the nodes and
gastric dorsum:

( 1 ) Postpetiolar disc convex, evenly reticulo-punctulate, with only
feeble anterior traces of longitudinal costulae. First gastric tergite
finely longitudinally striolate, opaque-sericeous throughout (in S.
tococae , the gaster is costulate at the base, but otherwise1 smooth and
shining) .

(2) Ground pilosity strongly reduced, virtually absent on nodes
and gastric dorsum (plentiful and conspicuous in S. tococae). Fine
erect flagelliform hairs of nodes and gaster shorter and apparently
fewer than in S. tococae.

The color is ferruginous yellow, the gaster somewhat brighter yel-
low than the rest of the body.

Holotype [MCZ] a unique worker from the wooded ravine next
to the inn near the top of Cerro Campana, Panama Prov., Panama,
17 January, i960, at an altitude of 800 to 850 m (G. B. Fairchild

1961]

Brown — Strumigenys

61

and W. L. Brown leg.). The worker was found walking on a rotten
stick lying on the leaf litter on the shaded slope of the ravine ; hasty
search of the litter and soil nearby failed to uncover more specimens.
Since the very closely related S. tococae was found at Belem, Brazil,
nesting in foliar sacs of the plant Tococa, it is possible that S. fairchildi
was also nesting in a plant cavity above the ground. Since the rela-
tionships of S. fairchildi were not recognized until it was critically
examined in the laboratory, the possibility of a plant-cavity habitat
was not investigated in the field.

Group of S. lanuginosa Wheeler

Two species, S. lanuginosa Wheeler and S. hindenburgi Forel, share
a number of traits that apparently indicate a fairly close relationship
between them : Mandibles lying close together at full closure, their
bases sharply narrowed from the outside; apical fork moderate in
length, with a single intercalary tooth; inner (masticatory) margin
concave near apex, the concavity with a short but acute preapical tooth
( S . hindenburgi has an additional minute denticle near the apical
third of the margin). Clypeus broadly triangular, with convex an-
terior margin. Antennal scape slender, tapered toward both ends,
very slightly curved at basal third.

Pronotum with humeral angles developed, bluntly tuberculate; an-
terior margin present (weak in S. lanuginosa) ; alitrunk in side view
with convex dorsal profile, broken only at the impressed metanotal
groove. Propodeal teeth of modest size, but acute, each subtended by
a low, concave infradental lamella ending in a ventral convexity.
Petiole distinctly pedunculate and with a short, dorsally rounded
node; postpetiolar disc convex; both nodes with complete and well-
developed spongiform appendages. Gaster normal in form, with a
strong anterodorsal spongiform margin and a thick anteroventral
spongiform pad ; basigastric costulae well developed.

Head, alitrunk and both nodes densely and finely reticulo-punctu-
late, opaque; postpetiolar disc usually with weak rugulosity or costu-
lation superimposed. Underside of gaster smooth and shining;
mandibles weakly shining in some lights, punctulate; legs and an-
tennae finely and densely punctulate. Sides of pronotum, anterolateral
surfaces of anterior coxae, and lower lateral surfaces of infradental
lamellae of propodeum with patches of fine reclinate hairs and fre-
quently encrusted with whitish material; apparently these represent
secretory areas.

Ground pilosity of head, promesonotum, posterior propodeun;
scapes and legs consisting of fine, abundant reclinate and arched-rec.lin-

62

Psyche

[June-September

ate hairs. Larger specialized hairs all fine, long, flagelliform, more or
less erect: a pair on each lateral occipital border (2 pairs in lanugin-
osa)I, a pair on the middle occiput, a pair on the humeri and another
pair on the mesonotum. Both nodes and both upper and lower sur-
faces of the gaster with a luxuriant growth of abundant, long, fine
flagelliform hairs, the longest of which, on the gastric dorsum, are as
long as or longer than the petiole. Each inner mandibular border
bears a row of fine oblique sensory hairs.

Males of neither species have yet been found. S. lanuginosa occurs
in Central America and southern Mexico, and has apparently been
introduced into the Bahamas, while S. hindenburgi is known from
southern Brazil and northern Argentina.

Struinigenys lanuginosa Wheeler

Strumigenys lanuginosa Wheeler, 1905, Bull. Amer. Mus. Nat. Hist. 21: 104,

fig. M, worker, female. Type loc. : Fort Charlotte, Nassau, Bahama Islands.

Syntypes in AMNH, MCZ, several examined.

Worker: TL 24-2.7, HL 0.55-0.60, ML 0.35-0.38, WL 0.58-
0.64 mm; Cl 79-80, MI 63-64. Measurements from 5 workers, in-
cluding a syntype, Panamanian and Mexican specimens.

Main distinguishing features:

( 1 ) Small size and slender build.

(2) Dorsolateral borders of head merely cariniform, not lamellate.

(3) Mandibles without a trace of a denticle basad of preapical
tooth.

(4) Basal costulae of gaster extended as fine, sericeous striation
(striolation) over the basal third or more of the first segment.
Gastric pilosity also more abundant and crowded than in S.
hindenburgi.

( 5 ) Two pairs of flagellate hairs on the lateral occipital margins.

(6) Postpetiole usually with traces of fine longitudinal striolation
or costulation overlying the punctulation.

Color light ferruginous, gaster often slightly more brownish. Man-
dibles and appendages lighter, more yellowish.

Female (dealate) : TL 2.9-3.0, HL 0.62-0.63, ML 0.36-0.38, WL
0.72-0.73 mm; Cl 80-84, MI 57-61 (from 2 syntypes). Males un-
known.

Distribution: Southern Mexico, Panama; in Bahamas, where prob-
ably introduced historically.

Localities for material examined: Bahama Islands, Nassau (W. M.
Wheeler leg.), type locality. Mexico, Veracruz: Cordoba (C. H.
Seevers leg.) , one worker under stone. Pueblo Nuevo, near Tetzonapa
(E. O. Wilson leg.), strays from leaf litter in tropical evergreen for-

1961]

Brown — Strumigenys

6 3

est, and foraging on surface of log in degraded rain forest. Panama
Canal Zone: Barro Colorado Island (J. Zetek leg.), a single worker.
This species is here reported from the American mainland for the
first time ; apparently it is widespread.

Strumigenys hindenburgi Forel

Strumigenys Hindenburgi Forel, 1915, Bull. Soc. Vaud. Sci. Nat., 50: 357,
worker. Type loc. : “Argentine,” La Plata from label on syntypes. Syntypes
in Coll. Forel, Museum d’Histoire Naturelle, Geneva, and MCZ, several
examined.

Strumigenys hindenburgi , Kempf, 1958, Stud. Ent. (n.s.) 1:555, record from
10 km sw of Agudos, S. Paulo State, Brazil, among dry leaves in wooded
gully.

Worker: TL 2. 8-3. 2, HL 0.60-0.69, ML 0.38-0.40, WL 0.63-0.71
mm; Cl 83-88, MI 58-64. Measurements from 3 syntypes and 6
additional Argentinian specimens representing 3 nest series.

Within the lanuginosa group, this species is distinguished by its
large size and relatively broad head, as well as the following charac-
ters :

(1) Dorsolateral borders of the head (dorsal scrobe borders) on
each side produced as a narrow but distinct lamellar margin that grad-
ually narrows posteriad and ends in a shallow concavity behind the
level of the eye.

(2) Mandibles each with an extra minute preapical denticle, best
seen in oblique view in dark silhouette against a bright background,
near the apical third of the shaft.

(3) Basigastric costulae extending less than half the length of the
first segment, the rest of which is smooth and shining.

(4) Lateral occipital margins each with only one flagellate hair,
arising from the concavity at the end of the lamellate dorsolateral
margin.

(5) Occiput, pronotum and postpetiolar disc with traces of feeble
reticulate rugulation superimposed on the basic densely punctulate
sculpture.

Color medium ferruginous ; legs, mandibles and antennae more
yellowish ; gaster mostly brown.

Pseudogyne or ergatoid female from Itatiaia: TL 2.7, HL 0.60,
HW 0.49, ML 0.34, WL 0.60 mm; Cl 82, MI 57. This small in-
dividual has the mesonotum developed in the direction of the full
female, with raised margins dorsad, and the differentiated scutellum
acutely projecting posteriad. The petiolar and postpetiolar nodes are
wide, as expected in females of this genus, but the gaster is not un-
usually bulky for a worker. A tiny apparent remnant of a median
ocellus occurs in the central vertex. T his specimen appears to me to

64 Psyche [June-September

be classifiable as a pseudogyne, or pathological worker-female inter-
mediate.

Distribution: Northern Argentina, extending into southeastern Bra-
zil.

Localities for material examined: Argentina: La Plata (C. Bruch
leg.), 3 syntypes. Salinas, near Tucuman (Kusnezov and Golbach
leg., no. 1677). Parque Avellanida, Tucuman (P. Wygodzinsky
leg.), a small series of workers. Brazil: Rio de Janeiro State, Itatiaia,
Lago Azul (R. Barth leg.), the single pseudogyne described above.

The Variation and Synonymy of Strumigenys louisianae
Strumigenys louisianae Roger

Strumigenys louisianae Roger, 1863, Berlin, ent. Zeitschr. 7: 211, worker. Type
loc. : “Louisiana.” Type in Zoologisches Museum der Humboldt Universita’t,
Berlin, not examined

Strumigenys unidentata Mayr, 1887, Verh. zool.-bot. Ges. Wien 37: 575 and
in key, p. 570, worker. Type loc.: “St. Catharina.” Lectotype, by present
designation, in Naturhistorisches Museum, Vienna, examined, new syn-
onymy.

Strumigenys fusca Emery, 1894, Bull. Soc. ent. ital. 26: 215, pi. 1, fig. 8, work-
er. Type loc.: Manicore, Amazonas. Holotype in Museo Civico di Storia
Naturale, Genoa, examined, new synonymy.

Strumigenys clasmospongia Brown, 1953, Psyche 60: 2, worker. Type loc.:
Petropolis, Rio de Janeiro State, Brazil. Holotype in Coll. W. W. Kempf
(ex Coll. T. Borgmeier), Sao Paulo, Brazil; paratypes in Coll. Kempf,
USNM, MCZ, several reexamined, new synonymy.

In my “Revisionary Studies” of 1953, I showed that S. louisianae
is a very variable species ranging from warm temperate North Ameri-
ca south into Bolivia and northern Argentina. However, variation
at that time was thought to involve mainly body size and proportions
of the head and mandibles. After prolonged study, a number of names
(see synoptic synonymy below) was placed in the synonymy of S.
louisianae ; types were compared in most of these cases. The variety
longicornis was also synonymized on the basis of its original descrip-
tion.

In another paper entitled, “Three new ants related to Strumigenys
louisianae Roger” (Brown, 1953a), I described S. clasmospongia , S.
producta and S. mixta , stating that “each of the three forms has been
compared with all other neotropical species known to me, both des-
cribed and undescribed, except S. fusca and S. unidentata, two species
to be placed among the species inquirendae .” Since that writing, I
have been able to study the (previously unavailable) types of S. fusca,
S. unidentata and S. unispinulosa var. longicornis on deposit in Vienna
and Genoa, through the kindness of Dr. Max Beier and Dott. Delfa
Guiglia respectively. From the first comparison, it was clear that my

1961]

Brown — Strumigenys

65

clasmospongia was only a larger, relatively narrow-headed variant of
unidentata. The unique type of S. fusca was more puzzling, being a
rather large louisianae- like worker with fairly broad head and longish
mandibles, but having the posterior half of the first gastric tergite
nearly completely smooth and definitely shining. Var. brevicornis
turned out as expected: a large louisianae female fitting fairly well
with the largest worker forms from South American samples of the
species, so that my earlier synonymy was confirmed.

In addition to these types, I have been able during the last few years
to examine an increasing accumulation of louisianae- related forms from
southeastern Brazil and northern Argentina, and scattered specimens
have even come in from central Brazil and Surinam, previously blanks
on the map. These samples are due mostly to the kindness of Father
Borgmeier and Father Kempf. Study of this new material has com-
pletely changed my ideas on relationships of the species within the
louisianae complex. First, the relatively slender, long-mandibulate
forms with shining gastric dorsum were shown to be common in
southeastern Brazil; it is these forms to which the names unidentata
and clasmospongia have been applied. Taken in combination, the dis-
tinctive <( unidentata characters” give the impression of a distinct
species inhabiting southeastern Brazil, but all attempts to define a
species with these characters have failed. The reason for this failure
is that the new material clearly shows that each of the characters go-
ing to make up the unidentata in its “typical” manifestation actually
has its own independent pattern of geographical and individual varia-
tion within louisianae. The long mandibles (MI up to 67) of the
southeastern Brazilian samples are approached by samples from Bo-
livia (MI 60-63), and some series from southeastern Brazil have
much shorter mandibles (MI as low as 56, and perhaps even lower in
some samples seen but not measured), in this respect being not far
from average for the species louisianae taken as a whole. The gastric
sculpture of the unidentata pattern, in which basigastric costulae of
moderate length are followed by a smooth or nearly smooth, shining
surface (though this surface may be obscured by secretion or other
foreign matter), is not confined to southeastern Brazilian specimens
with long mandibles; in fact, it is found in series with much shorter
mandibles from localities as far away as Tucuman, in northern Ar-
gentina; Goias, Amazonas and Surinam in the northwest and north;
and even in southern Mexico (Veracruz, Puebla). Furthermore, cer-
tain series even in southeastern Brazil contain workers with shorter
mandibles, some individuals of which have the reticulate, opaque
" louisianae sculpture”, while others from the same nest sample have

66

Psyche

[June-September

the gaster predominantly smooth and shining. Variation in the queens
is poorly known because not many of the “ unidentata” workers are
accompanied by females in the collections I have seen. In general,
loaisianae- complex queens have stronger gastric sculpture than the
workers accompanying them.

In view of the discordant nature of the variation in the only good
distinctive characters available, I am forced to consider fusca, uniden-
tata, and clasmospongia as synonyms of louisianae. It is possible that
the variation of this very plastic species is even greater in central and
northern South America, from which our samples are so few, and per-
haps even the large, very long-mandibulate producta is only another
extreme variant of louisianae. The type of fusca does show tendencies
in the direction of producta, but we shall need more material from
Western Brazil and Bolivia before we decide this question. Of course,
the possibility must not be overlooked that louisianae really is made
up of a number of cryptic species, inseparable by conventional mor-
phological study.

A fact of continuing interest is the absence of S. louisianae from the
forest on Barro Colorado Island in the Panama Canal Zone. Inten-
sive collecting by a number of mvrmecologists on the Island was re-
peated in January i960 by Dr. E. S. McCluskey and myself, making
full use of Berlese funnels and other modern collecting techniques,
but no one has yet found 5. louisianae on the Island or elsewhere in
Panama. This is especially strange in view of the fact that the species
is common in banana plantations on both the Atlantic and Pacific sides
of Costa Rica near the Panama border (El Palmar and Coto in the
Golfo Dolce, E. O. Wilson leg.). While we now have very inade-
quate ecological information, it does seem possible that S. louisianae
may be a species that has adapted to habitats marginal to the rain
forest of the South American continent, and that this has something
to do with its present wide distribution — the widest of any New-
World dacetine. In this sense, S. louisianae may fit Wilson’s (1959)
“Stage-I” category of expanding species. It is also of interest to note
that the species is much less variable (“more typical”) in the North
American extremities of its range than in the presumed evolutionary
center in South America. Furthermore, the “typical” characteristics
of short mandibles and reticulate gastric sculpture, while discordant
one with the other geographically, tend to prevail at the extremities
of the range in North America and South America as well, indicating
a centrifugal evolution and movement of these characters.

Belowr I have listed some of the available samples of S. louisianae
by geographical regions, with special emphasis on some of the more

1961]

Brown — Strumigenys

67

significant South American samples reviewed since 1953, and giving
certain measurements and proportions as known for the combined
samples from each region (n = number of workers in each combined
sample) .

Southeastern United States (8 localities in 5 states) : HL 0.52-0.61,
ML 0.27-0.23 mm; Cl 82-87, MI 52-57 (n = 15). Guatemala and
Costa Rica: HL 0.50-0.58, ML 0.27-0.33 mm; Cl 83-85 (n = 11).
Cuba and Puerto Rico: HL 0.52-0.60, ML 0.26-0.32 mm; Cl 83-87,
MI 51-56 (n = 11). Colombia (Rio Porce and Medellin, leg. N. A.
Weber) : HL 0.54-0.61, ML 0.32-0.34 mm; Cl 71-84, MI 55-56
(11 = 3). Surinam (La Poulle and Dirkshoup, leg. I. van der Drift) :
HL 0.49-0.53, ML 0.29-0.30 mm; Cl 83-86, MI 57-59 (n = 2).
Goias (Anapolis, leg. W. W. Kempf) : HL 0.47, ML 0.23 mm; Cl
85, MI 49 (n = 1). Southeastern Brazil (States of Rio de Janeiro,
Sao Paulo, Santa Catarina and Rio Grande do Sul) : HL 0.46-0.58,
ML 0.28-0.37 mm ; Cl 77-92, MI 56-67 (n ^ 22). Bolivia (Rosario,
leg. W. M. Mann) : HL 0.50-0.53, ML 0.31-0.32 mm; Cl 81-83,
MI 60-63 (n — 6). Holotype female of var. longicornis from Coroi-
co, Bolivia: TL 3.0, HL 0.66, ML 0.38, WL 0.77 mm; Cl 85, MI
58. Northern Argentina (3 localities) : HL 0.49-0.64, ML o. 26-0.35
mm; Cl 83-89, MI 52-56 (n = 6).

An additional record of interest is a sample of 5. louisianae from
Yuma, Arizona, “on cotton,” “HAGA” leg.

Descriptive Note on the Holotype of S. fusca

Holotype worker: TL 3.4, HL 0.67, ML 0.40, WL 0.71 mm;
Cl 82, MI 59. A large variant of S. louisianae ; mandibles farther
apart and relatively more slender than in North American louisianae
samples ; inner margins not so sharply concave near apex. Apical fork :
dorsal tooth about 0.12 mm long, ventral about 0.09 mm; two sub-
equal intercalary denticles. Preapical tooth about 0.05 mm long, and
its tip situated about 0.05 mm from dorsal apical tooth ; slightly curved
toward mandibular apex; larger and farther from apical fork than in
most S. louisianae. No other teeth or denticles on inner mandibular
border. Scape L 0.44, funiculus L 0.58 mm.

Promesonotum high, with convex profile, promesonotal suture (sul-
cus) visible in some lights. Metanotal groove weak. Propodeal teeth
acute, elevated, with narrow infradental lamellae approaching carini-
form; teeth about 4/5 as long as the distance between their basal cen-
ters. Petiolar peduncle longer than node; node broader than long.
Postpetiole robust, fully punctulate, opaque. Spongiform appendages
as usual for S. louisianae. Gastric costulae extending about 1/2 the

68

Psyche

[June-September

length of the basal tergite, interspersed with reticulo-punctulation ;
remaining half of tergite nearly completely smooth, distinctly shining.
Pilosity as usual in S. louisianae, but a little coarser than the average.
Color dark ferruginous, but not as dark as some specimens of the
“ unidentatci” conformation from the wet Serra do Mar, the coastal
mountain strip of Sao Paulo ; these latter samples approach black.

The fusca type is similar to producta as well as to more “typical”
louisianae , but differs in details of cephalic proportions and in sculp-
tural details. The body of the fusca type is more robust, and the hairs
of the pilosity are larger throughout. The preapical tooth is larger
and is situated a little farther from the apical fork.

Descriptive Note on the Lectotype of S. unidentata

Lectotype worker: TL 2.2, HL 0.55, ML 0.36, WL 0.54 mm; Cl
79, MI 66. Despite its slightly smaller size and wider head, there
is little doubt that this type and the species I described as clasmospon-
gia are conspecific. I have partially cleaned and reexamined the
postpetiole and gaster in the available types of unidentata and clas-
mospongia, and the postpetiole is now seen to vary from weakly to
moderately shining discad, while the gastric dorsum now appears as
smooth and shining, with reticulation nearly or quite absent, except
for the basal costulae. The frequent presence of a refractory hardened
secretion (?) on the gastric dorsum is misleading when describing
sculpture in a number of dacetine groups. The specimen now in the
Naturhistorisches Museum in Vienna has been selected as lectotype,
and is so labeled.

Synoptic Synonymy of Stru/nigenys louisianae

In the list below are found the names currently considered to be
synonyms of S. louisianae. Full page references for each synonym are
to be found in Brown, 1935b, p. 28, or, in the case of new synonymy,
with the species heading above.

S. louisianae Roger, 1863

= unidentata Mayr, 1887, n. syn.

^ unispinulosa Emery, 1890.

— unispinulosa var. longicornis Emery, 1894.

— fusca Emery, 1894, n. syn.

— louisianae var. obscuriventris Wheeler, 1908.

= bruchi Forel, 1912.

— inpdelis Santschi, 1919.

— eggersi var. cubaensis Mann, 1920.

= louisianae subsp. laticephala M. R. Smith, 1931.

— louisianae subsp. soledadensis Weber, 1934.

= louisianae subsp. guatemalensis Weber, 1934.

— louisianae subsp. costaricensis Weber, 1934.

= clasmospongia Brown, 1953, n. syn.

1961]

Brown — Strumigenys

69

References Cited

Brown, W. L., Jr.

1953a. Three new ants related to Strumigenys louisianae Roger. Psyche,
60: 1-5.

1953b. Revisionary studies in the ant tribe Dacetini. Amer. Midi. Nat.,
50: 1-137, cf. pp. 28-31.

1959. The neotropical species of the ant genus Strumigenys Fr. Smith:
group of emeryi Mann ( Hymenoptera ) . Ent. News, 70: 97-104.
Wilson, E. O.

1959. Adaptive shift and dispersal in a tropical ant fauna. Evolution,
13: 122-144.

A NOTE ON THE ANT GNAMPTOGENYS HARTMANI
WHEELER. — Workers and winged females of this ant were
received recently from Dr. M. R. Smith of the U. S. National
Museum and were determined by direct comparison with the type
in the Museum of Comparative Zoology. The sample represents a
nest taken in the soil of a banana plantation on Zapote Farm, La
Lima, Honduras, May 18, 1961 (E. Molendez leg.). This is the
first record of the species to come to light since the original description,
published in 1915, based on a single worker specimen fromi Huntsville,
Texas (C. G. Hartman leg.). The type locality in northeastern
Texas has always seemed anomalous for a genus otherwise confined
to a more strictly tropical climate and not known from any other
samples occurring natively within the continental United States, but
until the present find, the good possibility remained that G. hartmani
was an extralimital relict. It now seems more likely that the Texas
Record represents either a locality error or an adventive specimen taken
from bananas. No ecological data accompanied the original find. At
least, the occurrence of this (or other) species of Gnamptogenys in
Texas remains to be convincingly demonstrated. — W. L. Brown, Jr.,
Department of Entomology, Cornell University.

ANTHICUS TOBIAS MARSEUL,

ANOTHER TRAMP SPECIES
(COLEOPTERA: ANTHICIDAE)

By F. G. Werner
University of Arizona, Tucson

Among the rather numerous collections of Anthicidae examined
by the author during the past ten years there have been small numbers
of an unidentified species of Anthicus from a truly amazing variety of
localities. In the United States, which yielded the first examples seen,
specimens have turned up from the Atlantic to the Pacific and from
New England to Florida, including such places as Catarina, Texas and
the Sierra Ancha Mountains of Arizona. The author must admit that
he held these specimens to represent an undescribed species and had
prepared a description for publication and designated types. Then
a specimen was sent from Guam, in the Marianas, others from Oahu
identified as Anthicus mundulus Sharp, and, most recently, others
from Jamaica and Venezuela. By this time it had become apparent
that the species must have been introduced into part of the localities.
Structurally, it is quite unlike any other species in the New World,
especially in the details of the very elaborate structure surrounding the
male primary gonopore. So an Old World source was likely.

The true identity of the species was revealed in the British Museum
collection, which the author was privileged to study recently. Speci-
mens identified as Anthicus tohias Marseul from Aden, India and
Java are the same as those previously seen. The type specimen of
tohias has not been examined but the specimens at hand agree per-
fectly with the original description. Both Anthicus mundulus Sharp
and A. cervinus LaFerte, under whose names most specimens have
been misidentified, are quite different.

In the United States, at least, this is not a very abundant species.
In most cases only one or a very few specimens have been seen from
any one locality. It may be more abundant on Oahu, if one can judge
from the number of specimens that have accumulated in collections.
In order to help prevent further misidentification in any new areas
where the species may be found, a brief description and figure are
given here. The male genitalia are very distinctive. In most cases an
examination of the tip of the tegmen will suffice for the identification
of the species; none other has been seen with a similar conformation.
A quick perusal of the British Museum collection and the Pic collec-
tion in Paris failed to disclose any species that might be likely close

70

1961]

W erner — Anthicus tobias

71

relatives, at least on external features. However, the author is not
thoroughly familiar with the Old World fauna and may have
missed some species.

Anthicus tobias Marseul
(Figs. 1-3)

Anthicus Tobias Marseul, 1879, L’Abeille 17: 125. Pic, 1911, Coleop. Cat.,
pars 36: 77.

Rufous, the antennae, palpi and legs luteous; elytra with a ferru-
gineous to piceous area that extends from the basal 2/5 to 1/4 to the
apex, with the suture pale anteriorly, and with a very pale spot on
each elytron at about 1/5 from the apex. Prothorax and elytra
flattened.

Head subquadrate, but base rounded and slightly notched at the
middle. Surface smooth, with dense, rather small punctures through-
out, even on the midline. Pubescence moderately dense and short,
fine, decumbent. Eyes moderately large, prominent. Antennae slender
but slightly thickened apically. Prothorax widest at apical third,
flattened, with punctures and pubescence similar to those of head.
Elytra subparallel, flattened; omoplates slightly elevated; suture
elevated on apical half. Surface smooth, moderately densely punctured,
the punctures on the basal portion slightly larger and feebly asperate,
those behind fine and small. Pubescence golden in the pale zone,
brown over the dark markings, decumbent, fine, directed obliquely
laterally in the basal fourth, not conspicuous. Tactile setae very
short but erect. Male pygidium flattened, with truncate apex; hypopy-

.Mk

Figs. 1-3. Anthicus tobias Marseul. 1. Habitus sketch, with sculpture and
pubescence omitted. 2. Aedeagus, in ventral view as it lies in the abdomen.
3. Aedeagus, in left lateral view.

72

Psyche

[June-September

gium with a shallow, V-shaped notch. Aedeagus with a dorsal ridge
and subapical notch on the tegmen; gonopore armature very large and
complex, not inverted.

The total length of the specimens examined ranges from 2.6 to 3.1
mm. Both the pale area at the base of the elytra and the subapical
spots vary somewhat in size. The most similar-looking species in the
North American fauna is Anthicus cervinus , which has curved pubes-
cence on the elytra in most parts of its range, elytra that are not
flattened, and very different male genitalia.

Range: Except for the localities followed by a bibliographic cita-
tion, the author has seen specimens from each of the following locali-
ties: ASIA: Arabia (Pic, 1911) ; W. Aden Prot. : Lahej — XI-27-
1937 - — - Scott & Britton; Iraq (Marseul, 1897) >* Turkestan (Pic,
1 9 1 1 ) ; India (Pic, 1911) and U. P. : Fyzabad — Hingston. IN-
DIAN OCEAN: Mauritius (Pic, 1911). E. INDIES: Java: Pre-
ange-Tijembong — 4-A5 — Corporaal. PACIFIC OCEAN : Mari-
anas: Guam: Agana - — X-io-1952 — J. W. Beardsley; Hawaiian Is-
lands: Oahu: Ewa — X-1958 — It. trap; Waipio — - I-22-1946 and

VIII-1949 — It. trap — Pemberton; Manoa — IX-10-1949 — at It.

— O. H. Swezey. NORTH AMERICA: CANADA: Quebec:
Granby — VI-15-1911 — P. E. Mercier. U. S. A.: MASS.: Boston

— VIII-13-1911 — found dead on Boston Common — Fall Coll.;
Bedford — VII-15-1911 — C. A. Frost; Nahant — VI-3-1935 —
P. J. Darlington; Framingham — - VIII-12-1944 — C. A. Frost;
Dover — VIII-3-1949 — K. Christiansen. CONN.: Hamden —

VIII- 25-1939 — It. trap — N. Turner. WIS.: Wood Co.: Nekoosa

— VIII-23-1948 — It. trap — W. W. Barrett. IFL. : Chicago —

IX- 6-1909 — Fall Coll, and X-24-1921 — A. B. Wolcott; LaSalle
Co.: VIII-24-1936 — F. Werner, and Lowell — VIII-24-1948 —
It. trap — F. Werner & W. Nutting. PENN. : Delaware Co. — IX-
10-1941 — R. C. Casselberry; Williamsport — VIII-3-1949 — at It.
D. C. : Washington — VIII-15-191 1 — at It. — H. S. Barber. FLA. :
Jacksonville — VI-10 — R. L. Blickle; Broward Co.: Pineland —

X- 1956 — L. N. Bell. ALA. : Mobile — VI-23-1950 — E. O. Wil-
son. TEX.: Dimmit Co.: Catarina — VII-7-1948 — at It. — F.
Werner & W. Nutting. ARIZ. : Sierra Ancha Mts. — X-1-1925 —
D. K. Duncan. ORE.: Jackson Co.: Ashland — Black & Davis.
W. INDIES: Jamaica: Spanish Town — flying at dusk; Morant
Bay; and Gordon Town, all II- 1 937 — E. A. Chapin and R. E.
Blackwelder. S. AMERICA : Venezuela; Caracas — V-VI-1957 —
M. J. & S. Sargent.

CHEMICAL AND BIOLOGICAL CHARACTERIZATION
OL VENOM OE THE ANT SOLENOPSIS XYLONI
McCOOK

By Murray S. Blum1, J. E. Roberts, Jr.2, and A. F. Novak3

The reaction of human beings to the sting of the indigenous southern
fire ant ( Solenopsis xyloni McCook) is in marked contrast to that
produced by the sting of the closely related imported fire ant
( Solenopsis saevissima [Fr. Smith]). Whereas the sting of S. saevis-
sima is characterized by a painful edema and marked necrosis (Caro
et al. [1957]), we have found that the sting of S. xyloni seldom
results in more than a mild prurience. These facts strongly indicate
that the venoms of these two species of Solenopsis differ chemically.
The purpose of this present paper is to compare the chemical and
biological properties of these Solenopsis venoms in order to possibly
determine what is responsible for their different dermatological effects.

Materials and Methods

Venom was collected from major or media workers employing a
previously described method (Blum et al. 1958). The chemical and
biological properties of S. xyloni venom were studied by procedures
described elsewhere (Blum et al. 1958; Blum and Callahan i960).
A crystalline derivative of the main component in S. xyloni venom was
prepared from an ether extract of 450 poison glands dissected from
major workers. The derivative was isolated by the method of Blum
and Callahan (i960).

The dermatological effects of the sting of S. xyloni to human beings
were studied by observing reactions at sting sites.

Results and Discussion

The chemical properties of the venom of S. xyloni parallel those of
the venom of S. saevissima in nearly all respects. Like the venom of
S. xyloni , the venomous secretion of S. saevissima consists of an alkaline
two-phase system in which the suspended droplets represent the minor
phase (Blum et al. 1958). The main constituent in the venom of
S. xyloni is an amine which is chemically comparable to the amine
isolated from the venom of S. saevissima (Adrouny et al. 1959; Blum
and Callahan i960). The infrared spectrum of the venom of
S. saevissima is virtually superimposable on the spectrum of the venom
of S. xyloni and it is probable that the amine constituents which these

department of Entomology, Louisiana State University.

department of Entomology, Louisiana State University. Present Address:
Louisville General Hospital, Louisville, Kentucky.

department of Agricultural Chemistry and Biochemistry, Louisiana State
University.

73

1961]

Blum, Roberts , and Novak — Solenopsis

74

spectra represent are very similar. On the other hand, whereas the
venom of S. saevissima contains two rhodamine-complexing minor
components, the venom of S. xyloni contains only one.

The chemical similarities of the two venoms are paralled by their
biological properties. The venomous principles of S. xyloni exhibit
the same antimycotic and antibacterial activities as are found in the
venom of S. saevissima (Blum et al. 1958). The pronounced hemolytic
effect and insecticidal activity of S. xyloni venom compare to these
same properties in the venom of S. saevissima (Adrouny et al. 1959;
Blum et al. 1958). Thus the venoms of both of these fire ants feature
the same broad-spectrum activity against diverse types of cells.

The skin responses of human beings to the stings of these two fire
ants are similar only during the first few hours, both being charac-
terized by an immediate flare followed by a wheal. However, whereas
the sting of S. saevissima is always characterized by an umbilicated
pustule at the sting site (Caro et al. 1957), we have found that the
response to the sting of S. xyloni seldom results in more than a mild
prurience. In the few cases where minute pustules were observed,
they were on individuals who were quite sensitive to the sting of
S. saevissima. At least three explanations seem possible : ( 1 ) minor
structural modifications of the necrotoxin in the venom of S. saevissima
are associated with a large increase in necrotoxicity when compared to
its counterpart in the venom of S. xyloni (2) the concentration of
the necrotoxin in the venom of S. saevissima is greater than its counter-
part in the venom of S. xyloni, (3) the minor components contribute
to the necrotoxic action of the venom. These hypotheses remain to
be determined experimentally.

References Cited

Adrouny, G. A., V. J. Derbes, and R. C. Jung.

1959. Isolation of a hemolytic component of fire ant venom. Science
130:449.

Blum, M. S. and P. S. Callahan.

1960. Chemical and biological properties of the venom of the imported
fire ant ( Solenopsis saevissima var. richteri Forel ) and the isola-
tion of the insecticidal component. XI Int. Kongr. Ent., Vienna.
3:290-293.

Blum, M. S., J. R. Walker, P. S. Callahan, and A. F. Novak.

1958. Chemical, insecticidal and antibiotic properties of fire ant venom.
Science 128:306-307.

Caro, M. R., V. J. Derbes, and R. Jung.

1957. Skin responses to the sting of the imported fire ant ( Solenopsis
saevissima) . A. M. A. Arch. Dermotol. 75:475-488.

MASS INSECT CONTROL PROGRAMS:

FOUR CASE HISTORIES*

By William L. Brown, Jr.

Department of Entomology, Cornell University

PREFACE

Insect control is a vast subject. It encompasses many methods of
approach meant to protect a wide diversity of human resources, in-
cluding the lives and health of humans themselves. Upon the success
or failure of insect control programs have rested the fate of armies,
of great canals and populous lands. Yet, though man has registered
many practical successes against particular insect menaces, we do not
yet understand fully the underlying dynamics of insect populations
(or for that matter, of other animals, including man himself), and
until we do, perfect control will probably continue to elude us in
many cases.

However, there exist practical measures that have been used suc-
cessfully to control or eradicate many kinds of insects, even though

Figure 1. Insecticide sales by U. S. producers in recent years, projected
through to the end of 1961. Domestic consumption of insecticides actually
declined slightly during 1960 in the U. S., but exports more than made up
this dip. From Chemical Week, July 22, 1961, by permission.

*This study and the report were sponsored and supported by the Conserva-
tion Foundation, New York.

75

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we may not understand exactly how a particular measure takes its
effect. In recent years, developments in practical insect control have
come thick and fast, particularly in the field of pesticides. The de-
velopment since World War II of chlorinated hydrocarbons, carba-
mate and organic phosphate insecticides, distributed by mass aerial
spray techniques, has revolutionized control work and has raised insec-
ticide production and aerial application to the status of big businesses.
But, promising as it seemed in the immediate postwar years, simple
mass aerial broadcasting of toxic materials has not always led to efficient
control of the target pest. Furthermore, the extensive application of
this relatively unselective technique inevitably caused damage to in-
cidental targets — plants and animals or property valued by humans
— and there even arose a threat to human health itself.9* 20 As such
damage and threat of damage became more obvious, protest against
mass air-spraying increased in volume, and naturally the demand
grew for research into alternative means of control.

It is my intention now to attempt to illuminate the current status
and outlook of insect control methods in the United States by out-
lining four case histories of large-scale insect control programs. It
is difficult to say how representative these case histories may be,
considering the very diverse nature of insects and the damage each
kind does. All four of the, programs are large and expensive ones as
such operations go, all have been considered to be eradication programs
at one time or another, and all have been guided or conducted by
agencies of the United States Department of Agriculture (hereinafter
referred to as USDA).

Since these great programs affect or involve many people and many
diverse vested interests, they are all to some extent controversial.
Because controversy about them involves many contradictory findings
and interpretations, it is often difficult to gain a true and unbiased
conception of what is going on in a given instance. For this reason,
I have tried to draw my information from as large and varied a group
of sources as I could find (see Acknowledgements and References
Cited) . Let us now see if a resume of four programs — Gypsy Moth,
Fire Ant, Mediterranean Fruit Fly and Screwworm — will help us
to appreciate the problems of mass insect control.

THE GYPSY MOTH
Introduction

The Gypsy Moth, Porthetria dispar (formerly hymantria dispar ),
is a variable insect, a native of Eurasia, where it ranges from Portugal
and North Africa to Japan. The insect was imported to the Boston

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area from France in 1869 by a misguided naturalist who believed
that he could cross it with silkworms. Moths escaped from his breed-
ing colony, but it was not until 1889 that the first severe outbreak
defoliated fruit and shade trees in many towns of eastern Massachu-
setts. Control work was started by the state and apparently was
successful, for populations were so low by 1899 that control operations
were ended. The moth soon again built up extensive populations,
and control work was resumed in 1905, but it had spread by this
time to western Massachusetts and parts of Maine, New Hampshire
and Rhode Island. In 1906, Congress voted aid to the infested states
to help prevent the spread of the moth, but despite all efforts it con-
tinued to expand its range.

Biology and Nature of the Damage

The gypsy moth has a single generation per year. The winter is
passed in the egg stage, and in New England the larvae hatch in mid-
spring and feed through May and June, entering the quiescent pupal
stage in early July. The larvae feed on a wide variety of broad-leaved
trees and shrubs, especially oak, willow, poplar, birch, fruit trees and,
in heavy infestations, even hemlock and pine. Dense populations may
completely defoliate large jireas of forest, weakening many trees and
killing others outright.

The heavy-bodied female does not fly, but puts out a powerful scent
to which the strorTg-flying male responds, even to extremely minute
amounts carried on the air great distances, by flying upwind until
contacting the source individuals and copulating with them.18 The
female deposits her eggs on tree trunks, fences, rocks and other solid
objects. The young larvae spin silken threads on which they are
easily spread by the wind before they start to feed.

According to Campbell4 the strong fluctuations in abundance of
the moth are density-reactive, a most critical factor in this reactivity
being the larval behavior. At low densities, the caterpillars tend to
descend to the leaf litter to rest during the daytime, and feed mainly
at night out on the foliage. When density is intermediate, the larvae
rest during the day under loose bark on the tree trunks, a habit that
has been used to advantage in control work (bands of burlap placed
around trunks of infested trees are removed daily and the caterpillars
found beneath them are destroyed). At high densities, the larvae
remain on the foliage day and night, and are subject to heavy losses
due to disease, desiccation and attack by ichneumon-wasp parasites.
Population “crashes” are correlated with previous high densities of
larvae.

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Control Problems

Early control efforts by the State of Massachusetts and the Federal
Government included laborious and expensive methods such as hand-
creosoting of egg masses, shelter-band and tanglefoot trapping on tree
trunks, and various kinds of spray operations from the ground. For
many years, control and quarantine programs appear to have confined
the infestation to the area east of the “barrier” at the Berkshires and
Green Mountains. Occasional extralimital infestations appearing in
New Jersey, Ohio, Pennsylvania and Canada, particularly after egg
masses were spread widely by the hurricane of 1938, apparently were
eradicated before getting out of hand. Extensive introductions of
predatory and parasitic insects from Europe and Japan were made
beginning in 1905, and about ten such insects have taken hold in
North America. Much of the subsequent history of the infestation
was summarized in the report of the Gypsy Moth Eradication
Meeting11 held in Ithaca, New York, in September, 1957:

“Following World War II, DDT was found to be a specific
insecticide for the gypsy moth. At about the same time applica-
tion of insecticide by plane became a practical undertaking. It
was a new day for gypsy moth control. Heavy infestations
within the area of general spread were suppressed or brought
under control, and new infestations beyond the barrier were
detected and held in check. Pennsylvania eradicated with reason-
able effort and expenditure the gypsy moth on an area of 300,000
acres. Unfortunately more than 20 million acres were infested
in this country before a practical control was discovered.

For some unexplained reason, the gypsy moth infestations seemed
to explode* in 1950 and there was rapid spread beyond the bar-
rier zone. Following the outbreaks in 1953 and 1954, surveys
revealed the new areas of infestation west of the barrier zone
in New York, New Jersey and Pennsylvania, aggregating nearly
9 million acres. An isolated infestation found in the vicinity of
Fansing, Michigan, was immediately scheduled for eradication.
The occurrence of these infestations west and south of the barrier
posed a serious threat of spread to the hardwood forests through-
out the eastern and southern United States. The control and
quarantine programs that had successfully held the moth in check
for so long were no longer adequate. ...”

*The explosion might better be said to have fairly begun in 1951 or 1952;
see Figure 2. Its inception so soon after mass air spraying of DDT began
on an operational basis is a phenomenon which, curiously enough, seems to
have attracted little attention. It was first pointed out to me by Prof. F. M.
Carpenter of Harvard University. — W. L. B.

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ACRES
SPRAYED
BY AIR

Figure 2. Graphs to show the ups and downs of the struggle against the
gypsy moth in the U. S. Acreage showing substantial defoliation by gypsy
moth larvae each year (below) is compared with acreage sprayed from the
air (above) mostly with DDT at 1 lb per acre. Some suppression treatments
used only 1/2 or 3/4 lb of DDT per acre, and sevin has partly replaced
DDT in recent years. For details, see summaries by USDA in Appendix A,
upon which these graphs are based.

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[June-September

In spite of the difficulties involved, Federal and some state authori-
ties were still speaking in terms of “eradication” of the gypsy moth
in 1956 and 1957, while other state and local people were by this
time hesitant about backing an all-out eradication effort.

In 1957, after about three and one-half million acres had been
sprayed (two and one-half millions of them in New York State),
DDT residues were found on forage crops and in the milk of cows
that had grazed on treated areas in New York State, as well as in
eggs from poultry farms that had received spray.16 DDT tolerances
for milk are set at zero by the Federal Food and Drug Administration
and by health authorities in New York among other states.

When the DDT residues were found persisting on forage crops
and in the raw milk for periods up to a year, New York suspended
eradication efforts “. . . so that,” as the USDA’s Cooperative Plant
Pest Control Programs for 1958 put it, “the 1957 work could be
fully evaluated and any required ‘mopping up’ could be done; how-
ever, during the eradication season tests were made of several alternate
insecticides more suitable than DDT for use on pasture and forage
crops.”

Since 1958, New York has been doing a greatly reduced amount
of spraying by air, using in part the new insecticide sevin, a carbamate
having very low toxicity to mammals and birds, and one leaving no
residue in the milk. Unfortunately, sevin is not as good against the
gypsy moth as is DDT, it is highly toxic to honeybees, and it injures
plants to some extent.

Aside from the dairy-linked residue problem, DDT has received
rather good marks from most biologists checking the general ecological
effects of mass spray at one pound to the acre. A few fish, are some-
times killed, birds that catch insects on the wing depart, and certain
aquatic insects suffer, but the known damage does seem tolerable.
Long-term residual effects on soil organisms are, however, not well
known.

The chief short-range danger of mass aerial DDT campaigns lies
with the loose spray practices or accidents that result in duplication
(or worse) of spray strips in a given area. Field insect control men
often complain about the quality of pilots available for some spray
programs, and numerous incidents have occurred to illustrate the point
that some of the pilots are irresponsible or incompetent, or that they
are poorly directed. For this and other reasons, it seems certain that
operational mass spraying does not always give the same safe results
as are found for the neatly-sprayed test strips of some of the studies,
and landowners are often justified in complaining of double or triple

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doses of spray on their land. In view of these difficulties, DDT must
be considered as only a marginally safe compound even at the I lb
per acre dosage.

The issue of mass spraying has come to one court battle that at-
tracted considerable attention. A group of plaintiffs led by Dr. Robert
Cushman Murphy, the well-known ornithologist, sought injunctions
against mass spraying of DDT for gypsy moth on or near their land,
which was situated near New York City and mostly on Long Island.
Most of the plaintiffs were organic gardeners and nature-lovers, and
much of their testimony tended to be emotional in tone but rather
insubstantial as to verifiable facts. The government defended itself
with toxicologists and entomologists who presented a generally factual
picture, and the case was decided against the plaintiffs by the Federal
judge, although he warned the government to use more care in spray
operations. The main effect of the case appears to have been to make
the spray agencies hesitant about treating Long Island and many other
farm areas. Also, by agreement with New York health authorities, a
wide belt is left unsprayed around the large reservoirs of the metro-
politan water supply. Such areas can of course provide refuges for
the moth from which it is potentially able to recolonize adjacent
treated areas.

Thus, for various reasons, the large key “border state” of New York
has in fact been forced to abandon the “eradication” campaign, and
the Plant Pest Control Division of the USDA now speaks instead of
a “containment program” which would include chemical treatments
within the infested area and along its periphery to back up the con-
tinued quarantines.

Infestations in Pennsylvania and Michigan, thought on several
past occasions to have been eradicated or nearly so bv DDT spray,
still survive. Directly menaced are the hardwood forests of the
Atlantic Slope, the Appalachians and the Mississippi Valley.

What Can Be Done About the Gypsy Moth?

I gather from conversations and correspondence with entomologists
and foresters responsible for gypsy moth control at the state and local
level that they generally share an uneasiness about the use of air-
sprayed non-specific poisons such as DDT and sevin on forest and
watershed areas. Most of them expressed the hope that some substitute
control method eventually would be found. So far as we can see now,
potential substitute methods lie in four different areas: predator-
parasite manipulation, propagation of bacterial or viral diseases,

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baiting with attractants, and genetic disruption. In briefly discussing
these topics, we should not overlook the possibility that there may exist
entirely different modes of attacking the problem that have not yet
occurred to anyone.

Predators and parasites. As already mentioned, a number of
predaceous, parasitic and parasitoid insects, mainly beetles, flies and
wasplike types, have been successfully colonized in the United States
after being brought from Europe and Asia. Different ones attack
every stage of the moth, from egg through adult, but few of them are
strictly specific to the gypsy moth. The efficacy of the parasites is
now open to question, since they have obviously not prevented serious
outbreaks in areas where they are known to be established. Never-
theless, some natural enemies are known to be very effective at high
densities of the host, and their value in the absence of possibly disturb-
ing chemical control has not been thoroughly checked in recent years.
Furthermore, it is likely that the established introductions represent
only a fraction of the potentially useful arthropod enemies of the moth
existing in Eurasia or elsewhere. In theory at least, there remains the
possibility of keeping the moth at a tolerable population level by
means of natural enemies, especially if used in conjunction with other
biological control methods. Further research on natural enemies of
the moth would certainly be desirable.

Disease propagation. The gypsy moth larva is susceptible to certain
bacterial and viral diseases, among which Bacillus thuringiensis shows
enough promise to have stimulated large-scale tests by Federal and
state agencies. These tests, only partly completed, employ a “sticker”
of tung oil or one of the improved English Eovol products to fasten
the bacterial spores to the foliage. The suspension of spores in sticker
can be sprayed from the air, and presumably is not harmful to plants
or wildlife. So far, results have not been encouraging.

Attractants. The female gypsy moth, as already stated, can flutter
along the ground or over low plants, but she cannot truly fly for any
distance. The strong-flying males, like those of many moths, are
strongly activated, even over long distances, by scent released by the
female from the terminal segments or “tip” of her abdomen. Upon
sensing even minute amounts of this scent, the male responds by flying
upwind, in this way automatically approaching the scent-producing
female, and ultimately coming near enough to mate with her. The
scent obtained by extracting the female tips in benzol has been used
for years as a lure in metal or paper traps to survey suspected areas
in order to determine whether males, and therefore a likely infestation,
are present. The female tips are obtained by the laborious and
extremely expensive rearing of thousands of hand-collected female

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pupae, many of them imported from Europe and North Africa. Costs
have ranged up to a half dollar per tip in poor collecting years.

In i960, after producing several moderately effective synthetic
lures, M. Jacobson and his co-workers of the Entomology Research
Division, Agricultural Research Service, USDA, succeeded in isolating
the principal sex attractant from some half a million female gypsy
moth tips collected in Connecticut and Spain. The substance was
prepared synthetically and found to be an ester alcohol with 16 carbon
atoms in its main chain. In the course of preparing the natural lure,
a closely related substance (with 18 carbon atoms in its main chain)
was also found to act as a strong gypsy moth lure.17 This preparation,
named gypl-ure , has the advantage that it can be synthesized cheaply
and in quantity from ricinoleic acid, a common component of castor
oil. Tested in field traps, quantities of this substance as small as one
microgram proved equal in luring power to traps baited with the
natural lure. In 1961, as this is written, field trials are being carried
out to test the efficacy of gyplure-toxicant combination baits in re-
ducing moth populations. Included in this program; are “confusion”
tests with saturated levels of gyplure in granular and spray formula-
tions. Initial technical difficulties have been met, but it is hoped that
these can be cleared up during the 1962 season. It will be appreciated
that many hopes ride on these crucial trials.

Genetic methods. The success of the screwworm eradication pro-
gram (see below) has raised the possibility that the release of sterilized
males might be used to control or eradicate gypsy moth populations.
This possibility remains to be explored by further studies of the
moths’ mating behavior and physiology and the practicability of
rearing, sterilization and release procedures. Sterile male release
might be made much more effective after reduction of the population
by bait attractants or other means.

Other theoretical possibilities for control rest in the fact, discovered
years ago by R. B. Goldschmidt, that certain different native Old
World populations of P. dispar differ in their sex-determining mech-
anisms in such a way that crosses made between them produce inter-
sexes. It can be argued that the overall fitness of a population might
be cut by introducing north Japanese strains into the American
populations, which originated in France. The possibility is worth
investigation despite some theoretical difficulties.

THE IMPORTED FIRE ANT
Introduction

The fire ants belong to seven or eight New World species in the
gerninata group of genus Solenopsis. The group as a whole has a

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tropical warm temperate distribution throughout the Americas, from
southeastern and southwestern U. S. to central Argentina and Chile.
The species are quite closely related and are similar in their habits.
All form populous nests, at maturity containing 25,000 to more than
200,000 active and aggressive adult workers. The workers in a
mature nest vary considerably in size from large soldiers down to
much more numerous minor workers only 2-3 mm, long, and usually
only a single functional queen is present. Nest foundation follows
the pattern typical for ants, in which virgin winged females mate
with males during a nuptial flight, then quickly shed their wings and,
as young queens, burrow into the soil and begin the rearing of the
first brood in a small chamber. Later, as the nest grows, it usually
comes to be capped by an earthen mound sometimes two feet or more
high and often two or three feet in diameter.

Up to the First World War, only three of the fire ant species were
known to occur in the U. S., of which two, Solenopsis xyloni and S.
geminata (native fire ant) were found in the southeastern states. It
seems possible that the “native” fire ant is itself a post-Columbian
introduction, and it has been spread widely over the tropics of both
hemispheres by human commerce. In past years, S. geminata had
gathered to itself much the same reputation as a nuisance now gen-
erally assigned to the late-coming imported fire ant ( S . saevissuna)
that is the subject of this discussion. The imported fire ant arrived
at Mobile, Alabama in produce or ballast at or a few years after the
end of the First World War. At first the ant (then represented
solely, so it seems, by a blackish phase with a dull orange band at the
base of its gaster SB- the so-called “variety richteri ” common in
Argentina and Uruguay) spread only very slowly in Mobile and its
environs. At some time around the beginning of the 1930’s, a smaller,
light reddish form of saevissuna appeared in the Mobile area. This
phase corresponds to populations of the species common in southern
Brazil and Paraguay, and it seems most likely that its appearance
marks a second introduction of saevissuna into the Mobile Bay port
area.

Coincident with the advent of the red phase, the entire saevissuna
salient in southern Alabama entered upon a period of rapid expansion
that carried the main infestation across state lines by 1940. The
expansion apparently has not yet reached its full extent, although
infestations are or have been known to occur in ten states ranging
from Texas and Arkansas to North Carolina and Florida. Expansion
occurs in two main ways — by steady widening of the main infested
areas due to short-range aerial spread of winged females, and through

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colonization ahead of the main infested area by queens and colony
fragments transported by vehicular traffic. Nursery stock used to be
a prime source of new infestations, but since nursery treatments and
quarantine regulations have come into effect, fertilized females acci-
dently carried in automobiles are probably responsible for most
colonization.

Wherever the red phase has expanded to overcome the dark phase,
the two extreme forms have interbred to produce a series of inter-
mediates, and in most cases the red form soon comes to predominate
by a process of genetic swamping coupled with its greater success in
warfare between nests. In fact, it may not be too extravagant a
speculation to conclude that it was the injection of the red-form genes
into the existing dark population that sparked the spectacular spread
of the species in the last three decades. At present, the North Ameri-
can population consists mainly of light reddish ants, the dark phase
surviving mainly in peripheral situations and cool swamplands.

Wherever it spreads, S. saevissima tends to replace the populations
of S. xyloni and S. geminata in its path, though this is less true of
the dark-colored geminata occupying woodlands in Florida and per-
haps elsewhere26; saevissima in the U.S. generally avoids shaded situa-
tions. The imported fire ant is able to build up remarkably dense
populations. I have seen pastures in eastern Mississippi in which it
was literally possible to walk for a considerable distance by stepping
from mound to mound without touching a foot to the ground between.
Such situations are exceptional, and usually mark the entry of the
species into a new area, or else follow control measures that have
knocked out a stable population of old, large nests. When the old
nests are eliminated, large numbers (up to 185 per acre) of smaller
new ones take their places, but as they grow, nests are gradually
eliminated until the density is again relatively low (10-50 nests per
acre usually).

Studies made to date have not been critical enough to detect possible
widespread population fluctuations in untreated areas, but about a
century ago, Bates noted a radical change in a native population of
S. saevissima in the Amazon Basin.

A small number of parasites of this ant are known in its native
habitat, including several known or suspected inquilinous species of
ants and a phorid fly, but no real study has ever been made of this
phase of the ant’s biology. These parasites have been lightly dismissed
as a control possibility by previous writers, but it seems to me that the
whole subject of parasitism should be looked into. Parasites might do

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much better in the U. S. than in their native range, and even a minor
reduction in fire ant populations might reduce it appreciably as a
nuisance in some areas.

Nature and Extent of Damage

The kind and extent of the damage done by fire ants has been the
subject of much dispute. Generally, control agencies, and especially
the USDA-affiliated ones, have emphasized the deleterious effects
produced by the1 ant, while some zealous anti-insecticide writers have
written it off as doing negligible harm. Both groups admit that the
ant mounds do interfere with the harvesting of forage crops. Harvest-
ing machinery is often damaged by striking the hard mounds, and
field hands are stung by the ants — in some cases so badly that they
refuse to work infested fields. Occasionally, land values have fallen
somewhat in badly infested areas. The health threat must also be
considered in cities and towns, where the ants may infest lawns and
gardens and even sometimes enter houses. Small children and unusu-
ally sensitive adults have occasionally suffered grave illness, or in two
or three cases may even have died as a result of fire ant stings. Numer-
ous stings result in a rash-like group of pustules that can be very an-
noying for several days or more. Still, the fire ant as a health menace
must be ranked far below ordinary bees and wasps, which are respon-
sible for many times the deaths that fire ants cause during a given
period of years, in the same states. It is difficult to see how the ant
can be classed as a serious public health problem despite scare stories
in the press, television and in a USDA-sponsored film. Professor F. S.
Arant, head of the entomological contingent at Auburn University,
current president of the Entomological Society of America, and a
top authority on the fire ant, agreeing with Dr. J. L. George10 and
other state entomologists in the Southeast, calls the fire ant a “major
nuisance,” but deprecates its role as a crop pest. Studies made at
Auburn14 and elewhere in the South generally have borne out this
evaluation. It is interesting to note that the studies6’27 that have
found more or less serious damage done to crop plants were made
before 1953. These studies were mainly concentrated in south-central
Alabama, near the Mobile Bay center of fire ant spread, and were
based on personal investigation as well as uninvestigated farmer
reports. That some crop damage was done in this area in the late
’forties and early ’fifties is incontestable, but even then, the damage
does not seem to have been insupportable. That more recent studies
have failed to find serious crop damage is probably to be laid to a
gradual change in the habits of the ants or their population density,

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or both. Whatever is the case, it does seem that the damage currently
being done by the imported fire ant in the untreated sections infested
in this country is less than would seem to justify the massive campaign
that has been mounted against it. Agencies in all but two infested
states do not even grant the fire ant a place in their lists of the more
important plant pests. The USDA cites farmer support for the
program, and this support certainly exists at least in some sections.
But the enthusiasm of farmers for the spray programs is too often
based merely on a vague feeling that insecticides in general are a good
thing. When, as in large areas covered by the present program, the
farmers individually get the spray free, they tend to overlook possible
bad effects it may bring with the benefits. In any case, the satisfaction
of farmers is certainly no substitute for a careful and extensive
professional check of current fire ant damage. No such check has
been made by the USDA, or at least none has been reported upon
since 1952.

Control Operations

Control efforts directed against the imported fire ant were first
initiated on a small scale by the State of Mississippi in 1948, without
notable success. A survey of the infested area was begun by the USDA
in the fall of 1948, and, together with limited investigation of the
biology of the ant and control measures against it,6 ran until research
funds were stopped in 1953. This investigation did not deal with
aerial control measures, and little attention was paid to wildlife
damage. It is important to note that from 1953 until 1958, after
the USDA had started its mass spray program, it spent no money
for fire ant research.22 Meanwhile, several independent agencies had
done part-time research on various aspects of fire ant biology and
control, including medical studies of the effects of the venom on
humans at Tulane University, biological and control studies at
Auburn and Mississippi State Universities, and behavioral and other
investigations by Dr. E. O. Wilson and others (including the present
author) at Harvard University and in the field. The Fish and
Wildlife Service, although greatly hampered by lack of research
funds for this purpose, was giving some attention to the prospect of
mass broadcasting of insecticides as it could be expected to affect
wildlife.

Against this patchy research background, in March, 1957, the
USDA noted that it had requested the approval of Congress for
control of the fire ant, and Congress forthwith passed a special “Fed-
eral Plant Pest Act,” authorizing the USDA to take measures against

88

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[June-September

the ant. For the 12 months beginning July, 1957, 2.4 million dollars
was appropriated, to be matched by funds from state agencies, local
sources and/or individual farmers. (In practice, actual matching
appears to have been spotty at best, and the government has waived
farmer contributions in Georgia and parts of Florida since early in
the program.)

On April 18, 1957, after a brief correspondence with officers in the
Entomology Research Division of the Agricultural Research Service,
USDA, I received a letter from Dr. A. W. Lindquist, head of one
of the sections in the Division, which started in part as follows:22

“The idea of airplane spraying and dusting for control probably
stems from the fact that extensive areas are infested. This method of
application would of course be fine if it were effective. However, we
would want to see considerable research conducted to determine if it
would be effective and, if so, to determine what insecticides and special
precautions would be necessary for maximum results. As far as we
know, no research along these lines has been conducted.”

This answer may be compared with that received from Dr. M. R.
Clarkson,23 Acting Administator of the Agricultural Research Serv-
ice, dated January 3, 1958, stating in part:

“In planning field operations, all available results of applicable
research and practical experience are taken into account. Close liaison
has been established with the Fish and Wildlife Service of the Depart-
ment of the Interior and the states involved. Competent wildlife
observers have been assigned to the work and experience to date
indicates that a successful program can be carried out without serious
consequence to wildlife resources. . . . Both the Agricultural Research
Service and State Experiment Stations have expanded their research
program in a continuing effort to improve operational procedures.”
( Italics mine — W.L.B. )

In May, 1957, as a matter of record, Dr. Ross Leffler of the
Department of the Interior had written to Representative H. C.
Bonner, Chairman of the House Committee considering the bill, as
follows in part :

“Sufficient basic research has not been accomplished to predict losses
or to properly advise operating agencies on the means of obtaining
effective control and at the same time avoiding unnecessary fish and
wildlife mortality.”

With astonishing swiftness, and over the mounting protests of con-
servation and other groups alarmed at the prospect of another airborne
“spray” program, the first insecticides were laid down in November,
l957- The rate of application was two pounds of dieldrin or heptach-

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lor per acre, the insecticides being incorporated in granules of an inert
material to cut down wind drift and lessen loss by foliage interception.
It had been established that this formulation would be spread in the
upper soil layers when rain dissolved the granules, and that its effect
would last at least three years.1 Dieldrin was used at three pounds
per acre wherever another pest, the white fringe beetle, occurred as
well as the ant, thus treating for both pests at once. Where the ant
occurred alone, heptachlor was usually the choice. Dieldrin and
heptachlor are extremely toxic substances — about 4-15 times as toxic
to wildlife as is DDT.8 Many wildlife experts and conservationists,
as well as entomologists both basic and economic, felt a sense of
foreboding at the start of a program that would deposit poisons with
8-30 times the killing power of the common forest dosage of DDT
(one pound per acre in gypsy moth control).

The spray campaign got off to such a fast start that both state and
Federal agencies were caught without being able properly to organize
programs that year for assessing the effects of the poisons on wildlife,
so that results of such programs were delayed until after large amounts
of toxicants had already been laid down.

Now that some of these results are finally available, we can see that
they were acutely needed before the program was ever begun. The
misgivings of the wildlife people seem to have been justified on the
whole, since the kill of wildlife in sample treated areas appears to have
been high in most of those that were adequately checked.5’ 8> 10, 12> 21
The USDA disputes many of the claims of damage, but their own
statements often tend to be vague and general. It does seem to be true
that quail and perhaps other wildlife species will make a good come-
back on treated land after two or three years, provided that untreated
areas are available nearby to furnish replenishment stocks once the
treated land begins to recover. Still, most of the information on wild-
life repopulation comes from the accounts of hunters and other sources
not subject to proper checking, and we still have little in the way of
published studies by competent authorities on ecological recovery of
treated lands.

Wash-off into streams and inlets has led to heavy losses among
fish, crayfish and aquatic insects. Dieldrin at only one pound per acre
sprayed on a salt marsh at Vero Beach, Florida, killed all the fish
(including young tarpon) and Crustacea in the marsh and adjacent
waters, and the effect lasting for weeks.12 This particular test, meant
to control sandfly populations, applied only half of the dosage of
dieldrin originally used for fire ant control, and one-third the dosage
actually used on white fringe beetle together with fire ant.

90

Psyche

[June-September

Although the USDA claims that the evidence is inconclusive in
some cases, there does exist contrary information7, 10 indicating that
stock losses from fire ant poisons may sometimes be significant. Various
newspaper accounts, while sensational in tone and possibly exag-
gerated, add further to the impression that damage to cattle, horses,
poultry and household pets may on several occasions have been locally
serious. Even a few livestock deaths, if added to the time and effort
spent by farmers in carrying out awkward measures to protect their
animals from spray measures, must more than balance out any cumu-
lative loss that fire ants may have inflicted directly on farm stock since
the infestation began.

In 1959, the formulation was changed to a dosage of 1.25 lb of
dieldrin or heptachlor per acre, and more recently an alternative
dosage of a quarter pound per acre has been most widely used. This
latter dosage, used twice at three- to six-month intervals, was devel-
oped because of the growing concern about wildlife and the residue
problem. At this rate of application, wildlife apparently suffers much
less seriously, but the fire ant is also much safer than under the old
rate of two pounds per acre, and can probably come back in many
places a year or two after the “light treatment” has been applied,
according to the data of Blake, Eden and Hays1 for similar dosages.
Wildlife officials claim to have heard from Plant Pest Control officers
that there still exist stockpiles of the formulation yielding two pounds
of actual heptachlor or dieldrin per acre, and that this product was
still being used for treating junkyards as of March, 1961, but Dr. E.
D. Burgess of Plant Pest Control denies that this is so.

A serious blow was dealt the program in late 1958, when treat-
ments were only one year old ; Senator Sparkman and Congressman
Boykin of Alabama asked that the fire ant campaign be suspended
until its benefits and dangers could be evaluated properly. Then, in
the beginning of i960, the Food and Drug Administration of the
Department of Health, Education and Welfare lowered the tolerance
for heptachlor residues on harvested crops to zero, following the
discovery that heptachlor was transformed by weathering into a per-
sistent and highly toxic derivative, heptachlor epoxide, residues of
which turn up in meat and milk when fed to stock. Some state
entomologists now definitely advise farmers against the use of hep-
tachlor on pasture or forage.

At just about the time that the residue question arose, the Alabama
State Legislature refused to appropriate state funds for participation
in the program after hearing evidence from state entomologists and
some farmers that the fire ant is a nuisance rather than a direct source

1961]

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9i

of serious harm to crops or farm animals. (Alabama voted some
participation funds again in 1961.) Alabama was followed out of the
program by Florida in the spring of i960. According to a U. P.
release on March 26 of that year, Florida Plant Commissioner W.
G. Cowperthwaite announced, “Efforts to stamp out the fire ant
permanently in Florida have failed.” He said that “the all-out attack
on the pest is being abandoned. In its place a control program
centered on badly contaminated areas will be set up. We thought at
one time we could eradicate the fire ant, but it is impossible.”

It seems likely that Mr. Cowperthwaite’s words accurately express
the situation for the South insofar as the present means of control are
employed. The original plan set forth in 1957 called for eradication
of the ant on the North American continent, by rolling back the
infestation from its borders, applying eradication measures to more
central foci in the main infestation, and instituting an effective pro-
gram of treatment of especially dangerous sources of spread, such as
nurseries. Nearly four years and perhaps 15 million dollars after that
plan was announced, the fire ant is still turning up in new counties,
and is being rediscovered in counties thought to have been freed of
the pest in Arkansas, Louisiana, Florida and North Carolina. Un-
doubtedly, as the task of surveying for an elusive quarry continues,
more reinfestations will turn up, and further “spot control” will be
needed. Some two and one-half million acres, a little less than one-
tenth of the total acreage known to have been infested, have now been
treated with one or more of the formulations discussed above (July,
1961).

What Can Be Done About The Fire Ant f

Even before the aerial spray program began, independent research
workers had brought to the attention of the USDA authorities the
potentialities for fire ant control residing in the use of baits, both
poisoned and otherwise. New approaches to the use of baits were
being explored at the time at Harvard, and a good start was being
made at Auburn University; the two investigations have since brought
forth different but very promising results.

Difficulties in using most poison baits against ants include the
development of social “bait shyness,” a term that describes the fact
that ant colonies will often “learn” to avoid baits that have been taken
by, and presumably have killed, some of their foraging workers. It is
not known how bait shyness arises in the colony. Hays and Arant13
have developed a new peanut butter bait in which very low concen-
trations of a new, extremely slow-acting poison called Kepone® are

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Psyche

[June-September

mixed and squeezed into short lengths of paper soda straws. These
baits have proven to be extremely effective against the fire ant in test
plots in Alabama, probably because the Kepone takes five to seven
days to kill, and thus puts off bait shyness until the entire colony has
fed upon the poison. The USDA has also recently completed some
bait tests. The effect of these formulations upon wildlife has not yet
been fully tested, and there may be a hitch in this direction.

Perhaps even more promising is work done over the last few years
by E. O. Wilson at Harvard25 and M. S. Blum; and his associates at
Louisiana State University2 with the so-called “trail substance” of
the fire ant. This material, found in one of the sting glands of the
ant, is used by the ants to mark trails leading back to the nest from
food sources or other attractive objects. This liquid is released through
the sting, which is used like a pen to draw a trail on the ground. The
odor of the trail substance induces stereotyped foraging behavior, and
also serves as the marker along which the ants run. Apparently, each
species of fire ant has its own distinctive trail substance. At' the present
writing, the chemical composition of the trail substance is not known,
but like other natural products, it will eventually be worked out, and
synthesis of its components and related compounds should be possible.
The trail substance has the advantage that it is a necessary part of the
ants’ communication system, and it is extremely potent. Presumably,
it could be used to lead the ants to poison baits, or, more hypothetically,
it might be used as a “confusion lure,” broadcast in high concentra*
tions, leading the ants to forage fruitlessly in all directions.

THE MEDITERRANEAN FRUIT FLY
Introduction

The Mediterranean fruit fly (or “medfly,” Ceratitis capitata) and
other fruit Hies of greatest importance belong to a family (Trypeti-
dae) of the two-winged or true flies (Diptera): They are not to be
confused with the fruit flies of genetics, which are primarily yeast-
feeders of the genus Drosophila, belonging to another family of the
same order.

Biology and Nature of Damage Done

The adult true fruit flies vary from much smaller than a house-fly
to somewhat larger, and they usually have their wings “pictured”
with dark markings. In the usual case, the fruit fly female, after
mating, will puncture unripe fruit and deposit one or more eggs in
the incision. The larvae are whitish or yellowish maggots that feed in
the fruit on the branch, and then either drop to the ground, or leave
the fruit after it drops, and pupate in the soil. Infested fruit is, of

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course, rendered unfit for human consumption. Host fruits infested
are citrus, peach, mango and about 200 other fruits and vegetables.
Although some true fruit fly species are found in temperate regions,
most, including the medfly, are at home in tropical or subtropical
climates.

In a climate like that of Florida, the medfly can produce about 10-
12 generations per year, since the life cycle is completed in slightly
under one month in warm weather. The medfly is a native of Africa,
but it has spread to most of the world’s citrus-producing areas in
infested fruits carried by human commerce; the United States is one
of the few such countries that have managed to exclude it. Since 1912,
U. S. Plant Quarantine has intercepted the medfly over 1600 times
at various ports of entry in this country, and it became established
here twice, in 1929 and again in 1956, both times in Florida. On both
occasions, vigorous efforts by combined Federal and state forces eradi-
cated the fly before it could become established outside of Florida, and
at present writing, the pest has no known breeding population in the
continental United States.

The 1929 Campaign

On April 6, 1929, larvae were discovered in grapefruit at Orlando,
Florida, and by April 10, adult flies had been found and positively
identified as Mediterranean fruit fly. The Florida State Plant Board
and the USDA sprang into action immediately, shifting inspectors to
the area, and by May 1, 1929, a quarantine was invoked in connection
with a program aimed at prevention of spread of the pest and its
eventual eradication. Quarantine stations were set up on railways,
roads and ports on coastal waters and inland waterways. The quaran-
tine of automobiles moving north and south from, the infested area
proved difficult, but was strictly enforced — when necessary, with the
help of the National Guard. Between 410,000 and 625,000 vehicles
were examined each month, of which 6,900 to 13,100 were found
carrying contraband material, including fruits, vegetables, soil, nursery
stock, compost, etc.

Within the affected area, all actual infestations discovered and the
area surrounding each one for one mile were designated as “infested
zones,” while a “protective zone” extended for another nine miles
beyond every infested zone. Within the infested zones all known
fruits and vegetables were destroyed in order to deprive the flies of
breeding opportunities. Removal of host fruit was continued in the
infested zones, and no vegetables were planted there. Packing houses
were supervised in order to prevent shipping leaks through this
channel and to enforce sanitary measures against possibly infested

94

Psyche

[June-September

fruit lying around their premises. In both infested and protective
zones, the foliage was sprayed with a bait preparation containing
brown sugar and molasses plus a poison — lead arsenate or copper
carbonate.

The extent of the effort may be judged from these figures: the
treatment extended onto 1,002 properties in 20 counties with about
10,000,000 acres of land (containing nearly three-fourths of all the
bearing citrus land in Florida), including 120,000 acres of citrus and
160,000 of non-citrus crops. About 609,000 boxes of fruit were de-
stroyed in this area, and 25,000 outside it. Fifty thousand bushels of
host vegetables were destroyed, and about 300,000 pounds of lead
arsenate were used in the bait spray. Infested shipments were found
in ten localities in seven states outside Florida, owing to the fact that
three-fourths of the citrus crop had been marketed by the time the fly
was discovered.

It was found that kerosene and certain fermenting materials were
attractive to adult male flies, and glass traps containing these were
used to check on the presence of the pest.

By July, 1930, the medflv could no longer be trapped in the
continental United States. Its elimination took an expenditure of
about seven and one-half million dollars and the employment of a
peak work force of some 6,000 men. Reimbursement of those who
sustained losses through confiscation of fruit or other control measures
cost another seven million dollars. The “scorched earth” policy plus
effective quarantine and the crude bait spray had paid off; the medflv
had been eradicated for the time being on this continent.

The iq 56 Campaign

The second medfly infestation began when infested grapefruit was
found at Miami Shores in April of 1956. By June of that year,
infestations were found in 19 Florida counties. Again, Federal and
state forces were marshalled with admirable alacrity, but this time,
after a brief initial period of fruit-stripping in some of the southeastern
Florida counties, a new strategy was employed. In large part, this
plan was devised by L. F. Steiner, US DA fruit fly expert, who had
been working out control and detection methods for various pest fly
species in Hawaii. Fruit-stripping was abandoned, and quarantine
zones of one mile were established around each known infestation.
All fruit or produce moving out of these areas had to be fumigated
or processed immediately. New improved fumigation methods em-
ploying methyl bromide and ethylene dibromide were found quite
satisfactory for most fruit, and could be applied at a rate of only five

1961]

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95

cents a box. Some loopholes were exposed. For instance, mangoes,
which do not stand up well to fumigation, were sent unfumigated to
Chicago, but were found to have been transshipped to Louisiana, a
state vulnerable to the fly because of its mild winters.

Although over four and one-half million automobiles were examined
at roadblocks, the spread of the fly mainly followed the highways,
indicating that contraband fruit or adult female flies were moving by
car. Other minor routes of dispersal occurred through Indian reser-
vations, where mangoes were peddled after being transported by
canoe and otherwise away from the roads, and through the traffic
of guava pickers, who are independent and have their own pickup
stations.

Direct control methods employed a spray containing a bait of
protein hydrolysate (“sauce base” of the food industry) plus a poison
component, the organic phosphorus compound, wettable malathion,
mixed in just enough water to make up a spray that could be applied
by air. This bait attracted flies from distances of over 200
yards away, instead of the few inches or feet over which the 1929
sweetened bait had proved effective. The new bait lured and killed
almost all flies within 100 feet a few hours after their emergence, so
that swaths missed by the planes did not matter so long as they were
not excessively wide. By proper timing of sprays at seven to ten days
apart, the flies were prevented from ripening to sexual maturity after
eclosing from the pupal stage. Since the maggots were able to survive
(in grapefruit and oranges left on the tree) for up to 20 days after
reaching the final larval stage, the spray was continued for one full
generation (50-90 days) after the last fly find.

Detection methods depended primarily upon substances that would
lure male flies. Angelica seed oil in plastic traps with poison proved
to be a highly effective attractant for males, but the different lots of
the oil that were tried were found to be very uneven in their effective-
ness. Furthermore, this commodity was rare and expensive — $100
or more per pound. By early 1957, some 800 pounds of the oil (the
entire world production of ten years) had been used for fly baiting,
virtually exhausting the world supply. The last angelica seed oil
was offered on the world market at $500 a pound. Fortunately, at
just about this time the chemists came through with an effective and
relatively inexpensive substitute that they called siglure, containing
certain simple esters of cyclohexane carboxylic acid. It was learned
that the fruit flies tend to disperse from areas after fruit production
has ceased, and this was a good reason for leaving fruit on the trees
in infested areas. Fallen fruit was destroyed wherever possible.

96

Psyche

[June-September

An auxiliary control used in heavily infested zones was the applica-
tion of a formulation at the rate of five pounds of dieldrin per acre
to the soil under infested trees. This was aimed at pupating larvae
and adults leaving the pupal stage.

The program progressed steadily. Infestations were found in a
total of 28 counties, most of them south of the 1929 zone. While the
1929 infestation had affected mainly the major commercial citrus
groves of central Florida, the 1956 invasion was centered more in
the ornamental and dooryard plantings of residential areas in the
southern part of the state. This required the use of more of the
safer twin- and four-engined planes in the low altitude bait-application
flights.

One year after the first discovery of 1956, nine-tenths of the total
acreage had been treated, and only about 1 2,000 acres of new infesta-
tions remained to be discovered. One by one, during late 1956 and
early 1957, counties were released from the aerial spraying routine
after no more flies could be found in them, and in November, 1957,
the last known infestation was eliminated from an island off the coast
in Manatee County. The cost of the eradication program, paid jointly
by the state and Federal governments, was about $1 1 million, but only
small quantities of fruit had had to be stripped from the trees and
destroyed.

Eight hundred thousand acres were sprayed one or more times —
some of them up to a dozen times — for a total of six and one-half
million spray-acres. Twelve million pounds of malathion and a million
gallons of sauce base went into the bait spray, and 1,667,217 pounds of
dieldrin were used in the bait treatment, A maximum of 800 person-
nel was involved in the 1956 struggle, as compared to the 6,000 of
the 1929 campaign — labor costs of course having; risen steeply since
the earlier campaign. At the peak of the campaign, some 54,000
detection traps were in use all over Florida, and additional trapping
was done in other southern states and Cuba in areas where preferred
host fruits grow. About 12,000 fly1 specimens were caught, and none
of these came from states outside Florida. The Florida Legislature
has voted funds for continued lure trapping, using combined lures
for several fruit fly species in addition to the medfly. In June, 1958,
32,000 traps were still in use throughout Florida.

Harmful Effects of the Campaign

It seems reasonably clear that the two medfly campaigns were com-
pleted with little serious loss of wildlife or damage to non-infested
crops, domestic animals and human property. The 1956 program

1961]

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97

received good publicity in the press and on television and radio, and
most tropical fish producers were able to cover their ponds, while
paint and plastic testing laboratories could spread plastic sheeting
over their test plates. Housewives were advised to withhold wash
from clotheslines, and automobile owners to cover or be prepared to
wash their cars. Some damage was noted on cars with lacquer finishes,
but not on those with enamel, and the spotting proved to have been
caused by malathion. Some loss of tropical fish was also reported, but
not in ponds with deep enough water. Reported losses of birds,
mammals and beneficial insects were not confirmed upon investigation.
One C-84 twin-engine aircraft crashed at Boca Raton while ferrying
materials, killing a crew of five men.

Side benefits from the spray included control or depression of insect
pests such as houseflies, mosquitoes and the papaya fruit fly during
the period of application.

THE SCREW WORM
Introduction

The screwworm is the maggot (larva) of a large fly ( Callitroga
hominivorax , plus at least one other species occurring outside the area
concerned). The maggot lives in the flesh of warm-blooded animals
and gets its name from its fancied resemblance to a wood screw. All
sorts of mammals are attacked, but from the human standpoint in this
country, the damage it inflicts on cattle has been most important. The
screwworm has a year-round range in the American tropics and Sub-
tropics, from Texas and other border states south to Argentina. Each
summer, screwworm flies migrate northward to spread the infestation
into the midwestern states, and infestations are known to have been
introduced into Illinois, Iowa, New Jersey, South Dakota and other
northern states in livestock shipments carrying the pest. Each year
up to 1933, winter cold killed the infestation back to the southern
parts of the border states and to Mexico, where the winter weather is
mild enough to permit permanence of the fly population.

In the summer of 1933, screwworms appeared for the first time in
the southeastern United States, probably shipped in infested south-
western livestock, and before they could be controlled they had spread
into peninsular Florida. Here they found the climate mild enough to
support a year-round population, and thus a permanent infestation
became established in the Southeast. Each summer this infestation
spread outward from Florida into additional southeastern states, and
each winter it died back to Florida and the warmer parts of Georgia
and Alabama. During 1935-1937, the affected states in cooperation

9«

Psyche

[June-September

with the USDA applied the best known animal husbandry practices
and tried larvicides and repellents to treat and protect livestock
wounds directly. While these expensive measures did help to cut
livestock losses, enough larvae survived in neglected livestock and wild
animals to keep the infestation alive and dangerous. By 1957, the
State of Florida and the Federal Government were ready to support
the then new technique of eradication based on male sterilization, and
funds were appropriated to begin the campaign against the screw-
worm.

Biology and Nature of Damage

The screwworm is an obligatory feeder in the flesh of living
mammals. Each female fly lays her eggs in a mass of about 200 on
scratches or near exposed wounds on the animals, and the eggs take
12-24 hours to hatch. The larvae then enter the wound and feed
extensively on the muscle tissue. As tissue decomposition advances,
more and more female flies are attracted to infested wound areas, and
the maggot populations at such sites increase correspondingly. The
larvae burrow in the tissues for five to seven days, after which they
leave the wound and drop to the ground, where they burrow into the
soil to pupate. The pupal stage lasts a week or more, depending upon
the temperature. The pupa is vulnerable to low temperatures, and
freezing soil or prolonged cold kills it. After eclosing from the
puparium, the adult flies disperse and seek food. Flies have been
found to disperse to distances as great as 35 miles in one week. In the
summer, mating begins two days after eclosion, and four to six days
later the females have been mated and have laid fertile eggs. The
sexes reach adulthood in about equal numbers, and the females mate
only a single time, although the males normally mate several times.
(Some attention has been given to breeding males that will mate
a greater number of times.) Females segregated from males in the
laboratory to prevent fecundation oviposit as readily as do mated
females. In summer conditions, females live two to four weeks as
adults, and may deposit three, four or more egg masses during this
span.

Because oviposition is triggered only by the presence of a wound on
a suitable host animal, and because of predation of mature larvae by
insects, especially by ants, the number of adults produced is rarely
high. Uvalde County, Texas, has had the heaviest infestations in the
United States, with 100-500 flies produced per square mile per week,
but infestations south of the border may be even heavier.

Massive infestations of screwworm can quickly weaken and kill
even full-grown cattle, and very small animals often succumb before

1961]

Insect Control Programs

99

the flies can complete their larval growth. The pest has caused live-
stock losses of 20-40 millions of dollars annually, about half of this
figure in the Southeast.

Eradication , Operations

The story of screw worm eradication in the Southeast begins in 1936
with the work of Melvin and Bushland,3 who learned how to culture
the insect in the laboratory en masse on ground meat, blood and water
containing a small amount of formaldehyde to retard spoilage. Dr

E. F. Knipling, now heading entomological research in the USDA,
speculated in conversation in 1938 that the known habits of the
females suggested that they might mate only once, which if true meant
that laboratory-reared sterile males might be used to control isolated
populations such as the one in Florida. The idea was not followed
up until after the war, when Knipling directed that the mating habits
and physiology of screwworm flies be studied in detail, and that
attempts be made to find chemicals capable of rendering the males
sterile. In 1950, a general paper was published by H. J. Muller, in
which this famed geneticist pointed out that Drosophila fruit flies in
the laboratory were sterilized by irradiation. A colleague, A. W.
Findquist, passed this paper on to Knipling, who then contacted
Muller about the possibility of employing radiation sterilization on
screwworms. The reply encouraged Knipling to initiate experiments,
and Bushland and Hopkins eventually established that screwworms
were readily sterilized by irradiating pupae that had been held at 8o°

F. for five days. A dose of 2,500 r sterilized males, and 7,5°° r pre-
vented egg production altogether. Adult males emerging from irradi-
ated pupae proved able to mate normally with untreated females, but
the egg masses resulting were of course infertile. Determination of
critical doses proved to be laborious and time-consuming, but coop-
eration with cytogeneticists soon gave rise to important short-cuts in
the process, because damage could be assessed by cytological examina-
tion instead of waiting for the full life cycle to carry through in
order to get results.

Field tests run on Sanibel Island, two miles off the Florida coast,
proved that its screwworm population could be reduced by the release
of 100 sterilized males per square mile per week, a figure that sur-
passed the number of native males. But Sanibel is so close to the
mainland that it was easily reinfested, so eradication could not be
attempted there.

The conclusive eradication test was finally performed on the Dutch
island of Curacao in the south Caribbean Sea. Screwworms were

IOO

Psyche

[June-September

reared at Orlando, Florida, and irradiated in a cobalt-60 source
built at Oak Ridge. At first, flies were released by air at a rate of 100
males per square mile per week, but this proved only fractionally
effective because the swarming, unattended goats and sheep of
Curacao harbored a much larger screwworm population than had
been encountered in Florida. The release rate was accordingly in-
creased from 100-400 males per square mile per week, and the first
saturation of the island with sterilized flies caused substantially more
than half of the egg masses laid on test animals to be sterile. After a
month of continued releases, when another generation of adults
emerged, the native flies were so reduced in numbers that the percent-
age of sterile matings increased greatly. The emergence of the second
generation of wild flies saw the proportions so altered that practically
all matings were sterile ones. By generation III, only two egg masses
were found in goat pens on the island, and both of these were sterile.
No more screwworm eggs were found during the additional two
months that flies were released on Curacao, and release was terminated
in January, 1955, less than six months after the first flies were let go.

The Curasao experiment, heartening as it was, also showed the need
for improved procedures for mass production of sterilized males. At
a rate of 400 males per square mile, the 50,000 square miles of the
overwintering area in Florida was estimated to require 20 million
males weekly. The females produced equal the males in numbers and
are not easily separated from them in practice, so these doubled the
necessary weekly rate of release to 40 million flies. An additional ten
million flies had to be reared to make up for mortality of pupae and
to provide for breeding stocks. This came to a weekly grand total
of 50 million flies, in contrast to the 170,000 larvae raised each week
for the Curacao test.

To meet this demand, experts on insect rearing, irradiation methods
and mass production engineering cooperated to transform a large air-
plane hangar near Sebring, Florida, into a wonderfully efficient plant
capable of producing more than the needed number of sterile screw-
worm flies each week. This plant employed fully modern production
line techniques, with the larvae being carried through their feeding
life and thence to the pupal stage and the irradiation chamber on a
continually moving line of stacked trays suspended from a monorail.
Full safeguards were provided against possible escape of unsterilized
flies, and elaborate precautions set up to protect the employees from
radiation and from: the odor of the meat-blood larval food.

Designed, built and equipped on a “crash” basis in just nine months,
and at a cost of under a million dollars, the plant moved into full-

1961]

Insect Control Programs

IOI

scale production in July, 1958. By early spring of 1959, it was pro-
ducing for release at the phenomenal rate of 50-60 million flies per
week. The flies were placed in special cardboard cartons that could
be opened as dropped from the plane. About 20 light planes were used
at the peak of operations, each flying five to six hours a day over
prearranged flight patterns based on a few strategic release centers
spaced over Florida. Three long trap lines covering the state from
north to south provided information on the effectiveness of the opera-
tion, and a field force of about 50 livestock inspectors worked on
quarantine patrol duty. Stringent quarantine regulations were set up
to prevent infested livestock from entering the Southeast from across
the Mississippi.

The program had a swift and dramatic effect on the Florida screw-
worm population. By the middle of March, 1959, all attempts to
find egg masses or active screwworm infestations in Florida proved
negative. On June 13 of that year the USDA and the Florida Live-
stock Board could announce, “Southeast free of screwworms for 16th
consecutive week.” This record was marred in the following week by
the discovery of a single case of screwworm infestation in Highlands
County, Florida. The releases continued at a rate of about 42 million
flies a week, blanketing the area from southern Alabama and Georgia
south to Key West. After some weeks during which no signs of a
wild fly population were found, the rate of releases was dropped to
30 million flies per week and lower, and finally, on November 14,
1959, by releases were terminated. The total eradication of the south-
eastern screwworm population had been achieved.

In the months since the release ended, an infested dog has been
found in Florida — evidently brought in from the outside — and dur-
ing the spring and summer of 1961, infestations have appeared at
points along the Gulf Coast from the west, apparently originating
from infested livestock shipped from the Southwest. It seems that
these new threats to the Southeast can be handled with the available
weapons, and the long-range problem now is centered on rolling the
screwworm menace back across a defensible line in southern Mexico
or Central America, and holding it there by quarantine and possibly
by a constantly maintained belt of sterile flies.

COMPARISONS OF THE FOUR PROGRAMS

In comparing operations against the four pests we have just con-
sidered, it is well to recall once again that each insect is a separate
and distinct problem in control. Some insects have characteristics

102

Psyche

[June-September

that lend themselves to simple control, methods, while others are just
naturally tougher, faster-spreading or faster-breeding, and defy all
control methods that have been tried. However, it is also evident that
the four programs do differ considerably among themselves in basic
ways, especially in the resourcefulness and insight of their planning
and operating personnel, in the kind and amount of information upon
which control operations are based, and in the adaptability of the
operating plans to conditions as they are met while the campaign
proceeds. The first factor — personnel — is of course very difficult
for one outside of the agencies involved to evaluate, and in any case,
judgements are bound to be influenced by hindsight according to the
success of the particular program concerned.

The second factor for analysis is the nature and quantity of the in-
formation on which each program proceeded. Ideally, of course, a
control campaign is based, on a full knowledge of the target pest, its
life history, ecology, physiology and behavior; on a basic understand-
ing of the efficacy of various, methods that might be used against the
target; and on a reasonable assurance that these methods do not have
seriously harmful effects on valuable plants, animals, microorganisms,
inanimate human property, or on man himself. Such knowledge, of
which we can never get enough, is provided by previous investigations,
by pilot trials, and by continuing evaluation of operational results.
These activities, collectively known as research, are the counterparts
of intelligence-gathering in a military campaign. The public as well
as the technicians involved have come to take research for granted in
insect control programs, just as they confidently assume that the proper
tests of safety have been applied when a new antibiotic or vaccine is
issued by medical authorities.

If we look at the details of the four projects as they have been dealt
with in recent years, the differences among them, in research effort are
very striking. The research behind the screwworm program: has been
extensive, imaginative and persistent, and obviously it has paid off
handsomely. The second medffy campaign, unlike the desperate,
scorched-earth first one, was carried out with an efficiency grounded
on solid long-term research into the bionomics of fruit flies in general,
particularly that conducted by L. F. Steiner and his colleagues in their
Hawaiian installation. Here again, it is clear that previous research
was crucial in a successful eradication campaign.

The gypsy moth campaign has the longest history, and also the
oldest research program, of any of the four efforts considered here.
In the years before mass air-spraying, many kinds of measures were
tried against the moth, including the introduction of natural enemies

1961]

Insect Control Programs

103

already mentioned, as well as cultural methods (such as tree banding
and egg mass destruction) and poisons sprayed from the ground. Also
built up during the years was a store of knowledge concerning the life
history, foodplants, enemies and distribution of the moth, and particu-
larly a fund of information on the effect of the female attractant on
males. All this has proven, very useful in developing control methods.
Nevertheless, the recent work of Campbell (some results of which
are outlined above) indicates that there was and is much more of
importance to be learned about the behavior of gypsy moth populations
than has been generally appreciated. The preparation of gyplure
and other attractants in the last few years had doubtless been made
easier by technical developments in natural-product chemistry, but
perhaps even without these developments more could have been done
in the past with attractant research had more time and money been
spent on it. To sum up gypsy moth research, one might say that it
began rather well and then tended to get into a rut, from which it
has been pulled only during the last few years. The present research
program is expanding and striking out in new directions, and the
outlook now seems rather good for the eventual control of the moth.

As we have already seen, the fire-ant mass spraying program began
full blast in the fall of 1957. Considering the very high potency of
the poisons used and the great areas over which they were to be
sprayed, the research background of the fire-ant program was so
sketchy as to be virtually non-existent. USDA investigations ran
from 1948 to 1953, and consisted mainly of survey scouting for new
infestations plus routine life history, ecological and insecticide-testing
work. As already emphasized,22 no research was done by the USDA
from 1953 until after the mass spraying had gotten well under way.
The Gulfport Methods Improvement Laboratory was not opened
until 1958. Nevertheless, in their letters and releases,23 USDA
officials spoke of “expanding” the “continuing research effort,” thus
giving the impression that an unbroken chain of research studies
stretched back from the start of the spray program. The USDA
releases emphasize the liaison with the U. S. Fish and Wildlife
Service “from the outset,” and even serm to imply concurrence of
the Service in the mass spray program.24 As we have already seen
from Dr. Leffler’s letter,19 this concurrence could not possibly have
been granted at that time. The first meeting of USDA and Fish
and Wildlife officers on the fire-ant program took place, according to
the USDA, in Washington on December 12, 1957, about a month
after the spraying had started. The delay is important in view of
the time needed by wildlife researchers to set up and carry out a

104

Psyche

[June-September

complicated wildlife survey in an area about to be treated. In fact,
the Fish and Wildlife Service seems to have been presented with a
fait accompli upon which to make its studies.

What about outside research? In the years between 1948 and
1957, Dr. E. O. Wilson at Harvard had continued his work on fire-
ant variation, distribution and social behavior, and had discovered the
existence of a trail-forming chemical laid down by foraging worker
ants. Research on this substance was continued by M. S. Blum and
co- workers at Louisiana State University, and is still going on. The
active group at Auburn in Alabama studied fire-ant crop damage
(which had unaccountably dwindled practically to nothing by 1957)
and worked on promising bait formulations. The findings of these
groups swerved the spray program not at all. The Gulfport Labora-
tory is now working on baits and other angles of attack, but insofar
as their results have affected the operations to date, emphasis still
seems to fall on mass spray methods. No recent specific, detailed study
of the damage caused by the ant seems to have been reported, despite
the claims of competent state entomologists that crop damage is now
negligible. We are left, then, with no concrete information to counter
the claims of wildlife experts and state entomologists that the ant is
not a major pest deserving of the effort and funds expended upon it.
For research effort, the fire-ant program, must take low marks.

The last factor to be compared among the programs is their adapta-
bility to conditions met as operations proceed. This is so closely related
to the research facet of the respective program that we are not
surprised to find the flexibility of operations more or less closely
paralleling the quality and amount of research. The screw worm and
medfly programs made major adjustments smoothly and without delay
as the information available indicated they should.

The gypsy moth campaign has varied; sometimes the operational
response to changing conditions was rapid and efficient, while at other
times it lagged. Curiosity about the obviously great fluctuations in
abundance of the moth, and especially about the great peak following
the first extensive air spraying, are not reflected in the impassively
literal Annual Reports on gypsy moth control work. Even the over-
stepping of the Berkshire-Green Mountain barrier seems never to have
raised much doubt on the part of the government control officials that
the mass spray program in progress would eventually bring about the
eradication of the insect in North America, at least to judge from the
reports. But events have caught up with the program. The milk
residue problem in New York State first halted the program in much
of this key “frontier area,” and later forced a switch to the less effec-

1961]

Insect Control Programs

105

tive sevin for most districts. Finally, a new Methods Improvement
Laboratory is opening this year at Otis Air Force Base in Massachu-
setts, and one way or another we may hope to see some more sophisti-
cated control measures tried against the gypsy moth.

After five stormy years of air spraying, the fire-ant control program
goes on pretty much as before, but with greatly reduced dosage in
many areas. The reduction seems to have been forced in part by
serious wildlife kills and perhaps some destruction of livestock and
poultry, as well as by the threat of residues. Where the new double
quarter-pound treatment is being applied, damage to warm-blooded
animals is apparently not serious. It is, of course, effective against
the ants for a much shorter time, and it is doubtful whether its residual
effect is up to the task of preventing reinfestation of treated areas.
Recently, “mopping-up” activities have been required after treatment
in a number of places.

There is a question, already decided in the negative by some of the
infested states, whether the eradication campaign should continue in
its present form. Not without some logic, wildlife experts have called
the fire-ant program, “scalping to cure dandruff.” But this campaign
has so much momentum, fueled annually with 2.4 million dollars in
Federal appropriations, that even the defection of such key participant
states as Alabama and Florida has failed to halt it. As the possibility
of eradicating the fire ant by the present mass spray techniques recedes
into future decades, it will be interesting to see how many more years
Congress will vote to keep the present control machinery rolling.

CONCLUSIONS AND RECOMMENDATIONS

The case histories we have reviewed illustrate, I think, the point
that mass air spraying of non-selective insecticides can be disappointing
as control agents and are in some cases dangerous to the living human
environment as well, perhaps, as to man himself. These dangers are
usually discussed as “side effects,” a term which in itself reflects the
special viewpoint of many of the control men on the job. These are
“practical” people, absorbed in managing large teams with complex
apparatus, and often caught up in the direct urgency of “crash pro-
grams.” Their efforts are directed at a clear and simple goal — the
eradication or control of a particular insect. In the heat of such
campaigns, complaints arising from damage to humanly-valued re-
sources are likely to appear as mere incidental annoyances to the
control men, and the damage itself is minimized and shrugged off.
But the side effects of the control men may in reality amount to
catastrophes from other viewpoints, as in the case of the fire-ant

io6

Psyche

[June-September

campaign. When the cost of a campaign in dollars plus the losses in
wildlife, stock or other resources destroyed begins to balance or exceed
the benefits to be gained by eradication of the pest, then it is time to
give thought to cutting off or drastically modifying the program. In
such a case, side effects become main effects, and we should never
forget it. The dangers involved in the mass use of pesticides has
recently been dramatically recognized in Great Britain,15 where a
Parliamentary Investigation Committee of 43 Members has accused
the Ministry of Agriculture of negligence in insecticide research and
has recommended that pesticide use be intensively investigated and
rigidly controlled, and has called for the “immediate prohibition”
of heptachlor, dieldrin and aldrin.

Our case histories illustrate another point: alternative control
measures are increasingly available, and we should expect their devel-
opment to be accelerated. The medfly and screwworm campaigns are
shining examples of the results of real thinking and hard work, but
most of all they point up the value of new approaches and a sound
knowledge of the pest to be dealt with — in other words, they bear
the stamp of thorough research.

The issue is clearcut : in the face of a new and spreading insect
menace, do we rush out the planes and the poison, or do we first find
out what we ought to do and how it should be done, on the basis of
adequate information ?

The problem of urgency is sure to be raised in answering this ques-
tion ; otherwise, there could be only one answer. In the light of past
insect invasions, however, urgency has rarely been so great as to pre-
clude some kind of research assessment of the problem before mass
control could begin. Furthermore, research can be called upon to
provide a sound body of general background information and princi-
ples before the emergency occurs. Our insect control programs often
lack this kind of a background, as the makeshift fire-ant campaign
illustrates, but when they do have it, as in the case of the medfly, the
success of control efforts may be rapid and brilliant.

But in the USDA, entomological research is often hampered at the
basic level. Even in such fundamental fields as insect taxonomy and
morphology, USDA specialists are for the most part overworked and
overcrowded. Daily the cartons of insects submitted for identification
pile up on each man’s desk, and most of these highly qualifield research-
ers must work on their own time to get any basic investigations
completed. The same is often true of extension entomologists at the
state level. Permanent workers in the new and vital disciplines of
population dynamics and insect behavior have scarcely begun to be

1961]

Insect Control Programs

107

hired by the Federal Government or the states for work in their own
fields; yet, as our case histories demonstrate, these fields will surely be
pivotal in future control developments. Bright spots in the entomo-
logical research picture are the grants from, other governmental agen-
cies for the support of basic: research, mostly in the universities. But
such grants are no substitute for an adequate research establishment
within the USDA itself.

From all of these considerations, I think the recommendations must
be clear. They are as follows:

1. Every mass control campaign should have an adequate research
program functioning as far ahead as possible before control operations
get under way. The control work should be guided by the research
findings, and not the reverse, and every campaign should be reevalu-
ated frequently to see if a need for it continues.

2. The USDA quickly should be granted funds to expand all
permanent research facilities related to pest control. Special attention
needs to be given to basic fields such as systematics, physiology, be-
havior, ecology and genetics. The study of the natural-product
chemistry of insects should be stepped up.

3. Mass broadcasting of non-selective poisons, especially spraying
and dusting from the air, should be deemphasized and the development
of other measures, especially selective lures and sterilization tech-
niques, correspondingly augmented. Over lands other than intensively
cultivated agricultural blocks, mass insecticides should be used with
the greatest caution and only in real emergencies after other measures
have failed. Non-selective insecticides in general should be considered
only as stopgap remedies, pending the development of better means
of control for all types of land.

4. There should be established a strong permanent inter-agency
office to coordinate policies and activities related to pesticidal opera-
tions as they affect the biotic environment and human health. This
office should have ample funds to allot to the proper agencies for
research on specific problems. It would be made up of representatives
from the USDA Agricultural Research Service, the Fish and Wild-
life Service of the Department of the Interior, and the Food and
Drug Administration of the Department of Health, Education and
Welfare.

ACKNOWLEDGEMENTS

I am grateful to the many individuals and organizations who made
this essay possible, although I cannot mention them all by name here.
Particularly helpful were the information and criticisms offered by

io8

Psyche

[June-September

officers of the Plant Pest Control Division, USDA, and by the follow-
ing members of my own department at Cornell: Professors H. H.
Schwardt, R. A. Morse, A. A. Muka, T. Eisner and G. G. Gyrisco.
I also owe thanks to many of the authors listed in the References
section below for reprints of their papers and for other material I
received from them. It should be understood that those who helped
do not necessarily endorse the views here expressed.

Work on this report was initiated by and carried out with the
cooperation of The Conservation Foundation, of New York City.
I gratefully acknowledge’ the aid given me by the Foundation’s staff.

REFERENCES

1. Blake, G. H., Jr., W. G. Eden and K. L. Hays. 1959. Residual effec-

tiveness of chlorinated hydrocarbons for control of the imported fire
ant. Jour. Econ. Ent., 52:1-3.

2. Blum, M. S., J. R. Walker, P. S. Callahan and A. F. Novak. 1958.

Chemical, insecticidal and antibiotic properties of fire ant venom.
Science, 128:306-307.

3. Bushland, R. C. 1960. Male sterilization for the control of insects.

Advances in pest control research, Interscience Publ., New York,
3 : 1-25.

4. Campbell, R. W. 1959. Population dynamics of the gypsy moth.

Typed abstract.

5. Clawson, S. G., and M. F. Baker. 1959. Immediate effect of dieldrin

and heptachlor on bobwhites. Jour. Wildl. Mgt., 23:215-219.

6. Coarsey, J. M., Jr., and G.H. Culpepper. 1952. Research line project

I-h-8 2: Investigations on the control of the imported fire ant. Agr.
Res. Serv., Washington, mimeographed.

7. Conservation News, Washington, Sept. 15, 1958. Farmers protest fire

ant control program “throat-ramming,” p. +.

8. DeWitt, J. B., C. M. Menzie, V. A. Adomaitis and W. L. Reichel.

1960. Pesticide residues in animal tissues. Fish and Wildl. Serv.,
Washington, mimeographed.

9. George, J. L. 1957. The pesticide problem. The Conservation Founda-

tion, New York, 57 T 10 pp., mimeographed.

10. George, J. L. 1958. The program to eliminate the imported fire ant.

The Conservation Foundation, New York, 39 pp., mimeographed.

11. Guyton, T. L., D. R. Shepherd, F. A. Soraci and H. H. Schwardt.

1957. Gypsy moth eradication. Summary of a meeting held at Ithaca,
New York, Sept. 4, 1957 ; mimeographed.

12. Harrington, R. W., Jr., and W. L. Bidlingmayer. 1958. Effects of

dieldrin on fishes and invertebrates of a salt marsh. Jour. Wildl. Mgt.,
22:76-82.

13. Hays, S. B., and F. S. Arant. 1960. Insecticidal baits for control of

the fire ant. . . . Jour. Econ. Ent., 53: 188-191.

14. Hays, S. B., and K. L. Hays. 1959. Food habits of Solenopsis sacvis-

sima richteri Forel. Jour. Econ. Ent., 52:455-457.

15. Hillaby, J. 1961. Britain warned on wildlife risk. New York Times,

Aug. 5, 1961.

16. Huddleston, E. W., G. G. Gyrisco and D. J. Lisk. 1960. DDT residues

on New York dairy farms following the gypsy moth eradication
program. Jour. Econ. Ent., 53:1019-1021.

1961]

Insect Control Programs

109

17. Jacobson, M. 1960. Synthesis of a highly potent gypsy moth sex

attractant. Jour. Organic Chem., 25:2074.

18. Karlson, P., and A. Butenandt. 1959. Pheromones (ectohormones)

in insects. Ann. Rev. Ent., 4: 39-58, cf. p. 42.

19. Leffler, R. 1957. Letter to Congressman H. C. Bonner, dated May 1.

U. S. Dept. Interior, Washington.

20. Rudd, R. L., and R. E. Genelly. 1956. Pesticides: their use and

toxicity in relation to wildlife. Game Bull. 7, Calif. Dept. Fish &
Game.

21. Southeastern Association of Game and Fish Commissioners. 1958.

Proc. Symposium: The lire ant eradication program and how it affects
wildlife. Columbia, South Carolina, 34 pp. (Articles by Allen,
Tarzwell, Rosene, Baker, Lay, Glasgow, Newsom and Cottam.)

22. U. S. Dept. Agriculture. 1957. Letters from W. C. McDuffie (dated

March 28) and A. W. Lindquist (dated April 16) of the Agricultural
Research Service, Entomology Research Division.

23. U. S. Dept. Agriculture. 1958. Letter from M. R. Clarkson, Acting

Administrator of the Agricultural Research Service, dated January 3.

24. U. S. Dept. Agriculture. 1960. Memorandum from Plant Pest Control

Division to Conservation Foundation, New York, dated June 1, and
accompanying statement dated May 4.

25. Wilson, E. O. 1959. Source and possible nature of the odor trail of

fire ants. Science, 129:643-644.

26. Wilson, E. O., and W. L. Brown, Jr. 1958. Recent changes in the

introduced populations of the fire ant. . . . Evolution, 12:211-218.

27. Wilson, E. O., and J. H. Eads. 1949. A report on the imported fire

ant. ... in Alabama. Alabama Dept. Conservation, 53 pp. + 13 pi.,
mimeographed.

I IO

Psyche

[June-September

A ppendix A

[Data furnished by Plant Pest Control Division, Agricultural
Research Service, August 25, 1961.]

SUMMARY OF ACREAGE SPRAYED FOR GYPSY MOTH

CONTROL, SUPRESSION AND ERADICATION

(All DDT Except As Noted)

Application

Application

By Ground

By Aircraft

Equipment

Totals

(Acres)

(Acres)

(Acres)

1945

5,103

1,092

6,195

1946

62,201

19,427

81,628

1947

106,677

56,932

163,609

1948

212,260

53,650

265,910

1949

390,576

34,239

424,815

1950

582,895

17,205

600,100

1951

177,713

2,499

180,212

1952

202,109

15,032

217,141

1953

179,451

6,970

186,421

1954

i,37Ei99

29,817

1,401,016

1955

1,083,169

25,129

1,108,298

1956

926,073

I5,39i

941,464

1957

3,395,248

27,695

3,422,943

1958

516,150

18,426

534,576

1959

115,078"

35,343

150,421

i960

65,5382

33,369

98,907

1961

141,270s

i9,5834

160,853

Totals

9,532,710

411,799

9,944,509

By Aircraft

By Ground Equipment

19591

DDT 29,518 acres

Sevin 85,560 ”

All DDT

115,078 acres

I9602

DDT 54,103 acres

Sevin 11,435 ”

All DDT

65,538 acres

1961s

DDT 104,770 acres

4DDT

19,342 acres

Sevin 30,000 ”

Sevin

241 ”

Methoxychlor 6,500 ”

19,583 acres

141,270 acres

1 1 1

1961] Insect Control Programs

Summary of Gypsy Moth Defoliation

Calendar Years 1924 to i960

Year

A cres

Year

A cres

1924

825

1943

34.845

1925

48,560

1944

250,148

1926

80,822

1945

821,487

1927

140,920

1946

622,919

1928

262,514

1947

7,422

1929

55PI33

1948

32,467

1930

288,226

1949

78,673

1931

204,721

1950

5,368

1932

286,395

I95i

21,314

19 33

397,730

1952

293,052

1934

492,361

1953

1,487,077

1935

540,769

1954

491,448

1936

428,622

1955

52,061

1937

608,760

1956

43,158

1938

3G,954

1957

6,458

1939

492,640

1958

125

1940

485,636

1959

14,467

1941

468,021

i960

48,722

1942

44,577

1961

data incomplete

Moorestown, N. J.
August 16, 1961

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PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol.68

Established in 1874
December, 1961

No. 4

CONTENTS

Australian Carabid Beetles VII. Trichosternus, Especially the Tropical

Species. P. J . Darlington, Jr 113

Adrityla, a New Milliped Genus (Chordeumidea : Conotylidae) .

Nell B. Causey 126

Ants from Three Remote Oceanic Islands. Robert W. Taylor and Edward
O. Wilson 134

Studies on Carboniferous Insects of Commentry, France: Part III. The
Caloneurodea. F. M. Carpenter

150

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1961-62

President J. J. T. Evans. Harvard University

Vice-President C. Walcott, Harvard University

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Executive Committee R. W. Taylor, Harvard University

S. K. Harris, Boston University

EDITORIAL BOARD OF PSYCHE
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University; Associate in Entomology, Museum of Comparative
Zoology

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PSYCHE

Vol. 68 December, 1961 No. 4

AUSTRALIAN CARABID BEETLES VII.
TRICHOS TERN US, ESPECIALLY THE
TROPICAL SPECIES

By P. J. Darlington, Jr.

Museum of Comparative Zoology, Cambridge, Mass.

This is one of a series of papers describing new Carabidae of zoo-
geographic importance referred to in my account (1961b) of transi-
tion of Australian wet forest carabid faunas. Some other papers of the
series, including a list of my localities, are referred to below (p. 130).
The present paper is concerned with Trichosternus , especially with
the comparatively little known tropical species. Types of new species
are placed, at least for the time being, in the Museum of Comparative
Zoology; paratypes, with C. S. I. R. O. at Canberra and usually in
the Queensland Museum. Proportions given in the descriptions are
calculated from actual measurements.

Items of geographic or evolutionary interest derived from Tricho-
sternus are its total distribution ; the occurrence of two very distinct
(relict or primitive?) species, fax and montorum , high on Mt. Bartle
Frere (one reaching Mt. Bellenden Ker too) ; the distribution of T .
f rater and its apparent allies ( suhvirens and simplicipes of South
Queensland and nudipes to fisheri in the tropics) , which form a group
of mostly allopatric forms in which the male front tarsi have apparent-
ly been simplified in two different stocks and in which double invasion
or hybridization may have occurred in a very limited area on part of
the Atherton Tableland (pp. 122-125). Another case of hybridiza-
tion (of eungella and mixtus ) may have occurred on the Eungella
Range (p. 127). See map for distribution of species on and north
of the Atherton Tableland.

I cannot give an exclusive definition of Trichosternus. Characters
used by Sloane (1894 etc.) and Tschitscherine (1902) fail among
recently discovered species. Compared with Nurus, Trichosternus is
usually smaller, more lightly built, with S front tarsi usually dilated
and with 3 segments squamulose below, while in Nurus the cf front
tarsi are usually simple, rarely (e.g. in N. atlas Cast.) slightly dilated
and with 2 segments squamulose. However, exceptional Trichosternus

Psyche, 1961

Vol. 68, Plate 7

A\

I

I

• Daintree

V

Darlington — Trichosternus

1961]

Darlington — Australian Carabid Beetles

ii5

resemble Nurus in having cf tarsi simple ( T. subvirens Chd., sim-
plicipes SL, nudipes Dari., soror Dari.) or with only 2 segments
squamulose ( setosiceps SL). Compared with N otonomus, Trichos-
ternus is usually larger, often with prosternal process setose (rarely
in N otonomus) , often with alternate intervals of elytra raised, and
often without distinct 10th intervals at outer edges of elytra, but all
these characters fail in some species. A generic revision of the larger
Australian pterostichines is much needed. The Australian genera
should be compared with the New Zealand ones (Britton 1940) and
phylogenies should be worked out if possible, and the old types, many
of them in Europe, should be re-examined. However, I cannot under-
take all this now.

The known range of Trichosternus is the eastern edge of Australia
from the base of the Cape York peninsula (south of Cooktown) to
central New South Wales (perhaps a little north of Sydney), with
one species isolated in southwestern Australia (Darlington 1953) and
another on New Caledonia. All the tropical Australian species (ex-
cept cordatus ) live in rain forest, but some more-southern species occur
in savannah woodland, and some enter or are confined to south tem-
perate rain forest on the Dorrigo-Ebor and Mt. Royal plateaus.

Before considering the tropical Trichosternus, I give the following
tentative key to species south of the tropics, as a basis for comparison
( cf . Sloane’s key, 1899, pp. 567-569, and Tschitscherine’s, 1902, pp.

Explanation of Plate 7

Known distribution of Trichosternus in tropical Queensland. The finely
dotted line is the approximate eastern edge of high land (Atherton Table-
land etc.). Arrows indicate occurrence southward, at increasing distances
beyond the limits of the map, of the species indicated. The species are num-
bered in the order in which they are treated in the text. No. 1, on Mt.
B(artle) F(rere), is Trichosternus fax ; 2, obscuripennis , which extends north
beyond the limits of the map nearly to Cooktown; 3, montorum ; 4, nudipes,
on the Mt. Spec plateau ; 5, soror ; 6, f rater, which occurs north to Mt. Lewis
and south beyond the limits of the map on the Kirrama Range; 7, mutatus ;
8, fisheri; 9, eungella, and 10, mixtus, both on the Eungella Range; 11, corda-
tus, at the southern edge of the tropics; 12, spec, on the Mt. Spec plateau;
13, setosiceps , which is widely distributed on the south-central Atherton Table-
land and occurs also south of the limits of the map on the Kirrama Range ;
and 14, kirrama, on the Kirrama Range. Nos. 1, 2, 3, 13, 14 are very distinct,
phylogenetically isolated species. Nos. 4, 5, 6, 7, 8 are apparently interrelated,
chiefly allopatric members of the frater group (see text). Nos. 9, 10, both on
the Eungella Range, though structurally distinct, may be related to each other
and may hybridize (see text). No. 11 represents a primarily south temperate
rather than tropical stock. No. 12, on the Mt. Spec plateau north of Towns-
ville, is apparently related to another south temperate (New South Wales)
species.

1 1 6

Psyche

[December

523-530). I have specimens of all the 16 full species named in this
key except cyaneotinctus.

1.

2.

3.

4-

5-

6.

7-

8.

9-

10.

11.

Tentative key to Trichosternus of eastern
Australia south of tropics

Elytral intervals 3, 5, 7 with seta-bearing punctures (Mt. Royal
Range and highest part of Dorrigo-Ebor plateau; 2 species?)

australicus SI.

Only 3rd interval (if any) with seta-bearing punctures on elytral

disc 2

Mesosternum setose anteriorly 3

Mesosternum not setose anteriorly 7

Male front tarsi not dilated, not squamulose 4

Male front tarsi dilated, 3 segments squamulose below 5

Alternate discal elytral intervals scarcely differentiated ; greenish
(extreme SE. Q. & adjacent N. S. W. border) .... subvirens Chd.
Alternate discal intervals slightly elevated ; black or nearly so,
with cupreous or greenish margins (Bunya Mts., S. Q.)

sim plicipes SI.

Seventh elytral intervals sharply carinate at base ; prothorax rela-
tively broad at base; length c. 22 mm. (extreme NE. N. S. W.

and adjacent Q. border) angulosus Chd.

Seventh intervals roundly costate at base 6

Very large, c. 33-40 mm. (SE. Q. to Blackall and Bunya Mts.
etc.) renardi Chd.

Not so large, c. 25-32 mm. (N. S. W. S. to Dorrigo, Williams
River, Booral, etc., and N. to Stanthorpe-Dalveen area, S.Q.)
vigor si Gory

Three supra-ocular setae each side 8

Two supra-ocular setae each side 9

Purplish (savannah woodlands of N. N. S. W. chiefly west of

the rain forest zone) superbus Cast.

Greenish (unknown to me; may be another savannah woodland

species) cyaneotinctus Boisd. inch atroviridis SI.

Additional lateral prothoracic setae present (savannah woodland
and drier rain forest of S.Q., N. into edge of tropics; probably

includes he cate Tsch.) cordatus Chd.

Only usual 2 pairs lateral prothoracic setae present 10

Very large, 34 mm. or more 1 1

Smaller, 34 mm. (maximum size of cyaneus only) or less 12

Prosternal process setose; black; length 37-42 mm. (extreme NE.
N. S. W. and adjacent Q. border) perator Sh

1961]

Darlington — A ustralian Carabid Beetles

1 17

Prosternal process not setose; purplish; length 34-39 mm.

(Blackall Range, SE. Q.) porphyriacus SI.

12. Posterior-lateral prothoracic setae near base .... at least 3 species:
wilsoni Cast., nitidicollis Cast., and speciosus SI.
Posterior-lateral prothoracic setae about % of prothoracic length
before base 13

13. Smaller, length c. 21-27 mm.; black sometimes slightly bluish or
purplish (widely distributed in NE. N. S. W. and S. Q., from

Dorrigo to Blackall Range marginiferus Chd.

Larger, length c. 28-34 mm. ; wholly purple or blue ( Mt. Royal
Range, Dorrigo, etc.; possibly 2 species) cyaneus Cast.

The following key to tropical Australian Trichosternus differs from
the preceding key in form (ranges and authorities omitted) because
the tropical species are discussed individually. All the species have
rather long, strong, curved mandibles; rather short, irregular, sub-
parallel frontal foveae; well developed, emarginate mentum tooth;
palpi not strikingly modified; elytra margined at base; 8th (submar-
ginal) stria lined with very small ocellate punctures; lower surface im-
punctate or nearly so except for usual “fixed” punctures; and (so far
as my observations go) other usual characters of the genus, except as
otherwise stated.

Key to Species of Trichosternus of
Tropical Australia

1. Mesosternum setose anteriorly; dorsal elytral intervals equal or

nearly so (except in m onto rum) 2

Mesosternum not setose anteriorly; alternate elytral intervals
elevated (except scarcely so in kirrama) 9

2. Seventh intei vals of elytra not sharply carinate at base (at most

somewhat swollen) ; elytra without dorsal punctures 3

Seventh intervals sharply carinate at base; elytra with (small)
dorsal punctures 4

3. Subparallel, prothorax quadrate; humeri subdentate; color blue-

Pul'Ple fax

Elytia oval, prothorax subcordate; humeri not dentate; black,
elytia dull obscuripennis

4. Elytra with alternate intervals moderately raised; (form de-

piessed , coloi gieenish or bluish) montorum

Elytra with discal intervals equal or nearly so, except at extreme
base (f rater group) -

5- Male front tarsi not dilated, without squamules 6

Male front tarsi more or less dilated, always with some squa-

Psyche

[December

6.

7-

8.

10.

1 1.

12.

13.

mules below 7

Broader; greenish nudipes

Narrower; bluish or purplish soror

Greenish f rater

Black or purplish 8

Black or faintly purple; smaller (18-20 mm.) mutatus

Purple; larger (23-25 mm.) fisheri

Only 2 pairs supra-ocular and 2 pairs lateral prothoracic setae

(but extra apical ventral setae) present eungella

Extra supra-ocular and/or lateral prothoracic setae present ... 10

Prosternal process setose; if prosternal setae broken or not visible,
note 3 supra-ocular and 4 or more median-lateral prothoracic

setae each side, and odd elytral intervals costate mixtns

Prosternal process not setose 1 1

Only 2 pairs supra-ocular setae present, and odd elytral intervals

raised 12

Extra supra-ocular setae present, or odd elytral intervals (except

7th) scarcely raised and humeri scarcely dentate 13

Black cordatus

Purple spec

Only 1 pair median-lateral prothoracic setae present; odd elytral

intervals costate setosiceps

Extra lateral prothoracic setae present; odd elytral intervals (ex-
cept 7th) not costate Hr rani a

Trichosternus fax new species

Form as figured (Fig. 1), subparallel, not strongly convex; black,
head slightly metallic, pronotum with green or bluish marginal chan-
nels and copper-purple on disc brightest in baso-lateral areas, elytra
with marginal channels blue or greenish blue; pronotum shining, head
and elytra slightly duller. Plead c. 3/4 or slightly more width pro-
thorax ; eyes moderate, genae c. wide as eyes, rounded and somewhat '
narrowed to neck; antennae rather short (in genus), passing base of
prothorax by 1 or 2 segments; 2 supra-ocular setae each side. Pro-
thorax subquadrate, c. 1/3 wider than long at middle, not much nar-
rowed behind; base c. 1/10 or more wider than apex (variable);
apex broadly emarginate but angles not otherwise advanced, not mar-
gined; base subtruncate, slightly emarginate at middle, strongly mar-
gined at sides; sides weakly arcuate for much of length, broadly,
slightly sinuate before c. right, well defined posterior angles; lateral
margins narrow, slightly wider posteriorly, each with a seta about
1/3 from apex and on margin at basal angle; disc rather flat, middle

1961]

Darlington — A ustralian Carabid Beetles

I io

kirrama n. sp.

line deep, transverse impressions almost obsolete except for subbasal
foveae each side midway between middle and sides, separated from
margins by strong convexities ; disc and foveae not punctate. Elytra
c. 1/5 wider than prothorax, usually slightly narrowed anteriorly;
margin rectangular or almost acute at subdentate humeri ; elytra with
striae well impressed and punctate; intervals slightly convex but not
costate, 3, 5, and 7 slightly wider than others but not more elevated,
except 7th slightly elevated at base; narrow 10th interval indicated
posteriorly; dorsal punctures lacking. Mesosternum anteriorly and
prosternal process setose. Male front tarsi slightly dilated, 3 segments
squamulose; cT with 1, 9 2 setae each side last ventral segment.
Length 20-23; width 6.5-7. 7 mm-

Holotype cf (M.C. Z. Type No. 30,355) and 15 paratypes all
from Mt. Bartle Frere, North Queensland, west slope 3000-5000 ft.
(not found lower), Dec. 1957, collected by my wife, my son, and
myself, in mountain rain forest.

120

Psyche

[December

This new species is sufficiently distinguished from others in the
preceding key. It is not closely related to any other known species.
It may prove to be generically distinct from Trichostemus, or it may
be a relict of the ancestral stock of the genus.

Trichostemus obscuripennis (Macl.)

This is the northernmost Trichostemus , and it descends to rela-
tively low altitudes. Macleay (1887, p. 220) described it from the
Mossman River. Sloane had it from Helenvale south of Cooktown.
I have it from Mt. Finnigan south of Cooktown, Thornton Peak
near Daintree, Mt. Lewis near Mossman, and Black Mt. about 20
miles north of Kuranda. Although most of my specimens were taken
at 2000 or 3000 ft. altitude, several were found at or below 1000 ft.
in strips of rain forest along the trail between Thornton Peak and
the Daintree River.

T. obscuripennis is a very distinct species but hardly a separate
subgenus. Its lack of humeral teeth is approached in T. kirrama, and
lack of dorsal elytral punctures is matched by fax, although neither
of these species is close to obscuripennis in other ways.

Trichostemus montorum new species

Form as figured (Fig. 2), rather broad, depressed; black with
bluish, purplish, or sometimes greenish reflections, head and pronotal
disc moderately shining, elytra a little duller. Head about 4/5 width
prothorax ; eyes rather small, genae as wide or wider than eyes, irreg-
ularly arcuate and somewhat narrowed to neck; antennae rather long
(in genus); 2 supra-ocular setae each side. Prothorax cordate; 1/3
(z±=) wider than long at middle; base slightly (zb 1/10) narrower
than apex; sides rounded for about 3/4 of length, then rather strong-
ly sinuate before c . right, scarcely blunted posterior angles; side mar-
gins moderate, not much wider basally, each with a seta about 1/3
from apex and on margin at basal angle; apex not margined, broadly
emarginate but angles not otherwise advanced ; base broadly emar-
ginate at middle, slightly rounded toward sides, vaguely margined at
sides but not at middle ; disc depressed, middle line deep, transverse
impressions weak; baso-lateral foveae elongate but not strictly linear,
slightly nearer margins than middle, not quite reaching middle of
length, densely transversely rugulose, the rugulose areas meeting at
middle posteriorly, with middle of base longitudinally rugulose; disc
otherwise impunctate. Elytra oval, usually slightly narrowed anteri-
orly; margin somewhat obtusely angulate and finely toothed at hu-
meri ; elytra with finely punctate discal striae ; intervals more or less
convex, 3, 5, 7 increasingly elevated, 7th being sharply carinate for

Darlington — Australian Carabid Beetles

121

1961]

most of length (most strongly so near base) ; 10th interval indicated
but not well developed; each 3rd interval with o, 1, or 2 dorsal punc-
tures behind middle (often asymmetrical in number and position).
Mesosternum anteriorly and prosternal process setose. Male front
tarsi slightly dilated, 3 segments squamulose below; <A with 1, $ 2
setae each side last ventral segment. Length 23.5-27; width 7.9-9. 1
mm.

Holotype cf (M. C. Z. Type No. 30,356) and 39 paratypes all
from west slope Mt. Bartle Frere, North Queensland, 3000-5000 ft.
(none lower), Dec. 1957, collected by the Darlingtons, in mountain
rain forest. Also 6, not types, from east side Mt. Bellenden Ker,
3000-4500 ft., Dec. 1957, taken by myself, in mountain rain forest.

This seems to be a typical, rather generalized (primitive?) Trichos-
ternus, without known close relatives. It is of course distinguished
from other species in the preceding key.

Trichosternus nudipes new species

Form about average ; greenish black, the green most distinct at sides
and baso-lateral areas of pronotum and marginal channels of elytra;
head and pronotum shining, elytra duller. Head c. 3/4 (by measure-
ment) width prothorax; eyes small but abrupt, genae c. wide as eyes,
convex, somewhat narrowed to neck; antennae of moderate length; 2
supra-ocular setae each side. Prothorax subcordate, moderately nar-
rowed at base, c. 1/3 (±) wider than long at middle; base slightly
(less than 1/10) wider than apex; apex broadly emarginate with
angles (lateral channels) slightly advanced, not margined; base
broadly emarginate at middle, slightly rounded laterally, margined
laterally; sides broadly rounded for much of length, moderately sinu-
ate near c. right or slightly obtuse, scarcely blunted basal angles ; later-
al margins narrow, not wider basally, each with 1 or sometimes 2
setae each side 1/3 or less from apex and 1 seta on margin at basal
angle; disc not very convex, middle line fine, transverse impressions
weak; baso-lateral impressions deep, c. 1/3 length prothorax, separated
from lateral margins by convexities but extending inward somewhat
vaguely nearly to middle; disc of pronotum impunctate except base
irregularly rugulose. Elytra 1/3 or less wider than prothorax, sub-
oval, slightly narrowed anteriorly ; margins forming c. right or slight-
ly obtuse, finely raised-dentate angles at humeri; dorsal striae very-
fine, finely punctate; each elytron with 7 (including suture) rather
weak but almost entire costae, 7th carinate basally; 10th interval in-
dicated toward apex; 3rd interval usually 2-punctate, the punctures
at or before apical 1 /3 and near apex, but individual punctures some-

122

Psyche

[December

times absent or obscured. Mesosternum anteriorly and prosternal
process (sometimes slightly) setose. Male front tarsi narrow, without
squamules; cf with i, 9 2 setae each side last ventral segment. Length
22.5-26; width 7.9-9. 5 mm.

Holotype $ (M. C. Z. Type No. 30,357) and 87 paratypes all
from Mt. Spec plateau (Paluma Range), c. 40 miles north of Towns-
ville, North Queensland, 2000-3000 ft., Dec. 1957 and Feb. 1958,
taken by the Darlingtons, in rain forest; and 1, same locality, June 7-
10? 1955, E. O. Wilson.

This is similar to and perhaps related to T. subvirens Chd. of South
Queensland, but subvirens has an entire mentum tooth (emarginate
in nudipes) , posterior lateral pronotal setae just inside (not on) the
margin, and lacks or nearly lacks rugulose sculpture on the base of
the pronotum.

Trichosternus soror Dari.

I described this species (1953) from 3 specimens from Millaa Mil-
laa, on the Atherton Tableland. I did not find it at this locality dur-
ing my last trip, but found it on both the lower and upper western
slopes of Mt. Bartle, Frere, at 2000-3500 and 3000-5000 ft., and on
the east side of Mt. Bellenden Ker, at 3000-4500 ft.

Although this small, bluish species has simplified male tarsi, like
the preceding species, it may not be directly related to it but may be
independently derived from a common ancestor perhaps resembling
/ rater (below).

Trichosternus f rater new species

Slightly more slender than average of genus, rather convex; green-
ish or bluish black, head and pronotum moderately shining, elytra
somewhat duller. Head 4/5 or more width prothorax; eyes small,
genae about wide as eyes, rounded and somewhat narrowed to neck;
2 supra-ocular setae each side; antennae of moderate length. Pro-
thorax subcordate, c. 1/4 wider than long at middle; base slightly
narrower than apex; apex broadly enrarginate but angles otherwise
scarcely advanced, apex not margined; base emarginate at middle,
rounded toward sides, margined at sides ; sides broadly arcuate through
much of length, moderately sinuate toward base; basal angles c. right
or slightly obtuse, scarcely blunted ; side margins rather narrow ; not
much wider posteriorly, each with 1 seta near or before anterior 1/3
and 1 on margin at basal angle; disc with distinct middle line, less
distinct transverse impressions, and moderate baso-lateral foveae which
reach posterior margin but are separated from lateral margins by con-
vexities; surface of disc nearly smooth except with some (variable)

1961]

Darlington — Australian Carabid Beetles

12 3

transverse strigulation in baso-lateral impressions. Elytra slightly
more than 1/4 wider than prothorax, subparallel, slightly narrowed
anteriorly; margin forming obtuse (nearly right), slightly dentate
angles at humeri ; dorsal striae formed by fine vaguely punctulate im-
pressed lines in low spaces between intervals; latter subequal on disc,
slightly raised but not costate, except 7th strongly raised and costate
basally, 10th interval indicated posteriorly; each 3rd interval 1- or
2-punctate posteriorly, position of punctures variable. Mesosternum
anteriorly and prosternal process (sometimes inconspicuously) setose.
Male front tarsi moderately dilated, 3 segments squamulose below;
cf with 1, $ 2 setae each side apex last ventral segment. Length
20-23 ; width 6. 5-8.0 mm.

Holotype cf (M. C. Z. Type No. 30,358) and 73 paratypes all
from Kirrama Range, inland from Cardwell, North Queensland,
2000-3000 ft. altitude, Dec. 1957 and Feb. 1958, collected by the
Darlingtons, in rain forest. Also the following specimens, not types:
12, mountains above (SW of) Atherton, Atherton Tableland, 3000-
4000 ft., Dec. 1957 and Feb. 1958; 1, near Black Mt., c. 20 miles
north of Kuranda, probably near 2000 ft. altitude, May 1958; and
3, Mt. Lewis, near Mossman, probably near 3000 ft., Dec. 1957; all
collected by the Darlingtons, in rain forest. These localities are all
on the Dividing Range system, which has probably been the principal
path of north-south dispersal of wet forest Carabidae in North
Queensland.

T. frater may represent the ancestral stock, with dilated, squamu-
lose male tarsi, from which nudipes of the Mt. Spec plateau, subvirens
and simplicipes of South Queensland, and also (independently) soror
of the Atherton Tableland etc. have been derived. These 5 species
(including frater itself) are all allopatric and are generally similar
in form and technical characters except for the differences in the cf
tarsi. The two following new species may be local derivatives of the
same stock with cf tarsi still dilated and squamulose. See map for
distribution of tropical species of the frater group.

Trichosternus mutatus new~ species

Small; form about average for genus but rather convex; black,
sometimes with faint purplish reflections; moderately shining, elytra
slightly duller. Head 3/4 to 4/5 width prothorax; antennae rather
short, scarcely extending beyond base of prothorax; eyes moderate,
genae c. wide as eyes, rounded and narrowed to neck; 2 supra-ocular
setae each side. Prothorax subcordate, 1/4 (db) wider than long at
middle; base equal to or slightly narrower than apex; sides broadly

124

Psyche

[December

but not strongly rounded in much of length, weakly sinuate before
base; basal angles c. right or slightly obtuse, scarcely blunted; apex
subtruncate or very broadly emarginate with angles scarcely advanced,
apex not margined ; base subtruncate or slightly emarginate at middle,
slightly rounded-oblique toward sides, weakly margined only toward
angles; lateral margins narrow, scarcely wider basally, each with i
seta about 1/4 from apex and 1 on margin at basal angle; disc con-
vex, middle line well impressed, transverse impressions rather weak;
baso-lateral foveae rather weak and irregular, reaching posterior
margin but separated from side margins by convex areas, not punctate
and not much more rugulose than disc. Elytra slightly more than 1 /4
wider than prothorax, rather short, slightly narrowed anteriorly; hu-
meral angles nearly right, slightly toothed; striae represented by de-
pressed spaces between intervals; latter subequal on disc, convex but
not much elevated except 7th elevated and costate basally; 10th inter-
val indicated posteriorly; each 3rd interval with 2 or 3 dorsal punc-
tures posteriorly. Mesosternum anteriorly and prosternal process
(sometimes inconspicuously) setose. Male front tarsi moderately
dilated, 3 segments squamulose; c? with 1, $ 2 setae each side apex
last ventral segment. Length 18-20; width 6. 5-7.0 mm.

Holotype <$ (M.C. Z. Type No. 30,359) and 4 paratypes from
Mt. Fisher (see below), c. 4000 ft., Feb. 1958; and 5 additional para-
types from mountains above (SW of) Millaa Millaa, c. 3500 ft.,
Dec. 1957 ; all specimens taken by the Darlingtons, in rain forest. The
two localities are close together in the mountain mass between Raven-
shoe and Millaa Millaa on the Atherton Tableland, North Queens-
land.

This species differs from the preceding one (f rater) in smaller size,
color, smoother basal pronotal foveae, and other details. It differs
from soror in form of c? tarsi, in being a little broader, different in
color, and in other details. For further comments, see under following
species.

Trichosternus fisheri new species

Larger than preceding; black with strong purplish reflections, mod-
erately shining except elytra slightly duller. Head slightly more than
4/5 width prothorax; eyes moderate, genae c. wide as eyes, rounded
and narrowed to neck; antennae rather long (in genus), passing base
of prothorax by 2 or 3 segments; 2 supra-ocular setae each side. Pro-
thorax subcordate, c. 1/4 or less wider than long at middle; base c.
1/10 narrower than apex; apex subtruncate or very broadly emargin-
ate, with angles scarcely advanced; apex not distinctly margined; base

1961]

Darlington — Australian Carabid Beetles

125

slightly emarginate at middle, faintly rounded toward sides, rather
vaguely margined at sides; sides rather weakly rounded for much of
length ; moderately sinuate posteriorly before c. right, scarcely blunted
basal angles ; margins rather narrow, not much wider posteriorly ; each
with a seta about 1/3 from apex and on margin at basal angle; disc
convex, with fine middle line and weak transverse impressions ; baso-
lateral foveae deep but irregular, more or less transversely striolate;
surface of disc otherwise nearly smooth, except some striae radiating
from a point near base. Elytra nearly 2/5 wider than prothorax,
slightly narrowed anteriorly; margin forming strong but slightly
obtuse, slightly toothed angles at humeri; striae vaguely punctate in
clean specimens; intervals slightly raised but not strongly costate on
disc, subequal except 3rd and 5th slightly more prominent at base and
7th carinate especially basally; 10th interval scarcely indicated; 3rd
interval with usually 4 to 6 punctures, irregularly spaced. Mesoster-
num anteriorly and prosternal process (sometimes inconspicuously)
setose. Male front tarsi only slightly dilated but usually with some
squamae on 1st three segments (sometimes visible only on first seg-
ment — it is not clear whether they are worn off or originally missing
on other segments in this case) ; cf with 1, $ 2 setae each side apex
last ventral segment. Length 23-25; width 8. 1-8.9 mm.

Holotype cf (M. C. Z. Type No. 30,360) and 3 paratypes from
Mt. Fisher (see below), c. 4000 ft., Feb. 1958; and 2 paratypes from
mountains above (SW of) Millaa Millaa, c. 3500 ft., Dec. 1957; all
specimens taken by the Darlingtons, in rain forest. Both these locali-
ties are in the mountain system between Ravenshoe and Millaa Millaa,
on the Atherton Tableland, North Queensland.

This species differs from the 2 preceding ones (in different ways)
in details of form and color and size, in having cf tarsi less dilated,
and in having more punctures on 3rd elytral intervals. It and mutatus
appear to be confined to the same local mountain mass. Their occur-
rence together, localized between the ranges of frater on one side and
soror on the other (see map), suggests some sort of double invasion
or hybridization, although all the species in question now seem fully
distinct.

Trichosternus eungella new species

Rather small, broad, depressed ; black, virtually without metallic
reflections, moderately shining, elytra (except marginal intervals and
tops of costae) dull. Head 3/4 (zb) width prothorax; eyes moderate,
genae wide as or slightly wider than eyes, rounded and narrowed to
neck; antennae rather long (in genus), passing base of prothorax by

126

Psyche

[December

about 2 segments; 2 supra-ocular setae each side. Prothorax sub-
cordate, c. 2/5 or slightly more wider than long at middle; base
slightly narrower than apex; apex broadly emarginate with angles
slightly advanced, not margined; base broadly emarginate at middle
subtruncate at sides; sides broadly rounded through much of length,
rather strongly sinuate basally ; posterior angles right or slightly acute,
scarcely blunted; side margins moderate, not much broader basally,
each with a seta about 1/3 from apex and on margin at or very near
basal angle (additional setae rarely present anteriorly) ; disc with
well impressed middle line and anterior transverse impression, weak
posterior transverse impression but large irregular baso-lateral im-
pressions; latter strongly transversely strigulose. Elytra 1/5 (dz)
wider than prothorax, slightly narrowed anteriorly; margin forming
c. right (slightly obtuse) dentate angles at humeri; discal striae in
clean specimens indicated by rows of rather vague punctures; even
discal intervals scarcely convex, odd intervals costate; no distinct 10th
interval; each 3rd interval usually 2-punctate, behind middle and on
declivity. Mesosternum and prosternal process not setose. Male with
front tarsi slightly dilated, 3 segments squamulose; c? with 2 or 3,
9 with 3 or 4 (often asymmetrical) setae each side last ventral seg-
ment. Length 20-25 ; width 7. 8-8. 3 mm.

Holotype cf (M. C. Z. Type No. 30,361) and 53 paratypes all
from the Eungella Range, west of Mackay, Queensland, 2000-3000
ft. altitude, Nov. 1957, taken by the Darlingtons, in rain forest.

This is apparently related to T. wilsoni Cast, of South Queensland
etc., but eungella is slightly broader and more depressed, with stronger
elytral costae. For possible hybrids of this and following species, see
under latter.

Triehosternus mixtus new species

Form about average for genus, but prothorax strongly cordate and
elytra oval; black without metallic color, head and pronotum moder-
ately shining, elytra duller. Head large, 4/5 width prothorax; eyes
moderate, genae c. wide as eyes, rounded and somewhat narrowed to
neck; antennae rather long, passing base of prothorax by about 2 or
more segments; usually 3 supra-ocular setae each side (only 2 on
right side in one specimen, 4 on both sides in another). Prothorax c.
2/5 (=t=) wider than long at middle; base c. 4/5 or slightly more
width apex; apex strongly emarginate, angles (marginal channels)
advanced in arc of emargination ; apex not margined ; base slightly
emarginate, finely margined; sides rounded for much of length, strong-
ly sinuate about 1/6 of length before base, then subparallel to base;

1961]

Darlington — Australian Carabid Beetles

127

posterior angles right or slightly acute, scarcely blunted ; side margins
moderate anteriorly, slightly wider posteriorly, each with 4 to 9 setae
near and before middle (at least 4 on each side in all specimens) and

1 seta posteriorly well before angle; disc with usual middle line, weak
transverse impressions, and rather weak baso-lateral foveae separated
from base by transverse elevation ; foveae strongly transversely strigu-
lose, and base longitudinally strigulose. Elytra c. 1/4 wider than pro-
thorax; margin forming c. right or slightly obtuse, dentate angles at
humeri; striae in form of lines of small punctures (often obscured) ;
even intervals scarcely elevated but intervals 3, 5, and 7 carinate,
carinae of 3 and 5 not reaching base but 7th very strongly carinate
basally; no 10th interval; each 3rd interval 2-punctate posteriorly.
Mesosternum without but prosternal process with setae (all specimens
— most setae broken off in 2 specimens but vestiges visible). Male
front tarsi moderately dilated, 3 segments squamulose ; c? with usually

2 or 3, sometimes 4 (often asymmetrical), $ with usually 4, some-
times 5 setae each side last ventral segment. Length 24-29; width
8. 4-9. 5 mm.

Holotype cf (M. C. Z. Type No. 30,362) and 23 paratypes all
from Eungella Range, west of Mackay, Queensland, 2000-3000 ft.,
Nov. 1957, taken by the Darlingtons, in rain forest.

Superficially, mixtus is almost a miniature of cordatus but has extra
supra-ocular setae, posterior pronotal setae farther from base, and a
setose prosternal process. From eungella, with which it occurs, mixtus
differs in form, extra supra-ocular and lateral prothoracic setae, posi-
tion of posterior pronotal setae, and setose prosternal process. Most
individuals of both species are unquestionably distinct. However, I
have 5 specimens from the Eungella Range that have intermediate
characters and that may be hybrids. They are intermediate in form ;
like eungella in having only 2 pairs of supra-ocular setae; intermediate
in lateral pronotal setae (2 or 3 each side at and before middle, but
with posterior setae near base) ; and like eungella in having prosternal
process not setose. Only field study can determine the real status ^
these specimens.

Trichosternus cordatus Chd.

Although cordatus is primarily a South Queensland species, it ex-
tends into the tropics. It lives in drier woodland than most other
members of the genus except superbus Cast. I found it at Beechmont
(Queensland-New South Wales border) ; Benarkin ; Yarraman; Big-
genden (SW of) ; near Monto (in dry scrubby woods with Pamborus

128

Psyche

[December

viridis) ; and, in the tropics, at Byfield (N. of Yepoon) in semi-rain
forest, and c. 40 miles S. of Mackay in good savannah woodland.

Trichosternus spec new species

Form nearly of T . superbus, with strongly cordate prothorax and
oval elytra narrowed anteriorly; purple or purplish, not strongly shin-
ing, elytra slightly duller. Head large, slightly more than 4/5 width
prothorax; eyes small, genae slightly wider than eyes, subparallel an-
teriorly, rounded and slightly narrowed to neck ; antennae rather long,
passing base of prothorax by about 2 segments; 2 supra-ocular setae
each side (all specimens). Prothorax c. 2/5 or less wider than long
at middle; base slightly more than 3/4 width apex; apex broadly
emarginate except rounded toward sides, with angles (marginal gut-
ters) slightly advanced; apex not margined; base broadly emarginate
except oblique near angles, rather vaguely margined ; sides rounded
through much of length, strongly sinuate before basal angles; latter
right or slightly obtuse, not much blunted ; lateral margins rather
narrow anteriorly, broader posteriorly, each with several (usually 4
or 5) setae at and before middle and 1 before basal angle; disc with
middle line distinct, transverse impressions weak; baso-lateral foveae
rather deep but vaguely limited, with weak convexities toward lateral
margins, and closely transversely strigulose, the strigulations extending
onto disc across base. Elytra c. 1/4 wider than prothorax, widest
behind middle; forming c. right (slightly obtuse), slightly dentate
angles at humeri; striae indicated by weakly punctate impressions;
even intervals slightly convex, odd ones 3, 5, and 7 subcostate to
costate on disc, 7th strongest and sharply carinate anteriorly; no 10th
interval; each 3rd interval usually 2-punctate, with punctures behind
middle and on declivity. Mesosternum and prosternal process not
setose. Male front tarsi slightly dilated, 3 segments squamulose.
Length 24-30 ; width 8.5-10.3 mm.

Holotype d (M. C. Z. Type No. 30,363) and 33 paratypes all
from the Mt. Spec plateau (Paluma Range), c. 40 miles north of
Townsville, North Queensland, 2000-3000 ft., Nov. 1957 and Feb.
1958, taken by the Darlingtons in or on the edges of rain forest.

In spite of their wide geographical separation, this species and
super bus Cast, (of New South Wales) may represent one original
stock. The new species resembles superbus in form, color, and many
details, but differs from it in having only 2 (instead of 3) supra-
ocular setae on each side and in having better defined posterior pro-
thoracic angles.

1961]

Darlington — Australian Carabid Beetles

129

Trichosternus setosiceps SI.

Sloane (1923, p. 23) described setosiceps from the South Johnstone
River and Malanda, which are on or near the Atherton Tableland.
The species is widely distributed on the Tableland, except apparently
the northern part, and reaches the lower slopes of Mt. Bellenden Ker
and extends south to the Kirrama Range.

This species differs from all other Trichosternus in having 2 seg-
ments of each male front tarsus squamulose.

Trichosternus kirrama new species

Form as figured (Fig. 3), large, with strongly cordate prothorax
and oval elytra; black, head and prothorax moderately shining, elytra
dull. Head large, 4/5 or slightly more width prothorax; eyes small,
genae slightly wider than eyes, broadly rounded and narrowed pos-
teriorly to neck; antennae rather long, passing base of prothorax by
c. 2 segments; 2 or 3 supra-ocular setae each side (often asymmetri-
cal) ; frontal foveae as usual except area between them slightly de-
pressed and/or with longitudinal prominences outside them. Prothor-
ax strongly cordate, nearly 1/2 wider than long at middle; base c.
4/5 (±) width of apex; apex emarginate at middle, rounded toward
sides, with angles (lateral channels) advanced; apex not margined;
base broadly emarginate except subtruncate near angles, rather irregu-
lar, vaguely margined ; sides rounded through much of length, strongly
sinuate about 1/6 before base, thence subparallel to c. right, well de-
fined posterior angles; lateral margins moderate anteriorly, slightly
wider posteriorly, each 2 to 4 (rarely 5) setae at and before middle
and one a little before base; disc with usual middle line, transverse
impressions weak ; baso-lateral impressions irregular, transversely strig-
ulose, the strigulose area extending forward at sides and across base;
extreme basal strigae longitudinal. Elytra 1/5 or slightly more wider
than prothorax; margin somewhat obtusely angulate but only vaguely
toothed at humeri ; striae formed by lines of small punctures ; even
intervals slightly convex, odd intervals 3 and 5 slightly more raised
but not costate, interval 7 strongly raised, costate at base; no 10th
interval; each 3rd interval 2-punctate, behind middle and on declivity.
Mesosternum and prosternal process not setose. Male front tarsi
moderately dilated, 3 segments squamulose; 2 or 3 (often asymmetri-
cal) setae each side last ventral segment in both sexes. Length 30-33;
width 9.9-10.6 mm.

Holotype cf (M.C. Z. Type No. 30,364) and 11 paratypes all
from Kirrama Range, near Cardwell, North Queensland, 2000-3000
ft., Dec. 1957 and Feb. 1958, taken by the Darlingtons, in rain forest.

In form, appearance, and some characters (near obliteration of

130

Psyche

[December

humeral teeth, lowness of elytral intervals) this species approaches
obscuripeimis, but the arrangement of setae is very different. I do
not know the real relationships of either species.

References

Britton, E. B.

1940. The Carabidae (Coleoptera) of New Zealand, Part I, Ptero-
stichini. Trans. R. Soc. New Zealand, 69: 473-508.

Darlington, P. J., Jr.

1953. Australian carabid beetles II. Some new Pterostichini. Psyche,
60: 90-101.

1961a. Australian carabid beetles IV. List of localities, 1956-1958. Psyche,
67: 111-126.

1961b. Australian carabid beetles V. Transition of wet forest faunas
from New Guinea to Tasmania. Psyche, 68: 1-24.

1961c. Australian carabid beetles VI. The tropical and some subtropical
species of Pamborus, Mystropomus , and Nurus. Breviora, No.
142: 1-13.

Macleay, W.

1887. [7\ obscuripennis.'] Proc. Linn. Soc. New South Wales, ser. 2,

2: 220.

Sloane, T. G.

1894. [ Homalosoma , inch Trichosternus.] Proc. Linn. Soc. New South

Wales, ser. 2, 9: 417-425.

1899. \Homalosoma, inch Trichosternus. ] Proc. Linn. Soc. New South
Wales, 24: 567-572.

1923. [7\ setosiceps.] Proc. Linn. Soc. New South Wales, 48: 23.

Tschitscherine, T.

1902. [ Trichosternus .] Horae Soc. Ent. Rossicae, 35: 520-534.

ADRITYLA, A NEW MILLIPED GENUS
(CHORDEUMIDEA: CONOTYLIDAE)

By Nell B. Causey1
Introduction

This is the second of a scries of papers on the widely dissimilar
North American species formerly assigned to the genus Conotyla.
In the first (Causey, 1961), the genus Austrotyla was proposed for
coloradensis, humerosa , montivaga , and specus. The remaining species
will be reviewed and the basal region of the gonopods described in
subsequent papers.

A grant from the National Science Foundation (G- 14486) made
the completion of this paper possible.

I am grateful to Dr. Herbert W. Levi for the loan of the speci-
mens of Conotyla deseretae in the Museum of Comparative Zoology.

Genus Adrityla, new

Type species. Conotyla deseretae Chamberlin, 1910, by monotypy.

Diagnosis. Large bodied (length to 24 mm.), pigmented conoty-
linids of 30 body segments, distinguished especially by the modification
of legpair 10 (81), of which the coxa is greatly enlarged and lobate
and the telepodite is reduced to 2 or 3 minute segments. The sternum
of the anterior gonopods is divided, and the small sternites are fused
to the base of the gonopods. The latter are unsegmented and larger
than the coxites of the posterior gonopods.

Relationships. The inclusion of this genus in the subfamily Cono-
tylinae is justified by the following sexual characters of the male:
the unsegmented anterior gonopods, the retention of a division between
the coxal and prefemoral segments of the posterior gonopods, and the
absence of coxal glands on legpair 11 (82). The divided sternum of
the anterior gonopods suggests a close relationship with the Anstrotyla-
Taiyutyla line rather than with Conotyla , which has this sternum
undivided. The remarkable modification of legpair 10 surely occurred
independently in Adrityla, as it has in other widely separated
chordeumids.

Distribution. Utah.

Adrityla deseretae (Chamberlin), new combination
Figures 1-4

Conotyla deseretae Chamberlin, 1910, Ann. Ent. Soc. Amer. 3 (4) : 235-236, pi.

31, figs. 3-8; pi. 32, figs. 1-7. Loomis, 1943, Bull. Mus. Comp. Zool. Harvard

92(7): 381. Chamberlin and Hoffman, 1958, U. S. Nat. Mus. Bull. 212:98.

Tayetteville, Arkansas, and Gulf Coast Research Laboratory, Ocean
Springs, Mississippi.

132

Psyche

[December

Types. Present location of male holotype unknown. Two larvae,
undoubtedly metatypes, from Mill Creek Canyon, Salt Lake Co.,
Utah, and my plesiotypes (cf, 9) from South Willow Canyon,
Tooele Co., Utah, are in the Museum of Comparative Zoology.

Figures 1, 2. Adrityla deseretae (Chamberlin). 1. Segments 10 and 11,
dorsal view. 2. Left anterior gonopod, anterior view.

Type locality. The Wasatch Mountains, Salt Lake Co. and Utah
Co., Utah.

Range. The Wasatch Mountains in central and northern Utah
(Chamberlin and Hoffman, 1958).

Description. Length 22 to 24 mm., width 2.6 to 3 mm., the cf
larger than the 9- Body reaches its greatest width at segment 8 or 9.
Dorsum with a dark brown medial longitudinal band ; pleura with a
brown longitudinal band along the lower surface of the paranota;
venter, legs, and remainder of dorsum light brown to yellowish;
antennae dark brown. Ratio of length and width of collum approxi-
mately 34. Paranota with the typical conotylinid shape and position
are on segments 2 through 25 (Fig- 0- Segmental setae acute; on
typical segments the external setae are markedly longest, about 0.5
mm., and the medial and internal (nearest the dorsomedial furrow)

1961]

Causey — Adrityla

133

setae are about 0.35 mm.; the relative distances between them are
as follow:

med. seta — ext. seta 1 dorsal furrow — int. seta 1

med. seta — int. seta 3 int. seta — med. seta %

Mentum undivided. Clypeus, frons, and vertex with the usual
short, unequal setae. Eyes triangular, from 23 to 25 black ocelli
arranged compactly, but not touching, in rows of 1, 7, 6, 5, 4 (3),
2 ( 1 ) , Antennae slender, the length approximately 4.3 mm. in the cf
and 3.3 in the 9*

In the male, legpairs 1 and 2 are about two-thirds the length and
thickness of the following legs ; legpairs 3 through 7 are shorter and
thicker than the postgonopodal legs and segment 4 bears an incon-
spicuous rounded protuberance on the mesioproximal surface; the
ventral surface of the coxa of legpair 7 is conspicuously swollen ; the
ventral surface of the tarsus of all legs from the third to about the
middle of the body is pulvillar; legpair 10 (81) is highly modified by
the enlargement of the coxa and the coxal sac and by the reduction
of the telopodite; and legpair 11 (82) has the coxa slightly swollen
but lacks a coxal sac or coxal perforation ; the ectoventral surface of
segment 2 of legpair 1 1 bears a small, rounded, caudally directed lobe.

The anterior gonopod (Fig. 2) is an unsegmented, rectangular
piece irregularly serrated on the ventral margin and directed down
and slightly forward. In the coxosternal region it is contiguous with
its homologue along a movable suture. The spiracle opens into a
large depression near the anterior margin of the coxosternum.

The posterior gonopod (Figs. 3 and 4) consists of the usual three
segments : 1 ) a coxa with two elongated coxites, the mesial one in-
conspicuous and narrowly attenuated, and the more lateral one longer,
stouter, clavate, warty on the apical surface, and covered distad by
the larger coxite of legpair 10; 2) a slender prefemur (one-third the
length of the femur) with a setose lobe on the mesiodistal surface;
and 3) a large oval femur (the ratio of the length to the thickness
is 1/3) with an irregular pigmented pit on the caudal surface near
the apex. The sternum is thin in the midline and bent sharply ventrad ;
laterad it is fused with the coxae and thickened ; the spiracles open
from large pigmented pits near the ectocephalic angles.

Feg 10 consists of two regions: 1) an enlarged coxosternum with
one massive coxite that curves back over leg 1 1 and covers the apical
region of the larger of the two coxites of the posterior gonopod, and
2) a minute telopodite of 2 or 3 unequal segments that cannot be

134

Psyche

[December

1961]

Causey — A drityla

135

seen unless leg 1 1 is removed. The coxite is elongated, broadened
and rounded distad, smooth on the anterior surface, and thickly setose
over part of the more complex posterior surface; a subapical flange
is on the ectal surface, and near the base of the mesial surface is a
large perforation through which the enlarged, soft coxal sac projects.
The coxal sac is contiguous with its homologue in the midline. The
sternum is divided and coalesced with the coxa, forming a thick, short
coxosternum which is contiguous with its homologue along a movable
suture; the inconspicuous spiracles open on the ectal surface of the
coxosternum.

Appendages 72 and 81 (the posterior gonopods and legpair 10,
respectively) fit close together, but there is no fusion, and with
careful dissection they can be completely separated.

In the female, the setae on the vulvae are unusually long. The
legs are typical of the family.

Emendations. In the original description, Chamberlin (1910, p.
236) incorrectly interpreted the coxites of legpair 10 (81) as part
of the) anterior (71) and posterior (72) gonopods. The correct legends
of plate: 31, figures 4-7 are: figure 4, legpair 10, caudal aspect; fig-
ure 5, anterior gonopods and parts of posterior gonopods and legpair
10, cephalic aspect; figure 6, left gonopods and tenth leg, lateral
aspect; posterior gonopods, cephalic aspects. Only legpairs 4 and 5
were represented (PL 32, Figs. 2-6) as having a small lobe on the
mesioproximal surface of segment 4. I have found that legpairs 3
through 7 have segment 4 similarly modified ; possibly there is some
variation of this character. A large promentum was indicated in
plate 32, figure 7. There is no promentum; the small triangular
space anterior to the mentum is covered only by a membrane.

Variations. The ventral margin of the anterior gonopods is irreg-
ularly serrated in each of the 4 male specimens that I have examined.
The serrations vary between 3 and 14 or 15. A possible variation in
the lobation of the pregonopodal legs was mentioned in the preceding
paragraph.

Material Examined. UTAH : Mill Creek Canyon, Salt Lake Co.,
larvae (MCZ) [? metatypes]; South Willow Canyon, Tooele Co.,

Figures 3, 4. A drityla deseretae (Chamberlin). 3. Posterior gonopods and
left tenth leg, anterior view. 4. Left posterior gonopod and legpair 10, poste-
rior view. (Abbreviations: A71, A72, A81, sternal apodemes of anterior
gonopod, posterior gonopod, and leg 10, respectively; C72, C81, coxites of
posterior gonopod and leg 10, respectively; F, PF, femur and prefemur of
posterior gonopod, respectively; S71, S72, S81, coxosternites of anterior gono-
pod, posterior gonopod, and leg 10, respectively; T, telopodite of leg 10.)

136

Psyche

[December

July 5, 1953, 1 cf, 1 $, W. D. Hester (MCZ) ; American Fork
Canyon, Timpanogos Park, Utah Co., June 14, 1952, W. D. Hester,
3 cf , 8 $, larvae (NBC).

Literature Cited

Causey, Nell B.

1961. Austrotyla, a new milliped genus (Chordeumidea : Conotylidae:
Conotylinae) . Proc. Biol. See. Wash. 74: 251-266, Figs. 1-10.
Chamberlin, Ralph V.

1910. Diplopoda from the western states. Ann. Ent. Soc. Amer. 3(4):
233-262, pis. 30-43.

Chamberlin, R. V. and R. L. Hoffman.

1958. Checklist of the miliipeds of North America. U. S. Nat. Mus. Bull.

212, p. 212.

ANTS FROM THREE REMOTE OCEANIC ISLANDS
By Robert W. Taylor and Edward O. Wilson
Biological Laboratories, Harvard University

The three islands (Raoul, Clipperton, St. Helena) whose ant
faunas are described below have in common only extreme geographic
isolation. That ants occur on them at all confirms the idea that these
insects, with man’s help, have now populated every part of the earth
capable of supporting them. These and other remote oceanic islands
will undoubtedly attract more of the ecologist’s attention in the
future, since many animal taxa inhabiting them, including most or
all of the ant species, have only arrived within historical times and
present simple case histories of faunas in the first stages of local
adaptation.

We are grateful to Dr. J. S. Edwards, Dr. C. F. Harbinson, Mr.
Arthur Loveridge and Dr. B. A. Holloway of the Dominion
Museum, Wellington, New Zealand, for making these unusual
collections available. The study has been supported in part by a
research grant from the National Science Foundation.

Raoul Island, Kermadec Islands

The Kermadecs are a group of forest-clad volcanic islands lying
in the South Pacific between S. lat., 29.10 and 31.30 ; and W. long.,
177.45 and 179.00. The nearest sizable land mass is the North Island
of New Zealand, about 650 miles to the southwest, and the nearest
major Polynesian island is Tongatabu of the Tongan group, about
7001 miles to the north. Australia lies about 1,700 miles to the west.

The ants listed below were taken on Raoul or Sunday Island, the
largest of the group (11.25 sq. miles), or on its tiny outlier, Meyer
Island. Most of the material was collected during June 1956 for the
University of Auckland, New Zealand, by Dr. J. S. Edwards of
Cambridge University. A few records are from the collections of
the Museum of Comparative Zoology, Harvard University, or from
the Dominion Museum, Wellington, New Zealand. Unless otherwise
noted all collections were made by Dr. Edwards.

The collection includes a number of Berlese funnel samples as well
as hand-collected specimens, and although only three species are listed,
it is thought that they represent most, if not all, those present on
Raoul in 1956.

Ponera gleadowi Forel

The systematics and distribution of this ant have been discussed by
Wilson (1958). P. gleadowi is a widespread species, originally

137

138

Psyche

[December

described from India, and found throughout South East Asia,
Melanesia and Polynesia. It is apparently native throughout most
of its range, but may have been distributed in part by man (i.e., it is
a member of “Class 11“ discussed below) .

Material examined: raoul island: June 5, 1956; under rock,
watercourse, June 8, 1956 (nest with ergatomorphic male pupae) ;
ex Pohutakawa logs ( Metrosideros sp.), June 3, 1956. Crater:
June 4, 1956; ex moss caps on pumice boulders, June 4, 1956; ex
Pohutukawa log, June 4, 1956 (nest with alate queens). Denham
Bay: ex Narcissus litter Berlese funnel sample, June 7, 1956.

Denham Bay swamp : ex Pohutakawa litter Berlese funnel sample,
June 2, 1956. Hostel Bay: Berlese funnel sample, June 7, 1956;
Coxnocarpus litter Berlese funnel sample, June 8, 1956. Oneraki
Beach: beneath cow dung under trees, June 9, 1956 (J. S. Soeberg).
meyer island: Berlese funnel samples, June 7, 1956; June 16,
1956.

P oner a gleadowi is apparently the most abundant ant on Raoul
and was “commonly found (by Dr. Edwards) under stones or in
decaying logs or branches, frequently associated with termites.”
Moreover it was present in about half of the Berlese funnel samples
processed by Dr. Edwards. As indicated, winged queens are present
in the nests during June. No adult males were located, but several
pupae from the nest collected on June 8, 1956, were found to contain
almost fully developed ergatomorphic males. Except for genital
differences these are similar to the workers, and like them have twelve-
segmented antennae.

Tetramorium guineense (Fabricius)

This is one of the most common pantropical formicid tramp species.
It evidently originated in Africa and ranges widely throughout the
Old and New World tropics, and Northern Australia. It is wide-
spread in Polynesia (Wheeler, 1935) and has recently been found
established near Auckland, New Zealand. Dispersal by man has
clearly been of primary importance in the distribution of this species.

Material examined : raoul island: June 4, 1956; quarry, under
rock, June 5, 1956, (nest with alate females), meyer island:
Berlese funnel sample, June 7, 1956.

Monomorium antarcticu?n (White)

At present the name M. antarcticum is applied to what is almost
certainly a complex of species found in New Zealand and its outlying
islands, including the Chatham group, some 470 miles to the east

1961]

Taylor and Wilson — Ants

139

of South Island. It has undoubtedly been carried to the Chatham
Islands by man (Brown, 1958). (It is further likely that
Monomorium rapaense Wheeler (1936) of Rapa Island in the
Austral group, is conspecific with some New Zealand form belonging
to the present M. antarcticum complex. If this should be so the fact
that New Zealand and Rapa were both important whaling ports in
the nineteenth century may be significant with regard to the dispersal
of this form.)

Material examined: raoul island: Crater: nest in warm soil
beside fumarole, June 3, 1956. Four workers without further data
from the M. C. Z. collection.

The M. C. Z. specimens were studied by Wheeler in preparing
his 1935 paper and were probably collected by the W. R. B. Oliver
expedition to Raoul Island during 1907-08. Wheeler apparently
intended using these specimens as the types of a new species, to be
described in a later paper which, however, was never published due
to his death. It seems unlikely that they are specifically separate from
some North Island samples of the antarcticum complex, and they have
almost certainly been introduced to Raoul from New Zealand.

The Kermadec ant fauna is thus characterized by a lack of endemic
or native species and includes forms derived either from tropical
Polynesia, or the nearest large land area, and known to have capacities
for passive dispersal by man. The faunal tally of only three species
is probably well below the possible maximum carrying capacity of the
island (see Wilson, 1961). Although the present human settlement
on Raoul consists only of the staff of a weather station, there have
been several attempts to colowze it in the past. The parties concerned
are known to have travelled from Fiji and New Zealand, and to have
carried much produce as well as plants and seeds, thus providing
adequate passage for ants. In its relationships and depauperate nature
the ant fauna resembles those of other insect groups which have been
studied (see Miller, 1956 p. 420 for references).

Some interesting comparisons can be made with the faunas of Lord
Howe and Norfolk Islands, which are about the same size as Raoul,
and lie at approximately the same latitude, but much closer to
Australia.

The ants of these three islands may be grouped for analysis into
three classes.

Class I. Species which are probably native to the island concerned.
This category includes forms found also in Australia, and endemic
species (all of which are clearly derived from Australian stock).

140

Psyche

[December

Class II. Species native to, and sometimes widespread in the Indo-
Australian region which appear to have been distributed in part
by man.

As the islands under discussion are rather peripheral to the main
part of the Indo-Australian area it is difficult to estimate whether
“Class II” species listed from them are truly native, or whether they
have originated from propagules carried by man. This difficulty is
especially evident with respect to the Lord Howe and Norfolk Island
forms, but in our opinion, it is highly probable that the “Class II”
Kermadec species ( P oner a gleadowi and Monomorium antarcticum )
were carried to Raoul by man.

Class III. Widespread pantropical or cosmopolitan forms well
known as “tramp species” and distributed largely by man.

The numbers of species referable to each of these classes on the
islands concerned are shown in Table I.

Lord Howe

Norfolk

Raoul

Island

Island

Island

South latitude

30° 30'

290 48'

290 10'

Size (mi2)
Distance from

5

13

1 1.25

Australia (mi)
No. ants Class I

400

900

1700

(endemics bracketed

) 10(5)

8(0

— ■

No. ants Class II

3

1

2

No. ants Class III

1

3

1

Total number of ants

14

12

3

Table I. Certain geographical features of Lord Howe Island, Norfolk
Island, and Raoul Island ; with details of the class composition of their ant
faunas. Further explanation in text [data based in part on Wheeler (1927),
modified by recent revisionary studies].

Although native ants are apparently absent from Raoul they are
present on both Lord Howe and Norfolk Islands. The species
concerned are all of Australian origin, and more of them are present
on Lord Howe than on Norfolk, in spite of the considerably larger
size of the latter. Faunal size on these islands is thus negatively
correlated with their areas, a reverse situation from that observed by
Wilson (1961) in various Moluccan and Melanesian Islands.
Judging from the Melanesian faunal curves published by Wilson,
we believe that Lord Howe Island probably supports a saturated ant
fauna. Norfolk Island, however, would be expected to carry more

1961]

Taylor and Wilson — Ants

141

species than it does — and considerably more than Lord Howe. The
reason for the apparently sub-normal size of the Norfolk Island
fauna may be sought in the correlation between faunal size on the
two islands and the following two factors:

1. Ecological Diversity: The published reports of the terrain and
the state of the vegetation on these islands suggest that Lord Howe
is ecologically much more diverse than is Norfolk Island. In- spite
of its much smaller size it may, therefore, be capable of supporting
a larger and more varied ant fauna. Conversely, Norfolk Island may
have a lower carrying capacity due to its lower ecological diversity,
so that its fauna, although proportionately smaller than that of Lord
Howe, is nevertheless saturated.

2. Distance from Australia: The apparent correlation between

faunal sizes on these islands and their distances from Australia
contrasts with Wilson’s (1961) finding in Melanesia, where such
correlation is lacking. This situation may be due to the absence of
intermediate islands between those under discussion and Australia —
islands which might provide assistance as “stepping stones” to colonists.
According to this hypothesis the small size of the Norfolk ant fauna
is due to the fact that too few species have reached the Island to
saturate its carrying capacity.

If mere distance from Australia does have the filtering effect
envisaged it may be assumed that there is a maximum range of
natural overseas dispersal of the Australian fauna. As far as the
ants are concerned, Raoul, in lacking indigenous Australian elements,
may be considered to be outside this range.

It is not possible, at present, to evaluate the relative importance
of these two factors in determining the size of the Norfolk Island
ant fauna, but the problem would certainly be worthy of attention
by future field workers in these areas.

Lord Howe Island also has significantly more endemic ant species
than has Norfolk; each of these represents a stock which has diverged
from its Australian progenitor and in no case has splitting or radiation
taken place in situ on the island. The ecological diversity of the
island, coupled with the competitive pressures which must arise in
such a dense fauna, has probably caused this specialization of stocks,
with consequent phyletic change to the point of formation of an
endemic species in some lineages.

Clipperton Island

Clipperton Island is a lonely coral atoll about four miles in
diameter, lying at N. lat., 10.18 and W. long., 109.15. It is 1,800

142

Psyche

[December

miles due west of the Panama Canal, and about 670 miles southwest
of the nearest part of the Mexican coast. Most of the island is
barely 10 feet high, but it rises to about 70 feet at one point. The
vegetation consists of grasses, shrubs, and wild tobacco and there are
several coconut groves.

The island is now uninhabited but it was occupied by a Mexican
garrison from 1879-1917, and by about 100 phosphate miners from
1906-1917. Both of these groups received regular supplies from
Mexico. Clipperton was occupied by the United States Navy during
the Second World War.

The ants listed below were collected for the Museum of
Comparative Zoology by A. M. Peterson and C. F. Harbinson
during August 1958; the collection is almost certainly fully
representative.

Odontomachus insularis (Guerin)

This species is widespread in Central America and the West
Indies, and ranges north to Florida and Georgia. It also occurs on
Cocos Island and the Tres Marias Islands in the Eastern Pacific.
It is noteworthy that the Island of Clarion in the Revillagigedo group
is occupied by a second species, O. clara Roger {—haematoda
clarionensis Wheeler), which is widespread in Mexico and the West
Indies. A third far ranging species, O. bciuri Emery (= haematoda
div. auct.; nec haematoda L. ?), occurs on the Galapagos. All of
these exclusively New World species are to be distinguished from
the closely similar Indo-Australian O. simillimus Fr. Smith,
(=haematoda div. auct.; nec haematoda L.), which ranges eastwards
only to Tahiti. Three series collected.

Tetramorium simillirnum (Fr. Smith)

Like T. guineense this is a pantropical tramp species of African
origin, widespread both in Central America and Polynesia. Its total
range closely approximates that of guineense. Ten series collected.
Triglyphothrix striatidens (Emery).

A species of tropical Oriental origin, spread by commerce throughout
the Pacific, and established in North Africa and Queensland,
Australia. Seven series collected.

Like Raoul, Clipperton has a depauperate ant fauna, consisting
entirely of species which have probably been introduced by man.

Saint Helena Island

Saint Helena (S. lat., 15.57, W. long., 5.42) is a small volcanic
island 47 sq. miles in area, lying in the south Atlantic about i,8oo

1961]

Taylor and Wilson

143

— A nts

miles from South America, and 1,200 miles from Africa. The Island
differs in several respects from those discussed above, in that it is
well populated and is visited more regularly by shipping. Indeed,
it has at times been a major port-of-call for trade between Europe,
and South Africa and India. The ants listed below have probably
all been introduced to the island by man. They were collected for
the Museum of Comparative Zoology between 1958 and i960 by
Mr. Arthur Loveridge. The collection is probably less representative
than those already discussed, so that no faunal analysis can be
attempted at present.

Tetramorium simillimum (Fr. Smith)

The distribution of this species has been discussed above. One
series collected at Varneys.

Pheidole megacephala (Fabricius)

This is one of the most widespread and notorious of the commercial
tramp ant species, it is now virtually cosmopolitan in warm humid
areas. Sandy Bay: under rocks. Prosperous Bay: nest with alates,
under rocks, February 24, 1958. Varneys: alate males from a

storeroom cabinet, February 19, i960.

Solenopsis ( Diplorhoptrwn ) sp.

The subgenus Diplorhoptrum includes a number of taxonomically
confused, and in some cases widespread, cryptobiotic species, several
of which appear to be commercial tramps. A single series labeled
Varneys, 1000 feet.

Plagiolepis alluaudi Forel

This species, probably of East African origin, is a widespread
pantropical tramp species. A single series from Varneys.
Paratrechina bourbonica Forel

A widespread pantropical tramp species, apparently of African
origin. A single series from Varneys.

References

Brown, W. L., Jr.

1958. A review of the ants of New Zealand (Hymenoptera-Formicidae) .
Acta Hym., (Fukuoka), 1 (1): 1-50.

Miller, David

1956. Bibliography of New Zealand Entomology. 1775-1952. Bull. N. Z.
Dept. Sci. Ind. Res., 120: 1-492.

Wheeler, W. M.

1927. The ants of Lord Howe Island and Norfolk Island. Proc. Amer.
Acad. Arts Sci., 62: 121-153.

1935. Checklist of the ants of Oceania. Occ. Pap. Bishop Mus., 11 (11) :

144

Psyche

[December

1936. Ants from the Society, Austral, Tuamotu and Mangareva Islands.
Occ. Pap. Bishop Mus., 12 (18): 1-17.

Wilson, E. O.

1958. Studies on the ant fauna of Melanesia. IV. The tribe Ponerini.

Bull. Mus. Comp. Zool. Harvard, 119 (4): 320-371.

1961. The nature of the taxon cycle in the Melanesian ant fauna.
American Naturalist, 95 (882) : 169-193.

STUDIES ON CARBONIFEROUS INSECTS OF
COMMENTRY, FRANCE: PART III.
THE CALONEURODEA

By F. M. Carpenter
H arvard University

Introduction

This is the third in a series of papers redescribing certain of the
Carboniferous insects of the Commentry Basin, France.1 It treats
those Commentry species which now appear to belong to the order
Caloneurodea. This extinct order was erected by Martynov in 1938
for the Carboniferous family Caloneuridae and for related families
represented in Permian deposits of the USSR. It now includes addi-
tional families established by Carpenter (1943) for Permian species
from Kansas. At the present time seven families are known in the
order and of these only one, the Caloneuridae, is of Carboniferous age.
The Commentry specimens are by far the most important of the
Caloneuridae but Handlirsch’s and Meunier’s publications have given
us a superficial and confused knowledge of these fossils. The present
paper, which is based upon direct study of all the known Commentry
material, consists of redescriptions of the fossils and a revision of their
taxonomic assignments.

Survey of Commentry Species

A historical account of the Commentry collections and of the gen-
eral literature on the insects was included in the first part of this
series of papers (Carpenter, 1943). Among the species first described
by Brongniart (1885) from the Commentry shales was a single speci-
men, designated Caloneura dawsoni, and assigned to the family Pal-
aeacridiodea of the order Orthoptera. In his later monograph of the
Commentry insects (1894) Brongniart included drawings of five
additional specimens of dawsoni and gave a more extensive account of
the genus, which he placed in the same family, designated at that time
as Palaeacrididae. Handlirsch, in his 1906 treatise, established the
family Caloneuridae for the genus, assigning it to the extinct order
Protorthoptera.

This research has been aided by a grant (NSF-G14099) from the National
Science Foundation and by a previous grant (1938) from the Penrose Fund of
the American Philosophical Society. I am indebted to the authorities of the
Laboratoire de Paleontologie of the Museum National d’Histoire Naturelle in
Paris for placing at my disposal the unique collections of Commentry insects
in the Museum, both in 1938 and in 1961; and to the authorities of the British
Museum (Natural History) for allowing me to examine the Commentry
fossils in that institution.

145

146

Psyche

[December

In 1917 Lameere, following an examination of the Commentry
fossils in the Museum in Paris, placed in the Caloneuridae two species
(similis and royeri) which Meunier had originally described (1911)
in the order Paleodictyoptera. At about the same time (1919) Hand-
lirsch described four more species of Caloneura, all based on the Com-
mentry specimens illustrated in Brongniart’s monograph (1894).
Finally in 1925 Bolton described an additional species (subiilis) from
Commentry in the British Museum (Natural History) and also
figured a specimen of dawsoni included in the same collection.

As a result of my examination of all of these fossils and of addition-
al specimens in the Museum in Paris, not previously published upon,
I propose the following classification of the Commentry Caloneurodea :

Family Caloneuridae Brongniart

Genus Caloneura Brongniart (synonym: Confusio Handlirsch)
dawsoni Brongniart (synonyms: picta Handlirsch, major
Handlirsch, longicornis Handlirsch, robusta Handlirsch,
royeri Meunier).

Family Apsidoneuridae, new (type species, Apsidoneura flexa Carp.,
Permian Kansas)

Genus Homaloptila Handlirsch
similis Meunier

The Commentry species which Bolton (1925) described as Calo-
neura subtilis (British Museum [Natural History], type 17280) is
not, in my opinion, a caloneurid or even a member of the1 order Cal-
oneurodea. However, since the poor preservation of the specimen
prevents satisfactory determination even to order, I refer this species
to Insecta Incertae Sedis.

Family Caloneuridae Handlirsch

Fore and hind wings similar. Sc terminating well before the wing
apex; CuA and CuP very close together and nearly parallel; MP
forked nearly dichotomously ; 4 anal veins; cross veins numerous.
Body and legs slender.

This family is known only by the genus Caloneura. The Carbon-
iferous genera Caloneurella Carp. (Pennsylvania) and Pruvostiella
Handl. (Nord, France), both of which have previously been placed
here, should probably be included under Caloneurodea Incertae Sedis.

Genus Caloneura Brongniart

Caloneura Brongniart, 1885, soc. Amis Sci. Nat. Rouen., 21:59; 1894, Re-
cherches Hist. Ins. Foss.: 562. Handlirsch, 1906, Foss. Ins.: 141. Carpenter,
1943, Proc. Amer. Acad. Arts Sci., 75 :69.

Confusio Handlirsch, 1919, Denkschr. Acad. Wiss. Wien, 96: 37 (New syn-
onymy).

Fore wing with a somewhat broader costal space than the hind wing

1961]

Carpenter — Caloneurodea

147

and with the costal space abruptly narrowed basally; hind wing
narrower at base than the fore wing and with the costal margin nearly
straight basally. Rs with five or six main branches, one of these
usually having a short distal branch. MP usually forked, CuA and
CuP unbranched.

Type species: Caloneura dawsoni Brongniart.

Confusio was based upon royeri, which Meunier described in the
palaeodictyopterous genus Homaloneura, and which I consider a
synonym of dawsoni.

Caloneura dawsoni Brongn.

Figure 1

Caloneura dawsoni Brongniart, 1885, Soc. Amis Sci. Rouen, Bull., 21:59, pi.
[4], fig. 2; 1894, Recherches Hist. Ins. Foss.,: 562, pi. 36, fig. 5-11.
Lameere, 1917, Mus. Nat. Hist. Natur. Bull., 23 : 1 8 1 . Handlirsch, 1919,
Denkschr Akad Wiss., 96:35, fig. 36. Bolton, 1925, Brit. Mus. Fossil
Insects. 2 :1 5, fig. 5.

Caloneura picta Handlirsch, 1919, ibid., p. 35, fig. 37.

Caloneura major Handlirsch, 1919, ibid., p. 35, figs. 38, 39.

Caloneura longicornis Handlirsch, 1919, ibid., p. 35, fig. 40.

Caloneura robusta Handlirsch, ibid., p. 36, fig. 41.

Homaloneura royeri Meunier, 1911, Bull. Mus. Nat. Hist. Natur., 17:119, fig.
2; 1912, Ann. Paleont., 7:9, pi. 7, fig. 2; Lameere, 1917, Mus. Nat. Hist.
Natur., Bull., 23 : 1 8 1 -

Confusio royeri Handlirsch, 1919, Denschr. Acad. Wiss., 96:37, fig. 44.

This species was originally based on the single specimen figured in
Brongniart’s 1885 paper but was later known to Brongniart by five
additional fossils, all illustrated in his Recherches (1894). Five other
specimens (here designated as C1-C5), which have not previously
been mentioned in the literature, are contained in the Museum
National d’Histoire Naturelle in Paris. To these may be added the
specimen of dawsoni described by Bolton (1925), and the type of
royeri Meunier. Since I am convinced for reasons given below that
these thirteen specimens, all of which I have been able to examine,
are one species (dawsoni) , I have used them as a basis of the following
account of this insect.

Wings (Figure 1) : The fore wing, which is completely preserved
in several specimens, is from 45-48 mm in length, and from 10-12 mm
in width. The type specimen has a fore wing length of 45 mm and a
width of 10 mm. The hind wings of the several specimens are about
the same dimensions as the fore wings but are somewhat narrowed
basally. The costal area of the fore wing narrows abruptly just at the
level of the separation of CuP and iA. In the hind wing the costal
space is not narrowed at this point, continuing its full width as far as
it is preserved in all specimens.

148 Psyche [December

The general venational pattern is shown in figure 1, but this is
subject to much individual variation. The subcosta terminates on the
costal margin well before the end of Ri ; it does not extend to the apex
of the wing as shown by Bolton in his drawing of a specimen in the
British Museum. The number of main branches of Rs varies from
5 to 6 in the several specimens showing those veins, but usually one of
these branches has a short fork, which occurs most often on R2 or R3.
The branching varies in the fore and hind wings (and presumably in
the right and left wings) of individual insects. In the holotype speci-
men (36-5 )2, for example, this fork occurs on R3 in the fore wing but
on R2 in the hind wing. MP is less variable; it forks broadly at about
mid-wing. In only one specimen (C4) does either branch of MP
clearly fork again (see figure 1), but in several one or both branches

Figure 1. C aloncura danvsoni Brongn. Fore and hind wings based on the
holotype and other specimens in the Museum National, Paris. Sc, subcosta
( — ); Rl, radius ( + ); Rs, radial sector ( ); MPl + 2 and MP3 + 4,

branches of posterior media (' ) ; 1A, 2A, 3A, 4A, anal veins.

appear to have a small, terminal fork. CuA and CuP are consistently
unbranched ; the divergence of CuP and 1 A, near the base of the wing,
is either shallow or somewhat broader. 7 he four anal veins are con-
sistently present.

The most striking characteristics of the wings of dawsoni are the

2Since the Commentry insects in the Museum National are not numbered,
I am using here the figure number on Brongniart’s plates for the number of
the specimen. Hence, specimen 36-5 is the fossil illustrated on Brongniart’s
plate 36, figure 5.

1961]

Carpenter — Caloneurodea

149

markings, as mentioned by Brongniart in his original description. 1 he
main veins and most of the cross-veins are margined with dark pig-
ment ; the veins appear to be very thick but careful examination of the
wing shows that the actual veins are only of usual thickness. In
addition, nearly all cells of the fore wing contain a dark, irregular
pigment spot. The size of the spots varies in different wings but the
pattern formed in the wing as a whole is fairly constant. The hind
wings appear to be without both types of these markings ; at any rate,
no specimens that can clearly be recognized as hind wings show them.
In a few fossils some of the wings appeared blotched (as in the
holotype, 36-5), but the extreme irregularity of this and its absence
in other hind wings strongly indicates that it is the result of preserva-
tion.

Body structure: Very little is known of the body structure of

dawsoni. Several specimens show fragments and portions of the
thorax and four specimens (36-8, 36-9, 36-10, 36-11) include parts
of the legs and antennae but the abdomen is entirely unknown. The
legs are slender, nearly homonomous and apparently cursorial. The
fore and middle legs are about 34 mm long and the hind legs, 45 mm
long. The dimensions of the leg segments are given in Table 1. The
tarsi of all three pairs of legs are about the same length.

Femur

Tibia

Tarsus

Fore leg

12 mm

1 0 mm

12 mm

Middle leg

1 2 mm

10 mm

12 mm

Hind leg

18 mm

15 mm

12 mm

Table 1. Dimensions of leg segments in Caloneura dawsoni Brongn.

The femora and tibiae of the fore and middle legs are about the same
length, but the hind femur and tibia are slightly longer than those of
the other pairs. The antennae are obviously long and slender, though
they are not completely preserved in any fossil. In specimen 36-10
one antenna is preserved to the edge of the rock containing the speci-
men ; the preserved part is 55 mm long, which is considerably in excess
of the wing length.

So far as known, therefore, the body structure of dawsoni is similar
to that of the best-known Permian species, Paleuthy gramma tenui-
cornis Mart, (see Martynov, 1938b, p. 126).

Synonymy : As mentioned above, Brongniart figured in his R.e-

cherches (1894) five speciments of dawsoni which were not known
to him when he described the species in 1885. In 1919 Handlirsch,
basing his conclusion on Brongniart’s illustrations, established four

i50

Psyche

[December

additional species for these fossils: picta, major , longicomis , and
robusta. No specific characteristics were even mentioned for robusta
and those used for the others involved the nature of the branching of
Rs. Having examined all these specimens, I cannot accept any of
Handlirsch’s species as valid.

Meunier’s royeri, originally placed in the paleodictyopterous genus
Homaloneura , was transferred to the Caloneuridae by both Lameere
(1917) and Handlirsch (1919), the latter making the new genus,
Confusio, for it. Meunier’s drawing of the fossil (1911, p. 119,
fig. 2), although incorrect in many details, was more accurate than
Handlirsch’s (1919, p. 37, fig. 44), which was based solely on the
small photograph contained in Meunier’s later account (1912). The
wing, which I have examined at the Paris Museum, is characteristic
of dawsoni: Rs has six branches and MP is forked as usual in the
species, — not narrowly branched as in Meunier’s figure or un-
branched as in Handlirsch’s. CuP and iA are close together, as usual,
not widely separated as represented by Meunier. This specimen is
clearly dawsoni and since the costal margin is straight (not curved as
in Meunier’s figure), it is almost certainly a hind wing.

Specimens studied: The following is a list of the specimens of
dawsoni which are now known and which I have been able to examine.
In the Museum National, Paris; (1), no. 36-5, the holotype of daw-
soni, consisting of a nearly complete fore wing and two overlapped
hind wings. (2), no. 36-7 (type of picta Handl.), distal two-thirds
of a hind wing. (3),, no. 36-8 (syntype of major Handl.), a nearly
complete insect, with the fore wings overlapped and the hind wings
also overlapped; some parts of thorax and legs. (4), no. 36-9 (type
of longicomis Handl.), a wing base, parts of thorax, legs and anten-
nae. (5), no. 36-10 (type of robusta Handl.), wing bases, portions
of thorax, legs, antennae. (6), no. 36-11 (syntype of major Handl.),
nearly complete fore wing, basal third of hind wing, parts of thorax
and legs. (7), no. Ci, two overlapped wings, probably hind, bases
absent. (8), no C2, a well preserved fore wing, showing details of
fore wing and posterior margin, parts of thorax and legs. (9), no.
C3, distal third of a wing, probably hind. (10), no. C4, a very clear
fore wing, and basal parts of a hind, parts of thorax and legs. ( 1 1 ) ,
no. C5, basal half of a hind wing, posterior margin very clear. (12),
the type of royeri Meunier, a nearly complete hind wing, lacking only
the very base. In the British Museum (Natural History), London:
(13), no. I7292, consisting of at least three (probably four) over-
lapped wings; also parts of thorax and legs, poorly preserved. Bolton

1961]

Carpenter — Caloneurodea

I5i

(1925) was in error in describing a large precostal area in this fossil ;
he apparently did not realize that more than two wings were over-
lapped, resulting in a confused jumble of veins.

Apsidoneuridae, new family

Fore and hind wings similar. Sc extending to about the apex of
the wing; CuA and CuP very close together and nearly parallel; MP
forked broadly and unevenly, the anterior branch (MP 1+2) arch-
ing strongly away from MP 3 + 4; 3 anal veins; cross-veins numerous.
Body unknown.

This family is related to the Caloneuridae but differs in the length
of Sc, the number of anal veins and the nature of the fork on MP.
It is based on the type-genus Apsidoneura Carpenter (Permian, Kan-
sas), which was originally placed in the Permian family Paleuthy-
grammatidae. Study of additional specimens of Caloneurodea from
Permian strata of Oklahoma and of the type of similis from Commen-
try has led me to conclude that Apsidoneura and Homaloptila represent
a distinct family.

Genus Homaloptila Handlirsch
Homaloptila Handlirsch, 1919, Denkschr. Acad. Wiss., 96:36.

Rs with four main branches, one of these usually having a terminal
fork; MP 3 + 4 continuing in a nearly straight line the stem MP,
which arises from CuA at a considerable distance from the separation
of CuA from R. In Apsidoneura , MP arises from R or the junction
of R and CuA.

Type-species: Homaloneura similis Meunier. This is the only
species known in the genus.

Hojnaloptila similis (Handlirsch)

Figure 2

Homaloneura similis Meunier, 1911, Bull. Mus. Nat. Hist. Natur., 17:118,
fig. 1. 1912, Annal. Paleont., 7:8; fig. 6 ; pi 6, fig. 5.

Homalopt la similis Handlirsch, 1919, Denkschr. Akad. Wiss. 96:36, fig. 42,
43.

Fore wing: length, 30 mm., maximum width, 7 mm. Rs with four
main branches, at least two of which are forked distally. MP 1+2
arched away from MP 3 + 4 as characteristic of the family, but
curved strongly towards MP 3 + 4 distally. There is a tendency for
the formation of an irregular network of cells in the distal parts of Rs
and MP. Base of wing slender but not as slender as in Apsidoneura
flexa. Hind wing: similar to the fore wing in size and venation but
having a straight costal margin towards the base. The venational
pattern is shown in figure 2.

152

Psyche

[December

Holotype: Museum: National, Paris. This is a well preserved speci-
men showing a fore wing and hind wing, their bases overlapped but
otherwise independent. The veins are distinct and can be clearly seen
under good illumination.

Meunier’s drawing of this specimen is incorrect and misleading; he
completely overlooked in both wings the fork on MP and the proximi-
ty of CuA and CuP. Handlirsch’s drawing, based on Meunier’s

Figure 2. Homaloptila similis (Meunier). Fore and hind wing, based on
holotype in Museum National, Paris. Lettering as in figure 1.

published photograph, was equally inaccurate, showing the hind wing
with a broad anal area. However, even though the wings are slightly
overlapped basally, the margins of the wings are distinct.

The presence in the Commentry shales of this genus, allied to
Apsidoneura, is an interesting example of a Permian element in the
Commentry fauna. A similar occurrence was noted (Carpenter,
1943) in the Order Protodonata, the genus Meganeurula of Com-
mentry being virtually indistinguishable from the Permian genus
Typus .

References

Bolton, H..

1925. Insects from the Coal Measures of Commentry. Brit. Mus. (N.
H.), Fossil Insects, 2:1-56.

Brongniart, C.

1885. Les insectes fossiles des terrains primaires. Soc. Amis Sci. Nat.
Rouen, Bull. : 55-68.

1961]

Carpenter - — Caloneurodea

153

1894. Recherches pour servir a l’histoire des insectes fossiles des temps
primaires. These Fac. Sci. Paris, no. 821:1-494.

Carpenter, F. M.

1943. The Lower Permian insects of Kansas. Part 9. The Orders
Neuroptera, Raphidiodea, Caloneurodea and Protorthoptera.
Proc. Amer. Acad. Arts Sci., 75:55-84.

1943. Studies on Carboniferous insects from Commentry, France. Part

I. Introduction and families Protagriidae, Meganeuridae, and
Campylopteridae. Geol. Soc. Amer., Bull., 54:527-554.

1951. Studies on Carboniferous insects from Commentry, France. Part

II. The Megasecoptera. Journ. Paleont., 25:336-355.
Handlirsch, A.

1906. Die Fossilen Insekten. Leipzig.

1919. Revision der Palazoischen Insekten. Denkschr. Acad. Wiss.
Wein, Math.-Naturwiss. Klasse, 96:1-82.

Lameere, A.

1917. Revision sommaire des insectes fossiles du Stephanien de Com-
mentry. Mus. Nat. Hist. Natur., Bull., 23:141-201.

Martynov, A. V.

1938a. Permian fossil insects from the Arkhangelsk district. Part V.
The family Euthygrammidae and its relationships. Trav. Inst,
paleontol. Acad. Sci. URSS, 7:69-80.

1938b. Etudes sur l’histoire geologique et de phylogenie des ordres des
insectes. I. Palaeoptera et Neoptera — Polyneoptera. Trans, de
PInst. Paleont., 7:1-149.

Meunier, F.

1911. Nouveaux insectes du houiller de Commentry. Mus. Nat. Hist.
Natur., 17: 117-128.

1912. Nouvelles recherches sur quelques insectes du terrain houiller de
Commentry- Ann. Paleont., 7:1-19.

TP pp 155-56 removed
placed in front of v®
Binding Unit*

PSYCHE

INDEX TO VOL. 68, 1961

INDEX TO AUTHORS

Blum, M. S., J. E. Roberts, Jr., and A. F. Novak. Chemical and Biological
Characterization of Venom of the Ant Solenopsis xyloni McCook. 73

Brown, IV. L., Jr. A note on the Ant Gnamptogenys hartmani Wheeler. 69
Mass Insect Control Programs: Four Case Histories. 75
The Neotropical Species of the Ant Genus Strumigenys Fr.
Smith: Miscellaneous Concluding Studies. 58

Carpenter, F. M. Studies on Carboniferous Insects of Commentry, France:
Part III. The Caloneurodea. 150

Causey, Nell B. Adrityla, a New Milliped Genus (Chordeumidea : Conoty-
lidae). 126

Darlington, P. J., Jr. Australian Carabid' Beetles V. Transition of Wet
Forest Faunas from New Guinea to Tasmania. 1

Australian Carabid Beetles VII. Trichosternus , Es-
pecially the Tropical Species. 113

Evans, H. E. A Reconsideration of the Genus Epipompilus (Hymenoptera :
Pompilidae). 25

Levi, H. JV. and L. R. Levi. Some Comments on Walckenaer’s Names of
American Spiders, Based on Abbot’s Drawings. 53

Mockjord, E. L. The Rediscovery and Probable Phylogenetic Position of
Psilopsocus (Psocoptera). 38

Nutting, W. L. and A. B. Gurney. A New Earwig in the Genus Vostox (Der-
maptera: Labiidae) from the Southwestern United States and Mexico. 45

Taylor, Robert fV. and Edward 0. Wilson. Ants from Three Remote Oceanic
Islands. 134

Werner, F. G. Anthicus tobias Marseul, Another Tramp Species (Coleoptera:
Anthicidae). 70

157

INDEX TO SUBJECTS

All new genera, new species and new names are printed in capital type.

ADRItyla, a new milliped genus
(Chordeumidea : Conotylidae) , 131
Adrityla deserctae, 131
American spiders, 53
A new earwig in the genus V ostox
(Dermaptera: Labiidae) from the
Southwestern United States and
Mexico, 45

A note on the ant Gnamptogenys
hartmani Wheeler, 69
Anthicidae, 70

Anthicus toblas Marseul, another
tramp species (Coleoptera: An-
thicidae), 70
Ants, 58, 69, 75, 83, 134
Ants from three remote oceanic
islands, 134
APSIDONEURIDAE, 151
Arachnids, 53

A reconsideration of the genus Epi-
pompilus ( Hymenoptera : Pompili-
dae), 25

Australian carabid beetles V. Trans-
ition of wet forest faunas from
New Guinea to Tasmania, 1
Australian carabid beetles VII. Tri-
chosternus , especially the tropical
species, 113
Beetles, 1, 70, 113
Caloneura dawsoni, 147
Caloneurodea, 145
Carabid beetles, 1, 113
Chemical and biological characteri-
zation of venom of the ant Solcn-
opsis xyloni McCook, 73
Chordeumidea, 131
Coleoptera, 1, 70, 113
Conotylidae, 131
Dermaptera, 45
Diptera, 92
Earwig, 45
Epipompilus, 25
Epipompilus aztecus, 30
Epipompilus bifasciatus , 31
Epipompilus delicatus, 35
Epipompilus excelsus, 32
Epipompilus haupti, 32
Epipompilus innubus, 36
Epipompilus insolitus, 35
Epipompilus pulcherrimus , 29
Epipompilus williamsi, 29
Fire ant, 83
Fruit fly, 92

Gnamptogenys hartmani, 69
Gypsy moth, 76

Homaloptila, 151
Elomaloptila similis, 151
Hymenoptera, 25, 58, 69, 75, 83, 134
Imported fire ant, 83
Labiidae, 45
Lepidoptera, 76

Mass insect control programs: four
case histories, 75
Mediterranean fruit fly, 92
Millipeds, 131
Pompilidae, 25
Psilopsocus, 38
Psilopsocus nebulosus, 40
Psocoptera, 38
Screwworm, 97
Solenopsis xyloni, 73
Some comments on Walckenaer’s
names of American spiders, based
on Abbot’s drawings, 53
Spiders, 53
Strumigenys , 58
Strumigenys fairchildi, 60
Strumigenys fusca, 67
Strumigenys hindenburgi, 63
Strumigenys lacacoca, 58
Strumigenys lanuginosa, 62
Strumigenys louisianae, 64
Strumigenys micretes, 58
Strumigenys unidentata, 68
Studies on Carboniferous insects of
Commentry, France: Part III. The
Caloneurodea, 145

The Neotropical species of the ant
genus Strumigenys Fr. Smith: Mis-
cellaneous concluding studies, 58
The rediscovery and probably phylo-
genetic position of Psilopsocus
(Psocoptera), 38
Trichosternus , 113
Trichosternus cordatus, 127
Trichosternus eungella, 125
Trichosternus fax, 118
Trichosternus fisheri, 124
Trichosternus frater, 122
Trichosternus kirrama, 129
Trichosternus mixtus, 126
Trichosternus montorum, 120
Trichosternus mutatus, 123
Trichosternus nudipes, 121
Trichosternus obscuripennis , 120
Trichosternus setosiceps, 129
Trichosternus soror, 122
Trichosternus spec, 128
V ostox, 45

Vostox excavatus, 47

158

CAMBRIDGE ENTOMOLOGICAL CLUB

A regular meeting of the Club is held on the second Tuesday of
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The illustration on the front cover of this issue of Psyche is a
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PSYCHE

A Journal of Entomology

Volume 69
1962

Editorial Board

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

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

E. O. Wilson H. E. Evans

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

Cambridge, Mass., U. S. A.

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

Vol. 68, no. 4, Dec., 1961: March 13, 1962
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Of.

> PSYCHE

A JOURNAL OF ENTOMOLOGY

Established in 1874

Vol. 69

March, 1962

No. 1

CONTENTS

A Chemical Alarm Releaser in Honey Bee Stings ( Apis mellifera L.).
Ralph L. Ghent and Norman E. Gary 1

The Male of Platy patrobus lacustris Darlington (Coleoptera: Carabidae).

Carl H. Lindroth 7

The Spider Genera Steatoda and Enoplognatha in America (Araneae,
Theridiidae) . Herbert IV. Levi 11

A Permian Megasecopteron from Texas. F. M. Carpenter 37

A Remarkable New Genus of Lygaeidae from Sumatra (Hemiptera:
Heteroptera ) . James A. Slater 42

Notes on Fossil Cleoninae (Coleoptera: Curculionidae.)

John M. King solver 47

Pseudoscymnus , a New Genus of Asiatic Scymnini (Coleoptera:

Coccinellidae) Edward A. Chapin 50

Jlfj H@§ ..
f - mum

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1961-62

President J. J. T. Evans. Harvard University

Vice-President C. Walcott. Harvard University

Secretary A. R. Brady, Harvard University

Treasurer F. M. Carpenter, Harvard University

Executive Committee R. W. Taylor, Harvard University

S. K. Harris, Boston University

EDITORIAL BOARD OF PSYCHE
F. M. Carpenter (Editor), Professor of Entomology , Harvard
U niversity

P. J. Darlington, Jr., Curator of Recent Insects , Museum of Com-
parative Zoology

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

E. 0. Wilson, Associate Professor of Zoology , Harvard University
H. W. Levi, Associate Curator of Arachnology, Museum of Com-
parative Zoology

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

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The December 1961 Psyche (Vol. 68, no. 4) was mailed March 13,
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lished with the aid of a National Science Foundation Grant
(G 15817) to the Cambridge Entomological Club.

The Lexington Press. Inc., Lexington, Massachusetts

PSYCHE

Vol. 69 March, 1962 No. 1

A CHEMICAL ALARM RELEASER IN
HONEY BEE STINGS ( APIS MELLIFERA L.)1

By Ralph L. Ghent and Norman E. Gary
Department of Entomology
Cornell University

Chemicals that function to communicate alarm among the members
of hymenopteran colonies have been discovered in recent years by
several investigators. These chemicals, released by disturbed insects
and detected in the gaseous state by other members of the colony, have
been shown to elicit various forms of alarm behavior, differing accord-
ing to species, but always related to the defense of the colony. Certain
species of ants in particular have been shown to employ such “alarm
releasers” (Wilson, 1958; Butenandt and Lindauer, 1959; Brown,
i960; Ghent, 1961). Preliminary evidence presented by Huber
(1814) suggests that the honey bee (Apis mellifera) also possesses
such a mechanism.

It is a common observation among beekeepers that the probability of
being stung increases after the first sting. The frequency of stinging
often appears to rise exponentially: for example, a beekeeper working
with colony after colony becomes increasingly liable to be stung. This
phenomenon suggests that bees possess some means of labelling an in-
truder, presumably by applying an alarm releaser to the victim during
the act of stinging.

A characteristic, pleasantly sweet scent is noticeable at the site of
stinging. This scent, although not detectable from undisturbed bees,
becomes quite concentrated in a closed container of agitated bees and
is most perceptible from newly-inserted stings. One exception is that
the stings of recently emerged bees bear no such scent. When the bee
stings, the barbed sting and its basal motor apparatus are torn from the
departing worker and remain imbedded in the skin. The fragrant

This investigation was supported by a research grant (MY 3368) from
the Mental Health Division, U. S. Public Health Service.

Manuscript received by the editor October 16, 1961.

MSTlIUiiQ.it

I

Mini

2

Psyche

[March

substance associated with the sting would, therefore, seem to be an
efficient tagging mechanism. Subsequent stings would then replenish
and augment this chemical label.

An examination of the sting apparatus to determine the source of
this scent revealed several facts. The contents of neither the large
poison reservoir nor the so-called “alkaline” accessory gland have any
detectable odor. A minute amount of fluid, immiscible with water, is
held between the bulbous base of the sting shaft and the setose lobe
of membrane which folds over it. The sting odor is particularly asso-
ciated with this membrane, although it is sometimes detectable on other
basal structures as well.

No glandular tissue was found to be immediately associated with
the setose lobe covering the bulb of the sting shaft. There are, how-
ever, two masses of glandular cells, lying against the inner surface
of the quadrate plates, which secrete by individual ducts onto the
outer surface of these plates (Snodgrass, 1956). There is a contin-
uous space surrounding the sting base through which this secretion
can flow, eventually to collect beneath the setose lobe.

It may be significant that under natural circumstances the under
surface of this lobe is exposed only when the sting is partially extruded,
or during the act of stinging. The sting is frequently extruded by
alarm bees, particularly when bees in the winter cluster are disturbed.

Exposure of honey bees to stings pulled from freshly-frozen workers
results in a marked change in behavior. Pieces of filter paper, bearing
one or more stings, were introduced into an observation hive contain-
ing a normal colony, and the subsequent behavior observed. A sudden
agitation was first observed in the vicinity of the introduced stings,
spreading quickly outward to a radius of about 15-20 cm. After the
first sharp wave of agitation, during which most of the bees in the area
buzzed momentarily, a general orientation to the stings occurred, with
many bees converging on the paper. Individual bees standing near or
over the introduced stings assumed an abnormally high stance, with
the antennae constantly waving, and the wings partially extended, in
marked contrast to the normal posture of resting bees. These indi-
viduals successively accosted approaching workers, turning from side
to side and sometimes making short flights to do so. Apparently identi-
cal behavior may be observed in guard bees at the entrance of a hive.
Control pieces of filter paper, introduced into other parts of the hive,
or alternately with those bearing stings, elicited no such behavior if
the introduction was made carefully without mechanical disturbance.

An experiment was conducted to determine whether the presence

1962]

Ghent and Gary — -Alarm Rcleaser

3

of a fresh sting on an introduced object increases the probability of its
being stung. Captured workers were first induced to sting a disk of
suede leather 1.5 cm. in diameter. This was lowered on a wire into

Table 1. Response to suede leather disk bearing three fresh stings compared
to response to an untreated control disk.

(a) Disk Bearing Stings (b) Control Disk

No. of bees No. of addi- No. of bees No. of addi-
clustered on disk tional stings clustered on disk tional stings

Test #1

50

4

0

0

#2

12

3

0

0

#3

10

0

1

0

#4

5

1

0

0

#5

100

3

0

0

#6

30

1

0

0

Test of significance for difference in number of additional stings:
x2 — 12 with 1 degree of freedom
P < .001

an observation hive and held about 5 cm. above the comb for one
minute, then carefully removed. Allowing 30 seconds between intro-
ductions, this disk and other control disks bearing no stings were
alternately introduced in the same manner. The number of bees
which congregated on the disks during each test was estimated and
the newly deposited stings were counted after each removal. The data
in table ( 1 ) clearly exhibit greater attractiveness of the leather disk
bearing stings, as well as a significantly (P < .001) greater number
of stings retained in it.

These results seemed to indicate that the presence of a fresh sting
alone is sufficient to provoke stinging attacks. In subsequent observa-
tions, however, it was demonstrated that one or more secondary stimuli
are usually necessary to elicit stinging. Although stings invariably
attract bees, we observed very few cases of stinging unless the test
object was moved or jarred. Indeed, bees that congregated about an
introduced sting tended to fly at any moving object in the vicinity.
Since the loss of the sting is often fatal to the worker, there is an
obvious selective pressure against the stinging of immobile objects,
even though coated with an alarm releaser.

In subsequent experiments, it was found that extracts of stings
attracted worker bees, and induced a pattern of behavior which was
apparently identical to that of workers exposed to fresh stings.

The alarm reaction was further characterized in an experiment
relating the quantity of extract to the number of bees attracted to it.
Th ree hundred whole uncrushed stings, pulled from freshly-frozen
workers, were extracted in 1 ml. of methylene chloride. Measured

4

Psyche

[March

quantities were pipetted into vials (4.5 cm. long by 1.3 cm. in dia-
meter) with an inner lining of filter paper. Each vial was placed in a
cylindrical screen cage 10 cm. long by 3 cm. in diameter, which was

EXTRACT (ml.)

Figure 1. Attraction of worker bees to an extract of bee stings in methy-
lene chloride. The extract was pipetted onto filter paper and introduced in
screen cages laid over the combs of a colony. Bees attracted to each cage
were collected and counted.

open at one end. The cages were laid in groups of four across the top
bars of an open colony, immediately enclosed by an empty hive body
and cover, and left in this dark chamber for 3 minutes. Each was
then carefully transferred with the adhering bees into a closed con-
tainer. The bees thus collected were anesthetized and counted (Fig.
1 ) . Although the data show considerable variation, there was a
positive correlation between the quantity of extract per vial and the
number of bees attracted.

An extract containing about 5,000 stings in 10 ml. of methylene
chloride was distilled, beginning at 350 C. and slowly rising to 90° C.
The colorless distillate was collected in three approximately equal
fractions. The last two fractions, distilled at temperatures over

1962]

Ghent and Gary — Alarm Releaser

5

6o° C., had the characteristic bee sting scent. Some of this scent was
retained by the yellow residue.

Vapor phase chromatograms of the first fraction showed evidence
of only the solvent, methylene chloride. In both of the last two frac-
tions only two components in addition to the solvent were detected:
water and the scented substance presumed to be the alarm releaser.
Attempts to concentrate the odoriferous component by evaporation of
the methylene chloride were only partially successful. Vapor phase
chromatograms indicated that, although it was slightly concentrated
during evaporation, the greater part was lost.

60

• distillate
o solvent

to _ _

uj 50
H
3
Z

2
t^40

£30
oc

I—

u20

UJ
03

6

10 -

t COmm

100

10

50

DISTILLATE l.)

Figure 2. Attraction of worker bees to distilled extract of bee stings. The
distillate was pipetted onto filter paper and introduced in screen cages laid
over the combs of the colony. Bees attracted to each cage were collected and
counted.

The effect of the whole distillate on bee behavior was tested by the
same method as that used for the crude extract. Data on attraction

6

Psyche

[March

confirmed the presence of the alarm releaser in the distillate (Fig. 2).
The behavior of bees exposed to the distillate was indistinguishable
from that observed in the presence of fresh bee stings. Moreover, it
was evident that bees attracted to filter papers bearing the distillate
demonstrated an extraordinary tendency to sting upon the slightest
provocation.

We have not yet succeeded in concentrating sufficient quantities of
the alarm releaser to permit chemical characterization. Our observa-
tions indicate that extremely small traces of the scented substance of
bee stings are detectable both by humans and by honey bee workers,
and that the amount borne by single stings is in fact minute.

Alarm behavior in honey bees is governed by many factors, and the
presence of the alarm releaser is not essential to stinging. Bees often
attack moving objects or animals in the vicinity of the hive where no
alarm releaser could possibly have been applied previously. Move-
ment, odor, and texture probably all determine to some degree whether
the response is aggressive. Unless accompanied by a supplementary
stimulus, for example movement, the odor of the alarm releaser rarely
precipitates stinging. The function of the alarm releaser, besides serv-
ing as an efficient recruiting mechanism, must be considered as an
intensifier of the normal defensive responses of the colony.

Bibliography

Brown, W. L., Jr.

1960. The release of alarm and attack behavior in some New World
army ants. Psyche 66 :25-27.

Butenandt, A.. B. Linzen, and M. Lindauer.

1959. Uber einen Duftstoff aus der Mandibeldruse der Blattschneidera-
meise Atta sexdens ruhropilosa Forel. Arch. Anat. micr. Morph,
exp. 48 (suppl.) :12-19.

Ghent, R. L.

1961. Adaptive refinements in the chemical defense mechanisms of
certain Formicinae. Unpublished thesis, Cornell University.

Huber, F.

1814. Nouvelles observations sur les abeilles II. Transl. 1926 Hamilton,
111. : Dadant.

Snodgrass, R. E.

1956. Anatomy of the honey bee. 334 pp. Ithaca, N. Y. : Cornell Univer-
sity Press,

Wilson, E. O.

1959 (1958). A chemical releaser of alarm and digging behavior in
the ant Pogonomyrmex badius (Latreille). Psyche 65 :41-51.

THE MALE OF PLATYPATROBUS LACUSTRIS
DARLINGTON (COLEOPTERA: CARABIDAE)*

By Carl H. Lindroth
Zoological Institute, University of Lund, Sweden

The discovery of a new genus among the Patrobini from Lake
Superior (Darlington, 1938) was most unexpected. It was founded
on a single female from Batchawaung Bay, Ontario, in the Leconte
collection in the Museum, of Comparative Zoology at Harvard, and,
in the absence of a male, the author hesitated to state the true relation-
ship of the new genus. This female is the only specimen of the genus
Platypatrobus (species, lacustris Dari.) heretofore known.

Quite recently, in September, 1961, Dr. Darlington and I visited
the well-known coleopterist, Mr. C. A. Frost, Framingham, Mass.,
and, looking through his large collection, made the exciting discovery
of a male of Platypatrobus lacustris , received by him from Dr. A. E.
Brower. According to the label, the beetle had been taken at “black
light,” July 30th, a few years ago (exact year not stated) at Sinclair
in northernmost Maine. Its capture in a light trap and its well
developed hind-wings indicate that it is able to fly.

The new male, except for the two dilated basal segments of the
pro-tarsi, matches the type completely. The extra setae on prothorax
and elytra are considerably irregular in number and position : lat-
erally on the prothorax are 3 (left) and 4 (right) (in the type 4 + 3)
setae; on the elytra, 1st interval, 2 + 2 (type 1+2), 3rd interval,
12 + approximately 9 (type 11 + 12), 5th interval, 10 + 8 (type
5 + 6).

I was allowed to borrow the specimen and have dissected the
genitalia (fig. 1). The parameres are of the normal Patrobine type,
almost identical with those of the two related genera, Patrobus and
Diplous (Platidius) , that is, approximately symmetric with long,
narrow apical prolongations bearing 4 setae at tip. The accessory sub-
apical setae are inconspicuous, only 2 in number, and there is no
suggestion of the hairy membrane externally that is characteristic
of the septentriohis group of Patrobus.

The penis (median lobe) is non-sclerotized dorsally, as in Patrobus
and Diplous, but not entirely open, as in Deltotnerus , Platidiolus
(Patroboidea) , and related genera. The hook-shaped basal part is a
common feature of all Patrobini. The apex is long and slender as

* Manuscript received by the editor October 26, 1961.

7

8

Psyche

[March

in Patrobus longicornis and foveocollis, but without the subapical left
side tooth of longicornis and, compared with foveocollis, symmetric.
The shape of the apex, however, generally has little generic value in
carabid beetles.

The internal sac of the penis is less complicated than in most
Patrobus. The “apical plate” (Darlington; “Manschette”, Kiihnelt,
1941), forming the bottom (anterior part) of the eversible sac when
in repose, is slightly spiral, but not at all to the extent of the sibiricus
group of Diplous (Kiihnelt). It is not prolonged into a spine, as in
the North American Diplous (contrasted with the Siberian repre-

Fig„ 1. Platypatrobus lacustris Dari. Penis (c) and parameres (a, right;
b, left). The dark parts of the internal sac are not isolated sclerites but
thickened margins of lamellae.

sentatives of the genus). It lacks accessory spine(s), in contrast to
Patrobus. Presence of spines was used as a generic character of
Patrobus by Darlington, but there is no spine in the Palaearctic P.
assimilis Chd.

The most characteristic feature in the internal sac of Platypatrobus
is the dorso-basal part of the “apical plate”, which protrudes into

1962]

Lindroth — Platypatrobus

9

the softwalled section of the dorsum. In side view it resembles a
stalked button, but in dorsal view it is revealed as the side-face of
a horizontal though somewhat elevated and irregular lamella with
thickened margin. It seems to have no direct counterpart in any
species of Patrobus or Diplous.

In summary, the male genitalia of Platypatrobus provide no clue
to its taxonomic relationship with its two closest relatives, Patrobus
and Diplous. The intrageneric variation of the internal sac is so great
in both genera that no single consistently separating detail could be
discovered. And the male genitalia of Platypatrobus itself show little
peculiarity. Possibly, the general simplicity of the internal sac, with
complete lack of spine-like sclerites, could be regarded as a sign of
primitive organization. But it should be remembered that the same
applies to the depressus group of Diplous (Kuhnelt, 1941) as well
as to Patrobus assimilis.

The taxonomic position of genus Platypatrobus , therefore, has
still to be judged on external characters. In these, the genus is clearly
closer to Patrobus than to Diplous. This is shown by the protruding
eyes and the strongly constricted neck, as well as by several structural
details of the prothorax: the central furrow is deepened at base; the
basal foveae are deep and well defined ; the front margin is strongly
elevated (almost as in P. longicornis) , delimited basad by a deep
transverse impression which is coarsely punctured (as in P. septen-
trioms) and prolonged laterally to front-angles as an engraved line
(as in P. longicornis , septentrionis, and foveocollis) . However, as in
Diplous, there is no defined latero-basal carinula inside the hind angles.

There are also good characters separating Patrobus and Diplous
in the marginal region of the elytra1, not observed by earlier students
(Darlington, 1938; Kuhnelt, 1941 ; Lindroth, 1961) : (1) the raised
lateral bead is complete to apex in Patrobus, rather suddenly dis-
appearing well before apex in Diplous 2; (2) the 9th stria is better
developed in Diplous, still evident at the level of the meso-coxae,
whereas in Patrobus it disappears anteriorly well behind this point;
(3) the marginal row of setiferous punctures (on 9th interval) is
almost continuous in Diplous, consisting of about 20 (19-24) punc-
tures, whereas in Patrobus it is zb interrupted at middle and the
number of punctures is reduced (8-14). Jeannel (1941, p. 565ft)

Studied in the North American and (Patrobus) Scandinavian representa-
tives of the two genera.

This, however, is not quite constant in Diplous. An apparently undescribed
species from the Kolyma River district, E. Siberia (V. N. Kurnakov). has
the elytra margined to apex.

10

Psyche

[March

has used the last character as the main distinction between the “sub-
families” Deltomeritae (represented in North America by genus
Platidiolus, syn. Patroboidea ) and Patrobitae, in spite of the fact
that Diplous is referred to the latter group which he defines as having
constantly 8 setiferous punctures! In the three points mentioned
above (1-3), Platypatrobus agrees with Patrobus in points 1 and 2
but is intermediate in point 3, the marginal row consisting of 16
punctures on each side in the male investigated and being less inter-
rupted at middle than in Patrobus .

Platypatrobus , though generically distinct, is closely allied to
Patrobus and no “missing link” to Diplous. Its extreme rarity and
restricted distribution undoubtedly give the impression of a relict,
on the verge of extinction. On the other hand, it does not seem
possible to tell whether Platypatrobus is the phylogenetically older
genus. It should perhaps be regarded as “more simple” in general
construction (lack of prothorax carinula, nearly continuous marginal
row of setiferous punctures of elytra, simple internal sac of penis),
but evolution sometimes goes toward simplification. How often is it
actually defensible to state, without fossil evidence, what is “primi-
tive” and what “derivative”?

Literature Cited

Darlington, P. J., Jr.

1938. The American Patrobini (Coleoptera, Carabidae). Entomologica
Americana (Brooklyn) (new series), 18:135-183.

Jeannel, R.

1941. Coleopteres Carabiques, 1. Faune de France (Paris), 39:1-571.
Kuhnelt, W.

1941. Revision der Laufkafergattungen Patrobus und Diplous. Ann.
Naturh. Mus. (Wien), 51:151-192.

Lindroth, C. H.

1961. The Ground-Beetles (Carabidae, excl. Cicindelinae) of Canada
and Alaska, 2. Opusc. Ent. (Lund), Suppl. XX: 1-200.

THE SPIDER GENERA STEATODA AND
ENOPLOGNATHA IN AMERICA*
(ARANEAE, THERIDIIDAE)

By Herbert W. Levi

Museum of Comparative Zoology, Harvard University

The previous revisions (Levi, 1957a, 1957b) of the two genera
Enoplognatha and Steatoda considered only the North American
species. Since the revisions were published, large South American
collections have become available and the types of South American
species could be consulted.

The majority of species of both genera are found in the north
temperate zone and are fairly well known. The additional species
described here from the neotropical area are sometimes intermediate
between the two genera. Enoplognatha peruviana may lack the tooth
on the posterior margin of the chelicerae as in Steatoda species. The
males of several Steatoda (e.g. S. andina ) have the chelicerae enlarged
as is characteristic of Enoplognatha . South American Enoplognatha
species are found only in southern Peru and northern Chile ( Map 1 ) .
The genus has no representatives in Central America or the West
Indies. Steatoda species are found in all parts of South America, with
several endemic species and several that are widespread (S. ancorata ,
S. grossa, S. moesta) . Unlike Anelosimus species (Levi, in press)
Steatoda species cross the desert or mountain barrier into Chile
(Map 2).

The types of species could be examined through the hospitality and
cooperation of Dr. G. Owen Evans and Mr. D. Clark of the British
Museum (Natural History) ; Prof. G. C. Varley of the Hope Depart-
ment of Entomology, Oxford; Dr. L. Brundin of the Natural History
Museum, Stockholm; Prof. M. Biraben of the Museo de la Plata;
Prof. M. Vachon of the Museum National d’Histoire Naturelle,
Paris; Mr. J. Proszynski of the Polish Academy of Sciences, Warsaw;
Dr. L. Forcart of the Naturhistorisches Museum, Basel and Dr. M.
Beier of the Naturhistorisches Museum, Vienna. I would like to
thank the following for the loan of specimens for study: Prof. M.
Vachon, Museum National d’Histoire Naturelle (MNHN) ; Mr.
J. Kekenbosch of the Institut Royal des Sciences Naturelles, Brussels
(ISNB) ; Dr. W. J. Gertsch of the Ame rican Museum of Natural
History (AMNH) ; Dr. O. Kraus of the Senckenberg Museum,

* Manuscript received by the editor November 20, 1961.

12

Psyche

[March

Frankfurt (SMF) ; Dr. E. S. Ross of the California Academy of
Sciences (CAS) ; Dr. H. Exline (Mrs. D. Frizzell) for specimens
from northern Peru and Ecuador collected by herself and others;
Dr. R. X. Schick of California; Miss H. Zapfe (Mrs. G. Mann)
of Santiago de Chile; Dr. J. Abalos of Santiago del Estero and Prof.
H. Stahnke of Arizona. A trip to European museums to examine
types was made possible by a National Science Foundation Grant
(G-4317) and the research was supported by a grant from the
National Institutes of Health (E-1944).

Enoplognatha Pavesi

Enoplognatha Pavesi, 1880, Ann. Mus. Civ. Stor. Nat. Genova, vol. 15, p. 325.

Type species by original designation and monotypy Theridion mandibular e

Lucas.

Diagnosis. Medium sized to large theridiid spiders. Enoplognatha ,
like Steatoda, has a large colulus and a subspherical abdomen. Females
have a tooth on the posterior margin of the chelicerae. Males usually
have enlarged chelicerae, with enlarged teeth and have the paracym-
bium on the margin of the cymbium (Levi and Levi, 1962).

Species misplaced. Enoplognatha dubia Chamberlin, 1916 — Ane-
losimus studio sus (Hentz, 1850). Enoplognatha triangulifera Simon,
1902 = Aimlosimus recurvatus (Tullgren, 1901).

Map 1. Distribution of South American Enoplognatha species.
Key to American Species of Enoplognatha

1 a. Species found in South America 2

lb. Species found in north of Mexico 5

1962]

Levi — Steatoda and Enoplognatha

13

2a.
2b.
3 a.

3b.

4a.

4b.

5a.

5b.

6a.

6b.

7a.

7b.

8a.

8b.

9a.

9b.

10a.

10b.
1 ia.

1 ib.

12a.

Epigynum with a large central depression (Figs. 12, 13, 15) 3

Epigynum otherwise 4

Epigynum with central depression surrounded by a lip (Figs.

12, 13); ducts coiled (Fig. 1 1 ) ; Peru

E. ju nine n sis (Keyserling)

Epigynum with central depression lacking a lip posterior (Fig.

15); ducts with a sclerotized loop (Fig. 14) ; Chile

E. zapfei n. sp.

Fused ducts extending anterior and surrounding seminal recep-
tacles (Fig. 9) E. puno n. sp.

Fused ducts not extending anterior of seminal receptacles (Fig.

7) E. peruviana Chamberlin

Yellow- white coloration with black or red lines on carapace and
abdomen (1957a, fig. 4); introduced in northeastern and

Pacific coast states E. ovata (Clerck)

Gray, brown or black coloration 6

Males 7

Females 14

Lacking paracymbium on cymbial margin (Fig. 4) ; Arizona,

southeastern California E. maricopa n. sp.

With paracymbium on cymbial margin 8

Conductor with a stalked distal apophysis (1957a, fig. 13-16) 9

Conductor otherwise 10

Conductor apophysis widened distally and truncate (1957a, figs.

15, 16) ; California E. selma Chamberlin and Ivie

Conductor apophysis pointed distally (1957a, figs. 13, 14);

introduced in Oregon E. thoracica (Hahn)

Chelicerae with two subequal teeth on posterior margin (1957a,

figs. 24, 25) 11

Chelicerae with one large mesal tooth (1957a, figs. 46-48) 12
Abdomen with a spotted dorsal pattern ( 1957a, fig. 33) ; median
apophysis in ventral view only in distal half of alveolus (1957a,
fig. 26) ; widespread throughout United States and southern

Canada, Mexico E. marmorata (Hentz)

Abdomen with a foliate dorsal pattern ( 1957a, fig. 37) ; median
apophysis in ventral view extending into proximal half of
alveolus ( 1 95 7a> bg- 28) ; east of Rocky Mountains to New-
foundland E. tecta (Keyserling)

Ectal portion of conductor with long axis almost parallel to axis
of cymbium (1957a, figs. 42-45); western states, rare in
east E. joshua Chamberlin and Ivie

Psyche

[March

12b. Long axis of ectal portion of conductor at almost right angles

to axis of cymbium ( 1957a, figs. 38, 40) 13

13a. Embolus more proximal, conductor with a longer ectal portion

(figs. 38, 39) ; South Dakota to Utah at lower elevations

E. wyuta Chamberlin and Ivie

13b. Embolus more distal, conductor with a shorter ectal portion
(1957a, figs. 40, 41) ; Alaska to Greenland, in Rocky Moun-
tains above timberline and in spruce-fir forest, Minnesota,

Pennsylvania. New England E. intrepida (Snrensen)

14a. Epigynum with a light posterior transverse bridge (1957a, fig.
55) y western states, rare in east . E. joshua Chamberlin and Ivie

14b. Epigynum otherwise 15

15a. Epigynum with an oval depression having dark marks of

openings on the anterior of the depression (1957a, fig. 20);

California E. selma Chamberlin and Ivie

15b. Epigynum otherwise 16

1 6a. Epigynum with a transverse swelling, with a seam on the

anterior and openings on each end of seam (1957a, fig. 18);
introduced in Oregon E. thoracica (Hahn)

1 6b. Epigynum otherwise 1 7

17a. Epigynum with distinct dark marks indicating openings 19

17b. Epigynum with diffuse dark marks 18

18a. Swelling of epigynum divided by a transverse groove (1957a,
figs- 31, 32), spotted dorsal abdominal pattern (1957a, fig. 33) ;
widespread throughout United States and southern Canada,

Mexico E. inarm or ata (Hentz)

1 8b. Epigynum swelling undivided with only a slight depression
posterior ( 1957a, figs. 35, 36) , foliate dorsal pattern of abdomen
( 1 95 7a> fig- 37) y east of Rocky Mountains to Newfoundland

E. tecta (Kevserling)

19a. Dark mark of epigynum anterior to a light swelling (Fig. 2) ;

Arizona, southeastern California E. maricopa n. sp.

19b. Dark mark anterior to a transverse ridge 20

20a. Posterior edge of dark mark of epigynum straight (1957a, fig.

50) ; South Dakota to* Utah at lower elevations

E. wyuta Chamberlin and Ivie

20b. Dark mark dumb-bell shaped (1957a, fig. 52); Alaska to
Greenland, in Rocky Mountains above timberline and in spruce-

fir forest, Minnesota, Pennsylvania, New England

E. intrepida ( Sorensen )

1962]

Levi — Steatoda and Enoplognatha

15

Enoplognatha selma Chamberlin and Ivie

Additional record. California. Los Angeles Co.: Big Tujunga
Canyon, San Gabriel Mts., 3 June 1955 (R* X. Schick).

Enoplognatha marmorata (Hentz) and
Enoplognatha tecta (Keyserling)

Although these are common species in the eastern United States,
vve still have few observations on their habits. Mature E. tecta have
been collected on a garage door in Cambridge, Massachusetts and
numerous specimens were collected rolled up in leaves in herbaceous
vegetation along forest edge about 40 cm above ground in southern
Minnesota, in an area where E. marmorata was collected under boards
in gardens.

Additional records of E. marmorata. Canada. British Columbia:
Muskeg near Little Prairie (D. Rounds). Mexico. Hidalgo: Guer-
rero Mill (W. M. Mann).

Enoplognatha maricopa new species
Figures 1-5

Type. Male from Phoenix, Maricopa County, Arizona, 1 1 March
i960 (H. L. Stahnke) in the Museum of Comparative Zoology.

Description. Male carapace, sternum, legs red-brown. Abdomen
mottled purplish. Sclerotized portion of epigastric area and plate
above pedicel on abdomen red-brown. Carapace with a circular
thoracic depression. Anterior median eyes smallest, anterior laterals
largest. Anterior median eyes a little less than a diameter apart, their
radius from laterals. Posterior median eyes a little less than a diameter
apart, one diameter from laterals. Chelicerae heavy but not elongated,
with one large anterior tooth (Fig. 5). Total length 3.4 mm. Cara-
pace 1.8 mm long, 1.3 mm wide. First patella and tibia 1.4 mm;
second 1.3 mm; third 1.0 mm. Fourth femur 1.3 mm; patella and
tibia 1.6 mm; metatarsus 1.2 mm; tarsus 0.5 mm.

Female carapace, sternum and legs mottled brown or yellowish.
Abdomen with a black pattern on dorsum as in Enoplognatha wyuta
Chamberlin and Ivie (1957a, fig. 53). Anterior median eyes smaller
than others. Anterior medians one and one-half diameters apart, one
diameter from laterals. Posterior median eyes one diameter apart,
a little more than one diameter from laterals. Chelicera with one
anterior tooth and three denticles between tooth and base of fang.

i6

Psyche

[March

Posterior margin of chelicera with one small tooth (missing on one
side of one specimen). Total length one female 2.9 mm, another 3.8
mm. Carapace of the latter 1.5 mm long, 1.2 mm wide. First femur
1.6 mm; patella and tibia 1.8 mm; metatarsus 1.0 mm; tarsus 0.8
mm. Second patella and tibia 1.5 mm; third 1.1 mm; fourth 1.8 mm.

Comments. It is not certain that the male and female described
here belong together. However, since large collections of spiders
are available from southern and southeastern Arizona and no speci-
mens of Enoplognatha have been found, I assume that the distribution
of Enoplognatha maricopa is north from the type locality, where the
females were found, a poorly collected area.

Diagnosis. The palpus differs from other North American species
in that there is no paracymbium on the margin of the cymbium (Fig.
4). However, the transparent edge of the cymbium is difficult to see.
On the mesal side of the bulb is a sclerite, probably part of the conduc-
tor, which has two projections, one toward the venter and one toward
the distal end of the palpus (Fig. 3). The female differs from
Enoplognatha intrepida (S0rensen) in that it has two dark openings
connected by a black groove; posterior to the groove is a swelling
having a lighter center (Fig. 2).

Records. California. Placer Co., near Emigrant Gap, 8 July 1956,
2? (V. Roth, W. J. Gertsch, AMNH) ; Tahoe City, 8 July 1956,
? (W. J. Gertsch, V. Roth, AMNH).

Enoplognatha peruviana Chamberlin
Figures 6-8

Enoplognatha peruviana Chamberlin, 1916, Bull. Mus. Comp. Zool., 60:232,
pi. 16, figs. 8-11, 2. Female type from Urubamba, 3100 m alt., [Cuzco],
Peru, in the Museum of Comparative Zoology, examined.

Description. Carapace, sternum, legs red-brown. Abdomen black
with a dorsal pattern as in Enoplognatha tecta (Keyserling) , (1957a,
fig. 37). Anterior median eyes two-thirds diameter apart, one
diameter from; laterals. Posterior median eyes two-thirds diameter
apart, one and one-half diameters from laterals. Anterior median
eyes slightly smaller than others. Chelicera with three large teeth
on anterior margin, the first and third larger than the middle one;
none on posterior margin of the type. However, another specimen
seems to have the posterior tooth. Colulus relatively small with two
setae. Total length of female type 6.5 mm. Carapace 3.6 mm long,
2.4 mm wide. First femur, 2.7 mm; patella and tibia, 3.4 mm;

1962]

Levi — Steatoda and Enoplognatha

17

metatarsus, 2.0 mm; tarsus, 1.1 mm. Second patella and tibia, 3.0 mm;
third, 2.3 mm; fourth, 3.2 mm.

The male illustrated (Fig. 6) may belong to this species; this is
uncertain, as is the locality where it has been found.

Records. Peru. “Pampa Machei, 4400 m,” Feb. 1947, cf (Brecht,
AMNH). Cuzco: Cuzco, 3500 m, July 1947, 9 (W. Weyrauch,
AMNH).

Enoplognatha puno new species
Figures 9, 10

Type. Female from 32 km north of Desaguadero, Puno, Peru,
27 Feb. 1951 (E. I. Schlinger, E. S. Rcss), in the California Acad-
emy of Sciences. The specific name is a noun in apposition, after
the type locality.

Description. Carapace, sternum, legs dark brown. Abdomen
brownish black with an indistinct dorsal pattern as in E. peruviana.
Venter with some white pigment on each side. Anterior median eyes
much smaller (two-thirds their diameter) than oval posterior eyes.
Anterior median eyes two-thirds diameter apart, more than their
diameter from laterals. Posterior median eyes less than their shorter
diameter apart, two and one-half times their shorter diameter from
laterals. Chelicerae with three large anterior teeth and a minute
denticle on posterior margin. Total length 6.1 mm. Carapace 2.6
mm long, 2.0 mm wide. First femur 2.3 mm; patella and tibia 2.9
mm; metatarsus 1.7 mm; tarsus 1.0 mm. Second patella and tibia 2.5
mm; third 2.1 mm; fourth 3.0 mm.

Diagnosis. Only the epigynum (Fig. 10) and ducts of the internal
genitalia (Fig. 9) separate this species from E. peruviana ; the pattern
and the structure are very similar. It is possible that this species
belongs to a geographical race of E. peruviana.

Enoplognatha juninensis (Keyserling)

Figures 11-13

Lithyphantes juninensis Keyserling, 1884, Die Spinnen Amerikas, Theridiidae
2(1): 143, pi. 6, fig. 90, 9. Female lectotype here designated from

Maraynioc, Junin, Peru in the Polish Academy of Sciences, Warsaw,
examined.

Enoplognatha juninensis, Simon, 1894, Histoire Naturelle des Araignees, 1:
578.

The specimen examined from Pasco, Peru has the abdomen dark
gray with a light area on each side on dorsum ; the venter is also dark

Psyche

[March

gray with a light area on each side between epigynum and spinnerets.
The chelicerae have three large teeth on the promargin, a minute

Figs. 1-5. Enoplognatha maricopa new species. 1. Female genitalia, dorsal
view. 2. Epigynum. 3, 4. Left palus. 3. Ventral view. 4. Ectal view. 5. Left
male chelicera, posterior view.

Figs. 6-8. E. peruviana Chamberlin. 6. Palpus (doubtful determination).
7. Female genitalia, dorsal view. 8. Epigynum.

Figs. 9-10. E. puno new species. 9. Female genitalia, dorsal view. 10.
Epigynum.

Figs. 11-13. E. juninensis (Keyserling) . 11. Female genitalia, dorsal view.
12, 13. Epigynum.

Figs. 14-15. E. zapfei new species. 14. Female genitalia, dorsal view. 15.
Epigynum.

tooth on the posterior margin. The colulus is large, almost trans-
parent white, with two setae from the base.

Records. Peru. Pasco: near Huayllay, 4400 m, 10 Aug. 1953,

1962]

Levi — Steatoda and Enoplognatha

19

$ (H. W. Koepcke, SMF). Junin. Maraynioc, $ paratype with
9 paratype (BMNH).

Enoplognatha zapfei new species
Figures 14, 15

Type. Female from Putre, Province Tarapaca, Chile, Feb. 1948
(Aviles), in the Museum of Comparative Zoology. The species is
named after Miss H. Zapfe of Santiago de Chile.

Description. Carapace dark yellow, darker around anterior median
eyes. Chelicerae red-brown. Sternum, legs dark yellow. Abdomen
gray with darker gray mottled pattern on dorsum and a pair of
longitudinal lighter bands between genitalic area and spinnerets.
Anterior median eyes slightly smaller than others. Anterior lateral
eyes on slight tubercles. Anterior median eyes slightly less than one
diameter apart, a little more than one diameter from laterals.
Posterior median eyes three-quarters diameter apart, two diameters
from laterals. Chelicerae with three teeth on anterior margin, a blunt
indistinct tooth posterior. Total length 9.2 mm. Carapace 4.5 mm
long, 3.4 mm wide. First femur, 4.1 mm; patella and tibia, 5.0 mm;
metatarsus, 3.4 mm ; tarsus, 1.6 mm. Second patella and tibia, 4.3 mm;
third, 3.7 mm; fourth, 5.1 mm.

Diagnosis. This large species of Enoplognatha can be separated
from E. juninensis by the lack of posterior lip in the opening of the
depression of the epigynum (Fig. 15) and by lack of the internal
coiled duct. The ducts of E. zapfei are heavily sclerotized adjacent
to the seminal receptacles (Fig. 14).

Steatoda Sundevall

Steatoda Sundevall, 1833, Conspectus Arachnidum, p. 16. Type species desig-
nated by Thorell, 1869, On European Spiders, p. 93. S. castanea (Clerck,
1757).

Diagnosis. Medium-sized to large theridiid spiders, usually dark
in coloration. Colulus very large. Chelicerae armed with a tooth
or several teeth on anterior margin. But unlike Enoplognatha , it
has no teeth on the posterior margin of female chelicerae. Abdomen
subspherical. Males often with enlarged chelicerae and a sclerotized
ring around pedicel on anterior end of abdomen. The abdomen of
most species has a white line around anterior of dorsum, in addition
to other lines or spots (Levi and Levi, 1962).

Note. The common species of North America were described in

20

Psyche

[March

a previous paper (Levi 1957b). In this paper the ventral view of the
genitalia (cleared epigynum) was illustrated, not the dorsal as in
most of my papers on theridiid spiders. Gertsch (i960) has taken
issue with my treatment of species of the “fulva” group in the previous
paper and has named several new species. However, insufficient
ecological and life history data are available for the possible species
involved to permit making a final decision now (Levi, i960). Thus
the only species recognized are those of my previous revision.

Steatoda brasiliana Keyserling, 1884, Die Spinnen Amerikas
Theridiidae, 2(1) : 115, pi. 5, fig. 75, cf . Male type from Rio de
Janeiro, Brazil in the Naturhistorisches Museum, Vienna, examined
is a male Steatoda bipunctata (Linn.) from Europe (NEW SYN-
ONYMY). The female described by Keyserling in 1886 as S.
brasiliana is probably a specimen of S. inoesta (Keyserling).

Species misplaced : Lithyphantes juninensis Keyserling— Enoplog-
natha juninensis (Keyserling).

Steatoda rubra Keyserling, 1886, Die Spinnen Amerikas, Theri-
diidae, 2(2) : 239, pi. 20, fig. 294, $. Female type from Blumenau,
Brazil — — T heridion rubra (Keyserling).

Key to American Species of Steatoda

1 a. Epigynum with a prominent, wide, transverse bridge; areas
anterior and posterior to bridge shallow depressions (1957b,
figs. 43, 44, 46, 47, 49, 50) ; palpus with thread-like embolus
on venter, evenly curved, with a membrane inside embolus loop
from base supporting a portion of embolus (1957b, figs. 32-41) ;

North America S. fulva group

ib. Epigynum without transverse bridge or if with bridge, then
areas anterior and posterior to' it not depressed ; embolus other-
wise or if similar South American 2

2a. Seminal receptacles thickened, sclerotized, coiled ends of ducts
(1957b, figs. 88, 92, 98, 1 16); epigynum having a U-shaped
depression (1957b, figs. 105, 107-109) or a shallow circular
depression containing anterior a deeper depression on each side
of a septum (rarely absent) (1957b, figs. 93, 97, 99-103) or
epigynum as in 1957b, fig. 89; palpus with a prominent U-
shaped radix on mesal side (1957b, figs. 119, 124, 129, 134,
145, 155) ; North America S. bipunctata group

Map 2. Distribution of South American Steatoda species.

22

Psyche

[March

2b. Seminal receptacles oval or spherical; epigynum and palpus

otherwise or if similar, not North American 3

3a. Males 4

3b. Females 17

4a. Palpus with a large mesal projecting radix (1957b, figs. 59,

68) 5

4b. Palpus otherwise 6

5a. Embolus short, radix longer than half length of bulb (1957b,

fig. 59) > southern Canada to central Mexico

S. albomaculata (DeGeer)

5b. Embolus long, radix less than half length of bulb (1957b, fig.
68) ; southern Canada to central Mexico, common in north-

eastern states S. americana (Emerton)

6a. Palpal tibia one and one-half times length of cymbium (1957b,
fig- 76) ,* probably introduced and widespread in United States,

rare in South America S. triangulosa (Walckenaer)

6b. Palpal tibia as long as or shorter than cymbium 7

7a. Palpal embolus visible as a prominent loop supported by con-

ductor (1957b, fig. 73); probably cosmopolitan, Florida,

Caribbean S. erigoniformis (O. P.-Cambridge)

7b. Palpus otherwise 8

8a. Palpus with a transverse, ventral embolus base and a prominent

short thread-shaped embolus portion (1957b, fig. 74); cosmo-
politan ; coast states, wide-spread in Mexico and South America

S. grossa (C. L. Koch)

8b. Palpus otherwise 9

9a. Palpal embolus ventral, with a short distal hook as in 1957b,

fig. 18; southern Mexico to southern Brazil

S. moesta (O. P.-Cambridge)

9b. Palpus otherwise 10

10a. Palpus with biforked embolus (Fig. 22) ; northern Colombia

S. maria n. sp.

10b. Palpus otherwise 11

lla. Palpus with spine on base of embolus and complex U-shaped

radix (1957b, fig. 11; Figs. 44-48); southern Mexico to
southern Patagonia S. ancorata (Holmberg)

llb. Palpus otherwise 12

1 2a. Palpus with prominent ventral embolus (Fig. 18); chelicerae

tuberculate (Fig. 19) ; Venezuela, Ecuador, Peru to Chile

S. andina (Keyserling)

12b. Palpus and chelicerae otherwise 13

1962]

Levi — Steatoda and Enoplognatha

23

13a. Palpal embolus appearing as a tightly coiled structure supported
by the conductor (Fig. 30). Minas Gerais, Brazil

S. diamantina n. sp.

13b. Palpus otherwise 14

14a. Palpus with a long narrow radix (in ventral view) whose axis
is parallel to that of the cymbium (Fig. 27) ; Bolivia, Chile,

Argentina S. sabulosa (Tullgren)

14b. Palpus otherwise 1 5

15a. Palpal embolus small, on distal, ventral side of bulb (1957b,

fig. 14); Mexico S. quaesita (O. P.-Cambridge)

15b. Palpus with embolus on ectal side or hidden in ventral view.. 16
16a. Palpus with prominent ventral radix (1957b, fig- 27) ; southern

California, Texas, Mexico S. transversa (Banks)

16b. Palpus otherwise (1957b, fig. 30) ; Georgia, Gulf states, West

Indies, Mexico to Venezuela

S. quadrimaculata (O. P.-Cambridge)

17a. Epigynum with a wrinkled area (Figs. 39, 43) 18

17b. Epigynum with all parts smooth 20

1 8a. Epigynum with a shield having its anterior margin lobed
(1957b, fig. 67); ducts coiled (1957b, fig. 66); southern
Canada to central Mexico; common in northeastern states ....

S. americana (Emerton)

18b. Epigynum without lobed anterior margin; ducts not coiled .. 19
19a. Epigynum as in Figure 39; Chile S. porteri (Simon)

19b. Epigynum as in Figure 43; southern Mexico to southern

Patagonia S. ancorata (Holmberg)

20a. Epigynum with a light, transverse, oval, surrounded on sides
and behind by a raised area (Fig. 26) ; Bolivia, Chile, Argentina
S. sabulosa (Tullgren)

20b. Epigynum otherwise 21

2 1 a. Epigynum' with a median lobe extending from anterior (Figs.

17, 2l) 22

2ib. Epigynum without anterior lobe 27

22a. Posterior margin of epigynum heavily sclerotized and projecting

in middle (Fig. 21), northern Colombia S. marta n. sp.

22b. Epigynum otherwise 23

23a. Epigynum with a depression along posterior margin (1957b,

figs. 58, 72) 24

23b. Epigynum otherwise 25

24a. Posterior median depression bordered on each side by a thorn

24

24b.

25a.

25b.

26a.

26b.

27a.

27b.

28a.

28b.

29a.

29b.

30a.

30b.

31a.

31b.

32a.

32b.

33a.

33b.

34a.

34b.

Psyche

[March

( 1 957b, fig. 58) ; southern Canada to central Mexico

S. albomaculata (De Geer)

Posterior depression not bordered on sides (1957b, fig. 72);

cosmopolitan ; Florida, Caribbean

S. erigoniformis (O. P.-Cambridge)

Epigynum with two widely separated depressions (Figs. 34,

35) ; southern Brazil, Paraguay, Argentina

S. iheringi (Keyserling)

Epigynum otherwise 26

Depression of epigynum bordered all around (Fig. 17) ; Vene-
zuela, Ecuador, Peru to Chile S. andina (Keyserling)

Depression on each side of lobe not bordered behind (1957b,
figs. 84, 85) ; cosmopolitan, coast states, widespread in Mexico

and South America S. grossa (C. L. Koch)

Epigynum with a narrow, transverse bridge (1957b, fig. 81) ;
probably introduced and widespread in United States and South

America S. triangulosa (Walckenaer)

Epigynum otherwise 28

Epigynum with a median dark spot (1957b, fig. 17) ; southern

Mexico to southern Brazil S. moesta (O. P.-Cambridge)

Epigynum otherwise 29

Epigynum with a posterior directed lobe, posterior to the opening

(Fig. 32) ; Peru S. c hi tic hip c n. sp.

Epigynum otherwise 30

Epigynum with a slit-like opening on each side ( 1957b, fig. 20) ;

Mexico ' S. saltensis Levi

Epigynum otherwise 31

Epigynum with shallow longitudinal oval depression (Fig. 29) ;

Minas Gerais, Brazil S. diamantina n. sp.

Epigynum otherwise 32

Epigynum with a deep transverse depression (Fig. 24) ; Peru

S. variipes (Keyserling)

Epigynum otherwise 33

Epigynum with two black spots on posterior margin (1957b,

fig. 24) ; southern California, Texas, Mexico

S. transversa (Banks)

Epigynum otherwise 34

Epigynum with a pair of shallow depressions each with a black
spot (1957b, fig. 29); Georgia, Gulf States, West Indies,
Mexico to Venezuela .... S. quadrimaculata (O. P.-Cambridge)
Epigynum as in 1957b, fig. 22; Mexico .. S. autumnalis (Banks)

1962]

Levi — Steatoda and Enoplognatha

25

Steatoda moesta (O. P. -Cambridge)

Asagena moesta O. P. -Cambridge, 1896, Biologia Centrali-Americana,
Araneidea, 1 : 209, pi. 25, fig. 4-, $ . Female type from Guatemala in the
British Museum, Natural History.

Steatoda moesta, Levi, 1957, Bull. Mus. Comp. Zool., 117:3, p. 379, figs. 15-18,

9,$.

Distribution. Southern Mexico to southern Brazil.

Additional Records. Venezuela: “Caracas, Tovar,” 1888 (E.

Simon, MNHN). Peru. Hudnuco: 27 km N of Huanuco, Dec.
1954 (E. I. Schlinger, E. S. Ross, CAS). San Martin: Tarapoto
(MNHN); Moyobamba (MNHN). Brazil. Rio de Janeiro:
Mencles (Eidmann, SMF) ; Teresopolis (MNHN).

Steatoda erigoniformis (O. P.-Cambridge) , new combination

Theridion erigon forme O. P.-Cambridge, 1872, Proc. Zool. Soc. London,
p. 284. Female, male syntypes from Jordan in the Hope Department of
Entomology, Oxford, examined.

Asaganella erigoniformis, Schenkel, 1937, Festschrift E. Strand, 3:381, fig.
3, $ .

Lithyphantcs septemmaculatus Keyserling, 1884, Die Spinnen Amerikas,
2(1): 141, pi. 6, fig. 88, 9. Female syntypes from “Denver in Columbia”
collected by Marx. (Marx’s labels are often wrong and the Denver type
locality is undoubtedly an error). NEW SYNONYMY.

Distribution. Cosmopolitan. Florida, Caribbean, Panama, Vene-
zuela in America. Kaston (1948, Bull. Connecticut Geol. Nat.
Hist. Surv. no. 70:79) reports a specimen from Connecticut. O.
Kraus (in a letter) tells of a specimen from Japan in the Senckenberg
Museum. It also has been found in the Near East.

Additional Records. Panama Canal Zone: Gamboa, 7 Jan. 1958,
c? (A. M. Chickering) . British West Indies: Antigua: St. John,
1 95 5 (A. M. Nadler, AMNH ) . Venezuela. Aragua .-Tovar, 1888,
9 (E. Simon, MNHN). Carabobo: San Esteban, 1888, 9 $ (E.
Simon, MNHN).

Steatoda triangulosa (Walckenaer)

Aranea triangulosa Walckenaer, 1802, Faune Parisienne, 2:207. Type from
Paris, France, lost.

Steatoda triangulosa, Levi, 1957, Bull. Mus. Comp. Zool. 117(3) : 407, figs. 75,
76, 80-82, $ $.

Distribution. Central and southern Europe, southern Russia,
Mediterranean, United States from Massachusetts to Oregon, south
to southern Texas. The city-dwelling habit suggests that it has been

26

Psyche

[March

introduced in America. It has previously been reported from South
America, but only one specimen was seen.

Additional Records. Argentina. Santiago del Ester o: Santiago
del Estero, 12 June 1961, $ (J. Abalos).

Steatoda grossa (C. L. Koch)

Theridion grossum C. L. Koch, 1938, Die Arachniden, 4:112, fig. 321, $.
Female types from Greece.

Steatoda punctilineata Mello-Leitao 1939, Rev. Suisse de Zool., 46:61, figs.
30, 31, $. Two female syntypes from Leones, Argentina, in the Naturhis-
torisches Museum, Basel, examined. NEW SYNONYMY.

Steatoda grossa, Levi, 1957, Bull. Mus. Comp. Zool., 117(3): 404, figs. 74,
83-85, $ 8.

Note. Mello-Leitao (1939) figure 31 is printed upside down;
if it had been turned around the synonymy with the common S. grossa
would have been recognized earlier. Steatoda grossa females can be
confused with females of S. andina. Only details of the posterior
rim of the epigynum seem to separate females of the two species. The
male palpi, however, are very different.

Natural History. The species has been collected in subtropical rain
forests in Oxapampa, and under stones on guano islands.

Distribution. Cosmopolitan; along coast of United States, Mexico,
South America.

Additional records. Ecuador. T'ungurakua: Ambato, June 1943
(H. E., D. L. Frizzell). Peru. Isl. Don Martin (Guano Isl.) (L.
Pena, SMF) ; Piura: Negritos (H. E., D. L. Frizzell). ? Ancash:
Chimbote, Feb. 1953 (W. Weyrauch) . Pasco: Oxapampa, 1600 m,
E of Oroga (W. Weyrauch). Chile. Antofagasta: Taltal (H.
Zapfe). Coquimbo: La Serena (H. Zapfe) ; Los Vilos (H. Zapfe).
Aconcagua: San Filipe (L. Pena, ISNB). Santiago: Santiago (H.
Zapfe). Linares: Linares (L. Pena, ISNB). Malleco: Angol,
1950 (D. S. Bullock). Osorno: Osorno (L. Pena, ISNB).

Steatoda andina (Keyserling) , new combination
Figures 16-19

Lithyphantes andinus Keyserling, 1884, Die Spinnen Amerikas, Theridiidae,
2(1): 132, pi. 6, fig. 82, $ 8 • Male and female syntypes from Junin,

Amable Maria, Lima and San Mateo, Peru in the Polish Academy Sciences,
Warsaw. A female specimen determined by Keyserling, in the British
Museum, Natural History, examined.

Description. Carapace, sternum reddish brown, legs lighter.
Abdomen purplish black with a narrow white line around sides, crossed

1962]

Levi — Steatoda and Enoplognatha

27

in front by a median narrow white line. Very variable, sometimes
with a longitudinal white line or almost black. Venter with a lighter
W-shaped mark and lighter spots on sides. Eyes subequal in size,
anterior median eyes one diameter apart, one and one-half diameters
from laterals. Posterior median eyes about one diameter apart, two
diameters from laterals. Chelicera of female with a blunt, fleshy
tooth on anterior margin, that of male enlarged as in the genus
Enoplognatha , with a large tooth on anterior margin (Fig. 19); there
is no tooth, however, on posterior margin. Measurements of specimens
from Lima, Peru. Total length of female 8.3 mm. Carapace, 3.7 mm
long, 2.9 mm wide. First femur, 4.3 mm; patella and tibia, 5.1 mm;
metatarsus, 3.7 mm; tarsus, 1.6 mm. Second patella and tibia, 4.1
mm; third, 3.3 mm; fourth, 5.0 mm. Total length of male, 8.6 mm.
Carapace, 4.2 mm long, 2.7 mm wide. First femur, 4.7 mm; patella
and tibia, 5.5 mm; metatarsus, 4.3 mm; tarsus, 1.9 mm. Second
patella and tibia, 4.3 mm; third, 3.6 mm; fourth, 5.0 mm.

Variation. The epigynum (fig. 17) is variable in proportion. A
male from Chile has a shorter embolus than males examined from
central Peru.

Diagnosis. The more discrete posterior rim of the depression of
the epigynum (Fig. 17) separates this species from S. grossa (1957b,
figs. 84, 85) with which it may be confused. The palpus (Fig. 18)
is very different from that of S. grossa.

Natural History. Collected in stony grassland near Lake Junin,
shrubs in dry valley in Cajamarca and in Eucalyptus forest in Ancash.

Records. Venezuela. Aragua: Maracay (SMF). Ecuador.

Manabi: Manta (D. L. Frizzell). Guay as: Banos de San Vincente,
Santa Elena Peninsula (R. W. Landes) ; W of Guayaquil (R. W.
Landes) ; Colonche (R. W. Landes). Azuay: 22 km E of Cuenca
(E. I. Schlinger, E. S. Ross). Peru. Piura: El Alto (R. Wells);
Rio Quiroz (H. E., D. L. Frizzell) ; Negritos (H. S. M.) ; Parinas
Valley (H. E., D. L. Frizzell) ; Quebrada Mogollon (H. E., D. L.
Frizzell) ; Mancora (H. E., D. L. Frizzell). Cajamarca: Caja-
marca, 2700 m (W. Weyrauch) between Lives and Mirador, near
San Miguel de Pallaques, 1050 m (H. W. Koepcke, SMF).
Libertad: Otusco (E. I. Schlinger, E. S. Ross, AMNH). San
Martin: Hara, 32 km SE of Moyobamba (F. Woytkowski,
AMNH). Ancash: Puna near Huaras, 4300 m (W. Weyrauch,
AMNH) ; Huaras (H. W. Koepcke, SMF). Huanuco: 27 km S
of San Rafael (E. I. Schlinger, E. S. Ross) ; Tingo Maria, 670 m
(W. Weyrauch, AMNH). Lima: 3 km E of San Mateo (E. I.

28

Psyche

[March

Schlinger, E. S. Ross). San Mateo, 3000 m (W. Weyrauch,
AMNH). Lima (H. E., D. L. Frizzell); Canta, 2800 m (W.
Weyrauch) ; Matucana, 2300-2500 m (H. W. Koepcke, SMF) ;
Atocongo Lomas, rocky hills (H. W. Koepcke, SMF). Junin:
Tarma, 3100 m (W. Weyrauch, AMNH). Laguna Algacocha, near
Laguna Jumn, 4300 m (H. W. Koepcke, SMF). Laguna Junin,
4140 m (H. W. Koepcke, SMF). Cuzco: Cuzco, 4000 m (J. C.
Pallister, AMNH). Arequipa: Atiquipa, Chala, 200 m (W.

Weyrauch). Puno: 10 km S of Oroya (E .S. Ross, E. M. Michel-
bacher, CAS). Chile. Tarapacd: Putre, (Aviles) ; Oasis de

Minimine, (Aviles).

Steatoda marta new species
Figures 20-22

Type. Male from Sierra Nevada de Santa Marta, Magdalena,
Colombia (Dulm), in the Museum National d’Histoire Naturelle,
Paris (no. 18389). The specific name is a noun in apposition, after
the type locality.

Description. Carapace, sternum, legs rich dark red-brown. Abdo-
men purplish black with a median dorsal longitudinal white line and
a white line around the anterior edge of the abdomen. Anterior
median eyes slightly smaller than others, their diameter apart, one
and one-half to two diameters from laterals. Posterior median eyes
one and one-quarter diameters apart, two diameters from laterals.
Laterals separated by about their radius. Chelicerae of male cornicu-
late, with a blunt tooth. Female chelicerae smooth with a short tooth
on anterior margin. First and fourth legs subequal in length. Abdo-
men of male with four sclerotized round spots on dorsum. Total
length of male 7.2 mm. Carapace 3.0 mm long, 2.3 mm wide. First
patella and tibia, 3.5 mm; second, 2.9 mm; third, 2.5 mm. Fourth
femur, 3.0 mm; patella and tibia, 3.7 mm; metatarsus, 2.5 mm;
tarsus, 1.2 mm. Total length of female 8.0 mm. Carapace 3.2 mm
long, 2.5 mm wide. First patella and tibia, 3.5 mm; second, 2.9 mm;
third, 2.6 mm. Fourth femur, 3.3 mm.; patella and tibia, 4.0 mm;
metatarsus, 2.5 mm; tarsus, 1.1 mm.

Diagnosis. This species is very close to Steatoda andina (Keyser-
ling) but differs in having the embolus shorter with a spur, in having
a smaller conductor, and having a median apophysis of different shape
(Fig. 22). The female epigynum differs in having a scape in a shallow
depression. The depression is not bordered. The posterior edge of the

1962]

Levi — Steatoda' and Enoplognatha

29

Figs. 16-19. Steatoda andina (Keyserling) . 16. Female genitalia, dorsal
view. 17. Epigynum. 18. Left palpus. 19. Left male chelicera, anterior view.

Figs. 20-22. S. marta new species. 20. Female genitalia, dorsal view. 21.
Epigynum. 22. Palpus.

Figs. 23-24. S. variipes (Keyserling). 23. Female genitalia, dorsal view.
24. Epigynum.

Figs. 25-27. S. sabulosa (Tullgren). 25. Female genitalia, dorsal view.
26. Epigynum. 27. Palpus.

Figs. 28-30. S. diamantina new species. 28. Female genitalia, dorsal view.
29. Epigynum. 30. Palpus.

Figs. 31-32. S. chinchipe new species. 31. Female genitalia, dorsal view.
32. Epigynum.

Figs. 33-36. S. iheringi (Keyserling). 33. Female genitalia, dorsal view.
34, 35. Epigynum. 34. Type. 35. (Paraguay). 36. Female abdomen, dorsal
view (Paraguay).

30

Psyche

[March

epigynum is heavily sclerotized (Fig. 21). The scape is sometimes
subcircular, sometimes not constricted anteriorly (its sides being
nearly parallel), and is sometimes constricted at its posterior point
and widens again ; its shape seems to be variable in different specimens
collected together.

Records. Several $ paratypes from type collection.

Steatoda sabulosa (Tullgren), new combination
Figures 25-27

Lithyphantes sabulosus Tullgren, 1901, Svenska Exped. Magellanslandern
2(10) : 193, pi. 1, fig. 3, $ . Male type from Santa Cruz, in South Argentina,
in the Naturhistoriska Riksmuseum, Stockholm, examined.

Steatoda albiornata Mello-Leitao, 1940, Rev. Mus. La Plata, n.s., 2:36, fig. 34,
$. Female type from Valcheta, Rio Negro, Argentina in the Museo de la
Plata, examined. NEW SYNONYMY.

Description. (Specimens from Magellanes, Chile). Carapace,
sternum, legs yellow-brown to reddish brown. Abdomen purplish
black ; dorsum of female abdomen with pigment missing in some areas ;
white line around anterior of abdomen on dorsum and median, dorsal,
longitudinal white line which may be broken. Anterior median eyes
slightly smaller than others, two-thirds diameter apart, two-thirds
diameter from laterals. Posterior median eyes a little less than a
diameter apart, one diameter from laterals. The posterior median
eyes of the male are slightly farther from laterals. Male chelicerae
not large, with one large tooth, fang widened and short. Total length
of female 10.5 mm. Carapace 3.2 mm long, 2.9 mm wide. First
patella and tibia 4.0 mm; second 2.9 mm; third 2.7 mm. Fourth
femur 3.5 mm; patella and tibia 4.2 mm; metatarsus 3.6 mm; tarsus
2.3 mm. Total length of male 8.0 mm. Carapace 3.3 mm long, 2.5
mm wide. First patella and tibia 4.0 mm; second 3.6 mm;; third 2.9
mm. Fourth femur 3.5 mm; patella and tibia 4.2 mm; metatarsus
3.1 mm; tarsus 1.4 mm:.

The internal female genitalia (Fig. 25) are heavily sclerotized.
A female from northern Chile has the seminal receptacles slightly
wider apart and the ducts forming a V rather than a circle.

Record. Bolivia. La Paz: La Paz, (MNHN) ; 48 km N of
Potosi, 22 Feb. 1951 (E. S. Ross, A. E. Michelbacher, CAS). Chile.
Antofagasta: Tumbre, 3600 m, Cord. Antofagasta, Dec. 1955 (L.
Pena, ISNB). Santiago: Santiago (SMF) ; Los Valdes, 2000 m,
Cordilleras near Santiago (G. Mann, AMNH). Magallanes :
Laguna Amarga, Natales, Dec. i960, $ S (L. Pena).

1962]

Levi — Steatoda and Enoplognatha

3 1

Steatoda iheringi (Keyserling) , new combination
Figures 33-36

Lithyphantes iheringi Keyserling, 1886, Die Spinnen Amerikas, Theridiidae,
2(2) :240, pi. 20, fig. 295, $. Female type from Rio Grande do Sul, Brazil,
in the British Museum, Natural History, examined.

Lithyphantes canceilatus Mello-Leitao, 1944, Rev. Mus. La Plata, n.s., 3 : 325,
fig. 8, $. Female type from Jose C. Paz, Provincia de Buenos Aires,
Argentina in the Museo de la Plata, examined. NEW SYNONYMY.

Description. Specimen from Paraguay. Carapace yellow-brown,
cephalic area brown, sternum red-brown, legs yellow, ends of tibiae
darker. Abdomen with two series of dark spots on dorsum; sides
and area between posterior spots with some white pigment. Venter
darker with a small white mark just posterior to the epigynum.
Anterior median eyes slightly smaller than others, one diameter apart,
one diameter from laterals. Posterior median eyes three-quarters
diameters apart, one and one-half diameters from laterals. Chelicerae
with one tooth on the anterior margin. Legs quite thick, first equal
in length to fourth. Epigynum (Figs. 34, 35) with a transverse fold.
Internal genitalia difficult to study; the anterior ducts are very
transparent and in the single specimen available, could not be seen
completely. Total length of female type 2.9 mm. Carapace 1.30 mm
long, 1.08 mm wide. First femur, 1.04 mm; patella and tibia, 1.43
mm; metatarsus, 0.84 mm; tarsus, 0.52 mm. Second patella and tibia,
1. 10 mm; third, 0,95 mm; fourth, 1.50 mm.

The palpus illustrated by Figure 48 may belong to the male of this
species. It was collected with females of S. ancorata in Rio Grande
do Sul and belongs to the Keyserling collection in the British Museum.

Records. Paraguay: Taquararapa, Alto Parana, 1908, $ (AM
NH). Argentina. Buenos Aires, $ (Latarte, MNHN).

Steatoda diamantina new species
Figures 28-30

Type. Female from Mina Serinha, Diamantina, Minas Gerais,
Brazil, December 1944 (Mrs. E. Cohn), in the American Museum
of Natural History. The specific name is a noun in apposition, named
after the type locality.

Description. Carapace, sternum, legs red-brown. Abdomen black
except for one or two thin jagged white lines around sides and
anterior. A median, longitudinal, dorsal white line varies in thick-
ness and has several short pairs of lateral branches. Eyes subequal in
size, in female. Anterior median eyes two-thirds diameter apart.

32

Psyche

[March

Posterior median eyes their radius apart and one diameter from
laterals. Anterior median eyes of male slightly larger than others,
one-third diameter apart, one-quarter diameter from laterals. Posterior
median eyes one-third diameter apart, one diameter from laterals.
Chelicerae of male with one tooth on anterior margin. First femora
of male swollen and corniculate on venter. Total length of male 6.6
mm. Carapace 3.3 mm long, 2.4 mm wide. First femur 4.3 mm;
patella and tibia 3.9 mm; metatarsus 2.9 mm; tarsus 1.5 mm. Second
patella and tibia 3.2 mm; third 2.7 mm; fourth 3.9 mm. Total length
of female 6.1 mm. Carapace 2.9 mm long, 2.1 mm wide. First femur
3.5 mm ; patella and tibia 3.9 mm ; metatarsus 3.0 mm ; tarsus 1.4 mm.
Second patella and tibia 2.9 mm; third 2.3 mm; fourth 3.5 mm.

Diagnosis. This species is separated from other Steatoda by the
epigynum (Fig. 29), which has a shallow, oval to round depression,
the anterior end of which is dark and contains the opening. The
male' can be separated by the structure of the palpus (Fig. 30).

Records. Brazil. Santa Catarina: Nova Teutonia, lat 27° 11' S,
long 52° 23' W, ?, cf (F. Plaumann, SMF).

Steatoda chinchipe new species
Figures 31, 32

Type. Female from Rio Chinchipe, San Ignacio, 800 m elev.,
Cajamarca, Peru, July 1948 (W. Weyrauch) in the Museum of
Comparative Zoology. The specific name is a noun in apposition,
after the type locality.

Description. Carapace, sternum, legs brown. Abdomen evenly
purplish black with a wavy line around sides and anterior on dorsum ;
center of dorsum with a fine longitudinal white line having two
crosslines. Anterior median eyes smaller than other eyes. Anterior
median eyes three-quarters their diameter apart, one diameter from
laterals. Posterior median eyes their radius apart, slightly more than
one diameter from laterals. Lateral eyes slightly separated. Total
length 7.5 mml Carapace 3.2 mm long, 2.5 mm wide. First femur
3.5 mm; patella and tibia 4.2 mm; metatarsus 2.7 mm; tarsus 1.3 mm.
Second patella and tibia 2.9 mm; third 2.2 mm; fourth 3.7 mm.

Diagnosis. The epigynum differs from that of other Steatoda. It
has a raised circular area containing an anterior opening with a lip
on three sides (Fig. 32). The internal genitalia (Fig. 31) are heavily
sclerotized.

Records. Ecuador. Tungurahua: Banos, 7 May 1942 (H. E.

1962]

Levi — Steatoda and Enoplognatha

33

Frizzell) ; 32 km SE of Ambato, 8 Feb. 1955 (E. I. Schlinger and
E. S. Ross, CAS) ; Rio Pastaza between Banos and Mapoto, Aug.
1938 (W. C. Macintyre).

Steatoda variipes (Keyserling) , new combination
Figures 23, 24

Theridium variipes Keyserling, 1884, Die Spinnen Amerikas, Theridiidae,
2(1): 93, pi. 4, fig. 61, 9. Female lectotypes here designated from Amable
Maria [Junin], Peru in the Polish Academy of Science, Warsaw, examined.

Description. Carapace rich brown, sternum yellow-brown with
darker margin and darker spot in middle. Legs yellow-brown with
darker bands as wide as lighter areas. Abdomen probably mottled
black. Eyes about subequal in size, anterior median eyes one diameter
apart, one-quarter diameter from laterals; posterior median eyes one
diameter apart, one-third diameter from laterals. Muscle impressions
on abdomen and bases of setae slightly sclerotized. Colulus relatively
small with two setae. Total length of female lectotype 5.0 mm;
carapace 1.0 mm long, 1.9 mm wide. First femur 2.7 mm; patella
and tibia 2.9 mm; metatarsus 1.6 mm; tarsus 0.9 mm. Second patella
and tibia 2.3 mm; third 1.6 mm; fourth 2.4 mm.

Record: One $ paratype with type.

Steatoda porteri (Simon), new combination
Figures 37-39

Lithyphantes porteri Simon, 1900, Rev. Chileana, 4:50. Female type from
Chanarcillo, [Prov. Atacama], Chile, in the Museum National d’Histoire
Naturelle, Paris, examined.

Description. Carapace, sternum, legs yellow-brown; fourth leg
darker than others. Abdomen purplish black with dorsal white marks
(Fig. 37) and a small white spot posterior to genital groove on
venter. Anterior median eyes slightly smaller than others, one and
one-half diameters apart, one diameter from, laterals. Posterior median
eyes one diameter apart, slightly more than one diameter from laterals.
One broad tooth on anterior margin of chelicerae. Fourth leg slightly
longer than first. Epigynum (Fig. 39) with a wrinkled knob pointing
posterior. Total length 4.7 mm. Carapace, 1.8 mm long, 1.4 mm
wide. First patella and tibia, 2.1 mm; second, 1.7 mm; third, 1.4 mm.
Fourth femur 2.0 mm; patella and tibia, 2.4 mm; metatarsus, 1.6
mm; tarsus, 0.9 mm.

Record. Chile. Coquimbo : El Tofo, Sept. 1957 (H. Zapfe).

Figs. 37-39. Stcatoda porteri (Simon). 37. Female abdomen, dorsal view.
38. Female genitalia, dorsal view. 39. Epigynum.

Figs. 40-47. S. ancorata (Holmberg). Fig. 40. Male carapace and
chelicerae. Figs. 41, 42. Female genitalia, dorsal view. 43. Epigynum. 44-47.
Palpus. 44. (northern part of range). 45. (southern Peru). 46. (Bolivia).
47. (southern Chile).

Fig. 48. S. ? iheringi (Keyserling) , palpus.

Steatoda ancorata (Holmberg), new combination
Figures 40-47

Theridium ancoratum Holmberg, 1876, An. Agr. Rep. Argentina, 4:72, fig. 16.
Type from Argentina, lost.

} Lithyphantes vittatus Keyserling, 1884, Die Spinnen Amerikas, Theridiidae,
2(1) : 134, pi. 6, fig. 83. Juv. type from Minas Gerais, Brazil, in the Hope
Department of Entomology, Oxford, examined. NEW SYNONYMY.
Lithyphantes nigrofemoratus Keyserling, 1884, op. cit. 2(1): 139, pi. 6, fig.
87, $. Female type from Monte Rico, [Ayacucho], Peru, in the Polish Acad-
emy of Sciences, Warsaw, examined. NEW SYNONYMY.

Asagena alticeps Keyserling, 1886, op. cit. 2(2) : 4, fig. 136, $ . Male type from
N. Granada [Panama, Colombia, Venezuela] in the British Museum,
Natural History. NEW SYNONYMY.

Lithyphantes laetus O. P.-Cambridge, 1896, Biologia Centrali-Americana,
Araneidea, 1:181, pi. 22, fig. 12, $. Male type from Costa Rica, in the
British Museum, Natural History. NEW SYNONYMY.

[March

39

1962]

Levi — Steatoda and Enoplognatha

35

Asagena patagonica Tullgren, 1901, Svenska Exped. Magellanslandern 2:
194, pi. 15, fig. 4, 9. Female type from Ultima Esperanza and Puerto
Gallegus, Patagonia [? Magellanes, Chile] in the Naturhistoriska Riks-
museum, Stockholm, examined. NEW SYNONYMY.

Asagena melanomela Mello-Leitao, 1944, Rev. Mus. La Plata, n.s., 3:325,
fig. 5, $. Male type from Pergamino, [Buenos Aires], Argentina in the
Museo de la Plata, examined. NEW SYNONYMY.

Steatoda nigrofemorata, Levi, 1957, Bull. Mus. Comp. Zool., 117(3): 377,
figs. 11-13, 9 $ •

Note. This species is the most common Steatoda in Argentina, the
type locality of T. ancoratum. Specimens of this species from Argen-
tina determined by Mello-Leitao and by Biraben were named
Lithyphantes ancoratum. The synonymy of L. vittatus Keyserling
remains uncertain, however, as the type is a juvenile and no specimens
of S. ancorata have been examined from Minas Gerais, the type
locality.

The palpi are variable in structure, (Figs. 44-47). Since the palpal
sclerites are unusually complex, any slight change in position changes
the appearance of the palpus.

Natural History. Specimens have been found “near rockpile in
weathered shale” and “under flat rock” in Bolivia. In Mendoza,
Argentina it has been found in chaparral area.

Distribution. Southern Mexico to southern Patagonia.

Additional records: Venezuela. Dist Federal: Caracas (E. Simon,
MNHN) . Aragua: Tovar (E. Simon, MN HN) . Colombia. Valle:
10 km W of Cali (E. I. Schlinger, E. S. Ross, CAS). Peru.
Huanuco: Monzon Valley, Tingo Maria (E. I. Schlinger, E. S. Ross,
CAS); Huanuco (E. I. Schlinger, E. S. Ross, CAS). Ayacucho:
Ayacucho (W. Weyrauch). Cuzco: Cuzco, 3800m (W. Weyrauch) ;
Huadquina, 1600 m (W. Weyrauch). Puno: 95 km N of Puno
(E. S. Ross, A. E. Michelbacher, CAS); Puno (W. Weyrauch);
near Juliaca, 3900 m (H. W. Koepcke, SMF) ; Mazo Cruz, 3800 m
(L. Pena, ISNB) ; Camacani (L. Pena, ISNB). Bolivia. Chaco
(MNHN). La Paz: Timari, Nevada de Chicani (MNHN); La
Paz 4400 m (R. Walsh, MNHN) ; 65 km; NE of La Paz (R.
Walsh) ; Altiplano near H uayna Potosi Mtn., 5100 m (R. Walsh).
Chiquisaca: 26 km N of Camargo (E. S. Ross, A. E. Michelbacher,
CAS). Potosi: 45 km N of Potosi, 4300 m (E. S. Ross, A. E.
Michelbacher, CAS). Brazil. Parana: Bela Vista (MNHN). Rio
Grande do Sul. (BMNH). Paraguay. Asuncion (MNHN).
Caazapd: Pastoreo (D. Wees). Argentina. Jujuy: 8 km N of
Humacuaca, 2900 m (E. S. Ross, A. E. Michelbacher, CAS). Salta:
Salta (Reimoser) ; Cafayate (M. Biraben). Chaco: Resistencia

36

Psyche

[March

(MNHN). Cordoba: Alta Gracia (Bruch). Mendoza: 8 km SSW
Estacion Cachenta, Dept. Lujan, 1500 m (B. Patterson). Buenos
Aires: Necochea (M. Biraben) ; “Bahia” (MNHN). Chile. Anto-
fagasta: Tumbre, 3600-3700 m, Cord. Antofagasta (L. Pena, ISNB).
Magallanes : Cerro Castillo, Natales (L. Pena).

References

Gertsch, W. J.

1960. The fulva group of the spider genus Steatoda. Amer. Mus.
Novitates, no. 1982, p. 1-48.

Levi, H. W.

1957a. The spider genera Enoplognatha, Theridion and Paidisca in
America North of Mexico. Bull. Amer. Mus. Nat. Hist., 112(1) :
1-123.

1957b. The spider genera Crustulina and Steatoda in North America,
Central America and the West Indies. Bull. Mus. Comp. Zool.
117(3) :367-42+.

“1959” ( 1960) . Problems in the spider genus Steatoda. Systematic Zool.
8:107-116.

The American spiders of the genus Anclosimus. Trans. Amer.
Micros. Soc. in press.

Levi, H. W. and L. R. Levi

1962. The genera of the family T heridiidae. Bull. Mus. Comp. Zool.
127(1) :1-71.

A PERMIAN MEGASECOPTERON FROM TEXAS1

By F. M. Carpenter
H arvard University

A collection of six Permian insects recently received from Dr.
Sergius H. Mamay, of the U. S. Geological Survey, includes an
unusually interesting species belonging to the extinct order Megase-
coptera. The other specimens are fragments of cockroach wings,
which, in the light of our present knowledge, cannot be satisfactorily
placed in families and do not warrant description. The megasecop-
teron, however, is sufficiently well preserved to enable family and
generic diagnoses. It clearly belongs to the family Bardohymenidae,
of the suborder Eumegasecoptera.

Family Bardohymenidae

This family was based on Bardohymen magnipennifer Zal., from
a Permian outcrop along the Barda River, near Perm, Russia
(Zalessky, 1937). Two other genera, Sylvohymen Martynov
(Permian of Oklahoma and of Chekarda, Russia) and Cctlohymen
(Permian of Oklahoma) have also been placed here (Carpenter,
1947). The family includes species which are related to Proto-
hymenidae but which differ in lacking the coalescences of Rs and
MA, and of MP and CuA.

Actinohymen, new genus

Related to Bardohymen, but having the cross-vein between Ri and
R2 very strong, thickened at the costal end, and situated well basad
of the apex of the wing. Pterostigmal area thickened ; Rs with three
main branches.

Type-species: Actinohymen russelli > n. sp.

Actinohymen russelli, new species
Text-figures 1 and 2; plate 1.

Length of wing, as preserved, 28 mm; width, 8.5 mm ; estimated
complete wing length, 50 mm. Wing markings: pterostigmal and

irThis research has been aided by a grant from the National Science
Foundation.

Manuscript received by the editor January 4, 1962.

37

38

Psyche

[March

apical region margined with dark pigment; isolated triangular spots
at end of R4 + 5, MA, and CuA. Eight outer cross-veins, forming
a distinct row; CuP and iA forked distally. Other details of vena-
tion are shown in text-figure 1.

Holotype: No. 140898, Paleozoic Catalogue #29, U. S. National
Museum, Washington; collected by Dr. S. H. Mamay, in an outcrop
on the Emily Irish land grant, about 18 miles south-southeast of
Seymour, Baylor County, Texas. The fossil is very clearly preserved
and consists of about the distal half of a wing; the proximal half was
apparently broken away in the counterpart, which is missing. Slight
distortion of the anterior margin, just basal of the pterostigma, results
from a pronounced depression in the rock. Although incomplete,
this is the best-preserved specimen of a bardohymenid which has yet
been found.

Geological age: Lower Permian; Belle Plains Formation (Wichita
Group) ; this may1 be approximately correlated with the lower part
of the Wellington Formation of Oklahoma and Kansas. (See Dunkle
and Mamay, 1956; and Dunbar, et. al i960).

The species is named for Mr. Mart Russell, of Seymour, Texas, in
recognition of the cooperation, hospitality and interest shown to Dr.

Text-figure 1. Drawing of Actinohymcn russelli, n. sp., based on holotype.
C, costa ( + ) ; Sc, subcosta ( — ) ; Rl, radius ( + ) ; Rs, radial sector ( — ) ;
R2, R3, R4+5, branches of radial sector ( ); MA, anterior media (T);

MP, posterior media ( — ); CuA, anterior cubitus ( + ); CuP, posterior
cubitus ( ) ; 1A, first anal vein. The irregularities in the costal margin

have been restored.

Mamay and his associates during their collecting trips at the Emily
Irish deposit.

This remarkable fossil shows a number of interesting features,
mostly specializations of the anterior marginal area of the wing. As
in many Eumegasecoptera, such as Prothymenidae and Bardohy-

1962]

Carpenter — Permian Megasecopteron

39

menidae, the subcosta, and radius (Ri) are very close together and
also to the anterior wing margin (text-figure 2). The costa is
flattened and wide for its entire length. In the region of the
pterostigma the costa widens even more and is somewhat thickened,
but at the distal end of the pterostigma it disappears. The subcosta
appears to be flattened and to be contiguous with the costa and is
not readily distinguished from the latter. The radius (Ri) is con-
tiguous with the subcosta (in the preserved part of the fossil) or
with the costa beyond the end of the subcosta, except in the very distal
part of the wing. Beyond the short but distinct pterostigmal veinlet
the radius seems to fork, the more distal branch leading to the very
apex of the wing. The cross-vein at the basal part of the pterostigma
forms a heavy bar, which is especially thick at its costal end.

The flattened costa, subcosta and even parts of Ri bear several
irregular rows of setal bases or sockets (text-figure 2), as in some

Text-figure 2. Photograph of part of wing of Actinohymen russelli, n. sp.
(holotype), showing proximity of C, Sc and RI, and the origins of Rs and
MA. Setal bases can be seen along the costa.

Palaeodictyoptera, e. g. Dunbaria . It is curious that no setae are
preserved, especially since they are often visible on the wings of
Dunbaria.

The wing markings are not unlike those of other species of Megase-
coptera, especially the Carboniferous Aspidothorax triangularis
Brongn. (Commentry, France). Eumartynovia raaschi Carp.
(Permian, Oklahoma), though not at all closely related (z. e ., belong-
ing to the Paramegasecoptera) , has almost identical markings.

Psyche, 1962

Vol. 69, Plate 1

1962]

Carpenter — Permian Megasecopteron

41

References Cited

Carpenter, F. M.

1947. Lower Permian insects from Oklahoma. Part I. Proc. Amer.
Acad. Arts Sci. 76: 25 >4.

Dunbar, C. O., et al.

1960. Correlation of the Permian formations of North America. Bull.
Geol. Soc. Amer., 7 1:1763-1806.

Dunkle, D. H. and Sergius H. Mamay

1956. An acanthodian fish from the Lower Permian of Texas. Journ.
Wash. Acad. Sciences, 46(16) :3 08-3 1 0.

Zalessky, G.

1937. Etudes des insectes permiens du bassin de la Sylva et problemes
de revolution dans la classe des insectes. Prob. Paleont., 2-3 ;
601-607.

A REMARKABLE NEW GENUS OF LYGAEIDAE
FROM SUMATRA (HEMIPTERA: HETEROPTERA) *

By Jaimes A. Slater
Department of Zoology and Entomology
University of Connecticut

Many species of Lygaeidae possess fore femora that are strongly
incrassate and armed on the ventral surface with sharp spines. Many
workers have assumed that these powerful legs were associated with
predatory habits and indeed as recently as 1956 Miller illustrated
species of Blissinae ( Spalacocoris and Chelochirus) as examples of
legs modified for raptoral purposes. However, it has been evident for
a long time that this was at best an oversimplification and that very
strongly incrassate and heavily spinous legs were known in such sub-
families as the Pachygronthinae and Oxycareninae whose members so
far studied are entirely phytophagous, whereas in the predaceous
Geocorinae the fore femora are slender and not at all adapted for
seizing prey. The enlarged leg is best expressed in the great subfamily
Rhyparochrominae where nearly all of the many hundreds of species
possess enlarged and ventrally spined fore femora. Putshkov (1956)
and Sweet (i960) have shown that most, if not all, of the rhyparo-
chromines are seed feeders. Thus the function of these legs remains
unknown. They do not appear to be used in mating behavior or in
antennal cleaning, nor to any appreciable extent in carrying food etc.
Yet it seems unlikely that legs of this type would persist throughout
hundreds of species in many different genera distributed in several
different subfamilies and in all of the major zoogeographic regions
without having an important function. Solution of this problem should
be a matter of considerable interest to those concerned with the
question of correlation of form and function from inferred evidence.
Recently Sweet (in litt.) has noticed some remarkable threatening
behavior, displayed intraspecifically by several species of rhyparochro-
mines in defense of food, that involves the use of the fore femora.
This may offer a clue to an understanding of this interesting biological
problem.

Despite our lack of knowledge of the function of these fore legs
we do know enough of their occurrence in the family Lygaeidae to
conclude that they are not of random distribution. The presence of
incrassate fore femora is the predominant condition in the Rhyparo-
chrominae, Pachygronthinae and Oxycareninae whereas they are

* Manuscript received by the editor December 20, 1961.

42

1962]

Slater — New Genus of Lygaeidae

43

absent or infrequently developed in the Lygaeinae, Orsillinae, Ischno-
rhynchinae and Geocorinae. Thus it can be seen that the presence of
incrassate fore femora is at least roughly correlated with ventrally
located spiracles. The latter is the less specialized condition. The
correlation may well be more than coincidental and at least give some
indication that an enlarged femur represents the generalized condi-
tion within the family.

It is thus most interesting to encounter a member of the Ischnorhyn-
chinae with forelegs as strongly developed as in any of the subfamilies
whose members are usually provided with incrassate fore femora.
Except for the remarkable front legs this insect appears to be a con-
ventional ischnorhynchine. It possesses dorsally located spiracles, a
large claval commissure, a punctate clavus, hyaline membrane of the
fore wing, hind wing with hamus and intervannals present, a
“pruinose” body bloom, and a non-depressed posterior pronotal margin.

Macellocoris new genus

Fore femora strongly incrassate, armed below with four sharp,
prominent, elongate spines ; lateral margins of pronotum non-explanate
but sinuate, bearing a series of prominent setigerous tubercles; clavus
with a closely set row of punctures adjacent to claval suture and two
inner rows of very large coarse punctures; membrane transparent,
hyaline, exceeding apex of abdomen ; eyes prominent, in contact with
antero-lateral pronotal angles; apex of head attaining but not
exceeding first antennal segment; preocular distance greater than eye
length ; lateral margin of corium explanate.

Type species: Macellocoris incrassatus new species.

Macellocoris incrassatus new species
Plate 2

General coloration reddish-brown, pronotum lighter on either side
of midline posterior to calli and near anterior margin ; hemelytra
opaque whitish, the irregular punctures brown ; legs yellow with
anterior femora bright tan ; antennae with segment one, two except
extreme apex, basal one-half of three and basal one-fourth of four
yellowish with remaining antennal areas fuscous; venter reddish
brown ; mesal area of sternum black ; head with large, coarse, con-
tiguously placed punctures that give a rugose appearance; pronotum
and scutellum with rather small, deep evenly spaced punctures; claval
punctures very large and deep, those on corium irregular in size and

Psyche, 1962

Vol. 69, Plate 2

Slater — Macelloris

1962]

Slater — -New Genus of Lygaeidae

45

position with a definite row adjacent to the claval suture; surface
nearly glabrous, head, pronotum and scutellum with short semi-
decumbent setae in the punctures; appendages sparsely but definitely
pubescent.

Head non-declivent, moderately acuminate, tylus projecting well
beyond the juga; first antennal segment about attaining apex of tylus;
eyes large not produced, strongly in contact with anterolateral pro-
notal angles; length head .70 mm.; width across eyes .85 mm.,
interocular space .48 mm. ; pronotum evenly narrowing anteriorly,
lateral margins slightly sinuate, narrowly carinate, bearing 6-7 setose
tubercles, transverse impression absent mesally but faintly indicated
by depressed area near lateral margin, dorsal surface slightly convex
becoming less so in area of calli and anteriorly, length pronotum 1.05
mm., width pronotum 1.55 mm.; scutellum evenly tumid with
depressed base, no median carina, length .60 mm. ; corium with broad
slightly upturned explanate margin, this laterally rounded becoming
sinuate adjacent to apex of scutellum; apical co-rial margin sinuate,
concave along basal one-third ; distance apex clavus to apex corium
1. 10 mmu, distance apex corium to apex membrane .90 mm.; mem-
brane greatly exceeding apex of abdomen; distance apex abdomen to
apex membrane .75 mm.; middle and hind femora slightly enlarged,
with basal one-fourth slender, giving a clavate appearance; labium
elongate, attaining posterior margin of first abdominal sternite, first
segment exceeding base of head, second segment extending onto
anterior portion of mesosternum; length labial segments I .65 mm.,
II .62 mm., Ill .60 mm,, IV .35 mm.; antennae slender, segments
two and three terete, fourth segment narrowly fusiform ; length
antennal segments I .25 mm., II .75 mm., Ill .82 mm., IV .70 mm.
Total length 4.65 mm.

Holotype: Male. SUMATRA: Pematang S iantar, June 1937. (CT
& BB Brues). In Museum of Comparative Zoology (Harvard).

Acknowledgements

I should like to extend my appreciation to Drs. P. J. Darlington
and W. L. Brown (now at Cornell University), of the Museum of
Comparative Zoology, for the opportunity to study this interesting

Explanation of Plate 2

Macellocoris incrassatus new species. Drawing by Arthur Smith, British
Museum (Natural History).

46

Psyche

[March

insect. Appreciation is also extended to the University of Connecticut
Research Foundation for providing funds for the execution of the
plate by the distinguished Arthur Smith of the British Museum
(Natural History).

Literature Cited

Miller, N. C. E.

1956. The biology of the Heteroptera. Leonard Hill Ltd: London.
Putshkov, V. G.

1956. Basic trophic groups of phytophagous hemipterous insects and
changes in the character of their feeding during the process of
development. Zcol. Zhur. 35:(N. 1) :32-44.

Sweet, M. H.

1960. The seed bugs: a contribution to the feeding habits of the
Lygaeidae. (Hemipt. Heter.) Ann. Ent. Soc. Amer. 53:317-321.

NOTES ON FOSSIL CLEONINAE (COLEOPTERA:
CURCULIONIDAE) 1

By John M. Kingsolver
Illinois Natural History Survey

Dr. Samuel H. Scudder probably described more species of North
American fossil Rhynchophora than any other worker, yet he was
not a specialist in the weevils. His monograph of the fossil Rhyn-
chophora found in the middle Oligocene beds of Florissant, Colorado,
however, is one of the most extensive treatments of any group from
that site.2 Whereas most fossil beetle species have been described
on the basis of elytra, a high percentage of the Florissant weevils are
preserved so that the dorsal or the lateral aspect of the whole speci-
men is visible. In most examples from this site, according to the
illustrations, the rostrum is well preserved, and even antennal and
tarsal segments are intact but the body is usually compressed and
distorted and parts are often disarranged. Details of the mouthparts
are obliterated and the ventral surface of the body is seldom visible.
In these latter two areas lie some of the critical characters needed
for subfamilial and tribal differentiation in existing keys to extant
forms of the Rhynchophora.

There is a basis for comparison of the fossils with extant forms
where the modern classifications of groups are based on the characters
that happen to be well preserved in the fossil specimen. Fossil beetles,
however, are seldom preserved in enough detail to be of much value at
the specific level and in many cases at the generic level, except in
amber. Unless some diagnostic structure is particularly well preserved
in a specimen, most fossil beetles have not been of much value in
taxonomic studies.

Workers in the Rhynchophora should use caution in interpreting
Scudder’s illustrations. If reference to any of his fossil species is
contemplated, the type specimen or specimens should be checked, and
decisions should be based upon this examination instead of upon the
original description and illustration.

Through the kindness of Dr. F. M. Carpenter, I was recently
privileged to examine the type specimens of fossil species of Cleoninae

1This study was made possible by a travel grant from the Society of
Sigma Xi.

Manuscript received by the editor January 4-, 1962.

2Tertiary Rhynchophorus Coleoptera of the United States. U. S. Geological
Survey Monographs, Vol. 21, 206 pp., 12 pis., Washington, 1893.

47

48

Psyche

[March

which had been described by Scudder in 1893. The types are part of
the collection at the Museum of Comparative Zoology, Harvard
University, and were all collected at Florissant. This type study was
in conjunction with a survey being made to circumscribe the weevil
subfamily Cleoninae and to revise the included genus Lixus.

The characters which separate the Cleoninae from other subfamilies
of the Curculionidae are: (1) antennal scrobes originating on the
sides of the beak and curving abruptly downward to terminate beneath
the base of the beak, (2) tarsal claws connate at the base, (3) labial
palpi short and positioned on the ventral face of the labium at either
end of the ligular suture, (4) ligula attached to the terminal margin
of the prementum, (5) tegmen of the male lacking posterior dorsal
lobes, (6) anal veins 2dA2 and 2dA3 usually entire in the hind wings,
(7) eyes either vertically elongated, ovate or reniform. Obviously,
dissection of a specimen would be required to examine the genitalia
and the wings. The other five characters in order to be visible would
require a precise orientation of the beak and tarsal claws, yet the
combination of all these characters is necessary for inclusion of a weevil
in the Cleoninae.

In none of the six cotype specimens of Cleonus exterraneus Scudder,
the two of Cleonus degeneratus Scudder, or in the genotype of
Eocleonus subjectus Scudder are the labium or the tarsal claws visible.
Neither are the eyes nor the antennal scrobes sufficiently well preserved
or correctly positioned to permit accurate determination except in the
genotype specimen of Eocleonus subjectus. Two characters in this
latter specimen rule out the probability of its inclusion in the
Cleoninae. First, the visible eye is elongated horizontally instead of
dorso-ventrally as is the case in every modern, narrow-eyed Cleonine
weevil I have seen. Second, the antennal scrobe is directed toward the
eye as in some of the broad-nosed weevils. Not only are the characters
that are preserved in all of the examined specimens insufficient to
permit the species to be placed in the Cleoninae, but they are also
too vague to allow accurate placement in any other subfamily. There
is simply not enough detail preserved to give any substantial clues to
the correct taxonomic status of the specimens.

In his 1893 report, Scudder also described Cleonus foersteri and
Cleonus primoris, each based on a single specimen. The illustration
of foersteri indicates that this species most nearly approaches a true
Cleonus in the shape of the beak and eye, but other Cleonine characters
are not apparent. The location of the type specimen is not known.

1962]

Kingsolver — Fossil Cleoninae

49

The illustration of Cleonus pr i/nor is indicates that the specimen is
badly distorted and that it probably will yield few clues to its correct
placement in the Curculionidae. The type was stated by Scudder to
be in the Princeton University collections, but Dr. A. G. Fischer
assures me that the type is not presently in that collection and that
its whereabouts is unknown.

On the basis of my examination of the types, I suggest that the
three fossil species, Cleonus exterraneus, Cleonus degeneratus and
Eocleonus subjectus be relegated to incertae sedis status in the Curcu-
lionidae until such time as their true position can be determined, if
that is possible. Likewise, Cleonus foersteri and Cleonus primoris
should be placed in incertae sedis in the Curculionidae until the types
can be located and examined.

PSEUDOSCYMNUS, A NEW GENUS OF ASIATIC
SCYMNINI (COLEOPTERA: COCCINELLIDAE) *

By Edward A. Chapin
Museum of Comparative Zoology

In the course of work on a report on the Coccinellidae of Micro-
nesia, my attention was attracted by the figure of the antenna of
Scymnus kurohime Miyatake and the figures of receptacula of unusual
form of several species, mostly from Japan. Paratypes of S. kurohime
were generously presented to me by Mr. M. Miyatake and a series
of this species was found in a collection from Naha, Okinawa, made
by Mr. N. L. H. Krauss. Mr. Miyatake also supplied, at my request,
a series of S. hareja Ws. An analysis of the characters of these species
shows that they form a group which is intermediate between Scymnus
proper and Cryptogonus. It should be noted here that the eyes of
Cryptogonus and its close relatives are sparsely set with fine erect
setae, a character which has long been recognized as definitive of the
Scymnini.

When compared with Scymnus nigrinus Kugel. and Cryptogonus
orbiculus (Gyll.), the respective type-species, one finds the characters
of S. hareja and S. kurohime to be closer to Cryptogonus than to
Scymnus. In Scymnus the antenna is of the usual coccinelline form
and consists of eleven segments. The tarsus is also similar to the large
majority of coccinellids in being composed of four segments. In
Cryptogonus and Pseudoscymnus the antenna is very short and con-
sists of nine segments and is of an unusual form for the family.
The tarsus is truly three segmented, differing from those of most
coccinellids.

Genus Pseudoscymnus new genus

Body form and size of Scymnus Kugelann, upper surface set with
fine, short pubescence. Antenna nine-segmented ; basal segment stout,
almost as wide at its widest part as long; second segment stout barrel-
shaped, nearly equilateral, clearly separated from the basal; third
through ninth segments forming a fusiform club, the third longer
than wide, fourth through seventh wider than long, each wider than
the preceding, eighth segment usually shorter than wide and slightly
narrower than seventh, ninth segment a little more than half as

*Ma?iuscript received by the editor March 30, 1962.

50

1962]

Chapin — Pseudoscymnus

51

wide as eighth, subconical. The ninth segment bears at its apex
several long setae. Maxillary palp with the three segments nearly
equal in width, the terminal segment parallel-sided with apex sharply
oblique. Terminal segment of labial palp stout barrel-shaped with
truncate apex. Mandible with subapical tooth. Prosternum not
produced anteriorly to cover mouthparts, prosternal lobe rather
narrow, carinate. Abdomen with six visible sternites. Coxal arc
incomplete, much as in Nephus Mulsant. Tibiae simple, rather
slender. Tibial spurs absent. Tarsus three segmented. Claw with
subquadrate basal tooth. Elytral epipleura nearly flat and horizontal,
not distinctly foveolate.

Male — aedeagus symmetrical.

Female — receptaculum seminis with ramus short and stout,
nodulus long, drawn out in a slender, curved tube, or short and
stout, cornu curved, sausage-shaped. Sperm duct very short if nodulus
is elongate tubular, otherwise moderately long. Infundibulum absent.
Hemisternites intermediate between the “blade and handle” type of
the majority of the Coccinellidae and the “ovipositor” type of most
of the Scymnini.

Type-species — Scyrnnus hareja Weise.

I am including the following species in Pseudoscymnus —

Pseudoscymnus hareja (Ws.) 1879, Deutsch. ent. Zeit., 23: 150;
Miyatake, 1958, Japanese Journ. Appl. Ent. Zool., 2:251-
256. figs. 1 A-D, 3 A-B, 4 E-H, 5 A-C.

Pseudoscymnus kurohime (Miyatake) 1959, Mem. Ehime Univ.,
(6) 4: 136-138, figs. 50-61.

It is probable that the following four species also should be referred
to Pseudoscymnus —

Scyrnnus seboshii Ohta 1929, Ins. Matsumurana, 4: 1 1 ;
Miyatake, 1958, Japanese Journ. Appl. Ent. Zool., 2:251-
256, figs. 2 A-C, 3 C-D, 4 A-D, 5 D-F.

Scyrnnus sylvaticus Lewis 1896, Ann. Mag. Nat. Hist., (6)
77:36; Bielawski 1957, Trans. Shikoku Ent. Soc., 5:71,
figs. 5-1 1.

Scyrnnus pilicrepus Lewis 1896, op. cit., p. 36; Bielawski, 1957,
op. cit., p. 72-73, figs. 12-15.

Scyrnnus quinquepunctatus Weise, 1923, Arch. Naturg., 89.
A. 2: 188; Miyatake 1959, Mem. Ehime Univ., (6) 4: 138,
figs. 62-64.

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EN.

ec X5

PSYCHE

A JOURNAL OF ENTOMOLOGY

Established in 1874

Vol. 69 June, 1962 No. 2

CONTENTS

Defense Mechanisms of Arthropods. X. A Pheromone Promoting
Aggregation in an Aposematic Distasteful Insect.

T. Eisner and F. C. Kafatos 53

The Trinidad Cave Ant Erebomyrma (= Spclaeomyrmex) urichi
(Wheeler), with a Comment on Cavernicolous Ants in General.

Edward O. Wilson 62

A New Ant of the Genus Amblyopone from Panama.

William L. Brown, Jr. 73

A New Ant of the Genus Epitritus from South of the Sahara.

William L. Brown, Jr. 77

A New Damothus and a Key to the North American Dignathodontid
Genera (Chilopoda: Geophilomorpha : Digr^thodontidae)

R. E. Crabill, Jr T.. 81

cm i?:s.

U.s. Mi'll,

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1962-63

President L. M. Roth, Harvard University

Vice-President A. R. Brady, Harvard University

Secretary E. G. MacLeod, Harvard University

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Executive Committee C. Walcott, Harvard University

A. G. Humes, Boston University
EDITORIAL BOARD OF PSYCHE
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PSYCHE

Vol. 69 June, 1962 No. 2

DEFENSE MECHANISMS OF ARTHROPODS. X.

A PHEROMONE PROMOTING AGGREGATION
IN AN APOSEMATIC DISTASTEFUL INSECT.1

By T. Eisner and F. C. Kafatos2
Department of Entomology, Cornell University, Ithaca, N. Y.

A striking feature of many aposematic insects is their habit of main-
taining dense and often conspicuous aggregations (Cott, 1957). Rather
than spacing themselves more or less evenly throughout what is seem-
ingly a uniformly favorable habitat, they occur in distinct, sporadically
distributed clusters. Many meloid and coccinellid beetles, as well as
J a variety of pentatomid, coreid, and lygaeid Hemiptera, among others,
are well known for this habit. These insects possess chemical defense
mechanisms that protect them against predators, and their tendency
to advertise themselves to visually oriented predators such as birds by
pooling their aposematic resources in a collective display, appears to
have obvious adaptive value. Moreover, by restricting themselves to
a few relatively widely-spaced sites, the insects are exposed to but a
fraction of the total number of predators in the area. This is likely
to be of particular importance with respect to predators such as birds,
which are known in many cases to have well-delimited foraging terri-
tories, and each of which may be expected to inflict a toll upon the
insect population during the training period when the bird is learning
to discriminate against the insect. Clearly, the fewer the foraging
territories occupied, the greater will be the number of insects spared.

Hitherto no studies have been made on the mechanism by which
such aggregations are established and maintained. The purpose of

This study was supported by Grant E-2908 of the U. S. Public Health
Service. The work was done at the Southwestern Research Station of the
American Museum of Natural History, Portal, Arizona. We are indebted to
the director of the Station, Dr. Mont A. Cazier, for his generosity with equip-
ment and facilities, and to Miss Abby Rockefeller, who assisted ably in this
and related studies.

2New address: Biological Laboratories, Harvard University, Cambridge,
Mass.

Manuscript received by the editor January 9, 1962.

53

54

Psyche

[June

this note is to present evidence indicating that in at least one case, a
gregarious beetle of the family Lycidae, the clustering behavior of both
sexes seems to be mediated by a volatile attractant produced by the
males alone.

Lycids are widely distributed through the tropics and subtropics,
commonly are aposematic, are known to be distasteful to many verte-
brate and invertebrate predators, and frequently figure as dominant
Mullerian elements in mimetic associations. They form dense aggre-
gates, sometimes being found by thousands, closely spaced on the
exposed inflorescences of the host plants on which they feed and mate.

The particular species on which we worked, Lycus loripes (Chev-
rolat), is abundant on the grounds of the Southwestern Research
Station of the American Museum of Natural History, Portal, Ari-
zona, where this study was made. The locality has been described in
some detail elsewhere (Linsley et al., 1961), and it will suffice here
to mention that, at the time of the experiments (July 15-25, 1961),
the lycids were found almost exclusively on a patch ( ca . 80 x 160 ft.)
of sweet white clover (Melilotus alba) directly facing the principal
laboratory building of the Station (Plate 3, fig. 1). Over 3000 L.
loripes inhabited the patch at the time, distributed more or less irregu-
larly in dense clusters. These lycids are uniformly yellow-orange in
color (except for some black on the appendages), and are conspicuous
on the white flowers of the clover, particularly on those branches
where they are densely aggregated (Plate 3, fig. 2). They are sluggish
and do not take readily to flight when disturbed. They fly relatively
little even on their own initiative, and when they do, it is usually
during the noon hours.

Behind the laboratory building there was a second stand of Melt 'lo-
tus, roughly equal in area to the front patch, but almost entirely devoid
of lycids. This uninhabited patch became the test arena in which
formation and growth of aggregations was experimentally induced.

Initial tests, designed to see whether one could induce lycids to
aggregate after distributing them singly throughout the test area were
bound to failure, since these lycids, rather than being attracted to one

Explanation of Plate 3

Figure 1. The open field, densely overgrown with Melilotus alba, directly
in front of the main laboratory building of the Southwestern Research Station,
Portal, Arizona. On this field was found the main standing aggregation of
Lycus loripes.

Figure 2. A cluster of Lycus loripes on an inflorescence-bearing branch of
Melilotus alba. Within the main lycid aggregation, dozens of branches bore
one or more such clusters, as well as isolated individuals and mating pairs.

Psyche, 1962

Vol. 69, Plate 3

Eisner and Kafatos — Defense Mechanisms

56

Psyche

[June

another, were lured back to the main standing aggregate in the front
of the building. Instead of being scattered individually, the lycids
were then put out in a number of isolated incipient aggregations to see
whether these would tend to consolidate and attract newcomers. This
they were found to do. Five groups of L. loripes, each comprising 75
individuals of both sexes, were placed on single Melilotus plants (or
on compact clusters of branches) in such a way that three of the
groups were closely spaced and roughly equidistant (5-6 m.), whereas
the other two were spaced about 25 m. from each other and from the
center of the triangle formed by the other three. Within five days
(see Table I) the three closely spaced aggregations became consoli-
dated around one of the loci, which now numbered 390 individuals.
The two neighboring clusters had dwindled to 27 and 1 respectively.
Recruitment had taken place also from the two outlying clusters,
one of which had disappeared altogether, while the other now had
only 15 lycids. Although the individuals of each lot had originally

AGGREGATION

1

11

IH

IV

V

TIME (hrs.)

LYCID COUNT PER AGGREGATION

0

75

75

75

75

75

4

22

23

35

35

82

30

2

21

30

9

178

56

2

21

26

3

309

1 1 9

0

15

27

1

390

LYCIDS

ELSEWHERE

5

5

30

24

30

Table I. Fate of five artificially-established incipient aggregations (each
consisting of 75 lycids of both sexes) observed over a period of five days.
Aggregations III, IV, and V were 5-6 m. from one another. Aggregations 1
and II were 25 m. from each other and from the center of the triangle formed
by the other three. The times given for the various counts are measured from
when the aggregations were first put in the field. The column on right gives
the lycid count for the entire remainder of the Melilotus test patch behind the
laboratory building.

Explanation of Plate 4

Figure 1. One of the nets (enclosing Melilotus with lycids) used as a lure.
This particular net, which held males alone, was one that successfully attract-
ed other lycids (a few of the newcomers are seen on the net itself).

Figure 2. Group of male lycids, in the typical sequential arrangement they
often assume when confined by themselves.

Psyche, 1962

Vol. 69, Plate 4

Eisner and Kafatos — Defense Mechanisms

58

Psyche

[June

been labelled with a distinct color marking, these tended to flake
off after some days, and the exact course of the migration patterns
could therefore not be followed. However, judging from the number
of unmarked specimens present, which outnumbered by far the calcu-
lated maximum that could have lost its markings, it was clear that a
substantial number of lycids were new arrivals in the test area, prob-
ably stemming from the principal aggregation at the opposite side of
the laboratory building. Conversely, as expected, some marked indi-
viduals had been lured back to the front field.

It remained to be determined what particular attracting stimulus
is responsible for luring the lycids. Visual cues were eliminated by

DECOY GROUP

A

B

C

D

150 dd

150 dd

150 SS

15099

TIME(hrs.) NEWCOMER COUNT PER DECOY GROUP

dd

92

dd

92

dd

92

d d

99

20

39

7

65

13

4

2

4

3

45

41

1 6

25

8

0

1

0

0

72

29

1 7

S 9

5

0

S

0

0

D

150 <f<f

66

99

23

12

31

7

26

13

Table 11. Attraction of lycids to male and female decoy aggregates placed
in nets. The four decoy groups (A-D) were placed at the corners of a
square, roughly 25 m. to the side. The times given for the various newcomer
counts are measured from when the decoy groups were first staked out. At
72 hours, the females from group D were exchanged with the males from A.

1962]

Eisner and Kafatos — Defense Mechanisms

59

using porous nets of white cloth to enclose the groups of lycids, dis-
tributed as before on close clusters of Melilotus branches (Plate 4,
fig. 1 ) . The technique had the added advantage of preventing dispersal
of the decoy group, while at the same time facilitating the count of
newcomers assembled around the nets. Four nets were staked out with
150 lycids each, two of them containing only males, the other two only
females. Spacing was maximized within the area available, each net
being about 25 m. from its neighbors. The loci selected were known to
have supported no previous aggregations. Table II summarizes the
results. Only those lycids were counted as newcomers which were
either directly upon the nets, or on Melilotus branches within a few
feet around them. It is clear that the males, but not the females,
exerted immediate and persistent attraction for lycids of both sexes.
The fact that the newcomers included a preponderance of males
should not be taken to reflect a greater susceptibility of this sex to
the attracting stimulus, since the males were actually the more
numerous in the population at the time (actual counts made, based
on samplings from the main population site, showed the ratio to vary
between 3 : 1 and 6:1).

The possibility was ruled out that ecological factors, rather than
the males themselves, were the source of attraction. After the third
day, the males from net A were exchanged with the females fromi net
D. The other two nets were dismantled, and the lycids within them,
plus the assemblages that had accumulated around both original male
sites, were scattered at 1 m. intervals throughout the test area. Over
the next three days only one assemblage built up, this time around the
new male site (Table II). Clearly, the attracting stimulus is emitted
by the males, and it is most likely a diffusible chemical factor.

An additional observation is worth mentioning. On two separate
occasions, when artificially induced aggregations were allowed to
persist over a period of several days, and were subsequently removed,
the Melilotus branches that had harbored the beetles were found to
retain the potential to lure lycids, presumably as a result of residual
attractant with which the plants had been labelled. When the lycids
that would subsequently accumulate on these branches were sytemati-
cally removed once a day, and redispersed, the attractiveness, of the
plants dwindled to extinction within a few days.

The attraction of lycids to each other in the presence of males was
also demonstrated in captive specimens. Mixed lots of males and
females confined in glass-topped observation enclosures would soon
distribute themselves into individual mating pairs or small clusters of

6o

Psyche

[June

pairs. Females alone remained distributed singly and showed no
attraction for each other. Males, by contrast, clustered closely, usually
one on top of the other in overlapping sequence, as shown in Plate 4,
fig. 2.

A chemical attractant of the type involved here may appropriately
be called a pheromone. Pheromones, by definition (Karlson and
Butenandt, 1959), are substances secreted by one individual to the
outside, capable of eliciting specific behavioral or developmental
responses in another individual of the same species. Among insects,
these social chemical messengers include the sex attractants, the queen
substance of honeybees, the trail substances of ants and termites, the
releasers of alarm behavior in ants, etc. It is clear that the lycid attrac-
tant, aside from its obvious function in maintaining the aposematic
population densely congregated, also serves appropriately in bringing
together the sexes preparatory to mating. But since it lures both
males and females with apparently equal effectiveness, there must be
additional short-range stimuli operating within the aggregation to
insure that males and females will ultimately be properly paired. One
wonders what evolutionary justification accounts for the production
of attractant by the males alone, rather than by the females, or by
both sexes. Not enough is known about the life cycle of lycids, but the
possibility that the males are the first to emerge in the season and
hence are the ones that carry the aggregations through their incipiency,
is worth considering.

It is hoped, now that the groundwork of this problem has been laid,
that additional more precise experimentation on this unusual type of
attractant will be pursued. With L. loripes there are some especially
intriguing aspects to the problem. This lycid is the dominant Mulleri-
an element of an elaborate mimetic complex (Linsley et al., 1961)
that includes among others, a congeneric sibling species of lycid \Lycus
simulans (Schaeffer)], a cerambycid beetle ( Elytroleptus ignitus
LeConte), and a geometrid moth [ Eubaphe unicolor (Robinson)].
Whether the attractant produced by L. loripes , which far outnumbers
the others, exerts its action also on all or some of the mimetic associates,
thus insuring that these are lured to “safety” within the aggregations
of the dominant model element, remains unknown. Unfortunately all
of these mimetic forms were extremely scarce at the time of our experi-
ments, and could not be included for study.

Future work should concentrate also on some of the many other
gregarious aposematic insects known, in which similar attractant
mechanisms, or perhaps interesting alternatives, are likely to be at play.

1962]

Eimer and Kafatos — Defense Mechanisms

61

References Cited

Cott, H. B.

1957. Adaptive Coloration of Animals. Methuen & Co. Ltd., London.
Karlson, P., and A. Butenandt.

1959. Pheromones (Ectohormones) in insects. Annual Rev. Entomol.
4: 39-58.

Linsley, E. G., T. Eisner, and A. B. Klots.

1961. Mimetic assemblages of sibling species of lycid beetles. Evolution
15: 15-29.

THE TRINIDAD CAVE ANT EREBOMYRMA
( — SPELAEOMYRMEX ) URICHI (WHEELER), WITH
A COMMENT ON CAVERNICOLOUS ANTS
IN GENERAL*

By Edward O. Wilson

The Biological Laboratories, Harvard University

In 1922 W. M. Wheeler described a new pheidologetonine ant,
Spelaeomyrmex urichl, which he regarded as allied to Erebomyrma
but sufficiently removed to justify the erection of a separate genus.
The eleven syntypes, all minor workers, were collected in “Guacharo
Cave,” Trinidad, occupied by the oil-bird or “guacharo” ( Steatornis
caripensis Elumboldt). Their pale color, minute eyes, and long bristly
pilosity suggested to Wheeler that they represent a truly cavernicolous,
i.e. troglobitic, species. In addition, Wheeler speculated that “the
Texan E. longi Wheeler, which is certainly subterranean and has
been taken only once (during a nuptial flight), is really a cavernicolous
ant.” In 1938, however, Wheeler reported the collection of workers
of Spelaeomyrmex urichl by P. J. Darlington from leafmold in Cuba,
thereby casting doubt on the status of the species as a troglobite.

Certainly no more likely troglobite than S. urichl has been found
among the ants. Further information on its biology has promised to
be of exceptional interest. In the course of field work in Trinidad in
1961, the present author undertook to rediscover the species. The first
problem was to find the type locality. No fewer than six caves in
various parts of the island harbor the guacharo and could correctly
be called “Guacharo Cave.” The itinerary of the collector, the late
Prof. F. M. Urich, was not known. After the author had visited one
cave, on the Spring Hill Estate, without success, Dr. David Snow,
who was conducting an ecological study of the guacharos, suggested
the Oropouche Cave, near Cumaca, as the most likely locality. A
single visit to this cave, on April 18, resulted in the immediate finding
of two colonies of Spelaeomyrmex. Later, a surprising discovery was
made: a colony collected from savanna forest at Bernhardsdorp, Suri-
nam, in March 1961 and first identified as Erebomyrma, was on second
examination determined to be conspecific with Spelaeomyrmex urichl .
These findings have shed new light on the generic distinctness of

* Manuscript received by the editor January 9, 1962.

62

1962]

Wilson — Erebomyrma

63

Spelaeomyrmex, on its status as a troglobite, and on the ecological
singularity of cave ants in general.

Taxonomy
Erebomyrma Wheeler

Erebomyrma Wheeler, 1903, Biol. Bull, 4:137-148, minor worker,
queen, male. Type species (monobasic) : Erebomyrma longi Wheel-
er.

Spelaeomyrmex Wheeler, 1922, Amer. Mus. Novitates, 45: 9, minor
workers. Type species (monobasic) : Spelaeomyrmex uric hi Wheel-
er. NEW SYNONYMY.

Following is a list of the known species of Erebomyrma , with atten-
dant bibliography:

eidmanni Menozzi, in H. Eidmann, 193^ Arb. phys. Angew. ent.
Berlin-Dahlem, 3:47-48, fig. V ( 1-4), wqrker, soldier, queen. Type
locality: Mendes, Rio de Janeiro, Brazil.
longi Wheeler, 1903, Biol. Bull., 4:137-148, figs, 1-5, minor worker,
queen, male. Type locality: Denton, Texas.
moral Menozzi, 1931, Bull. Lab. Zool. Gen. Agr., Portici, 25:271-
272, fig. 7, worker. Original localities: Apaican, Vulcano; and
San Jose (Costa Rica). Borgmeier, 1949, Rev. Brasil. Biol., 9:207-
208, figs. 8-9, “ergatogyne” (= soldier), second record from San
Jose, Costa Rica.

never manni Mann, 1926, Psyche, 53:103-104, worker. Type locality:
Hamburg Farm, Reventazon, Santa Clara, Costa Rica.
peruviana Emery, 1905, Bull. Soc. Ent. Ital., 37:139 , nota, queen.
Type locality: Marcapata, Peru.

urichi (Wheeler), 1922, Amer. Mus. Novitates, 45:9-11, fig. iA-B,
minor worker. Type locality: “Guacharo Cave,” Trinidad.

(Spelaeomyrmex urichi). Other records: Cuba, Yucatan, Surinam.
The minor-worker characters cited by Wheeler as most strongly
separating urichi from longi, and hence Spelaeomyrmex from Ere-
bomyrma, are: larger head size; angulate (vs. round) humeri; pro-
portionately larger first gastric segment; and minor features in the
shape of the petiole, postpetiole, and legs. These differences probably
signify a specific distinction between urichi and longi, but they are not
great enough to justify a generic break, even by liberal standards. The
synonymy is supported by the fact that the newly-discovered queen
and male of urichi are closely similar to these castes in longi. One
possible difference of generic magnitude, unknown to Wheeler, still
remains: eidmanni, moral, nevermanni, and urichi are now known to

64

Psyche

[June

Figure 1. Soldiers of Erebomyrma urichi from Bernhardsdorp, Surinam,
and Oropouche Cave, Trinidad, and E. Inevermanni from Barro Colorado,
Panama, showing variation in thoracic and pedicellar structure in this little
known caste.

have a soldier caste, while none has been discovered in longi. However,
longi is known from but a single collection, made under circumstances
in which the scarce soldier caste could easily have been overlooked. It

1962]

Wilson — Erebo/nyr/na

65

seems a fairly safe conjecture that longi soldiers will be discovered
when whole colonies of that species are collected.

As just noted, the Oropouche uric hi and a colony taken at Bern-
hardsdorp, Surinam, are considered conspeeific. 1 he minor workers
of the two colonies are virtually identical. In the Oropouche soldier
the anterior face of the petiolar node is slightly more inclined posteri-
orly (thus forming a greater angle with its anterior peduncle), the
anterodorsal nodal angle somewhat more rounded, the propodeal
angles are more pronounced and acute, and there are some differences
in thoracic form and sculpturing (see text-figure). I he Oropouche
queens have slightly more developed propodeal spines and larger ocelli.
In other respects the soldiers and queens appear identical between the
two colonies.

Together, the Trinidad and Surinam uric hi differ markedly from
those in a series of Erebo/nyr/na collected recently at Barro Colorado
(W. L. Brown and E. S. McCluskey leg.) and tentatively determined
as nevermanni Mann. The urichi minor worker has distinctly nar-
rower, more erect propodeal spines; while the urichi soldier is much
smaller and with more pronounced propodeal spines (text-figure).

Ecology

Trinidad. The Oropouche Cave is the source of the Oropouche
River, which extends back into th£ cave as a clear stream several
meters in width. A colony of Erebo/nyr/na urichi was found approxi-
mately 30 meters inside the cave on the bank of the stream, in very
feeble light coming from the plainly visible cave mouth. Much of the
ground was covered by guano dropped from the large numbers of
guacharos nesting overhead. The arthropod fauna at this point was
rich and diverse, consisting of ants [Mesofionrera constricta Mayr,
Odonto/nachus haematodus (Linne), and Solenopsis (Diplorhoptrum.)
tenuis Mayr], as well as the Erebo/nyr/na entomobryid collembolans,
cave crickets, dermapterans, small flies, and mites. The ant species,
other than Erebo/nyr/na, have been collected outside caves in Trinidad.
Mesoponera constricta , which was the most common forager in the
vicinity of the Erebo/nyr/na nest, was also a dominant ant along the
trails through cacao plantations in the Cumaca area. Most of the
other insects appeared to be troglophiles. Fifteen meters farther in,
and in almost total darkness, a large Bufo marinus was found. In
short, at the site of the first Erebo/nyr/na nest, troglophilic (faculta-
tively cavernicolous) animals predominated.

A second group of foraging Erebo/nyr/na workers, almost certainly

66

Psyche

[June

representing another colony, was encountered between 200 and 300
meters from the cave entrance. This spot was reached only after
passing five major twists in the cave and was in apparently total dark-
ness. It was the final outpost of the guacharo nests. Here there were
no other ant species ; in fact, none of the other three found at the first
site ventured beyond the lighted portion of the cave. The arthropod
fauna was sparse in species and biomass and consisted wholly of blind,
white entomobryids, campodeids, isopods, and snails.

Just past the second site the cave ceiling dipped to within less than
a meter of the stream surface, and the passage continued tortuously
for another twenty meters or so. Beyond, the cave opened into a final
oblong chamber before dipping beneath the stream surface. In this
terminal room there were no guacharos; only bats had left a sprinkling
of guano on the floor. A careful search revealed no Erebomyrma
workers in the terminal chamber.

The nest at the first site, near the cave entrance, was located and
excavated. It was enclosed entirely within a large, smooth shale slab
partly buried in the cave soil and covered with a thin layer cf dense
clay. The rock was soft, naturally fractured, and could easily be
broken apart with a steel trowel. Columns of workers were observed
traveling from the guano piles to two entrance holes twelve centi-
meters apart on the upper edge of the rock. The holes were each about
two millimeters in diameter, and each was surrounded by low, incon-
spicuous piles of excavated clay. The nest, located only a few centi-
meters below the surface, consisted of several flat, irregular cavities
between five and ten centimeters wide and several millimeters in
height. Probably the great majority of workers and all of the other
adult castes were collected, both alive and preserved in alcohol. The
sample, censused the following day, had the following composition :
547 minor workers, 1 soldier, 10 dealate queens, 10 males. Also
present was a large quantity of brood, in all stages of development,
including one male pupa. It was estimated that the entire worker
population, including that part left foraging or missed in the nest,
was not less than 600 and not greater than 1000.

Previous to the excavation, the foraging workers were observed
briefly. Workers were found up to H/2 meters from the nest entrances,
but the great majority was within a meter’s radius. Most were hunting
singly or moving in loose files through guacharo guano, just as Urich
had found them forty years earlier. Workers returning to the nest
converged in two separate files, which, judging from the precision

1962]

Wilson — Erebomyrma

67

with which they repeated each twist and turn, must have been follow-
ing odor trails. Several were carrying objects: a mite, an entomobryid
collembolan, and two unidentified arthropod eggs. The mite and
entomobryid were freshly killed and had evidently been captured as
prey. This conjecture is supported by the fact that Erebomyrma
workers were most densely concentrated at points where large numbers
of entomobryids occurred. Later, in captivity, workers fed readily on
a wide variety of larger moths and flies presented to them, but only
after these had been killed and cut open. In the original nest over
a hundred unidentified globular objects resembling arthropod eggs
were found piled with the brood. These were cared for by the captive
colony in the artificial nest and may have been used sporadically for
food, although direct feeding was not observed. Similar structures
were found by Eidmann ( 1936) in the nests of Erebomyrma eidmanni.

Surinam. On March 14 a urichi colony was found in open, dry,
second growth forest at Bernhardsdorp, near Lelydorp. It was nest-
ing in a small rotting log partly buried in moist leaf litter in a well-
shaded part of the forest. The population consisted of a single dealate
queen, four soldiers, an estimated 500-1000 minor workers, and a large
quantity of brood in every stage of development. Adjacent to the
Erebomyrma colony was a large colony of the termite Armitermes
minutus Emerson (det. A. E, Emerson). The Erebomyrma colony
and a fraction of the Armitermes colony were placed alive in separate
but interconnected artificial nests. Within a few hours after establish-
ment Erebomyrma workers entered the still chaotic termite chamber
and began carrying off eggs. They were unopposed by the much
larger Armitermes adults, and in turn did not molest the Armitermes
adults or nymphs. While the response of the Erebomyrma workers
was clear-cut in this instance, it does not necessarily mean that the
species is termitolestic on undisturbed Armitermes colonies in nature.
The colony was kept alive for only a few days and there was no
opportunity to extend the experiment. In this connection it is worth
noting that Emerson (in Wheeler, 1936) found evidence that Solenop-
sis (Diplorhoptru?n) laeviceps Mayr collects and stores eggs of
Nasutitermes cavifrons (Holmgren) and N. costalis Holmgren in
British Guiana.

Behavior

The Trinidad colony was kept under observation in an artificial
nest for two months and notes taken on selected aspects of behavior.
The ethology of this species is of considerable interest because nothing

68

Psyche

[June

has been recorded hitherto on Erebomyrma , and, in fact, very little
information is available on the entire tribe Pheidologetonini.

The self-grooming movements of the minor worker are given as
follows, according to the classification proposed recently for ants as a
whole (Wilson, 1962): (1) oral leg cleaning, (2) antenna wiping,
(3) four-leg wiping, (4) three-leg wiping, (5) two-front-leg wiping,
(7) head wiping (8) abdomen wiping (front legs). The queen
repeated all of these movements and in addition (10) metapleuron
wiping, conspicuously missing in the worker. Other-grooming was
as frequent as in most other myrmicines. The workers were especially
attentive to the large queens, and a populous retinue followed each as
it moved about. As many as six were seen riding on one queen’s body
at the same time.

Worker-to-worker regurgitation was frequent. The queens appar-
ently were fed exclusively by worker regurgitation, and they never
approached the prey. In several emigrations studied, adult transport
was never observed. In one exceptional instance a worker carried a
newly eclosed worked by its left middle and fore legs clumsily and a
short distance over the brood pile, as though it were a piece of brood.
Discussion: The Ecology of Cavernicolous Ant Species

Penetration of caves by ants occurs chiefly in the tropics and is
limited mostly to the dimly lighted (twilight) zone near entrances.
The great majority of records belong to species that are obviously
either trogloxenic (penetrating as foragers from nests outside) or
troglophilic (facultatively cave-nesting). Further, these species are
typically those that are among the most abundant, widespread, and
adaptable members of the ant fauna in habitats outside the caves.
For instance, a collection of ants made in 1959 in the Batu Caves,
Malaya, by Mr. H. E. McClure and studied by the author, consisted
primarily of three identifiable species \Bothroponera tridentata (Fr.
Smith), Leptogenys dimmuta (Fr. Smith), Pheidole javana Mayr]
that are among the most widespread and common members of their
genera in the Oriental Region. Wheeler (1924) records Triglypho-
thrix striatidens Emery from 400 feet inside the entrance of Siju Cave,
Garo Hills, Assam ; this ant is a notably adaptable pantropical “tramp”
and by far the most widely distributed Triglyphothrix. Of 16 species
recorded from Yucatan caves by Wheeler (1938), 15 are identifiable,
and of these 13 have very extensive ranges in the New World tropics;
most are known further to be abundant in various habitats. The
remaining two species \Brachymyrmex cavernicolus Wheeler, Para-
trechina (Nylanderiaj pearsei Wheeler] belong to genera whose

1962]

Wilson — Erebomyrma

69

taxonomy and ecology are too poorly known to allow further general-
ization. Kempf (1961) records Labidus coecus (Latreille) from
“guacharo” caves in northern Peru and Venezuela. Both collections
were made in zones of total darkness, at 90 m. and 800 m. respectively
from the cave mouths. As Kempf points out, L. coecus is one of the
commonest and most adaptable army ant species and ranges, at widely
varying elevations, from the southern United States to northern
Argentina. Santschi (1914) records eight species collected by Ch.
Alluaud and R. Jeannel from caves at Tanga and Shimoni, Tangan-
yika. These can be roughly characterized as follows: Ponera dulcis
Ford, widespread but known from only a few records; Leptogenys
jeanneli Santschi, known only from the type collection but an unexcep-
tional member of the epigeic falcigera group ; O dontomachus haema-
toda L. var. troglodytes Santschi, probably the same as the very
widespread, abundant African ‘‘haematoda” or “haematoda stanleyi”
( Inec haematoda L. of the New World) ; Dorylus fimbriatus (Shuck-
ard), very widespread and abundant; Monomorium rhopalocerum
Emery subsp. speluncarum Santschi, probably equals rhopalocerum , a
widespread and apparently relatively common species, collected at
Shimoni only at a cave entrance; Strumigenys stygia Santschi, known
only from the type collection but otherwise a morphologically unex-
ceptional species of the rogeri group (see Brown, 1954) ; Mic costruma
marginata (Santschi), known only from the type collection taken at
cave entrance and not morphologically peculiar; Paratrechina (Nylan-
deria) jaegerskioeldi (Mayr), very widespread and abundant. Thus
this African cave fauna is made up of five more or less common widely
distributed species together with three species still known only from
the type caves. The latter exceptional group, however, belong to
genera (Leptogenys, Strumigenys, Miccostruma) in which rare, local
species are usual, so that no particular ecological significance can be
attached to the fact that their known range is at present so limited.

Cave ants do not as a group possess the usual morphological modi-
fications found in extreme cave dwellers. For the most part, they
show qo exceptional pigment or eye reduction when compared with
their congeneric relatives, and, in the cases where their habits are
kqQ\yn, they are no less epigeic in their foraging behavior. Conversely,
t% majority of the most highly modified hypogeic and subterranean
tropical ant taxa are unknown from caves, e.g., extreme species of
dmblyopone, Centromyrmex , Solenopsis (Diplorhoptrum) , Trano-
pelty, Jcropyga. We may conclude that the trogloxenes and
trog^pphiles are characteristically generally adaptable rather than pre-

70

Psyche

[June

adapted for cave life. Ants are not exceptional in this regard. Other
adventitiously cavernicolous animal taxa are often very varied in their
ecology; some other arthropod groups, such as the European copepods,
resemble the ants in that the cave species are typically widely distrib-
uted on the outside (Hesse, Allee, and Schmidt, 1951 ).

The question can now be raised, whether there are any ant species
that are troglobitic, i.e. limited to caves. Erebomyrina urichi is clearly
excluded on the basis of information just given, and the accumulated
evidence makes it improbable that any other member of the genus is
troglobitic. Paratrechma (Nylanderia) troglodytes Weber of Cuba,
originally recorded from a shallow cave, has been shown to be a
junior synonym of P. (N.) myops (Mann), which has been found in
soil away from caves in two montane localities in Cuba (Brown,
1955). A related species, microps M. R. Smith, occurs in places
remote from caves on Puerto Rico. Proceratium cavernicola (Borg-
meier), described from a queen found in the Chilibrillo Caves of
Panama, has been recognized as a synonym of P. micrommatum
(Roger), known from several non-cavernicolous series collected in
Central America and Cuba (Borgmeier, 1957; Brown, 1958).

There remain at least four ant species known only from single
collections made in caves. Brachymyrmex cavernicolus Wheeler
( 1938) was found beneath a stone near the mouth of Balaam Canche
Cave, Chichen Itza, Yucatan. It is pale yellow in color and has small
eyes, which perhaps suggest a normally cavernicolous habit. But it
belongs to a genus the taxonomy of which is in outstandingly poor
condition, so that the possibility exists that non-cavernicolous series
have been recorded in the literature under other names, as in the cases
of the Paratrechma and Proceratium species just mentioned. The
African cavernicoles Leptogenys jeanneli , Stru?nige\nys stygia, and 3
Smithistruma marginata have already been discussed. They belong j
to otherwise non-cavernicolous groups and show no apparent morpho-
logical adaptation to cave life themselves. Clearly, until stronger
evidence is produced, the existence of troglobitic cave ant species must
be considered in great doubt.

Enlarging on this point, let us ask why there are so few (if any)
troglobitic ants or, for that matter, troglobitic social insects of any
sort. The following hypothesis is proposed. The reproductive^unit
in social insects is the colony, and it follows that natural selection
operates on the colony as a unit. Further, the singly prefertilized
mother queen of a monogynous colony ordinarily contains thef whole
of the transferable genetic material, and, since she stores the spferm of

1962]

Wilson — Erebomyrma

7i

her mate (or mates), is genetically the equivalent of two or several
individuals, the female and sexual consorts in a normal population of
non-social animals. Hence, what may be referred to as the “equivalent
size” of the reproductive population, Nd, in an idealized deme of
social insects outside the nuptial season can be estimated as

na = -~~ (Q + mQ)

where Nt is the total adult population of the deme, including workers,
m is the average number of males that fecundated the queens assuming
approximately equal sperm contributions, Q is the average number of
mother queens assuming approximately equal egg contributions, and
Nc is the average mature colony size. Thus in a cave deme containing
10,000 adult individuals with an average mature colony size of 1000
and single fertilizations of single queens, the reproductive population
size equivalent would be only 20. We may note that a deme of 10,000
adults, the non-social condition, would perhaps be large enough and
contain enough genetic variability to be stable and self-maintaining.
Demes of this size are quite common in nature. But a deme-equivalent
of 20 adults, the social condition, must be below the critical threshold
or dangerously close.

In short, if species of social insects never become truly troglobitic
it may be simply because they are unable to maintain sufficiently large
cave demes. Due to limited habitable space, the Oropouche Cave could
not have contained more than) 20' colonies of Erebomyrma urichi and
probably held considerably fewer. Erebomyrma colonies are sometimes
polygynous, as we know from the single Oropouche example. If ten
queens per colony is accepted as the upper limit, and each were assumed
to be singly fertilized, the reproductive-population equivalent of the
Oropouche Erebomyrma is calculated not to exceed 400. Since the
excavated colony was the only obviously populous colony found, the
actual equivalent was probably far less.

But let us suppose that social insects do occasionally become troglo-
bitic; what characteristics might they be expected to have? In addi-
tion to the well-known morphological changes common to most
troglophilic animals, there are several features of social structure that
might be affected. There would likely be one or more of three devices
to increase Nd; namely, (1) decreased colony size, (2) increased poly-
gyny, and (3) increase in m (polygamy). The Oropouche colony of
Erebomyrma urichi was markedly polygynous, with the added result
that the estimated Nd of the colony was large (20). In fact, urichi,
although not a troglobite, may be pre-adapted for marginal cave exis-

72

Psyche

[June

tence. Finally, it is interesting to note that the predicted social modi-
fications are just as likely to be developed in any ant species sufficiently
ecologically restricted to be partitioned into very small denies.

Acknowledgements

The author wishes to thank Mr. Donald Dunston and Dr. and Mrs.
John Buxton for their valuable assistance in exploring the Oropouche
Cave. He is also indebted to Prof. A. E. Emerson for determining
the termite prey of the Surinam Erebomyrma; and to Prof. W. L.
Brown, Prof. K. Christiansen, Prof. W. S. Creighton, Dr. W. W.
Kempf, and Mr. R. W. Taylor for critical readings of the manuscript.
The field study was supported by a grant from the National Science
Foundation.

References Cited

Borqmeier, T.

1957. Myrmecologische Studien, I. An. Acad. Brasil. Cienc., 29: 103-128.
Brown, W. L.

1954. The ant genus Strumigenys in the Ethiopian and Malagasy
Regions. Bull Mus. Comp. Zool. Harv., 112: 3-34.

1955. Nylanderia myops (Mann), new combination (Hymenoptera : For-
micidae). Psyche, 62: 79.

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

Eidmann, H.

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

Hesse, R., W. C. Allee, and K. P. Schmidt.

1951. Ecological animal geography. Second edition. John Wiley and
Sons.

Kempf, W. W.

1961. Labidus coecus as a cave ant. Stud. Ent., 4:551-552.

Santschi, F.

1914. Formicidae in “Voyage de Ch. Alluaud and R. Jeannel en Afrique
orientale, 1911-1912,’’ (A. Schulz publ., Paris), Hymenoptera, II:
41-148, 30 fig.

Wheeler, W. Pyl.

1924. Hymenoptera of the Siju Cave, Garo Hills, Assam, I. Triglypho-
thrix Striatidens Emery as a cave ant. Rec. Indian Mus., 26: 123-
124.

1936. Ecological relations of ponerine and other ants to termites. Proc.
Amer. Acad. Arts Sci., 71 : 159-243.

1938. Ants f^om the caves of Yucatan. Carnegie Inst. Publ., 491: 251-
255.

Wilson, E. O.

1962. Biology of Daceton armigerum (Latreilje), with a classification
of self-grooming movements in ants. Bull. Mus. Comp. Zool. Harv.
(in press).

A NEW ANT OF THE GENUS AMBLYOPONE
FROM PANAMA1

By William L. Brown, Jr.

Department of Entomology, Cornell University

As known before i960, the genus Amblyopone in the New World
was restricted to temperate North America and the southern half of
South America. In my i960 review of the Amblyoponini (Bull. Mus.
Comp. Zool., 122: 143-230) I described as new A. orizabana from
Mt. Orizaba in southern Mexico, and the aberrant A. mystriops from
Guatemala. In a very recent paper, Kempf [1961, Studia Ent., Pet-
ropolis, Brazil (n. s.) 4: 489] has recorded A. degenerata Borgmeier
— previously known only from southern Brazil — as being collected
in Surinam). Now, a new species, to be described below, has been
found in lowland forest on the Isthmus of Panama, closing the last
significant gap in the distribution of Amblyopone in this hemisphere.
The genus is now seen to range from British Columbia and, the St.
Lawrence Valley south into southern Chile, and it seems likely that
it reaches Tierra del Fuego, even though no specimens have yet come
in from that far south. Ag it stands, Amblyopone is the most widely
distributed New World ant genus. While it is clear that the genus is
very sparsely distributed in the tropics, and that it reaches its best
development in cool temperate regions to the north and south, it does
seem likely that further collecting will show it to have a continuous
or near-continuous range in all but the driest and coldest parts of the
Americas.

Amblyopone tropicalis sp. nov.

Holotype worker: TL 3.0, HL (including clypeal teeth) 0.60,
HW 0.52 (Cl 87), WL 0.74, petiolar node L 0.26, W 0.35, post-
petiole W 0.38, scape L 0.34, outside straightline length of mandible
0.46 mm. ; measurements as in my i960 review.

Habitus that of the smaller Fulakora> group of Amblyopone,
especially A. orizabana Brown and A. chilensis Mayr. Head with
nearly straight (feebly sinuate) occipital border, sides feebly convex,
diverging anteriad, widest across anterior corners, which are furnished

3The work in Panama was supported by a Small Grant from the Milton
Fund of Harvard University.

Manuscript received by the editor December 15, 1961.

73

74

Psyche

[June

with small but acute genal teeth. Frontal lobes approximate, separated
only by a narrow linear groove. Anterior clypeal apron gently convex
in outline, with 5 large truncate teeth, of which the middle tooth is
much the largest, being composed of two median teeth completely fused
to about their apices, and projecting nearly twice as far as the smaller
teeth close on each side of them; corner (lateral clypeal) teeth still
shorter, each composed of a mesal and a smaller lateral element which
are fused at the base. Mandibles rather slender, their external margins
feebly convex (almost straight along basal 2/3), inner margins convex
except for apical quarter, each bearing 7 teeth : a triangular basal tooth,
followed by a spaced series of 5 acute, slightly recurved teeth, of which
each of the distal 3 or 4 has a shorter dorsal spur (difficult to see in
normal full-face view) representing the vestigial twin of a pair com-
mon in species of this group ; an indistinct reclinate tooth lies near the
narrow mandibular apex. The mandibles when closed cross each other,
but leave a fairly large triangular space between themselves and the
clypeal margin. Antennal scapes short and rather broad (thinnest near
their midlength) , feebly sigmoidal, reaching back to about the posterior
fifth of the head length. Funiculus 1 1 -segmented, conspicuously
enlarged apicad, but gradually so, without a definite number of
segments in the club ; all segments except first and apical broader than
long. No eyes detected.

Alitrunk feebly convex in profile; as seen from above, broadest
across the middle pronotum; promesonotal suture apparently flexible
and accompanied by a strong groove along the anterior mesonotal
border, at alitruncal midlength. Mesonotum transverse, forming
narrowest point of alitrunk; metanotal groove distinct but shallow;
propodeal dorsum broader than long, its sides diverging posteriad ;
declivity plane, rounding into dorsum, much broader than high.
Inferior borders of pronotum broadly rounded.

Petiolar node sessile, with vertical anterior and horizontal dorsal
faces both convex, the dorsal face broader than long as seen from
above. Ventral process of petiole a rounded, forward-thrust lobe with
a conspicuous oval thinned area or fenestra. Postpetiole broader than
petiole, but shorter, and also shorter than the succeeding segment,
which is the widest. Gastric apex not laterally compressed ; sting stout.
Legs robust; femora flattened and incrassate; tibial spur vestigial on
middle legs, but on each hind tibia there is a short but very thick,
curved, pectinate tibial spur and an adjacent, much smaller, straight,
slender spur.

Head densely reticulate-punctulate and opaque. Antennal scapes,

1962]

Brown — A mblyopone

75

mandibles and clypeus obscurely longitudinally striate-punctate, opaque
to subopaque. Lower sides of propodeum with the usual faint longi-
tudinal striae. Remainder of body smooth or nearly smooth and
shining, with fine spaced punctures, these most dense on petiolar dor-
sum and on two succeeding segments, where integument almost appears
loosely coriaceous in some lights, but is still definitely shining.

Pubescence appressed and subappressed, moderately dense, generally
distributed over body and appendages (not on sides of alitrunk) ; erect
pilosity short, mostly oblique, moderately abundant, becoming longer
on gastric apex, antennal apices and mandibles; very sparse on legs.
Color ferruginous yellow, the head capsule very slightly darker, appen-
dages somewhat lighter.

Paratype female (dealate) : TL 3.1, HL 0.65, HW 0.57 (Cl 88),
WL 0.91, petiolar node L 0.26, W 0.38, scape L 0.37, greatest
diameter of compound eye 0.1 1 mm.

Similar to the holotype worker, but with the usual differences of
caste: fairly large compound eyes (no eyes could be detected in the
worker), ocelli developed and with blackened calli, wing stumps
present and blackened ; meso- and metathoracic flight sclerites well
developed, but rather flat, continuing the weakly convex surface of
the alitrunk without major interruption. As usual for females of this
genus, the petiole and gaster are relatively a little wider than in the
worker, and the head, while still light ferruginous, is a trifle darker
than in the worker. Alitrunk also rather coarsely and closely punctate
above, but still distinctly shining.

The holotype worker and female paratype, the only adults taken,
were found on Barro Colorado Island, Panama Canal Zone, on
January 6, i960 [W. L. Brown, Jr., leg.], and deposited in the
Museum of Comparative Zoology. The specimens were found together
with a few larvae and pupae in a cavity in the underside of a small
rotten branch lying in moist leaf litter on the forest floor, in what
is variously described as rain forest or monsoon forest, close to
Snyder-Molino Trail and less than 100 meters from the Laboratory
Clearing of the Smithsonian Institution’s Canal Zone Biological Area.
These specimens were the very first ants I collected during a three-
week stay on the island. When first collected, they were mistaken for
Prionopelta, a related genus found very rarely on the island, but com-
mon elsewhere in Central and South America. After closer examina-
tion revealed their true identity, I searched energetically for the species
in likely habitats for the remainder of my stay, as did my companion,
Dr. E. S. McCluskey, but we never found it again. This is only one

76

Psyche

[June

of many ant species that have been collected on Barro Colorado a
single time, despite the very intensive collecting of its whole area by
several ant specialists, as well as the long-time Resident Naturalist,
James Zetek. It has been said that Barro Colorado is so well collected
for ants that scarcely anything in the way of new species can be collect-
ed there. That McCluskey and I, engaged in what was primarily an
ecological survey not particularly directed toward the finding of
novelties, were able to find this Amblyopone and at least two other
undescribed ant species in the square mile centered on the Laboratory
Clearing, speaks for the virtual inexhaustability of the island’s ant
fauna and for the richness of tropical forest faunas in general.

In the key to the New World Amblyopone in my i960 review (p.
191 to 192) , A. tropicalis runs to couplet 8, where its intermediate size
makes it fall ambiguously into either 8 or 9. Of the three species in
these two couplets \orizabana Brown, bierigi (Santschi), chilensis
Mayr], A. tropicalis is closest to orizabana in structure and to chilen-
sis in size, but differs from these in the reduced number of teeth in
the clypeal armament, in the strongly projecting, large median fusion
tooth, and in the nearly obsolete dorsal members of the mandibular
double teeth, rendering the tooth row apparently single as seen from
dorsal view. A. tropicalis is larger than orizabana , has relatively long-
er and more slender mandibles and a broader head.

A NEW ANT OF THE GENUS EPITRITUS
FROM SOUTH OF THE SAHARA*

By William L. Brown, Jr.

Department of Entomology, Cornell University

Recently Mr. G. E. J. Nixon, of the Commonwealth Institute of
Entomology in London, sent me a small series of a curious ant that
he had recognized as an aberrant and possibly undescribed species of
tribe Dacetini. Upon receiving the specimens, 1 found that the sample
represented a new Epitritus , the first member of the genus from Ethio-
pian Africa, and the first to be found in the tropics. I am grateful to
Mr. NiXon for making available this most interesting new species.

Epitritus laticeps sp. nov.

Figures 1-4

Holotype worker: TL 2.2, HL 0.47, HW 0.58 (Cl 123), ML
0.22 (MI 47), WL 0.49, scape L 0.29, funiculus L 0.43 mm, of
which about half is taken up by the apical segment. Measurements
and proportions are those standard in my other dacetine studies (see
Brown, 1953, Amer. Midi. Nat. 50: cf. pp. 7-15).

Shape of head, body and mandibles as shown in Figs. 1-4. Especially
characteristic of the known species of Epitritus and (judging from two
occipital lobes (which make the head distinctly broader than long) ;
the wide, basally lobiform antennal scapes; the large, conical labial
lobes; the 4 long straplike clypeal hairs and the peculiar mandibles,
lacking a long, spiniform dorsal “apical” tooth and with only a single
preapical tooth. The apex of the mandible is of the “inverted” type
characteristic of the known species of Epitritus and (judging from two
paratype workers of which the mandibles were opened) has 7 or 8
denticles, of which one or two in the middle are round-edged, and the
rest are acute. The basal lamella is small, with acutely rounded apex
(Fig. 3), and the ventro-medial margin below it is obtusely denticulate
or angulate (not shown in figures) . Eyes minute, with only 4-6 facets.
Antennal funiculus clearly 5-segmented.

Promesonotum seen from above broadly oval, almost circular, nar-
rowest behind, where it is terminated by the fine but distinct metanotal
groove, which crosses just behind the highest point of the swollen
mesonotum. Propodeum much narrower than pronotum, with high,

* Manuscript received by the editor December 15, 1961.

77

78

Psyche

[June

angular lamellae guarding the concave declivity, the declivity reaching
far up towards the metanotal groove.

Petiolar node distinct from its anterior peduncle, as seen from above
transverse, subrectangular, about twice as broad as long. Postpetiole

Figures 1-4. Epitritus laticeps sp. nov., paratype worker. Fig. 1, side view
of head and body. Fig. 2, dorsal full-face view of head and mandibles. Fig.
3, oblique dorsal view of opened left mandible. Fig. 4, ventral inside oblique
view of apical group of teeth or denticles. Drawn by F. A. McKittrick.

much broader than petiole and more than twice as broad as long,
convex above with a shallow median impression, articulated to the
entire width of the anterior gastric margin. Spongiform appendages
limited to narrow posterodorsal lamelliform collars on both nodes,
and absent from anteroventral face of gaster. Gaster laterally sub-
marginate at the extreme base ; short, coarse costulae fade caudad into
superficial, indistinct reticulation reaching nearly to the midlength
of the basal segment ; gaster otherwise shining and smooth.

Body reticulate-punctulate, opaque, except for the gaster, the inner
mandibular surfaces and the labral lobes, which are smooth and shin-
ing. Specialized pilosity, especially the large orbicular, cochlear hairs

1962]

Brown — Epitritus

79

of the head, and the spatulate ones of the scapes and clypeus, as shown
in the figures. Slender, erect spatulate-to-clavate hairs in rows of 4
or 6 on gastric dorsum, totalling about 32. Ground pilosity reduced
to minute stubby hairs, chiefly on promesonotum and nodes, and sparse,
short reelinate hairs on gastric dorsum and legs. T. ibiae and tarsi with
longer reelinate hairs, some of them spatulate. Gula with fine reelinate
hairs. Color medium ferruginous; head feebly infuscated dorsally;
appendages more yellowish.

Holotype [British Museum (Natural History)] and six paratype
workers [deposited with holotype and in Museum of Comparative
Zoology at Harvard College] taken in northern Nigeria near Zungeru
on the Kaduna Road, 19 December, 1956, from “base of dead tree’’
by W. A. Sands of the Termite Research Unit (Collection No. S
780). The paratypes are quite similar to the holotype, and range from
the same size down to the smallest specimen : TL 2.0, HL 0.46, HW

0.55 (Cl 120), ML 0.22 (MI 48), WL 0.49 mm. Cephalic index
range for the entire type series is 1 19-125. There is slight variation
in the size and shape of the small teeth or denticles in the apical com-
plex. Female and male unknown.

E. laticeps can be separated from its two congeners by means of the
following key :

1. Tooth at apparent (dorsal) apex of mandible small, about equal
to or slightly longer than the other largest teeth of the apical
group; promesonotum without conspicuous hairs of any kind

(Nigeria) laticeps Brown

Tooth at apparent (dorsal) apex of mandible long, straight and
spiniform, about 2 or 3 or more times as long as the largest of
the other teeth in the apical group ; promesonotum with numer-
ous large orbicular scale-like hairs like those of the dorsum of
the head 2.

2. Funiculus with 5 separate segments; mandible with two pre-

apical teeth (Japan: Kyushu, Honshu) hexamerus Brown

Funiculus with only 3 separate segments; mandible with 4 pre-
apical teeth (Mediterranean lands n. to Hungary)
argiol.us Emery

The finding of a species of Epitritus south of the Sahara marks the
genus as a zoogeographical curiosity of more than usual interest. Of
the two previously known species, E. argiolus is widespread in southern
Europe and North Africa, while E. hexamerus has been taken twice
in Japan. Evidently, Epitritus is a relict-distributed group that was
once more widely distributed in the tropical and warm temperate

8o

Psyche

[June

parts of the Old World. The insects are cryptobiotic in habits, and
are small and inconspicuous, so we may look forward to the discovery
of species elsewhere in the Old World.

Another interesting thing about the new species is its close conver-
gence in many morphological details to Talaridris ?nandihularis Weber
(tribe Basicerotini) of Trinidad and British Guiana (see Brown and
Kempf, i960, Stud. Ent., Petropolis, (n.s .)j: 233, 241-242).

A NEW DAMOTHUS

AND A KEY TO THE NORTH AMERICAN
DIGNATHODONTID GENERA
(CHILOPODA : GEOPHILOMORPHA :
DIGNATHODONTIDAE) 1

By R. E. Crabill, jr.

U. S. National Museum, Washington, D. C.

The genus Damothus was proposed by R. V. Chamberlin in i960
(p. 239) for the reception of a single species, month, which had been
collected in the Wasatch Mountains of Utah. While collecting arach-
nids and myriapods at Ophir in the Oquirrh Mountains of that State,
Dr. H. W. Levi unwittingly uncovered the second-known specimen
of the genus, which I judge to represent a new species, alastus, here
described. I wish to express my thanks to Dr. Levi for his kindness in
placing this and much other valuable material in my hands for study.

On the basis of all available information, the two most distinctive
features of the genus are the massively crassate and essentially tubular
ultimate legs of the male, and the presence of two basal denticles
on the prehensorial tarsungula. Indeed, the latter character alone will
distinguish Damothus from all other known chilopod genera of what-
ever order. After more is known about Damothus , it may well be seen
that two other features have significance diagnostically: the peculiar
shape of the 1st maxillary medial lobes; the relatively strongly-
developed labral sidepieces.

Comparing the Harvard specimen with Dr. Chamberlin’s original
description of month, I find the following to be significant distinguish-
ing features. D. month : (1) The first maxillae are without lappets.
(2) The coxopleural pores are concentrated along and mostly con-
cealed beneath the margins of the ultimate pedal sternite. (3) Ventral
pore-fields are absent. D. alastus, n. sp. : (1) The first maxillary
coxosternum bears a pair of conspicuous and relatively long lappets.
(2) The coxopleural pores are all exposed and are not concentrated
along and beneath the ultimate pedal sternite. (3) Small but con-
spicuous pore-fields are present on all pedal sternites except the last.

This study was undertaken with the assistance of a grant from the National
Science Foundation.

Manuscript received by the editor January 30, 1962.

8l

82

Psyche

[June

Dam o thus alastus new species
Plate 5

Holotype, male. Utah: Tooele County, Ophir, Oquirrh Mountains,
2000 m. 25 April 1961. Herbert W. Levi, leg., in cottonwood, sage.
Specimen preserved in the Myriapod Collection of the Museum of
Comparative Zoology, Harvard University.

General. Length: 11.5 mm. Pedal segments: 37, Body widest
over posterior third, anterior to which it is gradually acuminate. Color :
generally pale yellow ; the head and prehensors yellowish-orange.
Antennae. Length: (expanded in Hoyer’s) 1.65 mm. Shape: strictly
filiform, neither attenuate nor clavate. Setae gradually increasing in
number and decreasing in length on articles 1-14. Ultimate article
twice as long as penult ; its upper third with short, Hat, special sensory
setae on outside and inside surfaces. Cephalic plate. Length: 0.416
mm. Greatest width: 0.406 mm. Shape: sides evenly excurved; rear
margin perfectly straight. Clothed with straight, relatively short,
stiff setae. Frontal suture absent. Prebasal plate completely concealed.
Clypeus. Paraclypeal sutures broadly membranous, complete. Trans-
buccal sutures vague, passing only half-way to lateral margin. With
a pair of small and extremely weakly consolidated areas (plagulae)
on extreme posterior margin. Setae: postantennals, 1 + 1, very long;
midclypeals, 2 + 2, the inner pair much longer than the outer pair ;
prelabrals absent. Labrum. Midpiece very wide, armed over entire
width with long, hyaline, thin serratures. Sidepieces: strongly-
developed, well-sclerotized ; each with a few delicate, hyaline serra-
tures; widely separated centrally; separated from clypeus by wide
membranous strip. First maxillae. Coxosternum: without setae;
medially undivided ; very vaguely separated from medial lobes and
telopodites ; with a pair of concealed, relatively long lappets. Second
maxillae. Isthmus very wide from side to side but narrow antero-
posteriorly. Each coxosternite very weakly sclerotized, with few setae;
without special thickenings or similar appurtenances. Telopodite: with
strongly-developed dorsal and ventral basal condyles; the articles
separated by distinct sutures ; outer marginal setae extremly short and
robust; inner marginal setae much longer; apical claw long and thin,

Explanation of Plate 5

Damothus alastus sp.n. a. First and Second Maxillae. Left halves, all
setae shown, b. Ultimate Pedal and Postpedal Segments. Ventral aspect,
setae deleted, c. Left Sixth Leg, Tarsus and Pretarsus. Anterior surface, all
setae shown, d. Tarsungula and Intercalary Articles of Left Prehensor. Ven-
tral aspect, principal setae shown.

0.224 mm

Psyche, 1962 Vol. 69, Plate 5

Crabill — Damothus

84

Psyche

[June

smooth except for minute protuberance as shown in figure. Proster-
num. Without subcondylic sclerotic lines. Pleuroprosternal sutures
arching obliquely laterally, complete anteriorly. Anterior margin
without diastema or denticle. Prehensors. When closed, not sur-
passing anterior head margin. Trochanteroprefemur : basally bulging
on inner side; without a denticle; outside length, 0.198 mm; inside
length, 0.094 mm; basal width, 0.146 mm. Femoroid without den-
ticle. Tibioid with a distinct but small denticle. Tarsungula: rela-
tively short and robust ; basally with two large denticles ; dorsal edge
smooth; ventral edge over proximal half dissected into about 4 coarse
and rounded serrations; length, 0.208 mm. Poison calyx: of the simple
type, consisting of bunched digitiform appendices; situated in femoroid.
Poison gland situated entirely in the trochanteroprefemur. Tergites.
Without evident paramedian grooves. Tergites and intertergites
clothed with long, stiff, robust setae. Sternites. On the anterior
third of body each with a midlongitudinal, shallow depression. Pore-
fields: anterolaterals absent; each sternite from the first through the
penult with two small, subcircular fields on extreme posterior margin.
Pro- and metacoxal porefields present on the first through the penult
pedal segments. Setae ; few in number ; arranged in regular horizontal
rows. Legs. Clothed with stiff, long, robust setae. Pretarsi : very
long and thin, curved ; parungues acicular, short, approximately equal
in length. Ultimate pedal segment. Pretergite fused with its
pleurites, i.e. without sutures or divisions bilaterally. Tergite: greatest
width to length, 35 : 28 ; anterior corners rounded; sides straight and
posteriorly convergent ; rear margin broadly rounded. Presternite
with a vague midlongitudinal suture. Sternite with sides essentially
straight and convergent, its rear margin weakly rounded. Coxopleuron :
barely inflated ; ventrally with small, freely-opening, deeply-pigmented
pores; 5 on each coxopleuron. Ultimate leg: greatly swollen, essential-
ly tubular, notably much longer and more massive than the penults;
tarsus consisting of two articles, the second about half as long as the
first and conical in shape; pretarsus is a robust, dark claw; the whole
leg clothed with robust, stiff setae; ventral and inner surfaces of all
articles including and distal to the femur pierced by relatively large
glandular pores. Postpedal segments. Gonopod distinctly bipartite,
conical. Anal pores present and not concealed.

To assist in locating Damothus within the growing complex of
North American dignathodontid genera, I have presented a generic
key here: it is the first to be published since that of Attems of 1947,
p. 129. To some extent it had to be based upon information only

1962]

Crabill — Da/noth, us

85

available from the literature. Included are all of the genera now
known from North America including Mexico.

Excluded are three genera which had been previously reported from
the area under discussion: Leptodampius Chamberlin, 1938, p. 255;
Diplochora Attems, 1903, p. 281 ; Paraplanes V erhoeff , 1933, p. 22.
The original description of Leptodampius is not sufficiently full and
detailed to permit its confident placement within my key. The recent
examination of the holotype of fusata Attems, the type species of Diplo-
chora, shows it to be referable to T omotaenia Cook (new synonymy),
and to the subgenus Korynia Chamberlin. VerhoefPs Californian
Paraplanes calif ormicus, whose types I have studied at Munich, all are
referable to T omotaenia fusata (Attems) (new synonymy). The
details of these cases will be discussed in a separate paper soon to be
issued.

1 a. Each coxopleuron with two subsurface gland-pits, but without

freely-opening and exposed pores. ( Mexico)

Pagotaenia Chamberlin

ib. Each coxopleuron with freely-opening pores most or all of which

are exposed. Subsurface gland-pits absent. 2

2a. Prehensorial tarsungula with 1 or 2 prominent basal denticles

4

2b. Prehensorial tarsungula without a basal denticle 3

3a. Coxopleural pores numerous, opening over most or all of coxo-
pleural surface. Ultimate pedal sternite narrow and elongate.

(Idaho). Zantotaenia Chamberlin.

3b. Coxopleural pores few in number and opening only along and
under the margins of the ultimate pedal sternite which is wider

than long, (southeastern United States) A gathothus Bollman

4a. Tarsungula with 2 prominent basal denticles. (Utah)

Damothus Chamberlin

4b. Tarsungula with 1 prominent basal denticle 5

5a. Ventral pore-fields absent. (California)

I\I aloe hora Chamberlin

5b. Ventral pore-fields present 6

6a. Coxopleural pores concentrated along and beneath margins of
ultimate pedal sternite; this sternite always wider than long.
(United States, Missouri westward to the Pacific Coast; eastern
Asia) T omotaenia Cook ( sensu lato2)

2If T omotaenia is divided into subgenera on the basis of the presence or
absence of sutures on the ultimate pedal pretergite, then the oldest available

86

Psyche

[June

6b. Coxopleural pores freely dispersed at least over ventral surface of
coxopleuron ; not restrictively concentrated along and beneath
ultimate sternite margins ; this sternite usually longer than wide
or at most as wide as long. (Holarctic Region) .. Striganiia Gray

■ ■ ■ 7

7a. Ultimate pedal pretergite fused with its pleurites, i.e. not bilater-
ally impressed with sutures. S. (Linotaenia) C. L. Koch

7b. Ultimate pedal pretergite not fused with its pleurites, i.e. set off
from them by prominent sutures or fissures.

S. ( Striganiia) Gray3

References Cited

Attems, Carl graf.

1903. Synopsis der Geophiliden. Zool. Jahrb. (Syst.). 18 (2) :155-502.
1947. Neue Geophilomorpha des Wiener Museums. Ann. Naturh. Mus.
Wien. 5 5 :50-149.

Chamberlin, Ralph V.

1938. Three new geophiloid chilopods. Ent. News. 49: 254-255.

1960. Five new western geophilid chilopods. Proc. Biol. Soc. Wash. 73 :
239-244.

Verhoeff, Karl W.

1933. Schwedisch-chinesische wissenschaftliche Expedition nach den
nordwestlichen Provinzen Chinas. Arkiv for Zool. 26A(10) : 1-41.

name for the bisuturate specimens would have to be Diplochora Attems, 1903.
The non-suturate specimens would take the nominate generic name. The
explanation for this is complicated and beyond the scope of this paper; how-
ever, it is fully treated in another paper being published in Entomological
News.

historically three contending generic names have been applied to this
zoological entity. Some wrorkers continue to apply the wrong generic name.
The only correct generic name is Strigamia ; it is neither Scolioplanes nor
Linotaenia. The whole matter was thoroughly discussed and clarified by me
in a 1953 publication: see Entomological News, 64(7), pp. 169-172.

CAMBRIDGE ENTOMOLOGICAL CLUB

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The illustration of a spider on the front cover of this issue of
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Emerton and contained in the Museum of Comparative Zoology.
First published in the Transactions of the Connecticut Academy of
Arts and Sciences (1909, vol. 14, plate 7, fig. 5), it shows the dis-
playing posture of the male of Habronattus viridipes (Hentz).

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I

PSYCHE

A JOURNAL OF ENTOMOLOGY

Established in 1874

Vol. 69 September, 1962 No. 3

CONTENTS

The Cytotaxonomy of the Larvae of Some Mexican Fruit Flies in the
Genus Anastrepha (Tephritidae, Diptera). Guy L. Bush 87

Redescription of Lithophotina floccosa Cock. (Manteodea) with Some
Notes on the Manteod Wing Venation. A. G. Sharov 102

The Infrabuccal Pocket of a Formicine Ant: a Social Filtration Device.

T. Eisner and G. M. Happ 107

Australian Carabid Beetles XL Some Tachys. P. J. Darlington, Jr. 117

The Spider Genus Sosippus in North America, Mexico, and Central
America (Araneae, Lycosidae). A. R. Brady

129

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1962-63

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PSYCHE

Vol. 69 September, 1962 No. 3

THE CYTOTAXONOMY OF THE LARVAE OF SOME
MEXICAN FRUIT FLIES IN THE GENUS ANASTREPHA
(TEPHRITIDAE, DIPTERA)1

By Guy L, Bush2
U. S. Department of Agriculture

Introduction

During a study of host relations of the Mexican fruit fly, Anas-
trepha ludens (Loew), difficulty was encountered in obtaining
positive identification of tephritid larvae recovered from field infested
fruit. Existing larval keys based on morphological characters (e.g.
Phillips, 1946) were not adequate for differentiating between some
closely related Mexican representatives of the family. Accurate
identifications could only be obtained by rearing larvae to the adult
stage. This proved time consuming and increased the chance of
losing valuable host records when larvae failed to mature. For this
reason, a preliminary cytotaxonomic study was made on some of the
more common fruit infesting Anastrepha found in Mexico to see if
chromosome morphology would be of any use in identifying larvae.

This method of species differentiation is not new. It has been used
for many years by plant taxonomists to establish a more natural
classification within certain groups of plants (Darlington, 1956).
Its application to animal taxonomy has been somewhat restricted
owing, in part, to the difficulties of handling some animal material.
Many of these difficulties have now been eliminated through the use
of new and improved techniques. A great deal is now known about
the cytogenetics of animals and particularly of the insects. White
(1954) has presented an excellent summary of our present knowledge
of comparative cytology and its application to the study of animal
evolution and taxonomy, while Patterson and Stone (1952) have

This study was conducted under the auspices of the United States Depart-
ment of Agriculture while the author was employed by that organization in
Mexico, D.F., Mexico, from 1955 to 1957.

2Present address: Biological Laboratories, Harvard University.

Manuscript received by the editor Jan. 28, 1962.

87

88

Psyche

[September

discussed chromosome evolution in the genus Drosophila in detail.

Several investigations have been made concerning the cytology of
the Tephritidae. Metz (1916), after investigating the chromosomes
of Euresta melanogaster Loew [probably Dyseuaresta mexicana
(Wied.)], concluded that flies of the family Tephritidae were not
suitable for detailed chromosome studies. He did state, however,
that this species appeared to have a haploid number of six, though
he presented no figures. Keuneke (1924), on the other hand,
obtained clear metaphase complements from Tephritis arnicae L.,
which had an interesting XO instead of the normal XY sex deter-
mining mechanism found in most Diptera. This configuration resulted
in a diploid number of 1 1 in the male and 12 in the female. A reduced
number of chromosomes has also been reported for A. ludens by
Emmart (1935). A haploid number of 5 was found in spermato-
genesis, though the findings of the present study do not support these
observations for this species.

In more recent studies, Frizzi and Springhetti (1953) described
the karyotype of the olive fruit fly, Dacus oleae Gmel., as having a
haploid number of 6. This same modal number of 6, which seems
to be common in most higher Diptera (White, 1954), has also been
reported for six out of seven species of Queensland Dacinae by Davis
( 1 955 ) . One species had a haploid number of 7. Davis apparently
encountered some technical difficulty, as he was unable to observe
any details in the morphology of the chromosomes. Mendes (1958),
however, was able to find distinct morphological differences in the
chromosomes of two species of Brazilian tephritids, Anastrepha
fraterculus (Wied.) and Ceratitis capitata (Wied.), both of which
had the characteristic haploid number of 6. His description of the
karyotype of A. fraterculus is of particular interest in that he found
morphologically differentiated sex chromosomes. These distinguish-
able heterochromosomes were not found in the Mexican population
of this species by the author. The importance of these differences
will be discussed later.

Methods

The chromosomes of the following nine species of tephritids were
analyzed during the course of this investigation: Anastrepha ludens
(Loew) ; A. zuelaniae Stone; A. fraterculus (Wied.) ; A. mombin-
praeoptans Sein; A. distincta Greene; A. spatulata Stone; A. striata
Schiner; A. serpentina (Wied.) ; and A . aphelocentema Stone.

Larvae were reared from field collected fruit which was held in

1962]

Bush — Genus Anastrepha

89

racks over moist sand in well ventilated wooden boxes. A sample
of each collection was reared to the adult stage to confirm preliminary
identification. Some species, such as A. ludens , A. mornbinpraeoptans ,
A. f rater cuius , and A. serpentina , were also reared on a laboratory
diet of ground carrots and yeast (Finney, 1956). Eggs of these
species were collected from females which were induced to oviposit
in wax impregnated cheese cloth shells, formed and pigmented to
represent fruit (McPhail and Guiza, 1956). For most cytological
investigations only larvae in the prepupal stage were used. Other
larval stages had suitable but fewer metaphase plates.

The supraoesophageal and suboesophageal ganglion were used for
the evaluation of all karyotypes with the exception of those of A.
spatulata whose host and larva are not known, though the adult is
collected in large numbers at certain times of the year. Adult
spermatogonial metaphase plates were therefore used to establish the
karyotype of this species. Attempts were made to obtain suitable
oogonial metaphase plates, but these were unsuccessful. Larval and
adult tissues were dissected out in normal saline (0.75 NaCl) and
transferred immediately to a saturated solution of coumarin in
distilled water for six to ten minutes following the technique of
Sharma and Bal (1953) and Manna (1956). The majority of the
species, including those treated statistically, were pretreated in cou-
marin for seven minutes. Care had to be taken not to exceed ten
minutes as chromosomes tended to become condensed and unsuitable
for study (Fig. 8). However, the shortening effect of coumarin, if
used judiciously, makes it possible to obtain well flattened metaphase
plates that show the structural features of the chromosomes distinctly.
Without the use of coumarin, chromosomes remained bunched and
no structural detail could be observed.

Tissue that had been pretreated in coumarin was then transferred
either directly into aceto-orcein (2% orcein in 45% glacial acetic
acid) for 30 minutes to one hour, or hydrolyzed in iN HC1 for 30
seconds to one minute at room temperature prior to staining. Hydroly-
sis improved the over-all qualities of the preparations. Squashes were
then made in a drop of aceto-orcein on albuminized slides using
coverslips treated with a silicon anti-wetting agent, such as Desicote3,
and made permanent following the simple and rapid quick-freeze
method of Schultz et al. ( 1949) , as modified by Conger and Fairchild
(i953).

3Beckman Desicote 18772, Beckman Scientific Instruments Division, Fuller-
ton, California.

90

Psyche

[September

Photomicrographs were taken on 35 mm. Adox KB- 14 film with
the aid of a Micro Ibso attachment using a Zeiss 90X apochromatic
oil immersion objective of NA 1.3 and a Leitz iox ocular. All
films were developed with Neofin blau.4 Prints were made on No. 5
Kodabromide paper. Final magnification of all prints used in statisti-
cal analysis was 3750X.

Measurements of chromosome lengths were made from photomicro-
graphs after the method of Boyes and Wilkes (1953), as modified
by Robertson (1957), on A. fraterculus, A. mombinpraeoptans, and
A. distincta whose karyotypes could not be distinguished by visual
inspection. All measurements were carried out to the nearest 0.5 mm.
and the percent of the total complement length of each chromosome
pair calculated. A sine transformation was then made on the resulting
percentages to reduce any correlation between the means and their
corresponding variances (Snedecor, 1956). An analysis of variance
was made on both the longest and the shortest chromosome pairs which
were the only chromosomes that could be consistently identified with
certainty. There was not sufficient evidence to reject the null hypothe-
ses that in these three species the mean lengths of the long chromo-
somes are the same or that the mean lengths of the short chromosomes
are the same. (Short chromosome: F=2.28<F.05(2, 69)— 3-J3- Long
chromosome: F=2.5i<F 05(2 ;69)=3.I3.) The karyotypes of A.

fraterculus, A. mombinpraeoptans, and A. distincta therefore could
not be distinguished from one another on the basis of mensural
observations.

Description of Karyotypes

The terminology used throughout the following descriptions and
discussion of metaphase chromosomes is the same as that outlined
by White (1957) except for the terms used to designate the position
of the kinetochore or centromere. Major chromosome arms (MCA)
were considered only when they were clearly visible in the metaphase
plate as a point of flection or bend in the chromosome. This does
not rule out the possibility of missing a short arm that would be visible
only in anaphase configuration. Such chromosomes would be con-
sidered acrokinetic. A metakinetic chromosome has two major arms
with the kinetochore located near the center. Acrokinetic chromosomes
have the kinetochore located near the end of the chromosome giving
the appearance at metaphase of being one-armed. Dot chromosomes
are treated as though acrokinetic, although in future investigations

4Neofin blau, Tetenal-Photowerk, Hamburg, Germany.

1962]

Bush — Genus Anastrepha

91

these may prove to be metakinetic as has been shown in the IV
chromosomes of Drosophila melanogaster Meigen (Kaufmann, 1934).

The locality and the host fruit from which the karyotype was
described is also included in anticipation that future studies may
uncover chromosomal polymorphism or sibling species within this
genus. Whenever observable sex chromosomes were present, the
heterogametic sex was always the male, as is normal for Diptera.
This characteristic was checked by studying spermatogonial meta-
phase plates of adult males.

Anastrepha ludens (Loew)

Figure 1

The diploid number is 12. The MCA number is also 12 in both
sexes as all chromosomes are acrokinetic. No secondary constrictions
were noted. The male has a small dot Y chromosome about 1/4 to
1/3 the length of the rod-shaped X chromosome. Forty-seven meta-
phase plates were photographed from 16 larvae. A total of over 300
larval brain squashes from various localities were studied but not
photographed.

Source of cytological material. Cuernavaca, Morelos, Mexico.

Host plant. Mango ( Mangifera indica L.).

Collections of this species were made throughout the year from
many host plants other than mango. These included sapote domingo
( Mammea americana L.) ; avocado ( Persea americana Mill.) ; yellow
chapote ( Sargentia greggii S. Wats.) ; white sapote ( Casimiroa edulis
Llave and Lex.) ; sweet orange (Citrus sinensis (L.) Osbeck) ; sour
orange ( Citrus aurantium L.) ; and grapefruit (Citrus grandis (L.)
Osbeck) .

Collections were also made in the states of Colima, Veracruz,
Chiapas, Tamaulipas, Jalisco, Mexico, and Michoacan. No variation
in the karyotype from these localities was noted.

Anastrepha zuelaniae Stone
Figure 2

The diploid number is 12 in both sexes. The MCA number is 12
with all chromosomes acrokinetic. No secondary constrictions were
noted. The male has a rod-shaped Y chromosome about 2/3 the length
of the X chromosome. Thirty-nine metaphase plates were photo-
graphed from nine larvae. Sixty-three larval brain squashes were
studied but not photographed.

92

Psyche

[September

Figs. 1-9. Metaphase plates from the brain of: (1) Anastrepha ludens;
(2) A. zuelaniae; (3) A. fraterculus (A. mombinpraeoptans and A. distincta
are closely similar) ; (4) A. striata; (5) A. aphelocentema ; (6) A. serpentina
$ showing XiX2Y sex chromosomes ; (7) A. serpentina $ showing X1X1X2X2
sex chromosomes. (8) Extreme contraction of chromosomes of A. ludens
resulting from extended pre-treatment in coumarin. (9) Spermatogonial
metaphase plate from testes of adult A. spatulata. (Magnification of all
plates 1500X)

1962] Bush — Genus Anastrepha 93

Source of cytological material. Tamazunchale, San Luis Potosi,
Mexico.

Host plant. Volador ( Zuelania guidonia Britt, and Millsp.).

Collections were made from late May to early July 1957. The
larvae were found only in the fruit of the above host plant.

Anastrepha fratereulus (Wiedemann)

Figure 3

The karyotype of this species cannot be distinguished from those
of A. mombinpraeoptans and A. distincta at metaphase. The diploid
number in both sexes is 12. The MCA number is also 12, with all
chromosomes acrokinetic. No morphologically differentiated hetero-
chromosomes (XY) or secondary constrictions were observed. One
hundred twenty-eight metaphase plates were photographed from 32
larvae. Over 250 larval brain squashes were studied but not
photographed.

Source of cytological material. Monte Blanco, Veracruz, Mexico.

Host plant. Rose apple ( Eugenia jambos L.).

Collections were made from early July to late August, 1957.

Anastrepha mombinpraeoptans Sein
cf. Figure 3 (A. fratereulus)

The karyotype of this species cannot be distinguished from those of
A. fratereulus and A. distincta. The MCA number in both sexes
is 12. No morphologically differentiated heterochromosomes (XY)
or secondary constrictions were observed. Forty-six metaphase plates
were photographed from 14 larvae. More than 150 larval brain
squashes were studied but not photographed.

Source of cytological material. Cocoyoc, Morelos, Mexico.

Host plant. Hog plum ( Spondias mombin L.).

Collections were made from September through October, 1957.
Larvae which were reared from mangoes (M. indica) collected from
Veracruz from June through July, 1957 were also studied.

Anastrepha distincta Greene
cf. Figure 3 (A. fratereulus)

This species cannot be distinguished from either A. mombinprae-
optans or A. fratereulus. It has an MCA number of 12 in both sexes.
No morphologically differentiated heterochromosomes (XY) or sec-

94

Psyche

[September

ondary constrictions were observed. Forty metaphase plates were
photographed from seven larvae. Over 80 larval brain squashes were
studied but not photographed.

Source of cytological material. Cocoyoc, Morelos, Mexico.

Host plant. Inga inicuil Cham. & Schlecht.

Collections were made from August through October, 1957.

Anastrepha spatulata Stone
Figure 9

The diploid number is 12 in the male. The MCA number is 14,
as one pair of chromosomes is metakinetic. The male has a small rod-
shaped Y chromosome about 1/4- 1/3 the length of the X chromo-
some. Only eight adult males were available for study; three of these
gave suitable preparations for analysis. From these, four sperma-
togonial metaphase plates were obtained. Several photographs were
taken of one particularly good metaphase plate that regrettably did
not lie in one plane. A drawing made from a composite photograph
is therefore presented in Figure 9.

Source of cytological material. Canyon de Lobos, Morelos, Mexico.

Host plant. Unknown.

Adults were collected from January to March 1957 in glass traps,
using fermenting brown sugar as a lure (Baker et al., 1944). Each
adult was maintained for two weeks in the laboratory on a standard
laboratory diet (Rhode, 1957) prior to examination.

Anastrepha striata Schiner
Figure 4

The diploid number is 12 in both sexes. The MCA number is 16.
Two pairs of chromosomes are metakinetic. The shorter metakinetic
chromosomes have secondary constrictions on their longest arms. The
dot Y chromosome is present in the male and is about 1/4 to 1/3
the length of the X chromosome. Seventy-one plates were photo-
graphed from 20 larvae. Approximately 175 larval brain squashes
were studied but not photographed.

Source of cytological material. Tequila, Jalisco, Mexico.

Host plant. Guava (P sodium guajava L.).

Collections were made from Tequila during the month of August
1957- Other collections were made from guava in the states of Vera-
cruz, Chiapas, Morelos, and Michoacan in 1956 and 1957 with no
variation in chromosome morphology noted.

1962]

Bush — Genus Anastrepha

95

Anastrepha aphelocentema Stone
Figure 5

The diploid number in both sexes is 12. The MCA number is
22 as there are five pairs of metakinetic autosomes. The dot Y chro-
mosome is about 1/4 the length of the rod-shaped X chromosome
which has a proximal secondary constriction. Twenty-five metaphase
plates were photographed from five larvae. Approximately 35 larval
brain squashes were studied but not photographed.

Source of cytological material. Tamazunchale, San Luis Potosi,
Mexico.

Host plant. Socavite ( Lucuma standleyana Pittier).

This species was studied only from the area surrounding Tam-
azunchale during the months May through July, 1 957*

Anastrepha serpentina (Wiedemann)

Figures 6 and 7

The diploid number in the male is 1 1 and in the female 12. The
male (Fig. 6) has an MCA number of 20 as there are four meta-
kinetic pairs of autosomes, one of which in some metaphase comple-
ments has a secondary constriction on its longest arm (not visible in
photomicrograph of the male metaphase plate). There are also three
heteromorphic acrokinetic sex chromosomes designated XiX2Y. Both
the X1? which is the shortest, and the X2 have small proximal sec-
ondary constrictions which are not always visible. The long Y
chromosome is easily distinguished as it has a short arm separated
from the rest of the chromosome by what has been interpreted as the
kinetochore, though future studies may show this to be a secondary
constriction. The female karyotype (Fig. 7) has an MCA number
of 20 with four pairs of metakinetic autosomes and two acrokinetic
pairs of sex chromosomes. Since the Xx is much shorter than the X2
(Xi ca. 2/3 X2) , these two chromosomes are easily differentiated.
Thirty-nine metaphase plates were photographed from 13 larvae.
Approximately 75 larval brain squashes were studied but not photo-
graphed.

Source of cytological material. Monte Blanco, Veracruz, Mexico.

Host plant. Mamey ( Calocarpum mammosum (L.) Pierre).

Collections were also made from the states of San Luis Potosi and
Morelos in 1957. It should be noted here that one collection of this
species was made from Tapachula, Chiapas in 1956 from mamey.
Samples taken from this population did not demonstrate the com-

g6

Psyche

[September

pound5 sex determining mechanism. The diploid number was 12 and
the MCA number was 24. No heteromorphic chromosomes were
present. However, the collection was made at the beginning of the
study when methods were not yet perfected for making temporary
squashes permanent, and before photographic equipment was avail-
able. For this reason, only a few drawings were made. This dif-
ference in karyotype morphology may have resulted from inadequate
technique, but more likely it represents a different species.

Discussion

To date at least 19 species representing 10 genera in the family
Tephritidae have been investigated cytologically by several authors.
It is apparent even from the few species thus far studied that a great
deal of chromosomal variation exists within the family. Such varia-
tion not only includes characteristic positions of the kinetochore,
secondary constrictions, and chromosome length, but also involves
differences in chromosome number and sex determining mechanisms
as well. These differences can be put to good use in the identification
of immature forms and may possibly aid in establishing phylogenetic
relationships. It must be stressed that cytotaxonomy is seldom if ever
a “solve all” method of identification, and it is not surprising that
three species of Anastrepha show no distinct chromosome differences.
It is probable that as more species in this genus are investigated
chromosome patterns will be found similar to the ones reported here
as morphologically distinct. A combination of several criteria, in-
cluding chromosomal variations, gross morphology of the larvae, and
various ecological aspects of the species in question, may therefore
be necessary before accurate identification can be made.

With such limitations in mind, the following key is presented as
a tentative means of separating the larvae of six of the nine Mexican
Tephritidae investigated cytologically so far. Due to the similarities
of some female karyotypes, the key is based on the chromosome morph-
ology of the male karyotype whenever it is known. This makes it
advisable to study at least eight larvae (assuming a 1 :i sex ratio)
in a given collection to be fairly certain that all are not of the same
sex. No suitable means has yet been found to determine the sex of
immature forms in this family without resorting to karyotype analysis.

5Schrader’s (1928) terminology is followed here. A compound sex deter-
mining mechanism is one in which the X or the Y is represented by more
than one element in contrast to a multiple sex determining mechanism in
which there is an adherence of chromosomes belonging to different pairs.

1962] Bush — Genus Anastrepha 97

Cytotaxonomic Key to the Metaphase Plates of the Common
Species of Mexican Anastrepha (males only)

i. All chromosomes evidently acrokinetic (MCA=I2) 2

— All chromosomes not all acrokinetic (MCA^^ or more) . ... 4

2. No heteromorphic chromosome pairs present at metaphase

A. mombinpraeoptans , A. f rater cuius , A. distincta

— Heteromorphic chromosome pair present 3

3. Small dot Y chromosome present (Y<o.5 X) A. ludens

— Rod-shaped Y chromosome present but shorter than

X chromosome (Y>0.5 X) A. zuelaniae

4. MCA=i4 to 16 5

— MCA=i7 to 22 6

5. MCA=I4 A. spat ul at a

— MCA=i6 A. striata

6. MCA=20, XiX2Y sex mechanism present (2n=n)

A. serpentina

— MCA — 22, secondary constriction on X chromosome

A. aphelocentema

Though cytological evidence per se is not always suitable for dis-
tinguishing some species of Tephritidae, it can support and elucidate
certain phylogenetic relationships within the group. The cytogenetics
of certain Diptera has been extensively studied in the past three or
four decades so that many of the mechanisms of chromosome evolu-
tion in this group are now fairly well understood (Patterson and
Stone, 1952; White, 1954; da Cunha, i960). Since the number
of species of Tephritidae so far investigated is extremely limited, it
is as yet impossible to determine any conclusive generic or interspecific
relationships, though some interesting possibilities do appear, par-
ticularly at the interspecific level in the genus Anastrepha.

It is possible that A. mombinpraeoptans , A. distincta, A. ludens,
A. zuelaniae, and the Mexican and Brazilian forms of A. fraterculus
may form part of a chromosome complex representing a subgenus or
species group within the genus Anastrepha. This is supported by the
similarity in the morphology of the adults of these species. The
difference in karyotypes between the Mexican form of A. fraterculus
reported here and the Brazilian population described by Mendes is
interesting since this difference may represent a case of chromosomal
polymorphism or, more likely, sibling species. Biological data support
the latter (A. C. Baker et al., 1944; E. W. Baker, 1945) in that
slight but consistent morphological differences exist in the adults

98

Psyche

[September

from these widely separated areas. Such differences could be attributed
to geographical variation ; however, they also have distinctly different
host preferences. The Brazilian population has a wide host range
and is a destructive pest of citrus, while the Mexican population is
of no economic importance, infesting the rose apple and only occasion-
ally the guava.

The case of the compound sex determining mechanism encountered
in A. serpentina is also interesting as this type of system appears to
be rare in Diptera. Dobzhansky (1935) reported XiX2Y system in
Drosophila miranda Dobzh. believing it to be an example of deter-
minate disjunction. Cooper (1946), however, clearly showed that a
XiYX2 trivalent was actually formed during meiosis. Boyes (1952)
found the same type of trivalent formed in Hylemya fugax (Meig.).
It is possible that A. serpentina may also produce a trivalent, but the
preparations of gonadal tissue using the squash technique were not
suitable for establishing the interaction of the three sex chromosomes.

The different chromosome number of 2n=iO reported by Emmart
(1935) for A. ludens probably resulted from an incorrect interpreta-
tion of chromosome morphology in her study of meiosis in pupal and
adult testes. In the present study, larvae as well as adults were
studied from Cuernavaca, Morelos, the same locality from which
Emmart collected most of her material. A diploid number of 12
was always recorded. Meiotic figures in the testes without exception
had a characteristic haploid number of 6.

Little can be said about the other Anastrepha species at this time.
It is likely that a more thorough investigation of the karyotypes
within this genus will uncover many interesting phylogenetic rela-
tionships which can now only be hinted at on the basis of the present
study.

Spermatogonial metaphase plates can be put to good use in eval-
uating the chromosome morphology of those species of tephritids whose
larvae are unknown, as in the case of A. spatulata, or whose larvae
cannot be readily maintained in the laboratory. Such determinations
can also be used to obtain tentative identification of larvae collected
for the first time, and whose chromosome morphology is known only
from previously captured adults.

From the cytological data thus far accumulated for the family
Tephritidae, it appears that the variation between karyotypes is suf-
ficient to warrant more attention from the taxonomists of this group.
New methods of handling animal chromosomes, such as the many
pre-treatments now available, followed by simplified squash tech-

1962]

Bush — Genus Anastrepha

99

niques, have eliminated many arguments against inclusion of cyto-
logical data in taxonomic studies. Cytological information in many
cases offers the taxonomist who is interested in establishing better
phylogenetic relationships a tool which can often supplement and
strengthen his conclusions based on morphological data, as well as
provide information not available by any other means. In the family
Tephritidae this seems particularly true.

Summary

The karyotypes of nine species of Anastrepha (Tephritidae,
Diptera) are described on the basis of mitotic metaphase morphology.
The species include A. ludens, A. fraterculus , A . distincta, A. mom-
binpraeoptans, A. zuelaniae , A. spatulata, A . striata , A. serpentina ,
and A. aphelocentema. All species have a diploid number of 12, with
the exception of the males of A. serpentina where an XxX2Y sex
determining mechanism resulted in a diploid number of n. Only
six of the nine species investigated could be identified on the basis
of chromosome morphology. It is suggested that A. distincta , A.
mombinpraeoptans , and Mexican A. fraterculus , which have cyto-
logically indistinguishable karyotypes, as well as A. ludens, A.
zuelaniae, and the Brazilian form of A. fraterculus may represent
part of a chromosome complex within the genus Anastrepha. The
differences between the karyotypes of the Brazilian and Mexican
populations of A. fraterculus, along with differences in external
morphology and biology, suggest that these two forms may repre-
sent sibling species. In general, it is concluded that the metaphase
chromosomes of the family Tephritidae can be used for critical cyto-
taxonomic and phylogenetic studies.

Acknowledgements

1 would like to express my appreciation to Srs. F. E. Guiza, M.
Aguilar, and J. Ramirez for their assistance in rearing and main-
taining field collections and laboratory populations during the course
of this study. I wish also to express my gratitude to Drs. R. H. Foote,
M. S. Wasbauer, and Messrs. F. L. Blanc and E. G. MacLeod
for reading and discussing the manuscript with me. I would par-
ticularly like to thank Dr. Kenneth W. Cooper, Dartmouth College
Medical School, for his critical review and helpful comments con-
cerning various aspects of this work.

IOO

Psyche

[September

Literature Cited

Baker, A. C., W. E. Stone, C. C. Plummer, and M. McPhail.

1944. A review of studies on the Mexican fruitfly and related Mexican
species. U. S. Dept. Agr. Misc. Publ. 53 1 : 1-155.

Baker, E. W.

1945. Studies on the Mexican fruitfly known as Anastrepha fraterculus.
Jour. Econ. Ent. 38:95-100.

Boyes, J. W.

1952. A multiple sex-chromosome mechanism in a root maggot. Jour.
Hered. 43 : 194-199.

Boyes, J. W., and A. Wilkes.

1953. Somatic chromosomes of higher Diptera. I. Differentiation of
tachinid parasites. Can. Jour. Zool. 31:125-165.

Conger, A. D., and L. M. Fairchild.

1953. A quick-freeze method for making smear slides permanent. Stain
Technol. 28:281-283.

Cooper, K. W.

1946. The mechanism of non-random segregation of sex chromosomes
in male Drosophila miranda. Genetics 31:181-194.

Da Cunha, A. B.

1960. Chromosomal variation and adaptation in insects. Ann. Rev.
Ent. 5:65-110.

Darlington, C. D.

1956. Chromosome Botany. G. Allen and Unwin Ltd. 186 pp.

Davis, J. J.

1955. A note on chromosomes of some Queensland Dacinae (Trypetidae,
Diptera). Queensland Jour. Agr. Sci. 12:161-173.

Dobzhansky, Th.

1935. Drosophila miranda, a new species. Genetics 20:377-391.
Emmart, E. W.

1935. Studies of the chromosomes of Anastrepha (Diptera: Trypetidae).
I. The chromosomes of the fruit-fly, Anastrepha ludens Loew.
Proc. Ent. Soc. Wash. 37:119-135.

Finney, G. L.

1956. A fortified carrot medium for mass-culture of the Oriental fruit
fly and certain other tephritids. Jour. Econ. Ent. 49:134.

Frizzi, G. and A. Springhetti.

1953. Prime ricerche citogenetiche sul “Dacus oleae Gmel.”. Ricerca
Sci. 23 : 1612-1620.

Kaufmann, B. P.

1934. Somatic mitoses of Drosophila melanogaster. Jour. Morph. 56:
125-155.

Keuneke, W.

1924. liber die Spermatogenese einiger Dipteren. Z. Wissenschaftliche
Biol. 1:357-412.

Manna, G. K.

1956. Handling human chromosomes by a coumarin technic. Stain Tech-
nol. 3 1 : 45-50.

McPhail, M., and F. E. Guiza.

1956. An oviposition medium for the Mexican fruit fly. Jour. Econ.
Ent. 49:570.

Mendes, L. O. T.

1958. Observagoes citologicas em “moscas das frutas”. Bragantia 17:
29-39.

Metz, C. W.

1916. Chromosome studies on the Diptera. II. The paired association

1962]

IOI

Bush - — Genus Anastrepha

of chromosomes in the Diptera, and its significance. Jour. Exptl.
Zool. 21:213-279.

Patterson, J. T., and W. S. Stone.

1952. Evolution in the Genus Drosophila. The MacMillan Co. 610 pp.
Phillips, V. T.

1946. The biology and identification of trypetid larvae (Diptera:
Trypetidae). Mem. Amer. Ent. Soc. No. 12.

Rhode, R. H.

1957. A diet for Mexican fruit flies. Jour. Econ. Ent. 50:215.
Robertson, J. G.

1957. Somatic metaphase chromosomes in geographic isolates of the
carrot rust fly Chamaepsila rosae (F.) (Diptera: Psilidae). Can.
Jour. Zool. 35:453-458.

Schrader, F.

1928. Die Geschlechtschromosomen. Berlin: Borntraeger.

Schultz, J., R. C. MacDuffee, and T. F. Anderson.

1949. Smear preparations for the electron microscopy of animal chro-
mosomes. Science 110:5-7.

Sharma, A. K., and A. K. Bal.

1953. Coumarin in chromosome analysis. Stain Technol. 28:255-257.
Snedecor, G. W.

1956. Statistical Methods. 5th ed. Iowa State College Press. 534 pp.
White, M. J. D.

1954. Animal Cytology and Evolution. 2nd ed. Cambridge Univ. Press.
454 pp.

1957. Cytogenetics and systematic entomology. Ann. Rev. Ent. 2:71-90.

REDESCRIPTION OF LITHOPHOTINA FLOCCOSA
COCK. (MANTEODEA) WITH SOME NOTES ON
THE MANTEOD WING VENATION

By A. G. Sharov

Paleontological Institute, Academy of Sciences, USSR, Moscow

Dr. G. M. Byers recently sent to me for study the wing of an
insect from the Miocene deposits of Colorado. I determined this wing
to be the hind wing of a manteod, close to Lithophotina floccosa
Cock., which Cockerell described in 1908 from the same deposits. At
my request and with the approval of Dr. Hugo Rodeck (University
of Colorado Museum) Professor F. M. Carpenter brought the
holotype of L. floccosa to Moscow with him at the time of his visit
to the Paleontological Institute. A study of the holotype led me to
the conclusion that the hind wing which Dr. G. M. Byers sent most
probably belongs to the same species, i.e. L. floccosa. A comparison
with the venation of the fore and hind wings of the living manteod
Chaeteesa filata Burm., which belongs to the primitive relict family
Chaeteesidae, confirms this. L. floccosa also ought to be referred to
this family. Since the description of L. floccosa given by Cockerell
is very short, and since drawings are lacking and the photograph in
the article is poorly reproduced, a redescription of L. floccosa , with
a description of the hind wing of this species, is given below.

Family Chaeteesidae
Lithophotina floccosa Cock.

Figure 1

Lithophotina floccosa Cock., 1908, Canad. Ent., 40:343-344.

The fore wing of this species (Figure iA) was found in the
Miocene deposits of Colorado (Florissant) at Station No. 14. The
length of the wing is 21.5 mm, the width 7 mm. The wing is
narrowed at the base. Sc is long, taking up more than two-thirds
the length of the wing, ending at its widest portion. The branches
of Sc close to the base of the wing have the character of cross-veins;
in the center, together with the cross-veins which join them, they
form a double row of cells; and close to the apex they take on the
character of gently curving branches. At the apical portion of the

1 Manuscript received by the editor July 21, 1962. The editor is indebted
to Miss Helen V aitaitis for the English translation of the article.

102

1962]

Sharov — Lithophotina

103

wing, R has 4-5 branches in pectinated form, which are directed
toward the fore margin. MA at the center of the wing length forms
two branches, of which the fore branch divides again close to the
apex, and the hind one forms two short branches at the very apex.
The vein MP is merged with CuA and its base apparently forms
one of the cross-veins. MP + CuA dichotomizes and forms seven
branches.

Between R, M and their branches there is a double row of cells.
The anal portion in the holotype is missing; evidently it was torn
off while the wing was being carried by currents of water before it
was buried. The longitudinal and cross-veins are colored with brown
pigment.

The hind wing (Figure iB) was found by G. M. Byers in July
of 1953 in the same deposits at the Park County station. The length
of the hind wing is 18 mm. The different character of the venation
of the fore and hind wings, and also the ratio of the length of the
fore wing to the length of the hind wing of L. floccosa, is very close
to that of Chaeteesa filata Burm. (Figure 2), which is the reason
for placing the specimen of the hind wing found by G. M. Byers in
the same species, i.e. L. floccosa.

Sc ends at a distance less than two-thirds of the length of the
wing from the base. R has three oblique branches. Rs is simple and
does not branch. M is joined with R only at the base of the wing,
proximal to the beginning of MP. MA forms three branches in the
same sequence as in the fore wing. MP has the character of a cross-
vein but is stronger in comparison with the other cross-veins. MP +
CuA forms three branches. In the distal portion of the wing between
the longitudinal veins there is a double row of cells. Both the longi-
tudinal and the cross-veins are colored with brown pigment.

Although there is no doubt about the closeness of the Manteodea
to the Blattodea, the interpretation of the venation of the former,
particularly with respect to the hind wing, remains questionable (see
Smart, 1956, p. 550). In such cases the study of fossil material is
extremely desirable, and the fossil representative of the Manteodea
examined here actually permits a simple solution of some debatable
questions on the venation of these insects. (See figure 2).

Chopard (1949) and Ragge (1955) find an independent RS in
the fore wing of some Manteodea, the nature of which becomes clear
upon examination of the fore wing of Lithophotina, where R has
the same type of pectinated branching as in most Blattodea. In
Chaeteesa, as Smart (1956) has correctly noted, the remains of these

104

Psyche

[September

branches are short oblique veinlets on the end of R. In Mantis ,
Ameles and several other Manteodea only two of these branches are
preserved, the hind one usually being mistaken for Rs.

, Sc

Fig. 1. Drawing of Lithophotina floccosa Cock.; A. Holotype (fore wing) ;
B. Homeotype (hind wing).

The stem following R which is mistakenly identified in Blattodea
and Manteodea as M, in the opinion of the author, is this only in
the proximal portion. Its distal portion together with the branches
is really the fore branch of M, i.e. MA, whereas the hind branch,
MP, is merged with CuA. Only in some Paleozoic Blattodea
( Archimylacris pruvosti Laur., Aphtoroblattina shottoni Laur.) is
the base of MP in the fore wings represented as an oblique vein,
which in the rest of the Blattodea and in all known Manteodea is
missing or has the character of a cross-vein, just as has occurred in
the evolution of the Plecoptera (Sharov, i960).

The peculiarities of the venation of the hind wing of L. floccosa
confirm the correctness of the interpretation of R and Rs in the hind
wings of the Manteodea proposed by Ragge (1955) on the basis of
the study of the tracheation and accepted later by Smart (1956),
although with some reservations (p. 550). In L. floccosa Rs is
separated from M and cannot be considered as the fore branch of M.

1962]

Sharov — Llthophotina

105

Thus, there cannot be any doubt about the fact that, in the hind
wings of the Manteodea, the vein which is situated behind R is Rsy
which is characteristic also for the hind wings of the Paleozoic and
Mesozoic Blattodea.

In the hind wings of the Manteodea a free base of MP, which
Smart (1956) designates as the “oblique cross-vein”, is preserved.
Among the Polyneoptera a similar vein is preserved in the hind wings

(After Smart, 1956, with changed designations of the veins.)

of Plecoptera and Orthoptera. Its homology with the same vein
in the fore wings, in the cases where it is preserved, seems obvious.

Acknowledgements

1 wish to thank Professor F. M. Carpenter and Dr. Hugo Rodeck
(University of Colorado) for giving me the opportunity to study the
holotype of L. floccosa and for the photographs of this fossil insect,
which were used for the preparation of the drawings. I am very
grateful also to Dr. G. M. Byers for sending the specimen of the
hind wing of this insect.

Literature Cited

Chopard, L.

1949. Ordre des Dictyopteres. In P. P. Grasse, Traite de Zoologie 9,.
Masson, pp. 353-407.

io6

Psyche

[September

Cockerell, T. D. A.

1908. The first American fossil Mantis. Canad. Ent. 40:343-344.
Ragge, D. R.

1955. The wing venation of the Orthoptera Saltatoria with notes on
Dictyopteran wing venation. British Museum (Nat. Hist.), pp.
1-159.

Sharov, A. G.

1960. The origin of the order Plecoptera. Verh. XI. Int. Kongr. Ent.,
Wien, 1960, 1:296-298.

Smart, J.

1956. On the wing venation of C haeteesa and other mantids (Insecta
Mantodea). Proc. Zool. Soc. London 127: 545-553.

THE INFRABUCCAL POCKET OF A
FORMICINE ANT: A SOCIAL FILTRATION DEVICE1

By T. Eisner and G. M. Happ
Department of Entomology, Cornell University

The importance of regurgitative feeding as a means for distributing
liquid nutrient among workers, and from workers to larvae and queen,
has long been recognized, and it is now generally agreed that this
process is a most fundamental bond in the social life of many ants
(Le Masne, 1953; Wallis, 1961 ; Wheeler, 1923; Wilson and Eisner,
1957). Unlike honeybees, which also feed one another by regurgita-
tion, but which have their principal food reservoir outside their own
bodies in the honeycomb, ants store liquids exclusively within the crops
of the individual living workers. Crop storage and regurgitative feed-
ing are probably most highly developed in the specialized and success-
ful subfamilies Formicinae and Dolichoderinae. It is in these ants that
the crop is most capacious (witness the fact that ants with “replete”
castes are restricted to these subfamilies), and it is these that have a
special device, in the form of an elaborately refined proventriculus,
adapted to dam the posterior outlet of the distended crop (Eisner,
1957; Eisner and Brown, 1958).

In its basic features, the proventriculus of Dolichoderinae and For-
micinae is really no different from that of other ants and of Hymenop-
tera in general. It is a mechanical pump, consisting of a strongly
muscled bulb, with an anterior intake valve communicating with the
crop, and a posterior outlet valve leading to the midgut. When
nutrient is to be passed from foregut to midgut, the bulb is put into
operation, and through a series of rhythmic compressions and decom-
pressions, liquid is effectively pumped along. Whereas in most Hymen-
optera the intake valve of the proventriculus is a more or less mobile
portal, incapable of effective prolonged closure, and hence ill-adapted
to withstand for protracted periods the liquid pressure from a filled
crop, in formicine and most dolichoderine ants the portal valve is
permanently restricted to a narrow cruciform cleft through which
leakage of crop contents is virtually impossible. It is only during
proventricular pumping that liquid is passed through the clefts under

This study was supported in part by Grant E-2908 of the U. S. Public-
Health Service.

Manuscript received by the editor February 16, 1962.

107

108 Psyche [September

suction pressure from the activated bulb (Eisner, 19571 Eisner and
Brown, 1958).

Aside from its obvious advantage in making prolonged crop storage
possible, the permanent restriction of the proventricular portal poses
a problem. Solid particles passed into the crop have no place to go,
since they obviously cannot be pumped through the proventriculus.
What, then, happens to indigestible solids that are swallowed ? Or are
they perhaps not swallowed at all? Ants, as well as many other
Hymenoptera, have a so-called infrabuccal chamber, a ventral infold-
ing of the hypopharyngeal surface, that could conceivably act as an
effective solid-withholding device, guarding the opening to the crop.
Janet (1895a, 1895b, 1905) has described the anatomy of this struc-
ture, and has shown that in both ants and wasps, debris gathered while
cleaning themselves or their nestmates, as well as solid residue from
food ingested, often collects in this pocket and is ejected intermittently
as small discrete pellets. His observations were excellent, but left some
basic questions unanswered. The present study deals with an experi-
mental evaluation of the function of the infrabuccal pocket of a
formicine ant, Camponotus pennsylvanicus (DeGeer), in which crop
storage and regurgitative food transmission are known to be well-
developed social attributes (Plate 6).

Several laboratory-maintained colonies of Camponotus were avail-
able, but only nestmates, rather than internidally mixed lots, were
used for any one series of tests. For experimental purposes they were
fed honey mixed with various samples of corundum powder ranging
in particle diameter from iO/x to 300 [i.

EXPERIMENTS WITH INDIVIDUAL ANTS

An initial series of tests was designed to determine just how effective
a filtering device the infrabuccal pocket really is. Individual ants,
including only medium-sized workers, were confined in Petri dishes in
which they had access to single drops of honey-corundum mixture.
Eight corundum samples were tested (10, 20, 30, 80, 100, 150, 200,
and 300 /i) , each on 10-15 ants. The ants were starved for one or more
days before the tests, and, when introduced into the dishes, each would
promptly commence feeding and remain at the food source uninter-
ruptedly for up to several minutes. Only in a few exceptional cases
would an ant pause briefly partway through the meal and, after back-
ing away slightly and spreading its mandibles, would regurgitate a
small, typically kidney-shaped infrabuccal pellet, consisting of a densely
clumped packet of corundum plus small pieces of wood and other

1962] Eisner and Happ — Inf rabuccal Pocket 109

residual matter that the infrabuccal pocket must have contained
before the meal. As soon as the ants had gorged themselves and ceased
feeding they were drowned and their crops and infrabuccal chambers
examined under a microscope for corundum content. The crops were
dissected out and mounted intact in clearing medium, while the infra-
buccal pockets were induced to discharge their contents simply by
pressing the sides of the heads, causing the pockets to be everted. An
alternative technique was to examine crops and infrabuccal chambers
in intact corpses, after rendering their body cuticle transparent by
prolonged immersion in 10% aqueous KOH. The results were clear-
cut. Ants fed on the 200 p and 300p samples had no corundum
particles in their infrabuccal pockets and none in their crops : particles
of such caliber are evidently excluded altogether by the mouthparts
themselves, and they never even reach the infrabuccal chamber. Parti-
cles of the next smaller size tested (150/x) did get taken in, but only
as far as the infrabuccal pocket, which was invariably packed tightly
with them ; the crops were always clear. With the remaining samples
( 10-100 /i) the infrabuccal chambers were also replete with corundum,
but a substantial amount of particles had also been swallowed into the
crop ( Plate 7 ) . Evidently the narrow transverse slit by which food
gains entrance to the pharynx just above the infrabuccal chamber is
of such aperture as to bar particles larger than i^O/jl but not those of
iOO/x or less.

An additional experiment supported these findings. A group of ten
ants were fed individually on a honey sample as before, but this time
the mixture had particles of three sizes (10, 100, and 300/x). As
expected, the crops contained primarily iO/x-particles, the infrabuccal
chambers mainly 100/x-particles, and the 300/x-particles were not
recovered at alL

It is clear from the preceding that the filtering action of the infra-
buccal chamber is far from perfect, at least for particles smaller than
150 yx. Since the capacity of the chamber is limited, and since the
individual ant as a rule does not pause to rid itself of an infrabuccal
pellet every time the chamber is filled, but continues feeding even after
the chamber is full and no longer operative, one is led to believe that
in the normal course of food gathering a considerable amount of
particulate matter is likely to find its way into the crop of the indi-
vidual foraging ant. The experiments described next below were
designed to establish the fate of such particles, which must obviously
be voided in some fashion, or the crop would become solid-bound and
the proventriculus obstructed.

Psyche, 1962

Vol. 69, Plate 6

Eisner and Happ — Infrabuccal Pocket

1962] Eisner and Happ — Infrabuccal Pocket 1 1 1

EXPERIMENTS WITH GROUPS OF ANTS

Within the formicine ant society, such evidence as we have suggests
that regurgitative food transmission proceeds more or less continually
— at least during that part of the year when the colony is active.
Judging from the increased number of individual regurgitative
exchanges that can be witnessed in laboratory colonies that are given
renewed access to food following a period of deprivation, it is clear
that the arrival of returning foragers with new crop-loads greatly stim-
ulates the overall rate of intranidal exchange. Actual measurements
made with species of Formica, fed on food labelled with radioactive
tracer, have shown that the crop contents from single foragers may
become shared by an entire colony in a matter of hours (Wilson and
Eisner, 1957). Although no measurements have been made with
Camponotus , it is unlikely that the results with this close relative of
Formica would be much different. Evidently, an incoming crop-load,
in the course of being passed from ant to ant, and channelled, as it
were, through one infrabuccal chamber after another, could be expect-
ed to undergo progressive filtration and ultimately be completely
cleared of all debris. The following experiments proved that regurgi-
tative feeding does in fact provide a means by which the communal
crop supply is filtered and cleared.

Seven ants, fed to repletion on a honey- iO/a corundum mixture, were
each placed in a Petri dish with five unfed nestmates. A color marking
differentiated the laden ant from the others. Regurgitative donations
took place immediately, at the end of each of which the recipient
(previously unfed) worker was removed and killed before it in turn
had a chance to donate to others. Dissection of 25 recipients taken in
this fashion showed 22 of them to have corundum in their infrabuccal
pockets. Thirteen of these had their infrabuccal pockets packed full,
and some particles had already passed into their crops.

EXPLANATION OF PLATE 6

Fig. 1. Intact worker of Camponotus americanus Mayr, shown next to two
individuals whose appendages have been clipped, and whose gasters have
been dissected away to expose the digestive tract. Only crop (A), proventri-
culus (B), and midgut (C) are shown; the entire hindgut has been removed.
Notice the enormously distended crop in the freshly-fed forager (center),
contrasted with the crop of a starved individual (right). In Camponotus, as
in many other ants, the crop acts as a social stomach, capable of storing
amounts of nutrient far in excess of the demands of the individual forager.

Fig. 2. Regurgitative food exchange between two workers of Camponotus
pennsylvanicus (DeGeer). This is the process by which the liquid food
supply in the crops of incoming foragers is shared with the remainder of
the society.

Psyche, 1962

Vol. 69, Plate 7

Eisner and Happ — Infrabuccal Pocket

Fig. 1. Profile view of head of worker of Camponotus pennsylvanicus
(DeGeer), killed shortly after feeding and cleared in KOH, showing infra-
buccal chamber, plus pharynx and portion of oesophagus, densely packed
with corundum particles.

Fig. 2. Same as preceding, but of another individual, in dorsal view. The
infrabuccal chamber is full, but only few particles are seen in the oesophagus.

Fig. 3. Worker of Camponotus pennsylvanicus (DeGeer), killed shortly
after a meal and cleared in KOH, with cuticular shell of gaster dissected
away to expose the crop and proventriculus. Notice corundum particles in
the infrabuccal chamber, oesophagus, crop, and in the “calyx” (arrow) of
the proventriculus.

1962]

Eisner and Ilapp — Infrabuccal Pocket

113

Another experiment was designed to replicate more closely the
crowded conditions prevailing in actual nests. A total of 35 ants, fed
on honey- iO/x corundum and provided with color markings, were
divided into seven groups of five each, and each group was introduced
into an observation arena containing 50-75 unfed nestmates. Another
35 ants, similarly fed, served as controls, and were confined each ant
by itself. At the end of twelve hours both lots were killed, and a visual
estimate made of the corundum contents of their individual crops (one
ant of the first lot escaped). The results (text fig. 1) were strikingly
different in the two batches. Of the isolated ants, nearly every one had
the crop filled with corundum to one quarter or more of its capacity.
Of those that had been confined socially, about half had completely
clear crops, and in most of the remainder the crops were less than one-
quarter full with particles. Evidently, in the context of the society,
debris is rapidly filtered from the communal crop supply. One might
add that, whereas in each of the arenas containing the ants in groups
there were found at the end of the 12-hour period a dozen or more
infrabuccal pellets of corundum, the ants confined singly produced only
an occasional single pellet. The pellets recovered in the group-tests
stemmed not only from the five introduced food donors, but were
actually seen in some cases to have been ejected by residents that had
received food by regurgitation. (Ants confined singly over a period of
several days sometimes produced more than one pellet. Since during
this period they often regurgitated their particle-laden crop contents
onto the walls of their glass enclosures, the production of more than
one pellet may signify that their crop loads had undergone a series of
filtrations as a result of reingestion of the regurgitate. These observa-
tions may be of no more than incidental interest, since prolonged lone
confinement is obviously not the rule in nature.)

DISCUSSION

The above experiments show the infrabuccal chamber of Campono-
tus to be an organ of considerable social importance, in that it serves to
maintain the liquid communal crop supply particle-free. It would be
interesting to know whether the chamber is a similarly effective filter-
ing apparatus in other ants. In formicines, and in the more advanced
dolichoderines, the proventriculi of which are so constructed as to
preclude passage of solids, one would certainly expect this to be so.
Since the midgut of these ants is never likely to be exposed to the
abrasive action of particulate matter, it is noteworthy that a peritrophic

Psyche

[September

114

membrane has been found lacking in those species (including a Cam-
ponotus) of these two subfamilies that have been examined. The
primitive Myrmeciinae, the proventriculi of which have gaping portals
through which small particles could presumably pass, do have a mem-
brane (Waterhouse, 1953).

% CROP -LUMEN FILLED WITH PARTICLES

Text fig. 1. Frequency distribution of ants classified according to corundum
content of their crops. The ants were fed 12 hours earlier on honey laden with
10/i-corundum powder, and were confined immediately after the meal either in
isolation (black bars), or together with groups of unfed nestmates (striped
bars). See text for details.

One wonders whether, despite the effectiveness of infrabuccal filtra-
tion, enough detritus sometimes remains in the crop to interfere —
perhaps only temporarily — with the normal operation of the proven-
triculus: particles accumulated over the portal clefts might effectively

1962]

Eisner and Happ — Infrabuccal Pocket

1 15

block the passage of fluid during proventricular pumping. In this
connection it is of interest that some formicine proventriculi have a
special device that apparently serves to wipe the clefts of obstructing
solids (Eisner, 1957). In Camponotus , as well as in other formicines
with so-called “sepalous” proventriculi, such a device is missing, but
since the “calyx” of the proventriculus presumably undergoes rhythmic
constriction and dilation during the pumping cycle (the calyx is
enveloped by circular muscles), the space within it is likely to be
stirred sufficiently to prevent particles from accumulating over the
portal clefts on the sepals (Eisner, 1957).

In ants of some of the other subfamilies, the infrabuccal chamber
has been shown to serve special functions. Thus, in Atta and certain
other fungus growers (subfamily Myrmicinae), the chamber provides
the receptacle in which a supply of fungal spores is carried from the
parent colony by the departing nest-founding female that must ulti-
mately start a new fungus garden of her own (von Ihering, 1898;
Huber, 1905). In yet another subfamily, the Pseudomyrmecinae, the
larvae are fed with food pellets compacted in the infrabuccal pockets
of the workers, which deposit the pellets in a special postoral receptacle
(trophothylax) of the larva (Wheeler and Bailey, 1920).

In honeybees, the infrabuccal chamber is apparently inoperative as
a filter (Snodgrass, 1956). These insects rely on the intake and
digestion of pollen as a protein source for the subsequent manufacture
of brood food, and this special requirement can obviously be met only
in the absence of thorough preoral filtration. Interestingly, the pro-
ventriculus of honeybees is especially adapted to transmit dense pollen
suspensions to the midgut without becoming choked (Bailey, 1952) .

A representative comparative study of the infrabuccal chamber of
Hymenoptera has never been made. In the absence of such a study, it
is difficult to speculate on the evolutionary justification for the chamber
as it first arose within the order. But since adult Hymenoptera are
predominantly fluid feeders, one may reasonably presume that the
chamber functioned as a filter from the very outset [its filtering action
in at least some wasps has been demonstrated by Duncan (1939) and
Janet (1895b)]. To ants like Camponotus, as well as to all other
formicines and dolichoderines with an intranidal organization heavily
dependent on crop storage and regurgitative food transmission, the
infrabuccal filter is thus seen to represent an evolutionary preadapta-
tion of considerable importance. In the absence of an adequate preoral
filtration mechanism by which the communal crop supply is maintained
particle-free, the proventriculus could not have evolved toward pro-

1 1 6

Psyche

[September

gressive restriction of its aperture, and the crop might never have
achieved the extremes of storage capacity that it has in formicines and
dolichoderines.

References Cited

Bailey, L.

1952. The action of the proventriculus of the worker honeybee, Apis
mellifera L. J. exp. Biol. 29: 310-327.

Duncan, C. D.

1939. A contribution to the biology of North American vespine wasps.
Stanf. Univ. Pubh Biol. Sci. 8: 1-272.

Eisner, T.

1957. A comparative morphological study of the proventriculus of ants
(Hymenoptera : Formicidae). Bull. Mus. comp. Zool. Harv. 116:
439-490.

Eisner, T. and W. L. Brown

1958. The evolution and social significance of the ant proventriculus.
Proc. Xth int. Congr. Ent. 2: 503-508.

Huber, J.

1905. fiber die Koloniengriindung bei Atta sexdens. Biol. Zbl. 25: 606-
619, 625-635.

Ihering, H. von

1898. Die Anlage neuer Colonien und Pilzgarten bei Atta sexdens. Zool.
Anz. 21: 238-245.

Janet, C.

1895a. Etudes sur les fourmis. 8e note. Sur l’organe de nettoyage tibio-
tarsien de Myrmica rubra L., race levinodis Nyl. Ann. Soc. ent.
Fr. 63 : 691-704.

1895b. Etudes sur les fourmis, les guepes, et les abeilles. 9e note. Sur
Vespa crabro L. — Histoire d’un nid depuis son origine. Mem.
Soc. zool. Fr. 8: 1-140.

1905. Anatomie de la tete du Lasius niger. Ducourtieux et Gout, Limoges.
Le Masne, G.

1953. Observations sur les relations entre le couvain et les adultes chez
les fourmis. Ann. Sci. nat. Zool. ser. 11, 15 : 1-56.

Snodgrass, R. E.

1956. Anatomy of the Honey Bee. Cornell University Press, Ithaca, New
York.

Wallis, D. I.

1961. Food-sharing behaviour of the ants Formica sanguinea and For-
mica fusca. Behaviour 17: 17-47.

Waterhouse, D. F.

1953. The occurrence and significance of the peritrophic membrane,
with special reference to adult Lepidoptera and Diptera. Aus.
J. Zool. 1: 299-318.

Wheeler, W. M.

1923. Social Life among the Insects. Harcourt, Brace and Co., New
York.

Wheeler, W. M. and I. W. Bailey

1920. The feeding habits of pseudomyrmine and other ants. Trans.
Amer. phil. Soc. n.s. 22: 299-318.

Wilson, E. O. and T. Eisner

1957. Quantitative studies of liquid food transmission in ants. Insects
Sociaux 4: 157-166.

AUSTRALIAN CARABID BEETLES XI.
SOME TACHYS 1

By P. J. Darlington, Jr.

Museum of Comparative Zoology, Cambridge, Mass.

The preceding part of this series (Darlington 1962) dealt with the
Australian species of Bembidion. The present part deals with some
smaller Bembidiini of the genus Tacky s. Australian Tachys have been
treated twice by Sloane (1896; 1921), whose papers should be con-
sulted for references and synonymy. I shall now consider only selected
groups of the genus about which I have something new to say.

Australian species of Tachys are much more numerous and less well
known than those of Bembidion. They are phylogenetically diverse as
well as numerous and include several peculiar groups which may be
relict, or specialized. For example Tachys amplipennis Macleay has
the elytron fully striate with grooved striae and seems to lack the
apical striole that characterizes most Tachys. However the apex of
the 7th stria is deeply impressed, with a puncture beside it on the inner
side, and this part of the 7th stria is almost separated from the main
part of the stria in some individuals. This condition may be primitive
and may show how the apical striole originated. However, some other
Tachys have the apical striole attached to the 3rd stria (see Tachys
ectromioides group in the following pages, and also Tachys yarrensis
Blackburn, which will be treated in the next paper of this series) .

Species of Tachys are numerous, diverse, and frequently collected
almost everywhere on the continent of Australia, including Victoria,
but are comparatively few in Tasmania. Sloane (1920, 15 1) records
only four, well known Australian species from the island. In four
months’ collecting on Tasmania I encountered Tachys on only three
occasions: once in stone and gravel banks and bars of the King River
at the crossing of the Queenstown road, where I found T . ((<Bembi-
dion” ) hobarti Blackburn and a related new species; once on the bank
of the Mersey River, where I took one hobarti by washing gravel; and
once beside the Arve River in southern Tasmania, where I found one
specimen of T. australis Schaum under a stone.

The distribution of Tachys in Tasmania and southern Australia

Published with a grant from the Museum of Comparative Zoology at
Harvard College.

Manuscript received by the editor March 19, 1962.

n8

Psyche

[September

is complementary to that of the Trechini. The latter are numerous in
Tasmania and decrease rapidly northward on the mainland, where
Tachys increases in numbers. The two groups tend to be complemen-
tary ecologically too. Most Tachys live beside standing or running
water or in swamps, often in open as well as forested country. Most
Australian (including Tasmanian) Trechini live on the ground in wet
forest but not beside open water. [What I have just said about
complementarity of Tachys and Trechini is true and important, but
an oversimplification. The two groups overlap both geographically
and ecologically in ways that are too complex to describe here.]

Tachys hobarti group

In 1921 (p. 193) Sloane mentioned as unknown to him two sup-
posed Bembidion described by Blackburn: hobarti of Tasmania and
w'attsense of Victoria. I can now say that hobarti is a Tachys ■, not a
Bembidion. The oblique truncation of the outer angle of the anterior
tibia is (as Blackburn said) less marked in hobarti than usual in
Tachys , but absence of a scutellar stria and presence of a (modified)
apical stride mark the species as Tachys. I think that the Victorian
watts ense may be a synonym of hobarti. The former is described as
having 5 and the latter 6 dorsal striae on each elytron, but both condi-
tions occur in my series of hobarti from the King River. Sloane’s leai
may be the same thing, although I do not want to declare the synonymy
without comparing specimens. Sloane’s murrumbidgensis is a related
species. And 3 additional species that seem to be new, one of them
remarkable for reduction of elytral striation, are described below. All
these species, and perhaps additional ones still to be discovered, form
what may be called the hobarti group of Tachys. Although they are
certainly Tachys rather than Bembidion by current classification, the
species of this group are anomalous (primitive ?) in some ways and
should be specially considered by students of bembidiine phylogeny.

Characteristics of the hobarti group are: form subparallel (but ely-
tra considerably wider than prothorax) , moderately convex; upper sur-
face usually punctulate (scarcely so in lutus) . Head large (short but
wide, with neck very wide and not impressed) ; eyes of moderate size
but rather prominent; antennae rather short, with median segments
2 X or less long as wide, and segment 3 usually slightly longer than
2 ; clypeus truncate or broadly emarginate, impressed at middle anter-
iorly in some species; mentum not perforated at base, with a simple
tooth at middle. Prothorax subcordate, more or less lobed across base,

1962]

Darlington — Tachys

119

so that posterior angles are sometimes not quite basal (but I have
measured width of base of prothorax across the prominent sub-basal
angles) ; apex subtruncate; lateral margins narrow, each with usual 2
setae; disc with anterior transverse impression obsolete, middle line
moderately or lightly impressed but extending posteriorly behind
transverse sulcus, and latter variably impressed, sometimes interrupted
at middle. Elytra with rather prominent but more or less rounded
humeri; margins not serrate or faintly so; sutural stria entire and
deeply impressed; 5th stria deeply impressed at base (except in lutus) ;
8th stria absent excepting an apparent remnant deeply impressed near
apex and including 2 strong punctures; apical striole ending anteriorly
in or (usually) reduced to a conspicuous elongate puncture; 3rd
interval or stria 3-punctate. Inner wings fully developed. Abdomen
sparsely, inconspicuously pubescent ; front tibia with outer apical angle
less strongly oblique than usual in Tachys ; posterior tarsi rather short ;
males with 2 segments each front tarsus widely dilated, and males with
1, females 2 setae each side apex last ventral segment.

The known range of the hobarti group is Tasmania and temperate
southeastern Australia north to Rockhampton. All the species occur
in sand or gravel or under stones by rivers or brooks. Some of the
species superficially resemble Perileptus , with which they sometimes
occur.

Key to species of Tachys of hobarti group

1. Dorsal striae (except sutural) obliterated lutus

- Stria 5 and usually some other dorsal striae distinct at least in

part 2

2. Striae 1 and 5 well impressed but 2-4 light, irregular, sometimes

hardly traceable; clypeus not impressed anteriorly; rather shining;
size small (length c. 1.8 mm.) kingi

- Striae 2-4 distinct and individually traceable at least behind

anterior dorsal puncture, although less impressed than 1 and 5 ;
other characters variable, but none of the following species has all
the characters given above for kingi 3

3. Black; more shining, with microsculpture less distinct; clypeus not

distinctly impressed; length 2.3-2. 7 mm. (may include wattsense
Blackburn and leai Sloane) hobarti

■ Reddish; less shining, with microsculpture (microreticulation and
punctulation) more distinct; clypeus impressed at middle anteriorly

4-

120

Psyche

[September

4. Length c. 1.75-2.4 mm. m urru m h id gens is

- Length c. 2. 5-2. 8 mm. fitzroyi

Tachys lutus n. sp.

Figure 1

With characters of hobarti group as given above, but dorsal striae
(except sutural) obliterated ; form (Fig. 1) rather slender but convex ;
dark reddish piceous, appendages reddish testaceous; shining, virtually
without dorsal microsculpture or punctulation. Head .89 width pro-

thorax; eyes moderate in size and prominence; antennae with median
segments (not counting pubescence) hardly 2X long as wide; clypeus
subtruncate (slightly emarginate), not impressed anteriorly; frontal
sulci irregularly subparallel, extending onto clypeus; front slightly
convex, impunctate except faintly punctulate at 54 X in good light;
mentum with an entire tooth. Prothorax subcordate; width/length
1.2 1 ; base/apex c. 1.04; base/head .85; sides arcuate anteriorly,

1962]

Darlington — Tachys

121

oblique and strongly converging posteriorly, strongly but briefly
sinuate before posterior angles; latter forming small, sub-basal, rec-
tangular projections; apex truncate; base subtruncate; disc with
anterior transverse impression and middle line almost obsolete but
latter coarse basally; sulcus coarsely foveate. Elytra more than
wider than prothorax (E/P 1.57), probably widest about middle
(slightly spread), not margined basally (margin ending a little inside
humeri) ; margins not visibly serrate or setulose (at 54X ) ; sutural
stria entire, punctate anteriorly, groove-like posteriorly ; other dorsal
striae obliterated, including 5th> which is at most faintly indicated
toward base ; apical stride reduced to a coarse slightly elongate punc-
ture on declivity, nearer margin than suture; 3 dorsal punctures on
each elytron before anterior *4, near middle, and behind Y\.. Length
c. 2.5 ; width c. .95 mm.

Holotype (M. C. Z. Type No. 30327) $ from Termed (north of
Bateman’s Bay and east of the upper Clyde River), southern New
South Wales, October 1957, taken by myself. It was actually found
northwest of Termeil, well up in the hills, where I followed a system
of wood roads nearly to (south of) a conspicuous summit called the
Pigeon House. Here, in a small valley of eucalyptus woods, was a
small brook, nearly dry but with water still in the pools, which were
margined by sand bars. The single specimen of the present species was
taken by washing sand at the water’s edge.

The obliteration of the elytral striae (except the sutural) makes
this a distinct and easily recognizable species.

Tachys kingi n. sp.

With characters of hobarti group as described above. Small, moder-
ately elongate, moderately convex; dark rufous, head slightly darker,
appendages not much paler; shining, reticulate microsculpture faint
above but head and pronotum distinctly, sparsely punctulate. Head
.86 & .87 width prothorax (in <$ $ measured) ; antennae with middle
segments not quite 2X long as wide; clypeus truncate, not impressed
anteriorly (checked in all specimens) ; frontal sulci irregularly sub-
parallel, reaching and diverging on clypeus. Prothorax subcordate,
width/length 1.30 & 1.28; base/apex .89 & .93; base/head .81 & .79;
sides rounded anteriorly, oblique and strongly converging posteriorly,
rather strongly sinuate before angles ; latter sub-basal, forming nearly
rectangular projections; disc with anterior impression obsolete, middle
line lightly impressed, basal sulcus moderate, faintly punctulate, not

122

Psyche

[September

foveate unless at intersection of middle line. Elytra 1/3 or more wider
than prothorax (E/P 1.34 & 1.44) ; margins behind humeri faintly
subserrate and setulose ; sutural striae entire, stria 5 well impressed in
about basal half and reaching humerus, stria 2 less impressed, and
striae 3 and 4 irregularly and usually rather faintly indicated; apical
striole reduced to a conspicuous elongate puncture. Length c. 1.8;
width c. 0.7 mm.

Holotype cf (M. C. Z. Type No. 30328) and 10 paratypes all
from the King River just below the crossing of the road from Lake
St. Clair to Queenstown. The specimens were taken with hobarti,
under stones, on stone-and-gravel bars.

This species is about the size of typical T. murrumbidgensis. I do
not have specimens of the latter, but kingi evidently differs in having
elytral striae 2-4 less distinct, clypeus not impressed, and upper surface
more shining.

Tachys hobarti (Blackburn)

Bembidium hobarti Blackburn 1901, 123.
fBembidium cwattsense Blackburn 1901, 123.
fTachys leai Sloane 1896, 358, 370.

A rather elongate, black or blackish species, with characters of
hobarti group. Head .87 & .88 width prothorax; clypeus not or not
distinctly impressed anteriority (checked in all specimens). Prothorax
subcordate; width/length 1.3 1 & 1.34, base/apex .93 & .91 ; base/head
.82 & .81; posterior angles right-acute, nearer base than in preceding
species but separated from basal lobe by brief, strong sinuations.
Elytra much wider than prothorax (E/P 1.47 & 1.46) ; each with 5
or 6 discal striae (stria 6 variable). Length 2. 3-2. 7; width 0.8-1.0
mm.

This species was described from near Hobart, Tasmania. I redis-
covered it at the King River and later found a specimen by the Mersey
River, Tasmania, as noted in the preceding general discussion of
Tachys. If the synonymy suggested above is correct, this species has
been found on the mainland of Australia on the bank of the Watts
River, a tributary of the Yarra, east of Melbourne, Victoria (watt-
sense) , and at Tamworth, New South Wales (leai) .

Tachys murrumbidgensis Sloane

Sloane 1895, 407.

A small, piceous species, with clypeus impressed anteriorly (noted by

1962] Darlington — Tachys 123

Sloane). Measurements (of types, t. Sloane) : length 1.75; width
O.75 mm.

Described from 2 specimens from Narrandera, New South Wales,
and later (1921, 203) recorded by Sloane from “sand banks and
pebble beds” by the margins of the following rivers, all in New South
Wales: Murray (at Mulwala), Murrumbidgee (at Narrandera),
Cudgegong (at Mudgee), and Macquarie (at Narromine).

Tachys fitzroyi n. sp.

With characters of hobarti group as described above. Larger and
slightly broader than most species of group; dark rufous, appendages
not much paler; not very shining, reticulate microsculpture distinct
above but not deeply impressed, and whole upper surface sparsely
punctulate. Head .91 & .90 width prothorax; clypeus subtruncate or
slightly emarginate, impressed at middle anteriorly so that it is sub-
tuberculate each side at apex (in all specimens) ; frontal sulci sub-
parallel, diverging posteriorly, extending across clypeus as sharply
defined parallel grooves. Prothorax broadly subcordate; width/length
1.33 & i.37, base/apex .85 & .85, base/head .75 & .78; sides rather
broadly rounded for much of length, strongly converging posteriorly,
abruptly sinuate before posterior angles; latter sub-basal, forming
rather small c. rectangular prominences ; disc with middle line distinct,
basal sulcus moderate, not foveate but vaguely punctulate. Elytra
about 3/10 wider than prothorax (E/P 1.32 & 1.29) ; margin behind
humeri finely scalloped and setulose; sutural striae entire, stria 2
nearly entire but less impressed, stria 5 strongly impressed basally to
humerus, striae 3-4 light and slightly irregular but plainly traceable
except less distinct at extreme base, striae 6-7 at most faintly indicated ;
apparent apical striole present but irregularly impressed, ending anteri-
orly in a coarse impression; 3rd stria 3-punctate, anterior puncture
before of elytral length and almost joining 3rd to 4th stria, other
punctures near middle and behind % of elytral length. Length 2.5-
2.8; width c. 0.9- 1. 1 mm.

Holotype S (M. C. Z. Type No. 30329) and 22 paratypes all
from the Fitzroy River a few miles north of Rockhampton, Queens-
land, November 1957, taken by myself. They were in gravel and
cobble stone river bars and occurred with Perileptus, which they
resembled superficially.

This is probably the supposed large form of Tachys murrumbid-
gensis referred to by Sloane (1921, 203) as occurring with the smaller

24

Psyche

[September

typical form at Narromine. I have two reasons for considering it a
distinct species. First, the size range given by Sloane (loc. cit.) for
murrumbidgensis (1.75-2.75 mm.) is greater than expected in a popu-
lation of one species. And second, my series from the Fitzroy River
includes only the large form, which apparently therefore exists separ-
ately.

Tachys ectromioides group

Tachys ectromioides Sloane is a very distinct, large, rare species.
The reason for its rarity is probably its habitat. It, like the related
species described below, probably lives in debris, loose soil, or rotten
wood on the ground in heavy, damp woods, not associated with surface
water. This is a habitat where (in southern Australia) one expects
to find “Trechus” rather than Tachys , and in fact I mistook Tachys
bolus for a trechine when I collected it. Small Carabidae in this habi-
tat are rarely found by ordinary collecting methods. They can be taken
by sifting, but this is laborious and must be done persistently in just the
right place in order to get specimens. They can be taken more easily
and in greater numbers by shoveling debris and loose soil into quiet
water and catching the insects as they come to the surface. I have
not found T. ectromioides itself, but I have collected series of 2 new
related species by this method. The 3 species concerned may be con-
sidered to form the Tachys ectromioides group.

Important characters of the Tachys ectromioides group are: form
broad with base of prothorax broad; color variable; upper surface not
punctulate (but with reticulate microsculpture). Head rather long;
frontal sulci not extending onto clypeus but produced posteriorly and
vaguely joining depressed areas behind eyes, so latter on poorly defined
ocular hemispheres; antennae varying in length, segment 3 not or
slightly longer than 2; mentum with 2 deep impressions at base (but
not perforated) and with entire median tooth. Prothorax with pos-
terior angles costate. Elytra with humeri broadly rounded ; margins
ending inwardly about opposite ends 6th striae, not serrate or setulose ;
striation entire or nearly so but lightly impressed laterally and apically ;
8th stria parallel to margin, deep posteriorly, lighter or irregular
anteriorly; apical striole well impressed, long, approaching or joining
end of 3rd stria anteriorly, with a fixed puncture on inner side well
back; dorsal punctures present or absent, if present, 2 on each 3rd
interval. Inner wings long and folded, probably fit for flight in bolus,
perhaps not in bolellus. Lower surface almost impunctate, not (or at

1962]

Darlington — Tachys

125

most very inconspicuously) pubescent. Male with 2 segments each
front tarsus slightly dilated, inconspicuously squamulose; cf with i,
9 2 setae each side last ventral segment.

Key to species of Tachys ectromioides group

1. Elytra fasciate, brown on testaceous; dorsal elytral punctures

present, though small ; length c. 3 mm. ectromioides

- Elytra unicolorous ; dorsal elytral punctures absent 2

2. Bicolored, head and prothorax rufous, elytra castaneous; length

3.2-3. 6 mm bolus

- Color wholly castaneous; length 2. 6-2. 8 mm. bolellus

Tachys ectromioides Sloane

Sloane 1896, 356, 359; 1898, 477; 1921, 195 (prothorax), 198, 204.

Sloane described this species as with “. . . elytra testaceous, a very
wide dark piceous fascia across disc considerably behind base, apex
widely piceous . . .” and “. . . third elytral interval with two small
setigerous punctures — the anterior just before, the posterior just
behind discoidal piceous fascia . . .” The type of the species (now in
the Macleay Museum at Sydney) was said to be from Donnybrook,
Western Australia, but Sloane later (1898) suggested that this was
probably an error. The species has been found at the Richmond River,
northern New South Wales (Sloane 1898) ; on the Blue Mts., New
South Wales; and near Melbourne, Victoria (Sloane 1921, 204). I
have not collected it but have briefly examined a specimen at the
British Museum, unfortunately without looking for the dorsal elytral
punctures. The possible habitat of the species is suggested in discussion
of the group.

Tachys bolus n. sp.

Figure 2

With characters of Tachys ectromioides group as here defined; form
as figured (Fig. 2). Head and prothorax rufous, former darker
posteriorly and at sides, elytra dark reddish castaneous, appendages
rufous ; moderately shining but with distinct microsculpture isodiame-
tric on front, isodiametric or slightly transverse on pronotum, present
as very fine transverse lines on elytra, which are vaguely iridescent.
Head .66 & .65 width prothorax; antennae rather long, middle seg-

126

Psyche

[September

ments c. 3 X long as wide ; palpi slender, last segments rather long ( in
genus) but slender, subulate. Pro thorax strongly narrowed anteriorly,
slightly so posteriorly; width/length 1.32 & 1.33; base/apex c. 1.58 &
1.47; base/head 1.38 & 1.37; apex subtruncate, slightly lobed at mid-
dle; base subtruncate, broadly and slightly lobed at middle; sides
broadly arcuate through much of length, broadly but rather slightly
sinuate posteriorly; posterior angles c. right, well defined, costate;
lateral margins moderate anteriorly, slightly broader posteriorly, each
with usual 2 setae about 2/5 from apex and near basal angle; anterior
transverse impression of disc broad, not sharply defined ; middle line
rather coarse and well impressed, coarser basally and reaching base;
basal sulcus distinct but not foveate, interrupted at middle by longi-
tudinal impression; baso/lateral areas broadly depressed. Elytra broad
(E/P 1.59 & 1 .6 1 ) , somewhat rounded at sides, widest about middle;
striae moderately impressed, slightly, irregularly punctulate; dorsal
punctures lacking. Length 3. 2-3.6; width 1.3- 1.6 mm.

Holotype cf (M. C. Z. Type No. 30332) and 19 paratypes all
from Bellangry Forest, northwest of Wauchope, New South Wales,
about 3000 ft. altitude, April 1958, taken by myself. Also one speci-
men, not a type, from Barrington Tops, Mount Royal Range, about
4000 ft., October 1957, also taken by myself. All specimens were
taken among lumps of dirt and in loose soil in damp situations. The
first one at Bellangry Forest was washed from dirt and leaf litter
from the ground in wet forest. After finding it, I searched for addi-
tional specimens without success until I started kicking the actual
broken earth bank of a small brook into the water. Then the series
here recorded was collected without much trouble. The Barrington
Tops individual was taken among lumps of earth beside a seepage near
the top of the road that led up from the Allyn River toward (but at
that time not quite to) the plateau.

This species was very Trechus-MYe in life. It is somewhat Trechus-
like even under the microscope, although its technical characters
(subulate palpi, elytral striation, etc.) leave no doubt that it is a
Tacky s. It is evidently related to ectromioides , but differs as indicated
in the key. The absence of dorsal elytral punctures in this and the
following species is unexpected, but I have examined all specimens of
both the present and following species at about 100X under fluores-
cent illumination, which reduces surface reflection and makes minute
details clear, and have not found any trace of dorsal punctures in any
specimen.

1962]

127

Darlington — Tachys

Tachys bolellus n. sp.

With characters of ectromioides group as here defined. Color dark
reddish castaneous, appendages rufous; rather shining, reticulate
microsculpture of upper surface isodiametric on front, finer and strong-
ly transverse on disc of pronotum but isodiametric and in part actually
longitudinal in anterior-median area of pronotum, scarcely resolved
on elytra but probably present as very fine transverse lines, for elytra
slightly iridescent. Head .68 & .64 width prothorax; antennae rela-
tively short, middle segments c. 1 X or slightly more long as wide.
Prothorax strongly narrowed anteriorly, much less so posteriorly;
width/length 1.29 & 1.39; base/apex 1.46 & 1.43; base/head 1.27 &
1.34; apex subtruncate or very broadly emarginate; base subtruncate,
very slightly lobed at middle; sides rather broadly arcuate anteriorly,
nearly straight and moderately converging posteriorly, slightly or
scarcely sinuate before base; lateral margins moderate anteriorly,
slightly broader posteriorly, each with usual 2 setae at apical 2/5 and
basal angle; basal angles slightly obtuse (nearly right) , sharply defined,
costate; disc with rather vague anterior transverse impression, strongly
impressed middle line, broader basally, and reaching base ; basal sulcus
well impressed, interrupted at middle; baso-lateral impressions rather
large, deep, margined posteriorly, and margined exteriorly by strong
costae. Elytra broad (E/P 1.49 & 1.48), oval, widest near or slightly
behind middle; all striae indicated, but outer ones faint or almost
obsolete; dorsal punctures lacking. Length 2.6-2. 8; width 1.1-1.2 mm.

Holotype cf (M. C. Z. Type No. 30333) and 8 paratypes all
from the Williams River Valley a little above Barrington House, at
the foot of the Mount Royal Range, New South Wales, October 1957,
taken by myself. All the specimens were taken by washing wood-debris
from a rotten log lying on the ground in heavy gallery forest near
the river.

The present new species is sufficiently distinguished from bolus and
ectromioides in the preceding key.

(Some additional Tachys will be treated in the next number of
this series.)

References

Blackburn, T.

1901. [Australian Bembidiini.] Trans. R. Soc. South Australia 25:
120-124.

Darlington, P. J., Jr.

1962. Australian carabid beetles X. Bembidion. Breviora (in press).

128

Psyche

[September

Sloane, T. G.

1895. Tachys murrumbidgensis. Proc. Linn. Soc. New South Wales (ser.
2), 9: 407-408.

1896. [Australian Tachys.] Proc. Linn. Soc. New South Wales, 21 : 355-
377, 407-409.

1898. [Carabidae from Western Australia.] Proc. Linn. Soc. New South
Wales, 23: 444-520.

1920. Carabidae of Tasmania. Proc. Linn. Soc. New South Wales, 45:
113-178.

1921. [Australian Bembidiini.] Proc. Linn. Soc. New South Wales, 46:
192-208.

THE SPIDER GENUS SOSIPPUS IN NORTH AMERICA,
MEXICO, AND CENTRAL AMERICA
(ARANEAE, LYCOSIDAE)1

By A. R. Brady
Harvard University

Introduction. The genus Sosippus contains the only spiders in the
Nearctic Region of the subfamily Hippasinae, members of which are
unique among the Lycosidae in producing a large funnel-web resem-
bling that of the Agelenidae. The posterior spinnerets are more elong-
ate than in other Lycosidae, concomitant with their web building
habits. Although similar to the Agelenidae in these respects, they
represent typical Lycosidae in other characters. In Sosippus the eyes
are arranged in three rows: four small eyes on a vertical front form
the anterior row; two large posterior median eyes form the second
row, and two somewhat smaller posterior lateral eyes form a third
row. The trochanters are notched and the egg case is carried attached
to the spinnerets. These features are characteristic of all Lycosidae,
but are not found in the Agelenidae. The tarsi and metatarsi of leg I
and leg II are more densely scopulate in Sosippus than in most other
lycosids. Sosippus is found in tropical and subtropical America from
Costa Rica to the southern United States.

Porrima , found in South America, appears to be the closest relative
of Sosippus. Females of P. diversa (O. P. -Cambridge) and the male
holotype of P. harknessi Chamberlin resemble Sosippus in coloration
and especially in external genitalia (Figs. 12, 33), but are readily
separated by differences in the eye arrangement (Fig. 11). The Hip-
pasinae of the Neotropical Region, in addition to eight described species
of Porrima , are represented by two species of Birabenia and the mono-
typic genus Hippasella. C. F. Roewer (1959) splits Porrima into
three genera on the basis of the number of posterior cheliceral teeth and
slight differences in the eye arrangement. On the basis of great varia-
tion of these characters in Sosippus, it seems best to maintain the eight
species in question in the single genus Porrima until further study.

In the Ethiopian, Oriental, and Australian Regions the Hippasinae
are represented by 12 genera containing numerous species according to
C. F. Roewer (1959)- These Lycosidae have in common one feature

’Published with the aid of a National Science Foundation Grant of the
Department of Biology, Harvard University.

Manuscript received by the editor March 19, 1962.

129

130

Psyche

[September

that separates them from other lycosids, namely, the greater length of
the posterior pair of spinnerets. If the greater length of the spinnerets
is concomitant with web-spinning habits, then it would seem that this
assemblage might constitute a natural group deserving subfamilial
rank. If, however, the length of the spinnerets does not indicate a
web-spinning function, but is simply a structural convergence found in
otherwise diverse groups, it should not be used as a criterion to estab-
lish a subfamily. It may be that some species of Enprosthenops placed
in the Pisauridae also belong to this group since the genitalia are simi-
lar to those of Lycosidae and the eye arrangement resembles that of
Porrima.

Acknowledgements. This investigation was carried out as a gradu-
ate research program under the direction of Dr. H. W. Levi of the
Museum of Comparative Zoology to whom I am especially indebted
for encouragement, helpful advice, and constructive criticism. I thank
sincerely Dr. W. J. Gertsch of the American Museum of Natural
History, who placed the collections of his institution at my disposal.
I thank also Dr. H. K. Wallace of the University of Florida for
making large collections from Florida available for study. Measure-
ment of Wallace’s specimens was not undertaken because the paper
was near completion at the time of their arrival. Study of these speci-
mens elucidates certain facts that I will stress, and supports the con-
clusions already reached before their arrival. Locality data was uti-
lized and certain structural features were checked. Mr. J. A. Beatty
provided a number of specimens from Arizona and Sonora and sup-
plied ecological data for S. calif ornicus. I am grateful to Dr. G. Owen
Evans and to Mr. D. Clark of the British Museum, Natural Flistory,
for loan of the male of S. mexicanus. Collections in the field during
the summer of 1961 were made possible by a grant from the Sigma
Xi-RESA Research Fund. A National Institutes of Health Grant
(E-1944) helped defray some of the expenses.

Sosippus Simon

Sosippus Simon, 1888, Ann. Soc. Ent. France, 8 (6) : 206.

Types species: The problem of the type species has been discussed
by Bonnet (1958). I follow him for the sake of nomenclatural stabil-
ity. Simon (1888) established the genus Sosippus and designated
Doloinedes oblongus C. L. Koch as the type. At the same time he
described Sosippus mexicanus as a new species. In 1898 Simon trans-
ferred D. oblongus to the genus Lycosa (Diapontia) and established

1962]

Brady — Sosippus

13

S. mexicanus as the type species. Sosippus mexicanus has been assumed
to be the type for the last 64 years.

Characteristics. Anterior eye row, as seen from in front, procurved.
Lateral eyes subequal to the median eyes and mounted on distinct
tubercles. Anterior eye row wider than the middle row; the posterior
row wider than the anterior row (Fig. 10). Chelicerae robust, with
prominent bosses. Anterior cheliceral margin with three teeth on each
side. Posterior cheliceral margin with three or four cheliceral teeth
on each side, rarely five. Usually constant within a species, but some-
times variable, e. g. S. mimus. Labium longer than wide, as long as
wide, or slightly wider than long. Endites, heavily scopulate, slightly
converging in front of labium, less heavily scopulate. Carapace with
conspicuous longitudinal thoracic groove. Carapace of females highest
in the cephalic region, of males usually highest in the thoracic region.
Sternum always longer than wide. Fourth leg longest. Patella-tibia
IV longer than metatarsus (except in males of S. calif ornicus, which
have the metatarsus longer). Order of length of patellae and tibiae:
IV, I, II, III. Tarsi and metatarsi of legs I and II heavily scopulate.
Tibia I and II usually scopulate at distal ends. Males with legs longer
than those of females and more heavily scopulate.

Female epigynum characterized by a relatively narrow anterior
median septum connected to an expansive posterior blade (Fig. 19).
Male palpi with numerous sclerites (Fig. 36).

The variable nature of certain morphological characters is empha-
sized because some earlier authors considered such characters to be
diagnostic for the genus. Some are diagnostic at the species level.

Discussion. Spiders of the genus Sosippus represent a closely related
group of species as evidenced by their structural similarity and web-
spinning habits. It is probable that the group has diverged relatively
recently in geologic time. Two species groups might be established on
the basis of structural, similarities and distribution. One group con-
tains S. jloridanus, S. mimus , and S. texanus. The other group includes
S. californicus, S. mexicanus , S. agalenoides, S. michoacanus and S.
plutonus. The illustrations of the color patterns and the drawings of
the genitalia indicate the affinities within these two species groups.

C. F. Roewer, first in the Katalog der Araneae (1954) without
giving reasons, and then in 1959 attempted to separate Sosippus into
two groups, giving each generic rank. This division is based entirely
upon the number of teeth on the posterior cheliceral margin. Species
with four cheliceral teeth on each side were left in Sosippus and those
with three on each side were placed in the newly erected genus Sosip-

Psyche

[September

132

pinus. The division of the genus on this basis alone becomes untenable
since the number of teeth on the posterior cheliceral margin is ex-
tremely variable within certain species (S. mimus) . Similarities in
color pattern, eye arrangement, spination, relative length of leg seg-
ments, and especially the genitalic characteristics indicate that the eight
species considered in this paper should be maintained in a single genus.

Simon described S. mexicanus (type of the genus) as having four
posterior cheliceral teeth on each side. F. Pickard-Cambridge (1902)
reported that the most abundant species of Sosippus in Mexico, which
he felt surely was the one described by Simon, had only three cheliceral
teeth on each side. Of the two females of S. mexicanus examined, one
has 4-3 posterior cheliceral teeth. It is very possible that the specimens
of S. mexicanus that Simon had before him actually had four posterior
cheliceral teeth on each side, which, in the case of mexicanus , turns
out to be the exception rather than the rule. It is also very probable
that F. Pickard-Cambridge was describing the same species. Roewer’s
criterion of the number of posterior teeth of the chelicerae for defining
genera is completely artificial in the case of Sosippus and probably
other lycosid genera as well. J. Buchar (1959) has recently found
that the lycosid genus Trochosa in Central Europe shows considerable
variation within the same species in the number of posterior cheliceral
teeth. The similarities among the eight species of Sosippus far out-
weigh any differences that might be used to separate them into two or
more genera.

Incorrect Placement. Sosippus insulanus Bryant (1923), described
from Barbados, is an immature lycosid, evidently at the penultimate
stage of development. The coloration, scopulae of the tarsi and meta-
tarsi, and spinnerets are not like those found in Sosippus. Although
the true identity of this specimen can be ascertained only after associa-
tion with adult individuals from the same locality, it is best referred
to the genus Lycosa at the present time.

Measurements. Two sets of oculars with accompanying grids were
used in combination with low and high power objectives for making
measurements. From measuring a selected set of specimens several
times, it was determined that the higher power combination was accur-
ate to 0.02 mm and the lower power combination was accurate to 0.1
mm. In all cases the greatest dimension of the structure was measured,
e.g. patella-tibia length was measured as the greatest distance between
a line tangent to the most proximal part of the patella to a line tangent
to the most distal part of the tibia. Measurements were made under
conditions as uniform as possible. Conditions for the most important

1962]

Brady — So sip pus

133

measurements are specified below. A series of 20 measurements involv-
ing various components of the spider were made for each specimen.
The most diagnostic of these measurements are recorded for compari-
son in Table i.

The posterior median eyes (PME) and the posterior lateral eyes
(PLE), which form two rows in the Lycosidae, are referred to in this

Text Fig. 1. Measurement of the Posterior Ocular Quadrangle.

paper as the posterior ocular quadrangle (POQ). The measurement
of the POQ is illustrated in Text Figure i.

The distance A is the width of the anterior row of the POQ, the
distance B is the length of the POQ, and the distance C is the width
of the posterior row of the POQ. The length of the carapace was
measured as the distance from the line tangent to the posterior-most
part of the carapace to the line tangent to the anterior-most part of the
AME. Total length was measured from the most anterior part of
the AME to the tip of the anal tubercle, when this structure was
visible, or to the posterior tip of the abdomen. When the specimen
was stretched so that the lorum of the pedicle was visible (an abnormal
attitude in the living spider), the abdomen was measured and the
length of the carapace was added as the distance from the anterior

134

Psyche

[September

TABLE 1

Males:

POSTERIOR OCULAR QUADRANGLE

Species

N

Anterior
Eye Row

Anterior

Row

Posterior

Row

Length

S. calif ornicus

22

1.397 ± 0.075

1.190 ± 0.05 1

1.717 ±0.069

1.031 ±0.049

S. mexicanus

1

1.44

1.17

1.80

1.10

S. floridanus

3

1.29

1.12

1.62

1.04

1.17

1.05

1.52

0.99

1.24

1.10

1.57

1.04

S. mimus

4

1.47

1.22

1.79

1.00

1.50

1.25

1.84

1.17

1.49

1.22

1.84

1.05

1.50

1.27

1.89

1.14

S. t ex anus

2

1.7 5

1.45

2.17

1.25

1.79

1.42

2.17

1.25

Females:

S. calif ornicus

38

1.6 15 ± 0.563

1.311 ± 0.129

1.949 ±0.199

1.171 ±0.121

S. mexicanus

2

1.35

1.15

1.74

1.05

1.15

0.99

1.34

0.92

S. agalenoides

4

2.10

1.67

2.59

1.45

2.0 0

1.57

2.40

1.42

2.12

1.65

2.40

1.50

2.10

1.62

2.45

1.39

S. michoacanus

5

1.64

1.45

2.12

1.25

1.59

1.39

2.02

1.19

1.57

1.34

2.00

1.17

1.54

1.34

1.95

1.19

1.39

1.24

1.79

1.09

S. plutonus

1

1.34

1.19

1.70

1.07

S. floridanus

16

1.4 34 ± 0.1 1 3

1.238 ± 0.090

1.803 ±0.146

1.144 ± 0.080

S. mimus

4

1.59

1.30

2.12

1.20

1.54

1.32

2.00

1.22

1.37

1.17

1.72

1.04

1.97

1.49

2.29

1.39

S. texanus

10

1.794±0.313

1.459 ±0.240

2.218 ± 0.381

1.308 ± 0.185

All measurements are in mm with the mean and standard deviation calculated
where 10 or more specimens were available.

135

1962] Brady — So sip pus

TABLE 1

Males:

CARAPACE

Species

Length

Width

calif or nicus

6.90±0.50

5.01 ±0.14

mexicanus

6.4

4.6

floridanus

6.3

4.5

6.0

4.3

5.8

4.5

mimus

7.2

5.4

7.3

4.7

7.2

5.4

7.6

5.4

texanus

9.4

6.9

9.5

7.1

Females:

calif ornicus

7.34± 1.01

5.32±0.74

mexicanus

5.7

4.4

4.7

3.7

agalenoides

9.8

7.4

9.7

7.4

10.6

7.9

10.0

7.2

michoacanus

7.5

5.4

6.7

5.0

6.7

4.8

6.5

4.7

5.7

4.1

plutonus

6.1

4.3

floridanus

6.44± 0.65

4.59±0.51

mimus

7.5

5.5

7.3

5.2

5.7

4.3

8.8

6.2

texanus

8.61 ± 1.84

6.25 ± 1.33

(Continued)

LABIUM

Length

Width

Total Body
Length

1.037±0.065

1.021 ±0.067

1 3.92 ± 1.09

0.99

0.94

12.0

0.89

0.84

11.9

0.84

0.75

11.2

0.84

0.78

—

1.07

0.97

13.3

1.12

1.04

13.1

1.15

1.04

14.2

1.14

1.00

14.2

1.40

1.25

20.1

1.40

1.29

18.0

1.165±0.154

1.20± 0.157

16.16±2.46

0.99

0.94

13.4

0.78

0.78

12.6

1.62

1.39

20.1

1.57

1.50

22.9

1.77

1.62

25.0

1.65

1.50

22.2

1.29

1.17

15.1

1.14

1.07

13.5

1.15

1.05

13.2

1.10

1.04

14.3

0.99

0.95

11.4

0.94

0.95

11.8

1.029±0.124

0.988 ±0.099

13,25 ± 1.68

1.20

1.12

14.6

1.22

1.14

16.1

0.95

0.97

12.9

1.45

1.35

18.2

1.434± 0.305

1.327 ±0.232

17.88 ±3.25

136

Psyche

[September

TABLE 1 ( Continued )

Males:

SEGMENTS OF LEG IV
Patella-

Species

Femur

Tibia

Metatarsus

Tarsus

Total

S. calif ornicus

8.02± 1.96

9.43 ±2.31

9.95 ±2.36

3.76±0.79

31.16±2.2S

S. mexicanus

7.6

8.8

8.6

3.7

28.7

S . floridanus

6.4

7.4

6.7

3.3

23.8

6.1

7.1

6.7

3.0

22.9

6.2

7.5

7.0

3.2

23.9

S, mi mus

7.3

8.4

8.0

3.7

27.4

7.5

9.0

8.5

4.1

29.1

7.9

9.4

9.2

4.2

30.7

7.5

9.2

8.7

4.2

28.6

S. texanus

9.5

11.6

11.0

5.7

37.8

9.6

11.8

11.0

5.5

37.9

Females:

* S. calif ornicus

7.65 ±1.06

8.8 6 ± 1.16

8.26± 1.06

3.31 ± 0.37

28.11 ± 3.55

S. mexicanus

6.0

7.0

6.5

2.6

22.1

5.5

6.3

5.9

2.3

20.0

S. agalenoides

9.8

11.2

9.6

2.7

33.3

9.7

10.9

9.4

—

—

10.1

11.5

9.7

—

—

9.6

10.8

9.4

—

—

S. michoacanus 7.0

8.1

7.5

2.9

25.5

6.5

7.6

6.9

2.8

23.8

6.5

7.7

7.0

2.9

24.1

6.3

7.5

6.9

2.9

23.6

— .

—

—

—

—

S. plutonus

5.7

6.7

5.8

2.7

20.9

S. floridanus

5.72 ± 0.63

6.63 ±0.69

5.83 ± 0.59

2.84± 0.96

21.08 ±2.12

S. mimus

7.0

7.6

6.9

2.9

25.4

7.0

7.6

6.9

2.9

24.4

6.0

7.3

6.2

2.8

22.3

7.4

8.3

7.0

3.6

26.3

S. texanus

7.43 ±1.62

8.86 ± 1.79

7.30 ± 1.24

3.74±0.75

27.33 ± 5.38

All measurements are in mm with the mean and standard deviation calculated
where 10 or more specimens were available.

1962]

Brady - — Sosippus

137

part of the indention in the posterior edge to the tangent of the AME,
thus allowing for the abdomen over-hanging the carapace. For meas-
urements of the POQ the specimen was placed in a horizontal attitude
such that a definite space was visible between the PME and the AME
when viewed from above (as in Fig. 3 and not as in Fig. 4). This
gives the greatest length to the POQ. The anterior eye row was
measured by placing the specimen vertically in such a position that a
face view was obtained. The measurement of the AME was again
checked in this position. The measurement of leg segments was taken
from the prolateral aspect of the anterior pairs of legs and the retro-
lateral aspect of the posterior pairs of legs for all segments except the
femora. The femora of the anterior pairs of legs were measured from
the retrolateral aspect and those of the posterior pairs of legs from the
prolateral aspect to avoid breaking legs from specimens.

EXPLANATION OF FIGURES

The color descriptions and illustrations of S. texanus , S. jloridanus,
and S. calif ornicus were based on fresh specimens and represent these
species much as they appear in life. The color description and illustra-
tion of S. mimus (Fig. 3) is based on the holotype, which is in very
good condition. Sosippus michoacanus , S. agalenoides , S. mexicanus ,
S. mimus (Fig. 2), and S. plutonus were drawn from specimens that
have been in alcohol for some time, but have remained in a good state
of preservation. The relative condition of these specimens is indicated
by the order in which they are listed above. The description and
illustration of the type of S. plutonus probably deviates more from that
of the living spider than any of the rest since hair appears to have been
rubbed from the carapace and the abdomen is shrivelled.

Two drawings of the female genitalia were made for each species:
a ventral external view of the epigynum after all the hair had been
removed (thus revealing some internal structure through the integu-
ment), and a dorsal internal view with the genitalia removed and sub-
merged in clove oil for clearing.

Two views of the male palpi were drawn for each species: a ventral
view and a retrolateral view. The left palpi of the males were used
after gently scraping them free of hair (quite abundant in the living
spider) and spines (one or more at the ventral apex and several along
the retrolateral edge of the cymbium). These hairs and spines obstruct
the palpal sclerites and since the sclerites of the palpi are of much
greater diagnostic value, no attempt was made to indicate hirsuteness
or spination in the drawings of the male palpi.

38

Psyche

[September

SPECIES DESCRIPTIONS

Before analyzing the individual species a few comments should be
made concerning the treatment of certain sections.

Structure. Under this heading follows a description of structural
features not covered in the table of measurements and not defined
specifically under generic characters. Tibial spination is relatively
constant for each sex within a given species and there is a basic pattern
throughout the genus. Therefore, two tables are constructed showing
the typical patterns of tibial spination in S. calif ornicus and subsequent
species are compared to these.

When the difference between two dimensions is less than 0.05 mm
these dimensions are considered subequal.

Color. Color descriptions are based on specimens submerged in
alcohol and viewed at low power ( 10 X ) under a dissecting scope with
illumination from a microscope lamp. In fresh alcoholic specimens the
color is much the same as in the living spiders. The contrasting light
and dark markings, particularly on the carapace and dorsum of the
abdomen, are created by light and dark hairs. When these hairs
become rubbed off, the underlying color of the integument no longer
provides such marked contrast. All of the figures and descriptions are
based on specimens with the hairs present. In some cases, of course,
this is very difficult because of damage to the specimen.

Records. Locality records for states are listed geographically, coun-
ties alphabetically under states, cities and towns alphabetically under
counties. The records are for specimens examined except where noted.
The lower case “o” is used to indicate a juvenile specimen in the same
manner that the normal cf and ? signs represent a mature specimen.
The 00, cf cf and signs indicate more than one specimen collected
at a single locality.

KEY TO SPECIES

I a. Four posterior cheliceral teeth 2

ib. Three posterior cheliceral teeth 3

2a. Carapace with a distinct white median stripe beginning at second
eye row and continuing to posterior edge; and with two broad
white submarginal stripes as in Figure 1. Epigynum as in Figures

18-20. Palp as in Figures 40-43. Found in Florida only

S. florid an us

2b. Carapace without a distinct median white stripe running length
of carapace, and without distinct white submarginal stripes. Three
yellowish white stripes converging behind second eye row and
spreading posteriorly, but continuing for only a short distance, as

1962]

Brady — So sip pus

139

in Figures 2, 3. Epigynum as in Figures 13-17. Palp as in

Figures 34, 35 S. mini us

3a. Without a conspicuous white median stripe running length of
carapace and without distinct submarginal white stripes. Pattern

as in Figu res 2, 3, 4 or 5. 4

3b. With a definite median white stripe beginning behind second eye
row and continuing to posterior edge of carapace, and with dis-
tinct broad white marginal or submarginal stripes. Pattern as in

Figures 6, 7, 8, or 9. 6

4a. Black in color without distinct lighter markings as in Figure 5.
Epigynum as in Figures 25, 26. Found at high elevations in

Mexico S. plutonus

4b. Dark brown or gray with distinct white markings on carapace

and dorsum of abdomen as in Figures 2-4. 5

5a. Epigynum as in Figures 21, 22. Palp as in Figures 37-39. Found

in southern Texas. S. t exanus

5b. Epigynum as in Figures 13-17. Palp as in Figures 34, 35. Found

from eastern Louisiana to southern Florida S. mimus

6a. Abdomen with wide median brown stripe bordered by white lines
at the anterior end, these lines broken posteriorly as a series of
white dashes. No white chevrons crossing the median brown
stripe, as in Figure 6. Epigynum as in Figures 23, 24. Palp as

in Figures 46, 47 S. mexiccmus

6b. Abdomen with wide median brown stripe with indentations
accented by white spots anteriorly and with a series of white

chevrons crossing the median stripe posteriorly. 7

7a. Epigynum with broad median septum and greatly expanded blade

as in Figures 31, 32. S. michoacanus

7b. Epigynum with relatively narrow median septum and rounded

blade as in Figures 27-30. 8

8a. Epigynum as in Figures 27, 28. Palp as in Figures 44, 45.

S. calif ornicus

8b. Epigynum as in Figures 29, 30 S. agalenoides

Sosippus calif ornicus Simon
Figures 8, 27, 28, 43, 44. Map 1.

Sosippus calif ornicus Simon, 1898, Ann. Soc. Ent. Belgique, 42:25. Female
holotype from Lower California in the Paris Museum. Simon, 1898,
Histoire naturelle des Araignees, 2 (2) :323-325, fig. 331 $. Banks, 1913,
Proc. Acad. Nat. Sci. Philadelphia, 65:182, pi. 9, fig. 13 9. Comstock,
1913, The Spider Book, p. 622; 1940, op. cit., rev. ed., p. 639. Bonnet,
1958, Bibliographia Araneorum, 2(4) : 4093.

140

Psyche

[September

Sosippus pragmaticus Chamberlin, 1924, Proc. Calif. Acad. Sci., 12:674, fig.
117. Female holotype from San Carlos Bay, 8 Jul. 1921 (J. C. Chamber-
lin) in Museum of California Academy of Sciences. NEW SYNONYMY.
Sosippinus calif or nicus: Roewer, 1954, Katalog der Araneae, 2:313; 1959,
Exploration du Parc National de l’Upemba, Araneae Lycosaeformia II
(Lycosidae), p. 1002.

Structure. For comparison of certain diagnostic measurements of
S. calif ornicus to other species see Table I.

Posterior cheliceral margin with three teeth on each side. Of 39
females examined, 36 had 3-3 posterior cheliceral teeth, two 3-4; of
23 males, 22 had 3-3 posterior cheliceral teeth. A female and male
from Mecatan, Nayarit had 4-4 teeth on the posterior margin of the
chelicerae and two immature specimens from Acaponet, Nayarit have
the same number.

In the following table the denotation of tibial spines is from proxi-
mal to distal end, e.g. 1-2-2 indicates that there is one proximal spine,
one pair midway of the tibia, and one distal pair; 1-1 indicates one
spine one-third the length of the leg segment from the proximal end
and another the same distance from the distal end. Of the 39 female

specimens

Tibial

Spination

examined 18 had the
Dorsal

following tibial spination
Ventral Prolateral

Retrolateral

Leg I

0

2-2-2

I-I

0

Leg II

0

2-2-2

I-I

0

Leg III

0

2-2-2

I-I

1-1

Leg IV

0

2-2-2

I-I

1-1

Sixteen females differed from the above only in the ventral spination
of leg II which was 1-2-2. The five remaining specimens varied in
different respects.

Sixteen of the 23 males examined showed the following arrangement
of tibial spines:

Tibial

Spination Dorsal Ventral Prolateral Retrolateral

Leg I

0

2-2-2

1-1

1-1

Leg II

0

2-2-2

1-1

1-1

Leg III

0-1-0

2-2-2

1-1

1-1

Leg IV

0

2-2-2

1-1

1-1

The seven remaining specimens showed variable dorsal spination on
leg III and the ventral spination of leg II was 1-2-2 in three specimens,
otherwise the spination was constant except for malformities, e. g.
regeneration.

On all legs there appear on the dorsal surface of the tibiae, as well

1962]

Brady — Sosippus

141

as the metatarsi, and tarsi a series of trichobothria. These are extreme-
ly variable on all leg segments. The usual pattern is one or two large
proximal trichobothria and then a series of smaller ones decreasing in
size distally, although there may be one or two large trichobothria at
the distal end of the tibiae. Occasionally the basal trichobothria are
spine-like particularly on the tibia of leg III. A spine is distinguished
from a trichobothrium by its greater thickness at the base and the

socket within which it articulates. This basal socket leaves a scar
whenever the spine is broken off, whereas it is difficult to distinguish
an empty socket where a trichobothrium has been broken off.

Of 37 females measured the clypeus height was subequal to the di-
ameter of the AME in 32, in three clypeus height was less than the
diameter of the AME, and in two clypeus height was greater than
the diameter of the AME. Of 23 males examined the clypeus height

1 42 Psyche [September

was subequal to the diameter of the AME in 13, and less than the
diameter of the AME in ten.

Labium. Of 39 females measured : 26 have length subequal to
width, 1 1 wider than long, and two longer than wide. Of 23 males
measured: 19 have length subequal to width, two longer than wide,
and two wider than long.

Color. Female. Pattern illustrated in Figure 8. Carapace dark
brown with the eye region darkest. Eyes outlined with black. A
narrow pale brownish yellow median stripe beginning just behind the
second eye row and running the length of the carapace. Broad mar-
ginal stripes of the same color. Both the marginal stripes and the
median one densely clothed with white hair. Marginal stripes broad-
ening anteriorly and extending to edge of clypeus. Chelicerae dark
reddish brown, almost black.

Sternum brownish yellow. Endites darker reddish brown with
distal ends brownish yellow. Proximal segments of legs brownish
yellow without distinct contrasting markings, metatarsi and tarsi
darker brown.

Dorsum of abdomen with a broad dark brown median stripe from
base to tip, with two pairs of white spots at lateral indentations of
stripe anteriorly, and transverse chevrons clothed with white hair
posteriorly. Brownish yellow area adjacent to brown median stripe
on each side, thickly clothed with white hair; lateral edges of dorsum
darker brown, mottled with tufts of white hair. Venter brownish
yellow.

Male. Very similar to the female in coloration.

Diagnosis. Sosippus calif ornicus resembles S. agalenoides and S.
michoacanus in color pattern. S. michoacanus is darker in color, how-
ever, and both it and S. agalenoides can be separated from S. calif orni-
cus by the form of the epigynum. (compare Figs. 27, 28 with Figs.
29, 30 or 31, 32.) Sosippus plutonus is closest to S. calif ornicus in
the form of the epigynum, but is a very dark, almost black species and
smaller than S. calif ornicus (compare Fig. 8 with Fig. 5). Sosippus

Explanation of Plate 8

Fig. 1. Sosippus floridanus Simon, $ from Highlands Hammock State
Park, Highlands Co., Florida, 9 Jun. 1961.

Figs. 2-3. S. mimus Chamberlin. 2. Female from Archibold Biological Sta-
tion, Lake Placid, Highlands Co., Florida, 24 Jan. 1943. 3. Female holotype
from Mandeville, Saint Tammany Par., Louisiana, 1 May 1921. Note size of
abdomen, due to having recently constructed an egg case.

Fig. 4. S. texanus sp. n., $ paratype from Goose Island State Park,
Aransas Co., Texas, 15 Jun. 1961.

Psyche, 1962

Vol. 69, Plate 8

Brady — Genus Sosippus

144

Psyche

[September

mexicanus is separated from S. calif ornicus by the form of the epigy-
num and by the dorsal color pattern of the abdomen (compare Fig. 8
with Fig. 6).

Natural History. In southern Arizona, according to J. A. Beatty
(unpublished, 1961), S. calif ornicus is found primarily in riparian
woodland formations. These associations occur in or adjacent to drain-
ageways and their floodplains and are characterized by different vege-
tation than that of the surrounding non-riparian community. This
spider occurs at varying elevations depending upon the amount of
vegetation and moisture available. In the Santa Catalina Mountains
it was collected at about 850 m.

Sosippus calif ornicus constructs expansive webs with a central
funnel-shaped retreat leading to the base of vegetation, into crevices,
or under rocks. In El Coyote, Sonora it was collected from webs
leading under rocks in a dry stream bed at about 1000 m. A female
with egg case was collected from beneath a rock in a canyon filled
with Palm trees, 26 km east of Magdalena, about 1300 m elevation.

Distribution. Baja California, southern California, Arizona, south
to Nayarit (Map 1).

Records. Baja California. 9 holotype. California. Imperial Co. :
NE corner 9 (J. Anderson). Los Angeles Co.: Claremont (Baker).
Arizona. Maricopa Co.: Buckeye, 19 Feb. 1956 o (T. Barry) ; Mesa,
21 Man-23 May 00, 16 Jul.-n Nov. cf cf 9 $ (sev. coll.) ; Phoenix,
12-23 Feb. 1956 00 (sev. coll.) ; Santa Cruz River Vallejo, 9 May 1940
O (R. H. Crandall) ; Tempe, 14 Feb.-27 Mar., 20 Nov. i960 00 (sev.
coll.). Pima Co.: Baboquivari Mountains, Brown’s Canyon, 22 Apr.
1961 00 (J. A. Beatty), 9 Jun. 1952 cf cf 99 with egg cases (M.
Cazier, W. J. Gertsch, R. Schrammel), 4 Sep. 1951 9? (W. S.
Creighton), Rancho El Mirador, 4 Sep. 1950 9 (W. J. Gertsch);
Santa Catalina Mountains, Sabino Canyon, 10 Apr. 1959, 26 Sep.
1959, 20 Nov. i960 00, 26 Jun. -2 1 Aug. i960 cf cf 99 (J- A. Beatty),
6 Jun. 1952 cf cf (M. Cazier, W. J. Gertsch, R. Schrammel). Yuma
Co.: Yuma, 12 Feb. 1961 o (J. A. Beatty). Sonora. El Coyote, 28
km E of Rio Bavisbee, 1000 m, 19-24 Jul. i960 99 with egg case,
26 km E of Magdalena, 1300 m, 16 Jul. i960 9 (J- A. Beatty) ;
Minas Nuevas, 8 Aug. 1952 9 (P., C. Vaurie) ; Navajoa, 6 Aug.
1956 9 (V. Roth & W. J. Gertsch). Sinaloa. Culiacan, 19 Jun. 1939
cf (A. M., L. I. Davis). Nayarit. Acaponeta, 20 Nov. 1939 00 (C.
M. Bogert, H. E. Vokes) ; Mecatan, 23 May 1949 cf (G. M.
Bradt) ; San Bias, 6 Aug. 1947 9 (C., M. Goodnight, B. Malkin).

1962]

Brady — Sosippus

145

Sosippus mexicanus Simon
Figures 6, 23, 24, 46, 47. Map 1.

Sosippus mexicanus Simon, 1888, Ann. Soc. Ent. France, 8 (6):206. Female
holotype from Mexico in the Paris Museum. Simon, 1898, Histoire
naturelle des Araignees, 2 (2) :325, 327. F. P. -Cambridge, 1902, Biologia
Centrali-Americana, Araneidea, 2:332, p. 31, figs. 30, 31a-c. Banks, 1909,
Proc. Acad. Nat. Sci. Philadelphia, 61 :2 1 7. Bryant, 1948, Psyche,
55 (2) :55. Roewer, 1954, Katalog der Araneae 2:314. Bonnet, 1958,
BibJiographia Araneorum, 2(4):4093. Roewer, 1959, Exploration du
Parc National de l’Upemba, Araneae Lycosaeformia II (Lycosidae), p.
1004.

Structure. For comparison of certain diagnostic measurements of
S. mexicanus to other species see Table 1.

Posterior cheliceral margin with three teeth on each side. Of two
females examined one had 3-3 posterior cheliceral teeth, the other 4-3.
The male specimen, described by F. Pickard-Cambridge (1902), has
3-3 posterior cheliceral teeth. According to F. P.-Cambridge all the
specimens that he examined had 3-3 posterior cheliceral teeth.

Tibial spination in the two females examined was the same as in S.
calif ornicus with the following exceptions: ventral spines on leg II
1-2-2, dorsal spines on tibia III and IV 1-1. These spines slightly
smaller than the lateral spines.

Clypeus height of one female subequal to the diameter of the AME,
in the other greater than the diameter of the AME. Clypeus height
of the male subequal to the diameter of the AME.

Labium. Length subequal to width in the male and two females.

Color. Female. Pattern illustrated in Figure 6. Carapace brown,
darker in the eye region with the eyes circled in black. A thin pale
brownish yellow median stripe beginning immediately behind the
second eye row and continuing to the posterior edge of the carapace.
Broad marginal stripes of the same color starting at the edge of the
clypeus and extending the length of the carapace. The marginal
stripes, as well as the median stripe, clothed with short white hair.
Darker brown lines radiating from the thoracic groove. Chelicerae
dark reddish brown.

Sternum brownish yellow. Labium and endites darker reddish
brown. Legs brownish yellow without distinct darker markings.

Dorsum of the abdomen with a wide brown median stripe, enclosed
by two very light broken lines in the anterior region. Posteriorly the
lines are broken-up into a series of dots, the broken lines and dots
accented with white hair. Lateral to the light lines and dots the
dorsum is brownish yellow mottled with brown. Venter of the abdo-

146

Psyche

[September

men with a concentration of white pigment under the integument,
overlaid with yellow, producing a cream color.

Male. Median longitudinal stripe of carapace not as distinct as in
the female. Dorsum of abdomen with a series of paired white spots
outlining wide brown median stripe, otherwise similar to female in
coloration.

Diagnosis. This species is similar to S. agalenoides in the form of
the epigynum. It is separated from S. agalenoides most easily on the
basis of size in the limited number of specimens examined. Sosippus
mexicanus does not exceed 15 mm in total body length and S. agale-
noides is not less than 20 mm. Sosippus mexicanus also differs from
S. agalenoides in the dorsal pattern of the abdomen (compare Fig. 6
to Fig. 9) and tibial spination. Sosippus agalenoides is more hairy
than S. mexicanus , especially on the dorsum of the abdomen. The
venter of the abdomen in S. mexicanus is characterized by white pig-
ment underlying the integument, which is not found in S. agalenoides.
If the differences in tibial spination remain consistent upon examina-
tion of a larger series of specimens, this will provide an easy method of
separation.

Natural History. According to F. Pickard-Cambridge (1902),
“The spiders of this genus are very much like Agelenae , not only in
the shape of their body and the appearance of their legs, but to some
extent their markings. Still more do they resemble them in habits, for
the web consists of a very large sheet of fine silk spun over the bushes,
with a tube-like tunnel running down into some place of safety. Their
movements are exceedingly rapid, and one cannot at first sight believe
they are not true A gelenae ”

Distribution. Guerrero to Guatamala,

Records. Guerrero. Acapulco de Juarez, 1 Sep. 1940 (H. E.

Frizzell). Guatamala. <$ (Sarg). N. Banks (1909) reported this
species from Costa Rica, but the specimens in question are immature
and appear to be S. agalenoides , which Banks described from Costa
Rica.

Sosippus agalenoides Banks
Figures 9, 29, 30. Map 1.

Sosippus agalenoides Banks, 1909, Acad. Nat. Sci. Philadelphia, 61:217, pi.
6, fig. 33. Three female syntypes and one juvenile from Puntarenas, Costa
Rica, in the Museum of Comparative Zoology, examined. Roewer, 1954,
Katalog der Araneae, 2:313. Bonnet, 1958, Bibliographia Araneorum,
2(4):4093. Roewer, 1959, Exploration du Parc National de l’Upemba,
Araneae Lycosaeformia II (Lycosidae), p. 1004.

1962]

Brady — Sosippus

H7

Structure. For comparison of certain diagnostic measurements of
S. agalenoides to other species see Table 1.

Posterior cheliceral margin with three teeth on each side. Of four
female and one juvenile specimens examined all have 3-3 posterior
cheliceral teeth.

Tibial spination in each of the four females is exactly the same as
that shown in the table for S. calif ornicus.

Clypeus height is greater than the diameter of the AME in three
specimens, in one specimen it is subequal to the diameter of the AME.

Labium longer than wide.

Color. Pattern illustrated in Figure 9. Carapace light orange-
brown, darkest in the eye region. A thin median stripe, clothed with
white hair, begins slightly in front of the third eye row and continues
to the thoracic groove. Broad marginal longitudinal stripes, not as
distinct as in S. calif ornicus ; white hairs along these stripes heaviest
laterally just behind the clypeus, more diffuse posterior to this region.
The lateral edges of the carapace clothed with fine white hair. Cheli-
cerae dark reddish brown, almost black; boss on each side prominent,
with dense white hair surrounding it except at the point of articula-
tion.

Sternum light brownish yellow. Endites and labium darker reddish
brown, yellowish at distal ends. Legs light brownish, yellow with
metatarsi and trasi darker reddish brown.

Dorsum of the abdomen with a wide brown median stripe beginning
at the base and continuing to the spinneretes. The anterior end of
this stripe bordered by three white dashlines on each side, and inter-
rupted posteriorly by a series of white chevron markings. These
markings clothed with white pubescence. A series of dark brown dots
alternate crossing the dorsum with the white chevrons and continue
laterally for some distance. Lateral area of abdomen light brownish
yellow, venter lighter, almost cream colored.

Diagnosis. Sosippus agalenoides is most similar to S. calif ornicus
in coloration. It is separated from this species by the shape of the
epigynum (compare Figs. 29, 30 to Figs. 27, 28). Sosippus agalenoides
is similar to S. mexicanus in the form of the epigynum. It differs from
S. mexicanus in size and other characters discussed under that species.

Natural History. Banks (1909) does not supply any information
concerning the natural history of this species. Presumably it builds
the same type of sheet web and tubular retreat as found in S. cali-
f ornicus.

Distribution. Morelos, Oaxaca, Costa Rica.

148

Psyche

[September

Records. Morelos. Miacatlan, 4 Oct. 1942 O (C. Bolivar).
Oaxaca. Tehauantepec, 15 Dec. 1947 o, 6-10 Feb. 1948 o (T. Mac-
Dougall) ; Cerro del Armadillo, 7 Jan. 1948 $ (T. MacDougall).
Costa Rica. Puntarenas, 1909 9? (P. Biolley).

Sosippus michoacanus sp. n.

Figures 7, 31, 32. Map 1.

Type. Female holotype from Tzararacua Falls, 11 km from Urua-
pan, Michoacan, Mexico, 14 Jun. 1948 (A. M., L. I. Davis) in
the American Museum of Natural History. The species named after
the type locality.

Structure. For comparison of certain diagnostic measurements of S.
michoacanus to other species see Table 1.

Posterior cheliceral margin with three teeth on each side. Of five
females examined, all had this arrangement.

Tibial spination was the same as in S. calif ornicus with the excep-
tion that four specimens had 1-2-2 ventral spines on tibia II.

Clypeus height subequal to the diameter of the AME. Labium
longer than wide.

Color. Pattern illustrated in Figure 7. Carapace reddish brown,
overlaid with black pubescence giving it a darker appearance. Eye
region dark brown, almost black, with eyes encircled with black. Thin
median stripe beginning some distance behind the third eye row, con-
tinuing to posterior edge of carapace. This stripe not conspicuous, with
white hairs sparsely covering it, densest along the thoracic groove.
Broad submarginal stripes extending from clypeus to posterior edge of
carapace, clothed with white pubescence. Chelicerae very dark reddish
brown, almost black, with intermitent, long black hairs. Conspicuous
boss on each side, light brown.

Sternum light brown with gray overtones. Endites and labium
reddish brown, yellowish at distal ends. Femora brown with dusky

Explanation of Plate 9

Fig. 5. Sosippus plutonus sp. n., $ holotype from Tenango del Valle,
Mexico, Mexico, 25-26 Aug. 1946.

Fig. 6. S. mexicanus Simon, $ from Acapulco de Juarez, Guerrero, Mex-
ico, 1 Sep. 1940.

Fig. 7. S. michoacanus sp. n., 9- holotype from Tzararacua Falls, 11 km
from Uruapan, Michoacan, Mexico, 14 Jun. 1948.

Fig. 8. S. cahf ornicus Simon, $ from Brown’s Canyon, Baboquivari Mtns.,
Pima Co., Arizona, 9 Jun. 1952 (pattern and color same as specimens from
Sabino Canyon, Santa Catalina Mtns., Pima Co., Arizona, 26 Jun. 1960).

Fig. 9. S. agalenoides Banks, $ syntype from Puntarenas, Costa Rica, 1909.

Psyche, 1962

Vol. 69, Plate 9

Brady — Genus Sosippus

150

Psyche

[September

markings, uniformly gray on ventral surface, but forming bands on
the lateral and dorsal surfaces. Patellae and tibiae of same color, but
darker markings form more indefinite pattern. Metatarsi and tarsi
yellowish brown, without dusky markings.

Dorsum of abdomen with basal lanceolate reddish brown mark,
enclosed by typical wide dark brown stripe heavily pigmented with
black. Dark brown median stripe bordered by lighter brown area,
with lateral regions again becoming dark brown. Venter light yellow-
ish brown.

Diagnosis. Sosippus michoacawus has an abdominal pattern similar
to S. calif ornicus. It is much darker in color than S. calif ornicus and
is readily recognized by the female epigynum, which has a wide median
septum and greatly expanded blade (Fig. 32).

Natural History. No information concerning the habits is available.
It presumably constructs a funnel-web like that of S. calif ornicus.

Distribution. Michoacan and Guerrero.

Records. Michoacan, Tzararacua Falls, 11 km from Uruapan,
14 Jun. 1948 $$ (A. M., L. I. Davis). Guerrero. Mexcala, 2 Jul.
1941 $ (L. I. Davis) .

Sosippus plutonus sp. n.

Figures 5, 25, 26. Map 1.

Type. Female holotype from Tenango del Valle (Tenango de
Arista), 2400 m elev., Mexico, Mexico, 25-26 Aug. 1946 (H. Wag-
ner) in the American Museum of Natural History.

Structure. For comparison of certain diagnostic measurements of
S. plutonus to other species see Table 1.

Female holotype with 3-3 posterior cheliceral teeth. Of seven
immature specimens examined, six had 3-3 posterior cheliceral teeth,
one 3-4.

Tibial spination was the same as in S. calif ornicus with the follow-
ing exceptions: ventral spination of leg II 1-2-2, dorsal spination of
leg III 0-1-0.

Clypeus height equal to the diameter of the AME. Labium slightly
wider than long.

Color. Pattern illustrated in Figure 5. Carapace very dark reddish
brown with the eye region black. A few white hairs along the mid-
line might be indicative of a thin median white line once present.
Broad submarginal stripes, brownish yellow and covered with white
pubescence, beginning about the third eye row and continuing to the

1962] Brady — Sosippus 1 5 1

posterior edge of the carapace. Chelicerae black with lighter colored
boss on each side.

Sternum brown. Endites and labium dark brown, lighter at distal
ends. Coxae dark gray-brown on ventral surface with lighter amber
color at basal regions. Other leg segments gray-brown with lighter
amber colored stripes as follows : dorsal paired stripes at proximal and
distal ends of femora, stripe covering almost entire dorsal surface of
patellae, paired stripes at distal end of tibiae.

Abdomen (considerably shrivelled) dark brown, almost black, with
a series of paired lighter spots formed from tufts of white hair. Venter
brown.

Diagnosis. Sosippus plutonus is most similar to S. calif ornicus in
the shape of the epigynum. The median septum in S. plutonus (Fig.
26) is shorter than in S. calif ornicus (Fig. 28). Sosippus plutonus is
much darker than S. calif ornicus, being almost black, hence the specific
name.

Natural History. Sosippus plutonus presumably constructs a funnel
web similar to that of S. calif ornicus. Sosippus plutonus is probably a
montane species being found at extremely high altitudes in Mexico.

Distribution. Mexico, Mexico.

Records. Mexico. Tenango del Valle (Tenango de Arista), 2400
m elev., 25-26 Aug. 1946 00 $ (H. Wagner) ; Tenancingo, 2050 m
elev., 27 Sep.-7 Oct. 1946 00 (H. Wagner).

Sosippus floridanus Simon
Figures 1, 19, 20, 40-43. Map 1.

Sosippus floridanus Simon, 1898, Ann. Soc. Ent. Belgique, 42:25. Female holo-
type from Florida in the Paris Museum. Simon, 1898, Histoire naturelle
des Araignees, 2 (2):323, 325. Comstock, 1913, The Spider Book, p. 622,
op. cit. rev. ed., p. 639. Banks, 1904, Proc. Acad. Nat. Sci. Philadelphia,
56:121, 135. Chamberlin, 1908, Proc. Acad. Nat. Sci. Philadelphia, 60:293,
pi. 23, fig. 2, pi. 11, figs. 1 $, 4 9. Wallace, 1950, Florida Ent., 33:76.
Roewer, 1954, Katalog der Araneae, 2:314. Bonnet, 1958, Bibliographia
Araneorum, 2(4):4093. Roewer, 1959, Exploration du Parc National de
l’Upemba, Araneae Lycosaeformia II (Lycosidae), p. 1004.

Structure. For comparison of certain diagnostic measurements of
S. floridanus to other species see Table 1.

Posterior cheliceral margin with four teeth on each side. Of 37
females examined, 35 had 4-4 posterior cheliceral teeth; two had 4-5
posterior cheliceral teeth. Of 11 males examined 10 had 4-4 posterior
cheliceral teeth, one had 4-5 posterior cheliceral teeth.

Tibial spination in 16 females was the same as in S. calif ornicus
with the exception of ventral spination on leg IF In this position nine

Psyche, 1962

Vol. 69, Plate 10

Brady — Genus Sosippus

1962]

Brady — Sosippus

153

females had 1-2-2 spines, five had 1-1-2, and two females had 1-2-2
on one leg and 1-1-2 on the other leg. Tibial spination in three males
was the same as in S. calif ornicus except for 1-1 dorsal spines on leg
III and IV in S. florid anus.

Clypeus height in nine females is greater than the diameter of the
AME, in seven females clypeus height is subequal to the diameter of
the AME. In the three males examined clypeus height is subequal to
the diameter of the AME.

Labium longer than wide in 14 females, length equal to width in
two females. In the three males the labium is longer than wide.

Color. Female. Pattern illustrated in Figure 1. Carapace dark
brown, overlaid with black pubescence, black in eye region. Narrow
median longitudinal white stripe beginning at second eye row and
continuing to the posterior edge of carapace. Broad submarginal white
stripes originating at edge of clypeus and running to the posterior edge
of carapace. The white color of these stripes is due to presence of
short appressed white hairs. The fine white hairs and the black pubes-
cence provide contrasting pattern illustrated. If these hairs become
rubbed off the underlying integument is almost unicolorous, the
regions of the white stripes may remain somewhat lighter, however.

Chelicerae black with stout black hairs, lighter orange brown boss
on each side. Sternum brownish yellow. Endites and labium reddish
brown, lighter at distal ends. Coxae light brownish yellow on ventral

Explanation of Plate 10

Fig. 10. Frontal view, showing eye arrangement of Sosippus tcxanus, sp.
n., $ from Goose Island State Park, Aransas Co., Texas, 15 Jun. 1961.

Fig. 11. Frontal view, showing eye arrangement of Porrima diversa (O.
P.-Cambridge) from Rockstone, British Guiana, 1 July (determined by W. T.
Gertsch).

Figs. 13-17. <S\ mimus Chamberlin. 13-14. Female with 3-3 posterior chelic-
eral teeth from Archibold Biological Station, Lake Placid, Highlands Co.,
Florida, 24 Jan. 1943. 13. Genitalia, dorsal view. 14. Epigynum. 15-16.

Female holotype with 3-3 posterior cheliceral teeth from Mandeville, Saint
Tammany Par., Louisiana, 1 May 1921. 15. Genitalia, dorsal view. 16. Epigy-
num. 17. Genitalia, dorsal view of specimen with 4-4 posterior cheliceral
teeth from Alachua Co., Florida, 18 Apr. 1935.

Figs. 18-20. S. floridanus Simon, $ from Highlands Hammock State Park,
Highlands Co., Florida, 9 Jun. 1961. 18. Genitalia, dorsal view. 19. Epigy-
num. 20. Diagrammatic ventral view showing route of various ducts.

Abbreviations: S, seminal receptacle; B, copulatory bursa; P, copulatory
pouch (usually not heavily sclerotized) ; L, dorsal bursal ligament (difficult
to see unless the epigynum is tipped on edge). The route of this ligament is
difficult to trace, due to its transparency, and inaccuracy in its placement in
the drawings might occur, since it can hardly be seen from a dorsal view.

Figs. 21-22. S. texanus sp. n., $ paratype from Goose Island State Park,
Aransas Co., Texas, 16 Jun. 1961. 21. Genitalia, dorsal view. 22. Epigynum.

154

Psyche

[September

surface. Femora gray on ventral surface, gray color continuing around
leg segment at proximal and distal ends and toward the center as a
pair of wide bands against a brownish yellow background clothed with
white pubescence. Remaining leg segments brownish yellow, thickly
clothed with black hair.

Dorsum of abdomen with a reddish brown lanceolate mark at the
base, enclosed by a wide black median stripe. This broad stripe with
tufts of white hair forming two pairs of spots at indentations in the
anterior end and with chevrons clothed with white hair crossing it at
the posterior end. Lateral to dark median stripe are areas of brown
with scattered black spots. Venter of abdomen grayish brown, pro-
duced by black hair overlying brownish yellow integument.

Male. The white spots and chevrons on the dorsum of the abdomen
are not as conspicuous as in the female, otherwise the coloration and
markings are much the same.

Diagnosis. Sosippus floridanus is most similar to S. mimus. The
form of the epigynum in these two species is similar but consistently
different (compare Figs. 18, 19 to Figs. 13-17). The male palpi are
much alike, but in S. floridanus the tarsal segment is usually only twice
as long as wide, while in S. mimus the tarsal segment is usually more
than twice as long as wide (compare Figs. 40-43 to Figs. 34, 35) . The
most clear-cut difference between these two species is in the color pat-
terns (compare Fig. 1 to Figs 2, 3),

Natural Ilistory. Sosippus floridanus was collected from tubular
webs leading under the trunks of Palmetto bushes and into the bases
of tufts of high grass in areas of white, sandy soil. The webs were not
extensively developed, the tubular portion usually being hidden and
the sheet portion of the web consisting of radiating lines of silk forming
a very loose meshwork. Additional specimens were collected from
holes and ruts along a path of white sand that had once been cleared

Explanation of Plate 11

Figs. 23-24. Sosippus mexicanus Simon, $ from Acapulco de Juarez,
Guerrero, Mexico, 1 Sep. 1940. 23. Genitalia, dorsal view. 24. Epigynum.

Figs. 25-26. S. plutonus sp. n., 9 holotype from Tenango del Valle,
Mexico, Mexico, 25-26 Aug. 1946. 25. Genitalia, dorsal view. 26. Epigynum.

Figs. 27-28. S. calif ornicus Simon, $ from Brown’s Canyon, Baboquivari
Mtns., Pima Co., Arizona, 9 Jun. 1952. 27. Genitalia, dorsal view. 28. Epigy-
num.

Figs. 29-30. S. agalenoidcs Banks, 9 syntype from Puntarenas, Costa Rica,
1909. 29. Genitalia, dorsal view. 30. Epigynum.

Figs. 31-32. S. michoacanus sp. n., 9 paratype from Tzararacua 11 km
from Uruapan, Michoacan, Mexico, 14 Jun. 1948. 31. Genitalia, dorsal view.
32. Epigynum.

Psyche, 1962

Vol. 69, Plate 11

24

Brady — Genus Sosippus

56

Psyche

[September

for vehicles. In these situations the webs were not typical funnel webs
at all. However, most of these specimens were immature and this may
account for their under-developed webs.

Distribution. Florida.

Records. Florida. Alachua Co: i-io May 1934-37 cf c? $ (H. K.
Wallace, A. F. Carr) ; Gainesville, 12 Jun. 1935 $ (W. J. Gertsch).
Desoto Co.: 29 Jun. 1935 cf 99 (H. K. Wallace); 8 mi. W. of
Arcadia, 21 Mar. 1938 9 (W. J. Gertsch). Flagler Co.: 2 Sep. 1958
(H. IC Wallace). Highlands Co.: 1 Jun. 1958 $ (N. Causey);
29 Jun. 1935 9 (H. K. Wallace) ; Highlands Hammock State Park
near Sebring, 24 Mar. 1938 9 (W. J. Gertsch), 9 Jun. 1961 cf 9$
(A. R. Brady) ; Lake Placid, Archibold Biological Station, 25 Jum-4
Feb. 1943 00 (M. Cazier). Indian River Co.: 17 Mar. 1936, cf (H.
T. Townsend). Lake Co.: 21 Apr. 1933 cf (H. K. Wallace);
Emeralda Canal, 6 Mar. 1936 9 (Bishop coll.) ; Umatilla, Jul. 1934
9 (M. Broyles). Martin Co.: Port Mayaca on Lake Okeechobee,
29 Mar. 1938 cf 99 (W. J. Gertsch). Monroe Co.: Flamingo,
13 Mar. 1920 9 (W. S. Brooks). Lee Co.: 14 Apr. 1949 o 99 (H.
K. Wallace). Levy Co. : 9 Apr. 1937 00 9 ( H. K. Wallace) . Orange
Co.: 7 mi. E of Apopka, 29 Aug. 1944 9 (M. Nirenberg) ; Dec. 1934
cf (K. Boyer). Osceloa Co.: Runnymede 9 (N. Banks). Polk Co.:
Hobbs, 5.6 mi. W of Lake Wales, 13 Mar. 1937 00 9; 27 Jun. 1935
99 (H. K. Wallace). Sarasota Co.: 19 Jul. 1937 9 (M., A. Carr) ;
Englewood, 1-5 Apr. 1938 9 (W. J. Gertsch). Volusia Co.: Enter-
prise, 7 Jun. 1946 cf 9 (F. N. Young).

Sosippus mimus Chamberlin
Figures 2, 3, 13-17, 34, 35- Map 1.

Sosippus mimus Chamberlin, 1924, Proc. U. S. Nat. Mus. 63:27, pi. 6, fig.
43. Female holotype from Mandeville, Louisiana, 1 May 1921 (H. E.
Hubert) in Museum of Comparative Zoology, examined. Comstock, 1940,
The Spider Book, p. 639. Bonnet, 1958, Bibliographia Araneorum,
.2(4) :4093.

Sosippinus mimus: Roewer, 1954, Katalog der Araneae 2:313; 1959, Explora-
tion du Parc National de 1’Upemba, Araneae Lycosaeformia II (Lycosi-
dae), p. 1002.

Structure. For comparison of certain diagnostic measurements of
S. mimus to other species see Table 1.

Number of cheliceral teeth on posterior margin extremely variable.
Of 12 females examined, nine had 4-4 posterior cheliceral teeth, two
had 3-3 posterior cheliceral teeth, and one had 3-4 posterior cheliceral
teeth. Of nine males examined, two had 3-3 posterior cheliceral teeth,

157

1962] Brady — Sosippus

two had 3-4 teeth, three had 4-4 teeth, one had 3-5 teeth, and one had
4-5 teeth.

Tibial spination in four females examined was the same as in S.
calif ornicus with the following exceptions : ventral spination of leg II
1-2-2. Tibial spination in four males was the same as in S. calif ornicus
with the following exceptions: dorsal spination of legs III and IV 1-1,
one male lacked the dorsal spines on leg IV.

Clypeus height in three females greater than the diameter of the
AME, in one female clypeus height subequal to the diameter of the
AME. Clypeus height in two males greater than the diameter of the
AME, clypeus height in one male subequal to the diameter of the
AME, diameter of the AME greater than the clypeus height in one
male.

Labium longer than wide except that in one female it is slightly
wider than long.

Color. Female. Pattern illustrated in Figures 2, 3. Carapace dark
reddish brown (mahogany), darkest in eye region, with eyes circled
in black. Three longitudinal stripes beginning just in front of third
eye row and consisting of: one median stripe continuing to thoracic
groove, two lateral stripes passing inside of eyes of third row and end-
ing in the cephalic region. These three stripes yellowish white in
color. Lateral bands of the same color at anterior edge of carapace
and another pair of yellowish white bands below eyes of third row
running downward and diagonally backwards. No distinct stripe
along margins of carapace, diffuse yellowish white hairs instead. Black
lines radiating from thoracic groove, accented by yellowish white
pubescence.

Chelicerae black, clothed with large black hairs, with orange brown
boss on each side. Sternum yellowish brown. Labium and endites
orange-brown, lighter at distal ends. Leg segments yellowish brown
without distinct banding. Ventral surface of femora sometimes having
a gray appearance due to heavy clothing of black hairs. Dorsal surface
of femora clothed with short white hair.

Dorsum of abdomen with a wide brown median stripe from the
base of the abdomen to the posterior end, interrupted at intervals. A
pair of whitish spots at lateral edges of median stripe anteriorly, fol-
lowed by a series of five white chevrons crossing the median stripe ; the
ends of the chevrons merging with white spots at the lateral edges.
Bands of brown dots alternately crossing the median stripe with the
chevrons. Lateral areas of dorsum yellowish brown mottled with
darker brown. Venter of abdomen brownish yellow.

158

Psyche

[September

Male. The pattern in the male is very similar to that of the female.

Diagnosis. Sosippus mimus resembles S. floridanus in male and
female genitalia and is much like S. texanus in color pattern. Sosippus
mimus is separated from the former by characters discussed under that
species. Sosippus mimus is smaller than S. texanus (see Table i).
Although the males are difficult to separate by differences in the palpi,
the females are easily separated by differences in the epigyna (compare
Figs. 13-17 to Figs. 21, 22). It is possible that the specimens con-
sidered in this paper under S. mimus constitute more than one species.
There is considerable variability of certain characters in this species,
but these characters do not show geographic variation that would
permit separation, e. g. the holotype from Mandeville, Louisiana and
specimens from Lake Placid, Florida have 3-3 posterior cheliceral
teeth, while some specimens between these two localities have
4-4 posterior cheliceral teeth. The specimens considered under S.
mimus all agree in color pattern and genitalia. Additional collections
and field studies will provide a more definite answer.

Natural History. Sosippus mimus, according to R. V. Chamberlin
(1924), was collected from a funnel web much like that of other
species in the genus.

Distribution. Georgia, south into Florida and along the Gulf Coast
to eastern Louisiana.

Records. Georgia. Charlton Co. : Chesses Island, Pinebarrex,
15 Jun. 1922 9 (Wright). Florida. Alachua Co.: 13 Apr.-i8 May
I935"5C> cf 99 (H. K. Wallace). Columbia Co.: 27 Apr. 1935
9 (H. K. Wallace). Highlands Co.: Lake Placid, Archibold Biologi-
cal Station, 24 Jan.-4 Feb. 1943 00 9 (M. Cazier). Jackson Co.:
3 Apr. 1953 cf (H. K. Wallace). Levy Co.: 20 Apr. 1935 cf 99
(H. K. Wallace). Liberty Co.: Blountstown, 17 Apr. 1938 9 (W. J.

Explanation of Plate 12

Fig. 33. Left palpus, ventral view of Porrima harknessi Chamberlin, $
holotype from Huadquina, Peru, July, 1911.

Figs. 34-35. Sosippus mimus Chamberlin, $ from Blountstown, Liberty
Co., Florida, 17 Apr. 1938. 34. Left palpus, ventral view. 35. Left palpus,
retrolateral view.

Fig. 36. S. calif ornicus Simon, $ from Brown’s Canyon, Baboquivari
Mtns., Pima Co., Arizona, 9 Jun. 1952. Expanded left palpus.

Figs. 37-39. S. texanus sp. n., $ from Goose Island State Park, Aransas
Co., Texas. 37. Palpus, ventral view. 38. Palpus, ventral view (more retro-
lateral than 37). 39. Palpus, retrolateral view.

Abbreviations: C, conductor; E, embolus; H, basal haematodocha ; L,
lateral apophysis of conductor; M, median apophysis; A, mesal apophysis
of tegulum; T, tegulum.

Psyche, 1962

Vol. 69, Plate 12

Brady — Genus Sosippus

i6o

Psyche

[September

Gertsch). Marion Co.: Kerr Park near Lake Kerr, Oct. 1956 ?
with egg case (H. K. Wallace) . Saint Johns Co. : Hastings, Jul. 1927
cf (J. L. Scribner). Suwanee Co.: 25 Mar. 1933 c? (H. K. Wal-
lace). Mississippi. Forrest Co.: Hattiesburg, 2-6 Jan. 1942 00 (E. L.
Bell). Jackson Co.: Ocean Springs, 10 May 1931 c? (Dietrich).
Louisiana. Saint Tammany Par.: Mandeville, 1 May 1921 $ with
egg case (H. E. Hubert).

Sosippus texanus sp. n.

Figures 4, 21, 22, 37-39. Map 1.

Type. Female holotype from Goose Island State Park, Aransas Co.,
Texas, 15 Jun. 1961 (A. R. Brady) in the Museum of Comparative
Zoology.

Structure. For comparison of certain diagnostic measurements of
S. texanus to other species see Table 1.

Posterior cheliceral margin with three teeth on each side. Of 11
females examined all have 3-3 posterior cheliceral teeth. Two males
also have 3-3 posterior cheliceral teeth.

Tibial spination in 10 females is the same as S. calif or nicus with the
following exceptions in the case of ventral spines on leg II : six females
with 1-2-2 spines, three females with 1-2-2 spines on one leg and 1-1-2
spines on the other leg, and one female with 1-1-2 spines. 7\vo males
have the same tibial spination as S. calif ornicus with the exception of
the dorsal spination of legs III and IV which is 1-1 on each leg.

Clypeus height in seven females is greater than the diameter of the
AME, in three females the clypeus height is subequal to the diameter
of the AME. In the two males clypeus height is subequal to the
diameter of the AME.

Labium longer than wide in eight females and length subequal to
width in two females. Labium longer than wide in the two males.

Color. Female. Pattern illustrated in Figure 4. Carapace dark
brown, covered with black pubescence, black in eye region. Three
longitudinal white stripes beginning behind the second eye row where
they are convergent, consisting of one median stripe and two lateral
stripes passing on the inside of the eyes of the third row, ending at
edge of cephalic region. Marginal areas of carapace lighter, but no
definite stripe present. Sides of cephalic region with a white band
behind the clypeus, followed by a black band, then another white band
below eyes of third row running diagonally backward and downward.
Black lines, accented with white hair, radiating from the thoracic
groove.

1962]

Brady — So sip pus

161

Chelicerae black, clothed with long black hairs, conspicuous orange-
brown boss on each side. Sternum yellowish brown. Endites and
labium dark reddish brown, brownish yellow at distal ends. Ventral
surface of coxae yellowish brown. Femora gray-brown on ventral
surface. White hairs on dorsal surface of femora forming alternating
bands, consisting of : proximal gray-brown, white, gray-brown, white,
and distal gray-brown. Remaining leg segments dark brown with
tarsi and metatarsi darker. Heavy scopulae on tarsus and metatarsi
of legs I and II dark gray in color.

Text Fig. 2. Funnel web of S. texanus constructed in corner of cardboard
box with retreat behind large stick.

Dorsum of abdomen with a wide brown median stripe running the
length of the abdomen, with diffuse black hairs covering it. Two pairs
of white dots at the lateral edges of the stripe anteriorly, posterior to
these a series of six paired white spots connected by white chevrons
crossing the brown stripe. Usually the first chevron is represented by
two dashes and the arms of the second chevron do not quite join at
the midline. Alternating with the white chevrons are bands of darker
brown dots. The areas on each side of the median stripe are speckled

Psyche, 1962

Vol. 69, Plate 13

Brady — Genus Sosippus

Figs. 40-43. Sosippus floridanus Simon. 40-41. Male from Alachua Co.,
Florida, 8 May 1934. 40. Left palpus, ventral view, showing lateral apophysis
of conductor resting normally against mesal apophysis of tegulum. 41. Left
palpus, retrolateral view. 42-43. Male from Port Mayaca, Lake Okeechobee,
Martin Co., Florida, 29 Mar. 1938. 42. Palpus, ventral view showing lateral
apophysis of conductor lying below median apophysis of tegulum. 43. Palpus,
retrolateral view.

Figs. 44-45. S. calif ornicus Simon, $ from Brown’s Canyon, Baboquivari
Mtns., Pima Co., Arizona, 9 Jun. 1952. 44. Palpus, ventral view. 45. Palpus,
retrolateral view.

Figs. 46-47. S . mexicanus Simon, $ from Guatamala. 46. Palpus, ventral
view. 47. Palpus, retrolateral view.

1962]

Brady — Sosippus

63

with tufts of white hair intermixed with dark brown dots on a lighter
brown background. Venter light brown.

Male. The pattern in the male is very similar to that of the female.
There are fewer black hairs over the body and in over-all appearance
the male is more brown than the female, which appears dark gray 01-
even black.

Diagnosis . Sosippus texanus is most similar to S. mimus in colora-
tion. The white spots and chevrons on the abdomen of S. texanus are
usually more distinct. Sosippus texanus is also larger (see Table 1)
and slightly darker than S. mimus. The male palpal organs are very
similar, but the females are readily separated by differences in the
epigyna (compare Figs. 21, 22 to Figs. 14-17).

Natural History. Specimens of S. texanus were collected and
observed in the field at Goose Island State Park. This is a peninsular
area of dry sandy soil, characterized by dense stands of Live Oak
(Quercus virginiana) . Sosippus texanus was collected in an area at
the edge of a clearing from conspicuous funnel webs with tubular
retreats running under logs and holes in the ground. The sheets of
these funnel webs were not as extensive as those observed in Agelenop-
sis. Sosippus texanus was also collected from funnel webs at the base
of the trunks of these trees. Again the sheet portion of the webs was
not highly developed. The emphasis upon the tubular portion of the
webs may have been due to their situation, especially those webs at the
base of trees.

The spider rests at the mouth of the funnel and rushes forth with
great speed to grab insects that have touched the sheet portion of the
web. The prey consists chiefly of grasshoppers and ground beetles that
have fallen upon or run across the lines of radiating silk forming the
sheet. Upon grabbing the insect the spider retreats rapidly into the
tubular portion of the funnel so that the struggles of the victim are
impeded.

Spiders whose retreats dead-end into a hole or crevice construct a
second short tube that they utilized for an escape exit. Most spiders
seemed to have escape exists running off at angles from the main tube
rather than having an exit directly to the rear.

Several spiders from this locality were reared to maturity and were
observed for several months. One spider kept in a cardboard box,
provided with sand and sticks for a retreat, constructed a funnel web
like that found in the field at the base of Live Oak trees. This web is
shown in the accompanying photograph (Text Fig. 2).

Distribution. Southern Texas.

164

Psyche

[September

Records. Texas. Aransas Co.: Goose Island State Park, 7 Mar.
1959 00, 15 Jun. 1961 cT cf 9? paratypes (A. R. Brady). Cameron
Co.: Brownsville, 30 Nov. 1934 9 (S. Mulaik). Hidalgo Co.:
Resaca, 5 mi. SE of Brownsville, 26 Sep. 1937 9 (L. I. Davis, M.
Fones), Edinburgh, Sep.-Dee. 1933 9 (S. Mulaik), La Joya, 30 Oct.
1938 99 (L. I. Davis). Zapata Co.: Lopeno, 15 Apr. 1952 9 with
egg case (Willie).

Literature Cited

Banks, N.

1909. Arachnida from Costa Rica. Proc. Acad. Nat. Sci. Philadelphia,
61:194-234-.

Beatty, J. A.

1961. The spiders and scorpions of the Santa Catalina Mountain Area,
Arizona, (unpubl. thesis, Univ. of Arizona).

Bonnet, P.

1958. Bibliographia Araneorum. Toulouse, 2 (4) :3027-4230.

Bryant, E. B.

1923. Report on the spiders collected by the Barbados-Antiqua Expedi-
tion from the Univ. of Iowa in 1918. Univ. Iowa Study. Nat.
Hist., 10(3) :10- 1 6.

Buchar, J.

1959. Beitrag zur Bestimmuung der Mitteleuropaischen Arten der Gat-
tung Trochosa (C. L. Koch). Acta Univ. Carolinae, Biol., no.
3 :159-164.

Chamberlin, R. V.

1924. The spider fauna of the shore and islands of the Gulf of Cali-
fornia. Proc. California Acad. Sci., 12:561-694.

Pickard-Cambridge, F. O.

1902. Arachnida, Araneidea. in Biologia Centrali-Americana, 2:313-424.

Roewer, C. F.

1954. Katalog der Araneae. Brussels, 2 :923.

1959. Lycosaeformia II (Lycosidae). Exploration du Parc National de
l’Upemba. Brussels, 2:519-1040.

Simon, E.

1888. Descriptions d’especes et de genres nouveaux de l’Amerique
centrale et des Antilles. Ann. Soc. Ent. France, 8 (6) :203-216.

1898. Histoire naturelle des Araignees, Paris, 2(2) :193-380.

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PSYCHE

A JOURNAL OF ENTOMOLOGY

Established in 1874

Vol. 69 December, 1962 No. 4

CONTENTS

A Comparative Study of Oocyte Development in False Ovoviviparous
Cockroaches. Louis M. Roth and Barbara Stay 165

More American Spiders of the Genus Chrysso (Araneae, Theridiidae) .
Herbert W. Levi 209

The Neotropical Species of the Ant Genus Strumigenys Fr. Smith: Synop-
sis and Keys to the Species. W . L. Brown, Jr 238

The Egg-sac in the Identification of Species dC^^atrodectus (Black-
Widow Spiders). J. W. Abalos 268

Author and Subject Index for Volume 69

271

CAMBRIDGE ENTOMOLOGICAL CLUB
Officers for 1962-63

President L. M. Roth, Harvard University

Vice-President A. R. Brady, Idarvard University

Secretary E. G. MacLeod, Harvard University

Treasurer F. M. Carpenter, Harvard University

Executive Committee C. Walcott, Harvard University

A. G. Humes, Boston University
EDITORIAL BOARD OF PSYCHE
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W. L. Brown, Jr., Assistant Professor of Entomology, Cornell
University ; Associate in Entomology , Museum of Comparative
Zoology

E. 0. Wilson, Associate Professor of Zoology , Harvard University
H. W. Levi, Associate Curator of Arachnology, Museum of Com-
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PSYCHE

Vol. 69

December, 1962

No. 4

A COMPARATIVE STUDY OF
OOCYTE DEVELOPMENT IN FALSE
OVOVIVIPAROUS COCKROACHES*

By Louis M. Roth and Barbara Stay1
Pioneering Research Laboratories
Quartermaster Research and Engineering Center
Natick, Massachusetts

Recently Engelmann (i960) compared various internal and exter-
nal factors which affect the activity of the corpora allata in Leuco-
phaea maderae (Fabricius) and Diplotera punctata (Eschscholtz) . In
these two species the stimuli resulting from mating, food intake, gesta-
tion, and parturition differed in the degree to which they influenced
production of gonadotropic hormone.

In this paper we report our experiments on control of oocyte devel-
opment in several species of cockroaches that incubate their eggs
internally in a brood sac or uterus. We classify these species as false
ovoviviparous forms because the uterine eggs increase in water content
only (Roth and Willis, 1955) as opposed to false viviparous species,
like Diploptera, in which the embryos take up both water and solids
from the mother (Roth and Willis, 1955a). In both groups the
oviposition behavior is similar. The eggs do not pass directly from the
ovaries into the uterus but are first extruded to the outside of the body
and then retracted into the brood sac (Roth and Willis, 1954, 1958).

Cockroaches that incubate their eggs internally have two birth
products, the egg and nymph (Roth and Willis, 1958). Ovulation
and oviposition refer to the eggs being released from the ovaries,
oriented by the ovipositor, and covered by the ootheca. After the
eggs are in the uterus the females are pregnant (gestation) for a
certain period of time and give birth (parturition) to nymphs.

MATERIALS AND METHODS

Except for one series of experiments on Nauphoeta (see page 174),
all insects were reared on dog chow checkers and maintained at 240

* Manuscript received by the editor August 1, 1962.

Tresent address: Department of Zoology, University of Pennsylvania,
Philadelphia, Pennsylvania.

165

Psyche

[December

1 66

to 250 C. and 50 to 70% relative humidity. Engelmann ( 1957, 1959)
showed that yolk deposition and growth of the oocytes are correlated
with and dependent upon activity of the corpora allata in Leucophaea
and Diploptera and we have used oocyte development as an indicator
of endocrine activity. Measurements were made, with an ocular mic-
rometer, of oocytes that were dissected from ovaries in Ringer’s solu-
tion. Our measurements of the oocytes of Leucophaea are larger than
those reported by Engelmann (i960). This discrepancy is probably
due to the fact that he measured the oocytes after fixation (Engel-
mann, 1957). We measured one large oocyte per female; in establish-
ing the normal ovarian cycle or the sizes of the oocytes at a specific
period a number of females were usually dissected to give some indica-
tion of the extent of variation. Various operations ( allatectomy, nerve
cord severance, etc.) were performed on insects kept under carbon
dioxide anesthesia.

The species reported on in this paper are Pycnoscelus surinamensis
(Linnaeus), Byrsotria fumigata (Guerin), Blaberus craniifer Bur-
meister, Blaberus giganteus (Linnaeus), Nauphoeta cinerea (Olivier),
and Leucophaea maderae. There are two strains of Pycnoscelus suri-
namensis which differ physiologically. The bisexual strain cannot
reproduce parthenogenetically and the parthenogenetic strain females
when mated to males of the bisexual form show a reduction in fertility
and the resulting offspring are all females which reproduce partheno-
genetically (Roth and Willis, 1961). Practically all of the experi-
ments on Pycnoscelus were done on the parthenogenetic strain but a
few were performed on the bisexual form. A similar study on control
of oocyte development in Diploptera and two species of Blattella has
been reported elsewhere (Roth and Stay, 1961, 1962).

RESULTS AND DISCUSSION
Oocyte development in virgin and mated females

Pycnoscelus surinamensis : Biological data for the two strains are
given in table 1. The basal oocytes of the ovarioles of females less
than a day old are large and may already contain yolk. In fact yolk
may be present in the oocytes of some newly-emerged adults indicating
that perhaps gonadotropic hormone had already been released in the
nymphal stage. The ovarian cycle from emergence to the formation
of the second ootheca in the parthenogenetic strain is shown in figure
1. During gestation the oocytes remain small and increase only slight-
ly in length during the development of the eggs in the uterus. Yolk
deposition occurs after parturition and the oocytes increase rapidly in
size.

Table i — Biological Data For Two Strains of Pycnoscelus Surinam ensis

1962]

Roth and Stay — Cockroaches

167

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Psyche

[December

1 68

In the parthenogenetic strain the first ovulation occurs when the
female is about 13 days old whereas the second ovulation takes place
about 16 days after birth of young. This 3 day difference is explained
by the difference in size of the oocytes in the newly-emerged female
and in the female at parturition ; the oocytes are smaller after the
female gives birth and it takes about 3 days to attain the same degree
of development as they are at adult emergence. In Diploptera the
reverse is true and the second preovulation period is 3 days shorter
than the first although, as in Pycnoscelus the growth rate of the
oocytes is about the same during the first and second preovulation
periods. In Diploptera the oocytes at parturition are about the size
of those of a 3-day-old mated female which explains the shorter period
required for ovulation after parturition (Engelmann, 1959).

Pycnoscelus surinamensis

Fig. 1. Ovarian cycle of Pycnoscelus surinamensis (parthenogenetic strain).
Each point on the curve for oocyte development from 0 to 13 days is the
mean of 6 to 13 measurements (N — 134). Each point for the gestation period
from 13 to 68 days represents individual measurements (N = 99; when 2 or
more points were similar for a particular age only one is indicated). The
part of the curve representing the growth of the oocytes after parturition
(birth) is based on 1 to 3 individuals (N — 24) for each point. Vertical bars
— standard errors of mean values.

In the parthenogenetic strain of Pycnoscelus it is obvious that mat-
ing is unnecessary for development of the oocytes. The initial develop-
ment of the oocytes in the bisexual strain is similar to that found in the
parthenogenetic form but differs in that mating slightly stimulates the
growth rate and also is necessary for normal retraction of the ootheca
into the uterus. Mating a parthenogenetic strain female with a male

1962]

Roth and Stay - — Cockroaches

69

of the bisexual strain has no stimulating effect on growth of the
oocytes as indicated by age of the female at ovulation (Roth and
Willis, 1961).

Six parthenogenetic strain females, allatectomized when 1-2 days
old, did not oviposit within a month after the operations. Five of these
females had 2 pairs of corpora allata implanted at 29 to 30 days after
allatectomy. Four produced oothecae in less than 35 days and one
died after 44 days. At 1 1 1 days after allatectomy one female that still
had not ovulated received corpora allata implants and oviposited in
less than 21 days. This strain normally oviposits about 13 days after
emergence (table 1). The delay in oviposition after implanting cor-
pora allata may have been due to the presence of degenerating oocytes
in the ovaries since the oocytes already have yolk one to two days after
emergence (the age at which allatectomy was performed). In Leuco-
phaea, oocytes in resorption inhibit the corpora allata (Engelmann,

1957)-

Table 2 — Effect of mating on oocyte development and
oviposition in Byrsotria fumigata

OBSERVATION

MATED

VIRGIN

Total number observed

63

213

Number oviposited

53 (84%)

102 (48%)

Ootheca retracted normally

46

92

Ootheca incompletely retracted

7

5

Ootheca dropped

O

5

Number failed to oviposit

10 (16%)

hi (52%)

Oocytes large, well developed
or matured but degenerating

and being resorbed

41 2

ON

00

10

Oocytes small, abnormal in

shape, being resorbed

63

354

Oocytes small, normal in
appearance but only

slightly or not at all developed

0

8

1Three of the 4 females had sperm in their spermathecae ; one lacked
sperm.

2These females were 35 to 60 days old when dissected.

3A11 had sperm in their spermathecae.

4Twenty-one of these females were 32 to 60 days old. The other
14 were 11 to 24 days of age but since their oocytes were small and
abnormal they would not have oviposited.

170

Psyche

[December

8.0

76

72

6.8

6.4
6.0

5.6

5.2

4.8

4.4

4.0

3.6

3.2 -

2.8
2.4-

2.0

1.6 -

1.2 j
0.8
0.4 -

o

o

O o ■■

° U * O O

• 8> ° oaoooal ■

®*o ° ° ■ °B

• o o o ■■ ■

, o o

cm o8» o
o o

8 8oo
°o 00

® g • °

° 8

o o o

oo o
oo

o

o c
oo o

o o °o
no

o°o o

0.0

,a8o88°oo °

ro0D ooO
!§8° o°° °

Byrsotria fumigate

o = Oocytes, virgin females

• = Oocytes, mated females

■ = Degenerating oocytes, virgin females

■ ...I

i-l J..I— l-l-l.l. J,

10 15 20 25 3 0 35 40 45 50

AGE OF FEMALE (DAYS)

Fig. 2. Growth of oocytes in virgin and mated females of Byrsotria fumi-
gata. Each point represents one female. Females were mated when 1 to 17
days old.

1962]

Roth and Stay — Cockroaches

171

Byrsotria fumigata: The effects of mating on oocyte development
and oviposition in Byrsotria are shown in table 2 and figure 2. About
50 percent of the virgin females failed to oviposit but of these 1 1 1
females 61% had large well-developed oocytes that were degenerating
or were being resorbed (fig. 14E). It is evident from figure 2 that
after the thirty-fifth day of age the oocytes of many virgins degenerate
although most of them may reach a length of 5 mm. or more. About
16% of the virgins and about 10% of the mated females had small
abnormally-shaped oocytes that were being resorbed. It is unlikely
that lack of hormone is responsible for this type of abnormality since
Barth (personal communication) has dissected pheromone-producing
Byrsotria females which had small degenerating oocytes but accessory
glands filled with secretion.

In those females that mate, copulation has little, if any, effect on
the growth rate of the oocytes (fig. 2). Mated females oviposited at
26 to 41 days of age (x— 32.4+0.4 days; N;=z53) ; virgin females
oviposited 26 to 44 days after emergence (x— 34.3 ±0.4 days; N =
121). That there is little effect on the rate of growth resulting from
mating is further borne out by the fact that the females oviposit at
about the same age regardless of their age when mated. In our series
the females were with males continuously until they mated; copula-
tion occurred from 4 to 25 days after female emergence. The oocytes
may vary considerably in size in females between these age limits
(fig. 2). A female with large oocytes mated when 25 days old may
ovulate 10 days later whereas one with small oocytes mated at 4 days
of age may take 30 days to ovulate (fig. 3). This is quite different
from the effect of mating in Leucophaea (Engelmann, i960) where
the average interval between mating and oviposition is about the same
regardless of the age of the female when mated (fig. 3) because the
females tend to mate more readily when their oocytes reach a certain
size (see below). Barth (1961) found that Byrsotria females begin
to produce sex pheromone 10 to 30 days after the imaginal molt;
however, recently (1962) he has found that some females may mate
as early as 4 days after adult emergence.

It seems that in Byrsotria mating (perhaps the presence of sperm
in the spermathecae) serves as a stimulus to oviposition. This is
indicated by the fact that the oocytes in many virgin females apparent-
ly mature yet ovulation does not occur. The oocytes in virgin females
at ovulation vary in length from 5.90 to 7.60 mm. (x=6.79±o.o6 ;
N = 7). Although the mean ages at ovulation of mated and virgin
females are very similar a breakdown of the data (fig. 4) shows that

172

Psyche

[December

AGE WHEN MATED (DAYS)

Fig. 3. Relationship between age at mating and ovulation (as indicated by
oviposition) in 4 species of cockroaches. Each point represents one female.
(N — 356 for Leucophaea) .

38% of the virgins oviposited after the thirty-fifth day as compared
to 13% of mated females.

Fifteen virgin females allatectomized when 1 to 2 days old did not
produce oothecae within more than 50 days. At 52 to 210 days after
allatectomy, corpora allata were implanted ; 9 females ovulated in less
than 82 days and one oviposited in 128 days; 3 died without oviposit-
ing and two dissected after 150 days had small undeveloped oocytes.
One allatectomized female that oviposited after receiving corpora
allata implants had well developed oocytes although the ootheca was
in the uterus (fig. 14D). Of 25 sham operated females 16 oviposited
in less than 56 days. We don’t know how to account for the delay in
ovulation after implantation of corpora allata into allatectomized
females. Four pregnant females (i.e. with an ootheca in the brood
sac) had corpora allata implanted and were dissected at 35 to 41 days
of pregnancy. Their oocytes measured 4.90 to 6.81 mm. in length

PERCENT OVIPOSITING

1962]

Roth and Stay — Cockroaches

173

indicating renewed growth of the ovarian eggs as a result of the
implants.

Nauphoeta cinerea: The oocytes of virgins develop but unless
mating occurs the oocytes in many females may degenerate before they
reach ovulation size (fig. 5). Virgin females that ovulate do so in 31
to 47 days (x— 3 5. 8 ±1.2 days; N=i7). Mating results in stirnu-

Fig. 4. Effect of mating on oviposition. Solid circles — virgin females.
Open circles — mated females. The curves for Byrsotria are based on 53
mated and 121 virgin females. The curves for Rlaberus are based on IS
mated and 36 virgin females.

74

Psyche

[December

lation of the corpora allata so that the oocytes develop rapidly (fig. 5)
and oviposition occurs in 18 to 21 days (x= 1 8.9=1=0.40 ; N=8).
Copulation is completed in 17 to 30 minutes (x=20.4±0.8i min. ;
N=i7). Of 22 females kept with males continuously, 19 mated 5
days after emergence; the other 3 mated after 6, 8, and 10 days respec-
tively.

44 r
4.0;

3.6 7

3.2 7

2.87

2.4;
2D 7

1.6 7

1.2 7

0.8 o°

0.4;

0.0 p-

0 = Oocytes, virgin females

• = Oocytes, mated females

■ = Degenerating oocytes, virgin females

o

o 00
o

■ I

a. b I

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o o

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o

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1 1 1 1 1 1

Nauphoeta cinerea

I 1 1 1 . I

5 10 15 20 25 30 35 40 45 50 55

AGE OF FEMALE (DAYS)

Fig. 5. Growth of oocytes in mated and virgin females of Nauphoeta
cinerea. Each point represents one female. Except for 2 individuals mated
at 8 and 10 days of age, all others were mated when 5 days old.

In a series of experiments performed at Harvard University, tem-
perature was uncontrolled but usually higher than 240 to 25 0 C. ; the
insects were maintained on Purina Laboratory Chow. Both virgin
and mated females oviposited earlier than in the above experiment but
virgin females still oviposited later (24 to 35 days) than mated indi-

1962]

Roth and Stay — Cockroaches

175

Fig. 6. Effect of mating, starvation, and combined starvation and mating
on oocyte development in N auphoeta cinerea. The points are mean values;
fed virgins, N = 707; starved virgins, N = 133; fed mated, N — 58; starved
mated, N — 21. Females were mated when 4 to 6 days old. Vertical bars are
one standard error (only positive halves of standard errors are indicated
wherever errors overlapped) ; no vertical bars indicate standard errors of
±0.02 mm. or less.

176

Psyche

[December

viduals (15 to 18 days). The difference in rate of oocyte development
in virgin and mated females is shown in figure 6. The oocytes of
starved females that have mated develop at the same rate as fed mated
females (fig. 6).

Of 17 virgin females that oviposited, 10 retracted the ootheca com-
pletely into the uterus; several of these females aborted the egg cases
several days after oviposition. Four females partly retracted the egg
cases so that some of the eggs remained protruding from the abdomen ;
three dropped the egg cases while or after they were formed without
retracting them. In most virgin females (including those that retract
the ootheca normally) some mature oocytes remain in the ovaries.

Fig. 7. Ovarian cycle (first and second ovipositions) in Leucophaea
maderae. The points for the oocytes of unmated females (open circles) are
means of 5 to 21 individuals (N = 349). For the first preoviposition period,
females were mated (solid circles) when 16 days old and each point repre-
sents 1 individual. Each point for the gestation and post parturition periods
represents one individual.

Leucophaea maderae: At emergence the oocytes of Leucophaea are
o.97d=o.oi mm. (N = io). Mature oocytes at ovulation are 5-56±
0.12 mm. (N=20: 10 mated and 10 virgin females). The new basal
oocyte at oviposition is o.66±o.oi mm. (N=n) and at parturition
i.05d=0.0i mm. (N=4). Under our conditions gestation lasted 91.8

OOCYTE LENGTH (MM)

1962]

Roth and Stay — Cockroaches

177

±0.7 days (N=:35) and the second oviposition occurred 27.8+0.3
days (N = io) after parturition.

The ovarian cycle in this species is shown in figure 7. Mating
shortens the egg maturation period so that the female ovulates at a
more or less definite time (fig. 3) after copulation (Engelmann,
i960). It is almost impossible to predict what the extent of oocyte
development would be in virgins of known age (fig. 8). Only 25 of
381 mated females failed to oviposit. These were 16 to 40 days old
when mated and were dissected 30 to 62 days later. Eighteen had
large degenerating oocytes; 5 had small (1.01 to 1.73 mm.) and 2
had large but normal appearing oocytes. As Engelmann found mating
results in the rapid growth of the oocytes (fig. 7). Of the large
number of virgin females dissected (fig. 8) only 2 had oocytes that
were degenerating.

6.4 r

60 r

5 .6 r
5.2 7
4.8 7
4.4 j-

Leucophaea maderae

00 o

.0 oc

00 8 0

o

4.0 7

3.6 r
3.2 j-
2 .8 r
2.4 7

2.0 7

1.6 F-

0.8P

o o

0.47

o

o = Oocytes, virgin females
• = Oocytes, mated females

0.0„

10 15 20 25 30 35 40 45 50 55 60 65 70 75

AGE OF FEMALE (DAYS)

80 85

Fig. 8. Growth of oocytes in mated and virgin females of Leucophaea
maderae. Each point represents one female. Females were mated when 17
to 23 days old.

178

Psyche

[December

Of 47 virgin females that oviposited, only 1 1 retracted the ootheca
into the uterus; the others dropped the egg cases while they were being
formed. Virgin females frequently retain mature oocytes in their
ovaries and the egg cases are incomplete (Engelmann, 1957a).

Engelmann (i960) found that when females of Leucophaea had
constant access to males, mating occurred when their oocytes averaged
1 .08 ±0.0 1 mm. and none mated that had oocytes exceeding a size of
1.46 mm. He concluded (i960, 1960a) that the corpus allatum hor-
mone must be present in low titer for mating to occur, and as soon as
a certain titer is surpassed, the females did not mate even with ready
access to males. We exposed females of various ages to males for
relatively brief periods (the longest time females were with males was
2 days), and measured the oocytes of those that did and did not mate.
One hundred and fifteen females between 14 and 52 days of age were
mated and their oocytes varied in size as follows:

1.08 mm. to 1.95 mm. (x= 1.43 ±0.02, N = 83 )

2.00 mm. to 2.97 mm. (x=2. 34+0.06, N=2i)

3.1 1 mm. to 3.72 mm. (x=3.30±o.o8, N= 9)

4.90 mm. to 5.88 mm. (x=5. 39^0.49, N= 2)

A breakdown of the data into two age groups when mated was as
follows :

Size of oocytes (mm.)
when mated

Age (days) when mated and
number mated

14-25

26-52

1.08 - 1.95

50

33

2.00 - 2.97

8

13

3- 11 -3.72

0

9

4.90 - 5.88

0

2

As pointed out earlier our measurements are larger than Engel-
mann’s because in our experiments the oocytes were dissected and
measured in Ringer’s solution whereas he measured fixed oocytes. The
majority of the females mated when their oocytes averaged i.43±0.02
mm. This value probably corresponds to Engelmann’s mean of 1.08
±0.01 mm. However, 28% of the females mated when their oocytes
were more developed. Thirty-six females that failed to mate when
exposed to males along with the above females that copulated had
oocytes that ranged from 1.01 to 1.68 mm. (x= i.i9±0.02 mm.;
N = 25) and 2.05 to 5.88 mm. (x=4. 54^=0.38 mm.; N=n).

There was a slight but not very significant shortening of the inter-

1962]

Roth and Stay — Cockroaches

179

val between age at mating and age at ovulation when older females
mated (Engelmann, i960). Engelmann suggested that this shorten-
ing of the period needed for egg maturation could be explained by the
presence of larger amounts of reserve substances that would allow for
more rapid growth of the eggs and might not be due to the presence
of larger oocytes at the later mating. Our findings confirm Engel-
mann’s in that Leucophaea tend to mate more readily when their
oocytes reach a certain size. However, some females mate even though
their oocytes have grown beyond this critical size and the shortening
of the period between mating and ovulation is undoubtedly due to the
presence of large oocytes in these older females ; some females mate
even when there is a high titer of corpus allatum hormone (as indi-
cated by large oocytes).

Engelmann (i960) found that when the nerve cord of Leucophaea
was severed o to 2 days after mating, oocyte maturation occurred
about a week later than normal mated females. When the cord was
severed 3 to 19 days after mating, the females oviposited at the same
age as normal mated females indicating that an intact nerve cord is
necessary for at least 2 days after mating for the mating stimulus to
be effective. When the nerve cords of virgin females were severed
and they were not mated, ovulation occurred at the same time as
females that had their nerve cords severed O to 2 days after mating.
Engelmann concluded that severance of the ventral nerve cord in
virgins either stimulates the corpora allata or cuts off an inhibitory
center for the corpora allata but he favored the latter hypothesis.

We severed the nerve cords of females prior to mating them and
found that in most cases the spermatophore was not inserted properly.
Of 27 females that mated after their nerve cords were severed, only
8 had spermatophores that were apparently transferred by the male
normally. Four females had spermatophores that were visible in the
genital region but they had not been inserted properly in the bursa.
In one mating the spermatophore was dropped by the male without
being transferred to the female. Fourteen females had no spermato-
phores after mating and originally it was believed that none had been
transferred by the male. However, it was discovered that in some
females the male pierced the wall of the uterus and inserted the sper-
matophore in the body cavity near the right ovary (fig. 13C). This
was found in 7 females but may have occurred in 6 others that appar-
ently had no spermatophore inserted but were not dissected because
we did not realize that the spermatophore could be inserted into the
body cavity. One female had no spermatophore after mating, based
on dissection. It seems that the female takes an active role in the

= Oocytes, mated females o = Oocytes, virgin females ■ = Degenerating oocytes, virgin females

180

Psyche

[December

AGE OF FEMALE (DAYS)

Fig. 9. Oocyte development in mated and virgin females of Blaherus.
Each point represents one individual. Females of B. craniifer were mated
when 1 to 16 days old. Females of B. giganteus were mated at 19 to 35 days
of age and all were ovipositing when their oocytes were measured.

1962]

Roth and Stay — Cockroaches

1 8 1

proper positioning of the spermatophore in the bursa copulatrix, and
an intact nerve cord is needed for proper muscular movements of the
female genitalia. Of 1 1 nerve-cord-severed virgin females that ovi-
posited 7 dropped their egg cases when they were formed and 4 re-
tracted the ootheca into the uterus but aborted some time later.

Blaberus craniifer: The growth of the oocytes of virgins is rapid
but mating affords sufficient additional stimulation (fig. 9) so that
ovulation occurs about a week earlier than in unmated individuals
(fig. 4). Eighteen mated females oviposited in 19 to 29 days (x=
23.9±0.6 days) ; sixteen oviposited normally, 1 dropped its ootheca
and 1 failed to retract the egg case completely. Virgin females ovi-
posited in 18 to 41 days (x=32.0d=o.8 ; N = 36). Stimulation from
mating results in either an additional production of gonadotropic hor-
mone or it may possibly serve as an oviposition stimulus. The relation-
ship of age at mating and age at oviposition of the female is similar to
that found in Byrsotria (fig. 3). The older the female when mated
the shorter the interval to ovulation indicating that the oocytes of
these older females are large when mating occurs.

Of 40 virgin females that oviposited 23 (58%) failed to retract the
egg case completely and some of the eggs protruded beyond the abdo-
men. This may be due to the fact that in some females the eggs are
not aligned properly in the ootheca and may even be arranged in 3
rows (rather than 2) which may make it difficult to retract the egg
case completely into the uterus. Generally, in most virgin females that
ovulate, the proper amount of colleterial gland secretion does not flow
out over the eggs since the accessory glands are usually quite full even
after the egg case is formed. Sometimes not all of the eggs are laid
and mature oocytes remain in the ovaries and are eventually resorbed.
Thirteen females retracted the ootheca normally into the uterus. Four
females dropped the ootheca although some eggs remained in the
uterus. Perhaps this is related to the lack of proper amount of col-
leterial gland secretion being poured out around the eggs; the result
may be the formation of a weak ootheca which cannot support the
weight of the eggs as they are extruded some distance beyond the end
of the abdomen prior to their being retracted. In addition to the above
40 females, 5 unmated females that did not oviposit in 46 to 51 days
had oocytes that had obviously matured (based on size) but were
degenerating.

Six allatectomized virgin females that had not ovulated had cor-
pora allata implanted at 62 to 82 days of age; all 6 oviposited within
31 days after implantation. Nine allatectomized virgin females kept
for 66 to 238 days failed to ovulate.

182

Psyche

[December

All the nerves to the corpora allata were severed in 1 1 virgin
females and the glands were left in the animals; ten oviposited in
21.8zhO.49 days which is similar to ovulation in mated females. This
would indicate that the brain tends to inhibit the corpora allata in
virgin B. craniifer and mating overcomes this inhibition.

Blaberus giganteus: The oocytes of virgins grow and yolk is de-
posited but after about a month they may degenerate unless mating
occurs (fig. 9). In general mating appears to be necessary for comple-
tion of oocyte development, at least more so than in B. craniifer.
Fourteen females kept with males until mating occurred, mated at
8 to 35 days of age and oviposited when 35 to 51 days old (x=42.6
±1.3 days). Of 8 virgin females, not shown in figure 8, kept for 51
to 68 days, only 2 oviposited when 51 days old, and in both individuals
the oothecae were dropped and not retracted ; the 6 females that did
not oviposit had small abnormally shaped oocytes that failed to de-
velop.

The relationship between age when mated and age at ovulation
(fig. 3) appears to be similar to Byrsotria and B. craniifer rather than
Leucophaea. The females of B. giganteus which have continuous
access to males, mate over a rather wide age range, and their oocytes
may vary considerably in size at the time of mating.

Thirteen of 14 virgin females that had all the nerves to the corpora
allata severed at O to 19 days of age ovulated in 35.1 ±1.2 days after
the operations ; one oviposited 153 days after the operation at 163 days
of age. Severing the connectives to the corpora allata apparently
removed the inhibition from the brain.

The effects of mating vary in degree among the species of cock-
roaches that incubate their eggs internally or carry them externally
during the incubation period. In the summary given below, data from
Engelmann, (1957, 1959, i960), Roth and Willis (1961) Roth and
Stay (1961, 1962), and the present study have been used.

I. Effect of mating on oocyte development.

1 . Oocytes of virgins may degenerate :

a. before reaching ovulation size (N auphoeta cinerea and
Blaberus giganteus)

b. before or after reaching ovulation size (Byrsotria fumi-
gata)

Mating prevents degeneration of the oocytes in the above 3
species. The oocytes of virgins generally do not degenerate in
Blaberus craniifer (rarely), Diploptera punctata Leucophaea
maderae, Pycnoscelus surinamensis (parthenogenetic and bi-
sexual strains), Blattella germanica , and Blattella vaga.

1962]

Roth and Stay — Cockroaches

183

2. Mating increases rate of oocyte development so that the first
preoviposition period is less than in virgin females. Preovi-
position period shortened on an average of about:

a. 1 day (Pycnoscelus surinamensis, bisexual strain; Blattel-
la).

b. 9 or more days (Blaberus craniifer, Blaberus giganteus) .

c. 17 days (Nauphoeta cinerea).

d. 30 or more days (Leucophaea ?naderae).

e. Majority of virgins do not oviposit for months or not at
all. Oviposition occurs about 10 days after mating ( Di-
pl optera punctata).

3. Mating apparently has little effect on the rate of oocyte
development but may stimulate oviposition (Byrsotria fumi-
gata).

4. Mating has no effect on rate of oocyte development or on the
length of the preoviposition period (Pycnoscelus surinamensis
— parthenogenetic strain mated to males of the bisexual
form ) .

II. Effect of mating on ovulation and oviposition.

1. Ovulating virgins frequently retain mature oocytes in some
part of the reproductive tract so that not all of the eggs are
laid (Blaberus craniifer, Blattella vaga, Byrstotria fumigata,
Leucophaea maderae , Nauphoeta cinerea, Pynoscelus surina-
mensis bisexual strain).

Mated females usually oviposit all of the mature oocytes.

2. Ootheca is incompletely formed and oviposition is abnormal
in a large percentage of virgins.

a. Ootheca usually dropped when formed (Leucophaea ma-
derae, Pycnoscelus surinamensis [bisexual strain], Blattel-
la vaga).

b. Ootheca dropped or partly retracted into the uterus (Nau-
phoeta cinerea).

c. Ootheca usually only partly retracted into the uterus so
that some of the eggs protrude from the end of the ab-
domen (Blaberus craniifer).

Mating in a large percentage of females results in normal forma-
tion of the ootheca and complete retraction of the egg case into
the uterus in the above species.

3. Ootheca may be retracted normally into the uterus in a high
percentage of virgins (Byrsotria fumigata, Diploptera punc-
tata, and Nauphoeta cinerea).

4. Mating has no effect on normal oviposition ( Pycnoscelus

184

Psyche

[December

surinamensis — parthenogenetic strain mated to males of the
bisexual form).

From the preceding summary one finds two extremes of dependence
upon mating for stimulation of the corpora allata. In Diploptera , the
majority of females require mating for maturation of the oocytes and
its effect is the most striking since ovulation occurs about 10 days
after mating, whereas virgin females may go for months without ovi-
positing or they may never do so (Engelmann, 1959, i960; Roth and
Stay, 1961). At the other extreme is the parthenogenetic strain of
Pycnoscelus surinamensis where mating is unnecessary and the oocytes
mature in virgins about 13 days after emergence. In this species some
newly-emerged females already may have yolk in their oocytes. Be-
tween these two extremes are species which show varying degrees of
dependence on external mating stimuli for overcoming inhibition of
the corpora allata. The oocytes in virgins grow but unless mating
occurs the ovarian eggs do not mature and may degenerate before
reaching ovulation size. This is particularly true in Nauphoeta,
Byrsotria, and Blaberus giganteus and apparently in these species the
corpora allata in many virgin females secrete an insufficient amount of
hormone for the oocytes to mature; and in many of these females the
partly developed oocytes are not maintained but degenerate unless the
corpora allata are stimulated by mating.

Various species show different degrees of dependence on mating for
normal formation and retraction of the ootheca into the uterus. This
is of particular interest, for the ability of virgin females to place the
ootheca in the brood sac is a prerequisite to the evolution of partheno-
genesis in false ovoviviparous cockroaches (Roth and Willis, 1961).
Not all females of a species behave similarly which explains why some
forms are included in more than one category in the above summary.
It is this variation in behavior which may make possible the evolution
of parthenogenesis in bisexual species of cockroaches. From the few
species studied one can arrange the forms in a series to show the gradu-
al evolution of retraction of the ootheca into the uterus in virgin
females, although we do not imply that one gave rise to the other.
Almost invariably in the bisexual strain of Pycnoscelus surinamensis
the ootheca is dropped at formation in virgins. In Nauphoeta the
ootheca is dropped at formation, partly retracted, or completely re-
tracted. In Blaberus craniifer the ootheca is usually only partly
retracted into the uterus. In Byrsotria the ootheca of virgins that
ovulate is usually normally retracted into the brood sac. Although
parthenogenesis is uncommon in false ovoviviparous cockroaches
(other than the parthenogenetic strain of Pycnoscelus) it does occur

1962]

Roth and Stay - — Cockroaches

i B s

rarely. Nine females of Nauphoeta cinerea had eggs that developed
parthenogenetically and in 8 individuals the eggs hatched; two un-
mated females that developed from unfertilized eggs gave birth to 3
nymphs (Roth and Willis, 1956). We have encountered only one
case of parthenogenesis in Leucophaea (20 undeveloped eggs and 5
well developed embryos with pigmented eyes in an ootheca 89 days
after ovulation) and one in Byrsotria (2 well developed embryos, 55
days after oviposition) ; Barth (personal communication) has reared
a single adult female of Byrsotria that was produced parthenogenetic-
ally. Only one unmated female of the bisexual strain of Pycnoscelus
was found that had a developed embryo in one of the eggs of the
ootheca (Roth and Willis, 1961). Parthenogenesis in false ovovivi-
parous cockroaches depends upon ( 1 ) the ability of virgin females to
mature their oocytes, ovulate, and form and retract the ootheca into
the uterus, and (2) the capacity for unfertilized eggs to develop.
Although parthenogenesis cannot occur unless the above requirements
are met, the insects must first be capable of retracting the ootheca into
the uterus for unless this occurs the eggs desiccate since the ootheca
does not prevent water loss in cockroaches that incubate their eggs
internally (Roth and Willis, 1955).

No experiments were performed on the species, in this study, to
determine the mechanism of stimulation during mating. However, in
Leucophaea (Engelmann, i960) and Diploptera (Engelmann, 1959,
i960; Roth and Stay, 1961) it is a mechanical stimulus that activates
the corpora allata and it is probably similar in Pycnoscelus (bisexual
strain), Nauphoeta , and Blaberus.

Food intake and maturation of the oocytes

Food intake stimulates maturation of the oocytes in Leucophaea
(Scharrer, 1946; Johansson, 1955; Engelmann, i960) and Blattella
germanica and B. vaga (Roth and Stay, 1962) but is unnecessary
for oocyte development in Diploptera (Engelmann, i960; Roth and
Stay, 1961 ) . The effect of starvation on oocyte development in several
species used in this study was determined ; all females were isolated
from food at emergence.

Pycnoscelus surinamensis : Fifteen females of the parthenogenetic
strain were starved without water. All oviposited in 1 4.1 ±0.4 days,
which was about 1 day more than in unstarved females (table 1).
Nine virgin females of the bisexual strain starved without water ovi-
posited in 1 4.3 ±0.5 days, which was similar to unstarved individuals
(cf. table 1). Food is unnecessary in both strains to activate the cor-
pora allata or for maturation of the eggs for the first ovulation.

1 86

Psyche

[December

Nauphoeta cinerea: The effect of starvation on oocyte development
in Nauphoeta is shown in figure 6. The oocytes of starved virgin
females develop more slowly and to a lesser degree than those of fed
virgin females. When starved females were mated the oocytes matured
in the same period as fed mated females. In Leucophaea, females that
were starved but mated failed to deposit yolk in the oocytes and Engel-
mann (i960) concluded that the brain properly integrated the differ-
ent afferent stimuli (inhibitory during starvation and activating from
mating) into messages to the corpora allata and the endocrines were
not activated. Nauphoeta differs from Leucophaea in that the oocytes
of starved females become well developed and mating adds sufficient
stimulation to the corpora allata for the oocytes to mature normally
in spite of the absence of food.

Blaberus craniifer: Seven virgin females of Blaherus craniifer were
starved (with water) for 22 to 39 days. In 6 females the oocytes
measured 4.821+10.19 mm.; one female had oocytes that did not de-
velop (1.38 mm.). The oocytes of fed females 22-38 days of age were
5.61 ±0.1 1. Although there may be a slightly slower rate of growth
of the oocytes in starved females, food is unnecessary for initiating
activity of the corpora allata.

Byrsotria fumigata: Twenty-four virgin Byrsotria were starved
(with water) for 20 to 45 days. Six females starved 20 to 24 days
had oocytes 4.22 ±0.3 5 mm. long. Eight starved 29 to 40 days had
oocytes 4.86=1=0.36 mm. in length (several had oocytes that had begun
to degenerate). Two females had small undeveloped oocytes (0.88
=±=0.01 mm.) and 8 had small, round, abnormally shaped oocytes.
Thirty-three virgin females were starved without water for 26 to 50
days. Nine (starved 26 to 43 days) had oocytes 5.01 ±0.30 mm. long.
Twelve had large oocytes that were degenerating. Five females ovi-
posited in 34 to 38 days; four had undeveloped oocytes (1.14=1=0.08
mm.) and 3 had small abnormally shaped oocytes. The oocytes of
virgin females fed for 20 to 24 days were 4.85 =1=0.32 (N=i7) and
for 29 to 40 days, 6.36=1=0.22 (N = 39). Although the oocytes of
starved females may not grow quite as rapidly as unstarved individu-
als, neither food nor water are necessary for growth of the oocytes in
Byrsotria.

The degree to which cockroaches depend upon food intake for stim-
ulation of the corpora allata varies among the species. The forms may
be arranged in a series showing complete dependence to complete
independence upon food for oocyte development. The effects of star-
vation may be summarized as follows:

1. Oocytes do not develop (Leucophaea, Blattella germanica, and
Blattella vaga).

1962]

Roth and Stay — Cockroaches

187

J2. Oocytes develop but at a slower rate and to a lesser degree than
in fed females (N auphoeta) .

3. Oocytes develop at a normal or slightly slower rate than fed
females (Blaberus craniifer , Byrsotria) .

4. Oocytes mature about as rapidly asTed females ( Diploptera ,
Pycnoscelus surinamensis — bisexual and parthenogenetic
strains).

Inhibition of the corpora allata during pregnancy

During the first gestation the basal oocytes, in all of the species in-
vestigated in this study, usually remain undeveloped except for a small
increase in length ; some N auphoeta females may have oocytes con-
taining yolk at parturition. Yolk deposition occurs in these basal
oocytes only after parturition (except in Diploptera and some N au-
phoeta). This has already been pointed out in Pycnoscelus (fig. 1).
Very similar cycles occur in Blattella (Roth and Stay, 1962), Leu-
cophaea (Engelmann, 1957) and Diploptera (Engelmann, 1959;
Roth and Stay, 1961). However in Diploptera the oocytes begin to
show deposition of yolk about 3 days before parturition (Engelmann,
1959; Roth and Stay, 1961). Although complete ovulation cycles are
not given for Blaberus, Byrsotria , and N auphoeta, measurements of
the new oocytes at ovulation, and at parturition show that inhibition
of the corpora allata during gestation also occurs in these forms. In
Leucophaea (Engelmann, 1957, i960), Diploptera (Engelmann,
1959 ; Roth and Stay, 1961), and Pycnoscelus (Roth and Stay, 1959)
removal of the ootheca results in resumption of growth of the oocytes
prematurely, indicating that the ootheca in the uterus, in some man-
ner, inhibits the activity of the corpora allata. The following experi-
ments were performed to investigate the nature of inhibition of the
corpora allata during gestation.

Pycnoscelus surinamensis: The odthecae were removed from 84
females of the parthenogenetic strain, 62 (74%) of which subsequent-
ly ovulated. Of the 22 females that failed to oviposit 25 to 37 days
after the operation, 15 had oocytes that showed essentially no develop-
ment (0.64 to O.79 mm.) and 7 had oocytes with definite yolk deposits
(0.84 to 2.39 mm.) ; one female had oocytes that had apparently
matured but had not been laid and were being resorbed. There is an
inverse relationship between the age of the ootheca at the time it is
removed from the uterus and the time required to ovulate again.
Less time is required to ovulate again, the older the uterine eggs are
when removed (fig. 10). This relationship also has been found in
Blattella and Diploptera (Roth and Stay, 1961, 1962). One

1 88

Psyche

[December

of the factors which might account for this may be that the oocytes
increase in size during gestation so that at the time an older ootheca
is removed the oocytes are larger when again subjected to gonadotropic
hormone. The larger oocytes may contain greater amounts of reserve
substances allowing for a more rapid maturation of the eggs.

Since the period between the first and second ovulations is about
70 days (table 1) it is evident from figure 10, showing the relatively

O

o

30

> • •
• ••

25 j
20 1

Pycnosce/us surinamensis

15 -

10 7
5 1

• •

I 1 1 1 1 I 1 ' 1 1 l 1 1 1 1 l 1 i i 1 I 1 1 1 1 l 1 1 1 1 1 1 1 1 1 I 1 1 1 1 l i 1 1 1 I 1 1 1 1 l 1 1 1 1 1

5 10 15 20 25 30 35 40 45 50 55 60

AGE (DAYS) OF OOTHECA WHEN REMOVED FROM UTERUS

Fig. 10. Relationship between the age of the ootheca at the time it was
removed from the uterus of Pycnoscelus surinamensis (parthenogenetic strain)
and the time required to form a new ootheca. Each point represents one
individual. The points at 53 to 58 days on the x axis, are for females that
gave birth normally; all of the other points are based on females that had
their oothecae removed manually.

rapid development of the oocytes (as indicated by oviposition) after
removal of the ootheca, that the oocytes are inhibited by the presence
of the eggs in the uterus.

Virgin females of the bisexual strain of Pycnoscelus almost invari-
ably fail to retract their oothecae into the uterus (Roth and Willis,
1961 ) . Fourteen virgin females that had dropped their oothecae when
they were formed, oviposited again in 28 to 39 days (x=32-9±i.i
days). The normal interval between the first and second ovulations
in this strain is about 67 days (53 days of gestation plus 14 days
postparturition, table 1 ) and the absence of uterine eggs in the brood
sac resulting from aberrant oviposition hastened the development of

1962]

Roth and Stay — Cockroaches

189

the oocytes. Three mated bisexual form females that failed to retract
their oothecae also formed the second egg case in 30.3 ±2.7 days.
Three mated females that had their oothecae removed 2, 5, and 7 days
after oviposition ovulated again in 32 to 33 days. Both strains of
Pycnoscelus are similar in that the presence of an ootheca in the uterus
inhibits the development of the oocytes.

Leucophaea maderae: The oothecae were removed from 102 preg-
nant females at different stages in pregnancy to determine the time
required for the next ovulation. Forty-three females oviposited during
the period of the experiment and these results are plotted in figure 1 1 .
As in Pycnoscelus the time required to ovulate after removal of the
ootheca varied with the age of the ootheca when it was removed ; the
younger the ootheca the longer it took to mature the oocytes. Of the
remaining females, 45 showed little or no growth of the oocytes; those
whose oothecae were removed O to 1 8 days after ovulation had oocytes
i.o6d=o.03 mm. (N=22), 62 to 82 days later and 23 females whose

AGE (DAYS) OF OOTHECA WHEN REMOVED FROM UTERUS

Fig. 11. Relationship between the age of the ootheca at the time it was
removed from , the uterus of Leucophaea maderae and the time required to
form a new egg case. The points plotted at 82 to 97 days on the x axis are
for females that gave birth normally; all of the other females had their
oothecae removed manually. Each point represents one female.

190

Psyche

[December

oothecae were removed 23 to 76 days after oviposition had oocytes
1. 1 9 ±0.04 mm., 34 to 63 days later. Fourteen females that had their
oothecae removed O to 77 days after ovulation had developed oocytes
3.72dbo.30 mm. long, 34 to 71 days later. It is unknown why about
45% of the females failed to show oocyte development after removal
of the odtheca ; the presence of degenerating oocytes that were not laid
in the first ovulation may account for some of these cases.

Byrsotria fumigata: In Byrsotria gestation lasts from 71 to 82 days
(x— 76.2+11.4; N = 6). The basal oocytes at parturition vary in
length from 1.43 mm. to 1.71 mm. (x= 1. 53+0.04 mm.; N=io).
The second ovulation occurs 21 to 30 days (x— 24.8 ±1.6; N = 5)
after parturition. Oocyte development during pregnancy in mated
females is inhibited and no yolk is deposited until after the young are
born.

Five mated females had their oothecae removed at various periods
during pregnancy. One whose ootheca was removed 27 days after
ovulation oviposited 45 days later. Two females whose oothecae were
removed 28 and 40 days after oviposition had practically mature
oocytes, 5.98 mm. and 6.22 mm. (fig. 14B) respectively, 32 days later.
The oocytes ( 1.23 and 1.29 mm. long) of two females whose oothecae
were removed on the first and thirty-first day of pregnancy failed to
develop when examined after 75 and 32 days. The oothecae of 10
virgin females were also removed with the following results. Three
females whose oothecae were removed 29 to 34 days after ovulation
had mature oocytes that were being resorbed 53 days later. One
female whose ootheca was removed 38 days after oviposition ovulated
again 39 days later. Six females whose oothecae were removed from
1 to 24 days after oviposition failed to develop their oocytes (x=i.53
±0.05 mm.) when examined 35 to 59 days after the operations. In
the mated and virgin females that failed to develop oocytes after
removal of the oothecae, several unlaid degenerating oocytes were
present from the previous ovulation which may account for the results.

Virgin females of Byrsotria that deposit their unfertilized eggs
normally in the brood sac frequently carry these oothecae for a longer
period of time than mated females. When the undeveloped eggs are
finally extruded the ovarian oocytes may be large and contain consid-
erable yolk in spite of the fact that an ootheca was present in the
uterus during the entire “pregnancy” period. Thirteen females that
carried their unfertilized eggs for 71 to 90 days had oocytes 1.5 1±
0.04 mm. which is normal for the size of the oocytes at parturition of
mated females. However, the oocytes of 14 virgins that had carried
their oothecae for 87 to 97 days had oocytes that varied in length from

1962]

Roth and Stay — Cockroaches

191

2.86 to 6.12 mm. (x=4.6odb0.23 mm.). One mated female that
aborted an ootheca with undeveloped eggs after carrying for 79 days
had oocytes 3.72 mm. long. It is apparent that toward the end of the
“gestation” period in virgin females or once the time at which parturi-
tion should normally take place is passed, the inhibition of the corpora
allata (due to the presence of the ootheca in the uterus) breaks down
and these endocrines again secrete the gonadotropic hormone. Eleven
virgin females that aborted their oothecae 91 to 104 days after ovi-
positing, were kept until they ovulated a second time. Five of the
females oviposited in 21 to 30 days (x=25.6d= 1.6) which is the
same as mated females indicating that their oocytes at the time of
aborting were relatively undeveloped. The other 6 virgin females
ovulated in 11 to 18 days (x= 14.8=!= 1 .4) undoubtedly because their
oocytes were already well developed when the egg cases containing
undeveloped eggs were extruded from the uterus.

Blaberus craniifer: The oocytes of this species at emergence are
about 1.39 to 1.44 mm. (N = 2) in length. The mature oocytes are
about 6.12 to 6.37 mm. (N=3) and at oviposition the new basal
oocytes vary from 1.02 to 1.16 mm. (x= 1.09 ±0.03 ; N = 7). At
parturition the oocytes are 1.34 to 1.85 mm. (x= i.56±0.07 ; N =
6). Gestation lasts 73 to 87 days (£=79.2 ±2.4; N = 5). After
birth, a second ovulation occurs in 16 to 27 days (x=22.odz 1.9 ;

N = 5)\

Six virgin females had their oothecae removed on the day of ovi-
position. One oviposited again 47 days later. The others were dis-
sected 44 to 60 days later and all had well-developed oocytes (3.96=11
0.43 mm.). One female whose ootheca was removed 8 days after
ovulation had oocytes 5.88 mm. long, 54 days later. Two females
whose oothecae were removed 73 days after oviposition (i.e. close to
parturition in mated females) had oocytes 3.23 mm. and 3.82 mm.
long, only 10 days later. Removal of the ootheca in B. craniifer results
in renewed development of the oocytes.

The principal evidence for Engelmann’s (1957) hypothesis that a
hormonal factor from uterine eggs inhibits the corpora allata via the
brain was his claim that implantation of uterine eggs into the abdo-
men of females of Leucophaea inhibited oocyte development, and
nerve cord severance of pregnant females only had a slight but tem-
porary effect on growth of the oocytes. However, more recently,
Engelmann (i960) found that severance of the nerve cord in preg-
nant females results in growth of the oocytes indicating that nervous
stimuli may also be responsible for inhibition of the corpora allata
during pregnancy.

192

Psyche

[December

We have repeated these and performed additional experiments on
the following species of cockroaches :

Pycnoscelus surinamensis (parthenogenetic strain) : Some of the
experiments on this species were briefly described elsewhere (Roth and
Stay, 1959). The ootheca was removed from the uterus of each of 10
females 1 to 16 days after oviposition and one-half of each ootheca
was implanted into the body cavity of the donor female. Twenty-three
days after the operation the oocytes ranged in length from 2.12 mm.
to 3.19 mm. (x=2.70±o.io mm.) clearly larger than the oocytes
of females that have been pregnant for 24 to 39 days which vary from
0.59 to 0.66 mm. Implantation of uterine eggs into the abdomens of
females that had their oothecae removed did not prevent subsequent
growth of the oocytes. Two of the 10 females had oocytes that had
practically matured and the oocytes of the remaining 8 females were
approaching maturity (2.97 to 3.36 mm., cf. table 1) and undoubted-
ly would have matured in about the time one would expect ovulation
following removal of the ootheca (cf. fig. 10). One-half of young
oothecae were implanted into the body cavities of 6 females one day
old or less; after 11 days the oocytes were 2.65 to 3.14 mm. (x—
2.91=1=0.10) in length. The oocytes of untreated 11-day old females
averaged 2.93=1=0.06 mm. (N=io). These results show that uterine
eggs when implanted into the abdomen of a recently emerged female
have no effect on the initial development of the oocytes. Nor does
implantation of uterine eggs into the abdomen of a female that had
her ootheca removed inhibit subsequent development of the oocytes.

The oothecae of 20 pregnant females were removed 13 to 25 days
after ovulation and a wax “ootheca” about the size and shape of a
normal ootheca was inserted into the uterus. Examined 20 to 37 days
later all had small oocytes (fig. 13A) similar in size to those found in
females that were pregnant for 36 to 52 days (table 3). However

Explanation of figure 12

Fig. 12A. Pycnoscelus surinamensis (parthenogenetic strain). Ootheca
(upper) and ovaries (lower) of a female that had been pregnant 60 days
and whose nerve cord was severed on the thirty-second day of pregnancy.
When dissected 28 days after the operation, the embryos in the ootheca began
to hatch. The oocytes in the ovary had matured but were being resorbed.
(Note the abundant colleterial gland secretion [arrow]). Vertical line — 3
mm.

Fig. 12B, 12C. Blaherus craniifer. B. Mated female whose nerve cord was
severed 26 days after oviposition. The ootheca (0) containing well developed
embryos (note pigmented eyes) was being aborted 34 days after the operation.
The oocytes (arrow) were 5.88 mm. long. C. Virgin female that had carried
an odtheca with undeveloped eggs for 93 days (well beyond the normal gesta-
tion period). The oocytes were 3.92 mm long. Vertical line — 5 mm.

1962]

Roth and Stay — Cockroaches

193

Table 3 — Effect of inserting a wax “ootheca” into the uterus and
subsequent nerve cord severance on oocyte development in
the parthenogenetic strain of Pycnoscelus surinamensis

194

Psyche

[December

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1Numeral in ( ) — number of days, prior to measuring oocytes, nerve
cord was severed.

1962]

Roth and Stay — Cockroaches

195

Fig. 13A-B. Pycnoscelus surinamensis (parthenogenetic strain) . A. Ovaries
(0) undeveloped as the result of the presence of a wax “ootheca” in the
uterus (u). The ootheca was removed on the thirteenth day of pregnancy,
and replaced with wax. The female was dissected 37 days later. B. Oocytes
which developed in a female that had a wax “ootheca” in its uterus for 37
days. The ootheca was removed 13 days after ovulation and replaced by
wax. Twenty-four days later the nerve cord was severed and the female was
dissected 13 days later.

Fig. 13C. Leucophaca maderae. Dissection of a female that mated after her
nerve cord had been transected. The spermatophore (s) was inserted by the
male into the body cavity near the right ovary (arrow).

196

Psyche

[December

Table 4 — Effect of nerve cord severance on oocyte development in
the parthenogenetic strain of Pycnoscelus surinamensis

DAYS AFTER
OVIPOSITION NERVE
CORD WAS SEVERED

DAYS AFTER
OPERATION
OOCYTES WERE
MEASURED

OOCYTES (MM.)
MEAN ± S.E.

N

<1

24

2 .08 ± 0.07

3

4

25

2.94

1

<1

26

2.57=1=0.30

2

o1

29

3.062

1

<1

31

3.52zbo.i62

2

<1

32

2.94

1

27-32

23-33

Oocytes matured

5

Controls

and degenerating3

(sham operated)

1-4

25

0.67=b0.0i

4

<1

29

0.65 ±0.02

4

1

38

o.66±o.02

5

1Operated on just after the female retracted the ootheca.

2Oocytes matured.

3The uterine eggs of these females were completely developed and
parturition was imminent. The eggs began to hatch (fig. 12A) from
3 of the 5 females after their oothecae were removed from the uterus.

when the nerve cords were severed in five females that had been carry-
ing a wax “ootheca” in the uterus for 24 days, the oocytes were well
developed (fig. 13B) 8 to 13 days later (table 3).

The effect of nerve cord severance in pregnant females on develop-
ment of the oocytes is shown in table 4. The oocytes could mature
(2.97 to 3.36 mm., table 1), in females carrying oothecae once the
nerve cord was severed. The time required for the oocytes to mature
in pregnant nerve-cord-severed females was essentially the same as
that taken by females after their oothecae were manually removed.
When removed at O days a new ootheca was formed in about 28 to 31
days. When removed after 27 to 32 days of pregnancy ovulation
occurred about 22 to 25 days later (fig. 10). The five females that
had their nerve cords severed 27 to 32 days after oviposition all had
mature oocytes that were degenerating or being resorbed 23 to 33 days
later at the time the uterine eggs were ready to hatch (fig. 12A).
Apparently oviposition could or did not occur while an ootheca was

1962]

Roth and Stay — Cockroaches

197

in the brood sac, and the mature oocytes degenerated. In addition to
the fifteen females shown in table 4, two females had their nerve cords
severed prior to ovulation and oviposited normally; 24 and 25 days
later their oocytes had grown considerably and were 2.18 mm. and
2.72 mm. respectively. The nerve cord may be severed at any site
between the second and sixth segments to eliminate the inhibition of
the corpora allata during pregnancy. Two females had their nerve
cords severed between the second and third abdominal segments 4
days after oviposition; 29 days later their oocytes were 2.75=1=0.01
mm. Six females had their cords severed between the third and fourth,
fourth and fifth or fifth and sixth abdominal segments, 4 days after
oviposition; 22 days later their oocytes were 1.8911=0.24 mm. long.
Six pregnant females taken from cultures (histories unknown) had
their nerve cords severed between the fourth and fifth, or fifth and
sixth segments; 20 days later their oocytes measured 2.28=1=0.33 mm.

Unmated females of the parthenogenetic strain oviposited normally
in 98 percent of 248 individuals examined (Roth and Willis, 1961).
Twenty-two females had their nerve cords severed prior to oviposition.
Of these, 15 (68%) ovulated in the normal period of time and de-
posited eggs in the uterus ; 8 oviposited all their eggs and had normal
oothecae but the other 7 had small abnormally shaped oothecae and
from 1 to 12 mature oocytes remained in their ovaries. The remaining
7 of the 22 females operated upon failed to retract the ootheca into the
brood sac; in 4 of these one or more mature oocytes remained in the
ovaries but in the others all the eggs were laid. Of 9 females that
were sham-operated when 1 to 4 days old, all oviposited normally and
no mature oocytes remained in their ovaries. Apparently an intact
nerve cord is necessary for normal deposition of mature oocytes and
for normal formation and retraction of the egg case in some females
of P. surinamensis (parthenogenetic strain). Some center, possibly in
the brain, may be involved in this behavior.

Virgin females of the bisexual strain almost invariably fail to retract
their oothecae into the uterus (99% of 138 females, Roth and Willis,
1961) and drop the incompletely formed ootheca. Thirteen virgin
females had their nerve cords severed when 1 to 9 days old. All ovi-
posited abnormally, which is the typical behavior of virgins of the
bisexual strain; 10 dropped their oothecae and all had mature oocytes
left in their ovaries. The other 3 carried their oothecae extruding
from the abdomen but failed to retract them; 2 had some mature
oocytes left in the ovaries but the third had none. Virgin females of
the bisexual strain with severed nerve cords behaved like unoperated
virgin females in oviposition and deposition of mature oocytes.

198

Psyche

[December

In Blattella pressure on the oothecal chamber by the ootheca ap-
pears to be responsible for the inhibition of the oocytes, the stimulus
being transmitted via the nerve cord (Roth and Stay, 1959, 1962).
When the ootheca is in the uterus of Pycnoscelus the ovipositor
is bent forward and is held in that position by the egg case. This
suggested the possibility that the gonapophyses might be involved in
transmitting nervous stimuli to the brain which then inhibits the
corpora allata. Two experiments on Pycnoscelus were performed to
test this hypothesis. Glass beads (3-3.5 mm. in diameter) were insert-
ed into the vestibule of 7 females 1 to 2 days of age. A small drop of
ferrule cement on the anal segments prevented the beads from being
extruded ; the beads exerted pressure on the ovipositor. The oocytes
were measured at 3 different periods. At 5 days of age they were 1.77
±0.13 mm. (N=4), at 7 days, 2.20 mm. (N=i), and at 13 days,
2.85 ±0.01 mm. (N==2). Normal females at 5, 7, and 13 days of
age had oocytes 2.03 ±0.08 mm. (N=9), 2.28zbo.o8 mm. (N =
15), and 2.94±0.04 mm. (N=6) respectively. The presence of a
bead and the resulting pressure on the ovipositor of recently-emerged
females had essentially no effect on the development of the oocytes.

To determine if release of pressure by the ootheca on the ovipositor
during gestation would result in resumption of oocyte development,
the oothecae of pregnant females were partly extruded, a portion of
the egg case was cut off and the remainder was pushed back into the
uterus. This was done to 8 females 11 to 12 days after oviposition
and their oocytes were measured on the fifty-fourth to fifty-sixth days
of pregnancy when the females gave birth or parturition was immi-
nent. Five operations were successful in that the ovipositors were

Explanation of figure 14

Fig. 14. Reproductive organs of Byrsotria fumigata.

A. Normal mated female sham operated (nerve cord) when pregnant
38 days and dissected after 70 days of pregnancy. The eggs (arrows) in
the ovaries are undeveloped; U = uterus containing ootheca.

B. Mated female whose ootheca was removed 40 days after oviposition
and dissected 32 days later. The eggs in the ovaries have almost matured
(6.22 mm. long).

C. Mated female whose nerve cord was severed at 39 days of pregnancy
and dissected 32 days after the operation (71 days pregnant). The eggs
(arrows) in the ovaries have nearly matured (5.88 mm. long) ; U — uterus
containing ootheca.

D. Virgin female allatectomized at one day of age. After 52 days, cor-
pora allata from two females 9 to 10 days of age were implanted. Oviposition
occurred 28 to 35 days after implantation of corpora allata. The eggs
(arrows) in the ovary are almost full grown (5.88 mm.) although an ootheca
remains in the uterus (U).

E. Ovary of a virgin female 43 days old. The oocytes are large and
degenerating (A-D=:X2; EZZX4).

1962]

Roth and Stay — Cockroaches

199

Table 5 — Effect on oocyte development of various implants into the
abdomens of virgins of Byrsotria fumigata

200

Psyche

[December

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1962]

Roth and Stay — Cockroaches

201

Table 6 — Effect of severing the nerve cord on development of the
oocytes, in Byrsotria females that were carrying oothecae

DAYS AFTER
OVULATION NERVE
CORD WAS SEVERED

DAYS AFTER
OPERATION OOCYTES
WERE MEASURED

OOCYTES (MM.)
MEAN dz S.E.

N

Virgin Females

O1

56-59

3.69±o.39

4

18

27,33

4.03 =b 1. 1 1

2

28

33

5.39

I

O1

43-62

i.34±O.Oi

13

21-27

33

1 .39 dz O.07

4

Controls ( sham
operated)

O1

43-57

i.37±o.02

7

Mated Fe?nales

12, 19

4E 43

6.0ld=0.02

2

27-28

29-32

3.68±o.48

4

30-36

31-32

5. 02 ± 0.40

6

42-44

32

5.50+10.71

3

21-39

30-37

i.39d=o.02

5

Controls (sham
operated)

12, 30, 38

32, 43

1.43 ±0.02

4

xThe nerve cords of these females were severed prior to oviposition
and therefore they may be considered to have had the cords cut when
the female was ovipositing.

freed for the period of the experiment. The oocytes of these females
measured o.78±o.oi mm. In 3 females the remaining portion of the
ootheca in the uterus continued to apply pressure on the ovipisitor and
their oocytes measured 0.7 7 ±0.05 mm. As controls 6 females were
sham operated, i.e. their oothecae were partly extruded and pushed
back, without being cut off, into the uterus 11 to 13 days after ovula-
tion. They all gave birth at 54 to 56 days of age and their oocytes
measured 0.741+10.01 mm. These experiments indicate that relieving
the pressure of the ootheca on the gonapophyses during pregnancy had
no effect on oocyte development.

One mated female of the bisexual strain that had oviposited normal-
ly failed to give birth in the usual period of time (53 days). It was

202

Psyche

[December

dissected after 62 days of pregnancy and the oocytes were 1.56 mm.
long and contained yolk. The uterine eggs were degenerating and
were undeveloped but the oocytes had developed although the egg
case had been in the uterus. This failure of endocrine inhibition
during “pregnancy” was also found in Blaberus and Byrsotria (see
below) .

Byrsotria fumigata: The effect of various implants into the abdomi-
nal cavities of virgins is shown in table 5. Portions of egg cases of
Byrsotria and Leucophaea and entire oothecae of Pycnoscelus failed
to inhibit the development of the oocytes in Byrsotria.

Severance of the nerve cord in pregnant virgin and mated females
resulted in resumption of oocyte development in some females (cf. figs.
14A and C) although an ootheca was in the uterus (table 6). How-
ever, the oocytes developed only in 7 of 24 virgins as compared to 15
of 20 mated individuals. All of the virgins that failed to develop
oocytes had many degenerating oocytes that had not been laid during
the initial ovulation which may account for the negative results in
many of these females. Of the 7 virgin females that developed their
oocytes after nerve cord severance, 5 had no old degenerating oocytes,
one had one old oocyte and the last had several oocytes that had
remained from the previous oviposition. In addition to the 24 virgin
females that had been operated upon after ovulating (table 6), 11
females had their nerve cords severed 7 to 24 days after emergence
and 8 others were sham operated when 5 to 20 days old. These
females failed to oviposit and were dissected 31 to 38 days after the
operations. Of the nerve-cord-severed females 7 had matured degener-
ating oocytes and 4 had small oocytes with some yolk but these had
degenerated. Of the sham operated females 5 had mature degenerated
oocytes, 2 had small degenerated oocytes and 1 had oocytes that failed
to develop. The oocytes in females that had been operated on prior to
oviposition were essentially similar to those found in unoperated virgin
females.

Experiments were performed on several females to determine the
effect of removing the ovipositors or relieving the pressure of the
ootheca on the gonapophyses. The ovipositors were cut off of 9
virgin females 6 to 26 days after oviposition. The oocytes were
measured after the females had carried their oothecae for 75 to 84
days. In 8 females the oocytes measured 1.62 ±0.06 mm. indicating
no growth other than might be expected in unoperated females (1.53
dzO.04 mm. at parturition). One female whose ovipositor was cut
off 13 days after ovulation had oocytes 6.37 mm. after 82 days of
pregnancy. The oothecae of 13 virgin females were partly extruded

1962]

Roth and Stay — Cockroaches

203

manually, part of the egg cases were cut off and the remainder pushed
back into the uterus in an attempt to free the pressure normally exert-
ed on the ovipositor. In 6 successful operations the ovipositors were
freed 9 to 12 days after ovulation, and at 73 to 75 days of “pregnancy”
their oocytes were 1.47 ±0.03 mm. Seven females in which the opera-
tions (7 to 18 days after ovulation) did not free the ovipositors, had
oocytes 1.49=1=0.05 mm. after carrying their oothecae for 71 to 76
days; one female that was unsuccessfully operated upon 8 days after
ovulation had oocytes 3.77 mm. 64 days later. These experiments
indicate that removing the ovipositor or releasing the pressure of the
ootheca on the ovipositor during the gestation period does not influence
the development of the oocytes. The two individuals in which the
oocytes grew may be explained by the fact that inhibition of the cor-
pora allata in some virgins of Byrsotria may break down during gesta-
tion.

Blaberus craniifer: Parts of oothecae (about 5 mm. x 10 mm.) of
B. craniifer were implanted into the abdomens of 9 virgin females less
than 1 to 3 days old (one female had an entire ootheca implanted).
Eight females dissected 15 to 30 days later had well developed oocytes
4.84=1=0.28 mm.; one female dissected 28 days after the implant
showed no growth of oocytes (1.48 mm. long). The oocytes of
unoperated virgins 15 to 30 days old were 5.10=1=0.05 mm. The
length of mature oocytes are about 6.25=1=0.07 mm. (N— 3) ; the new
basal oocyte at ovulation is 1.10=1=0.03 mm. (N = 5). Uterine eggs
implanted into the abdomens of virgin females did not inhibit oocyte
development in B. craniifer.

Six mated females had their nerve cords severed on the twenty-
second to twenty-sixth days of pregnancy and were dissected 34 to 39
days later. Four of these females (operated on the twenty- fifth
to twenty-sixth day of pregnancy) had oocytes 4.09=1=0.84 mm., 34 to
38 days later (fig. 12 B) ; two females operated on the twenty-second
and twenty-third days of pregnancy showed very little oocyte develop-
ment (1.74=1=0.12 mm.), 34 and 39 days later (the oocytes at par-
turition are 1.56=1=0.07 mm. long; N = 6).

Two virgin females of B. craniifer carried their oothecae for 93
and 107 days, which is longer than the normal gestation period (about
79 days) of mated females. When the undeveloped uterine eggs were
extruded the oocytes measured 3.92 mm. and 3.68 mm. respectively
(fig. 12C). Inhibition of the corpora allata in B. craniifer apparently
can break down in the late stage of “pregnancy” in virgin females,
as it does in Byrsotria and in Pycnoscelus.

Blaberus giganteus: Eight pregnant females were taken from cul-

204

Psyche

[December

tures (histories unknown), their nerve cords were severed and their
oocytes were measured on the day they gave birth or aborted their
odthecae. Five females gave birth and 3 extruded odthecae containing
well developed embryos in 19 to 33 days after the operations. In every
female the oocytes grew, as a result of nerve severance, and measured
4. 241+10.33 mm. At ovulation the mature oocyte is 5.86±0.O4 mm.
(N=ii) and the new basal oocyte is o.97±o.oi mm. (N=6).
Normally at parturition the oocytes are 1.67+10.03 mm. long (N =
10). Gestation in this species lasts about 95 to 103 days. Nerve cord
severance at least 33 days before parturition eliminated the inhibition
of the corpora allata resulting from the presence of the egg case in
the uterus.

Leucophaea maderae: The odthecae of 7 females were removed and
part of the egg cases were implanted into the abdomens of the female
donors. Five females which had their egg cases removed and im-
planted 10 to 15 days after oviposition, ovulated 61 to 65 days later.
This is about the time one would expect ovulation after removal of
the ootheca (fig. 11). One female had oocytes 2.18 mm. long 65 days
after an implant (made 10 days after ovulation). One female whose
ootheca was removed and implanted 22 days after ovulation had
oocytes 5.14 mm. long 41 days later. Four females whose odthecae
were removed 14 to 40 days after ovulation and had a wax “ootheca”
inserted into the uterus showed no yolk deposition in the oocytes
(1.05+10.05 mm.) 58 to 65 days later.

The implantation of uterine eggs into the abdomens of females did
not prevent the oocytes from maturing. The results with wax “odthe-
cae” insertions indicate that the corpora allata may be inhibited by
pressure of the ootheca in the uterus.

To determine whether there was a hormonal influence on oocyte
development in Leucophaea, Engelmann (1957) removed the eggs
from the uterus and implanted about one half of the ootheca into the
abdominal cavity. He found that the eggs (in the ootheca) still
affected the corpora allata when they were implanted into the abdo-
men (as they did when in the uterus). To rule out any possible effect
of a mechanical pressure on the abdomen, or the effect of other sub-
stances resulting from decay of tissues ( i.e. decaying implanted uterine
eggs) he implanted paraffin blocks, muscle tissue, or agar blocks of
about the size of half an ootheca after removal of the egg case. These
implants did not inhibit the corpora allata and Engelmann concluded
that the arrest of the corpora allata was not caused by mechanical
pressure. However, it should be pointed out that pressure exerted by
an implant in the abdominal cavity may be quite different from pres-

1962]

Roth and Stay — Cockroaches

205

sure exerted in the uterus by the growing eggs (or by an implant into
the uterus). In his more recent work (i960) Engelmann found that
nerve cord severance did in fact result in renewed growth of the
oocytes in pregnant females and that nervous stimuli are primarily
responsible for inhibition of the corpora allata during pregnancy.
However, he found a statistically significant delay of egg maturation
after severance of the nerve cord, compared with animals from which
egg cases were removed (35.2=1=0.7 versus 39.1 ±1.4 days in animals
operated on 29 to 37 days after ovulation; 64.7=1=1.9 vs. 73.4=1=1.5
days in animals operated on o to 1 day after ovulation). He concluded
that other factors play an important role in inhibiting the corpora
allata during pregnancy. By injecting O.i ml. of clear supernatant
fluid from homogenized uterine eggs every fifth day for 30 days, he
inhibited the corpora allata of Leucophaea. However, the injection of
muscle homogenate resulted in a similar inhibition and Engelmann
suggested that a non-specific substance inhibited the corpora allata
during pregnancy.

Although Engelmann has shown a delay in ovulation in females
that had nerve cords cut compared to females from which oothecae
were removed and has demonstrated that extracts of uterine eggs and
muscle tissue have an inhibitory effect on the corpora allata, he has
not demonstrated that there is a substance normally produced by the
uterine eggs which acts to inhibit the corpora allata. Our experiments
do not corroborate Engelmann’s finding that a substance from uterine
eggs inhibits the corpora allata. We find that removing eggs from the
uterus and implanting them into the abdomen (in Rycnoscelus, Byrso-
tria, Blab eras craniifer and Leucophaea ) removed inhibition of oocyte
-development, i.e. oocytes developed in the ovaries. We also find that
cutting the nerve cord of pregnant females allows the oocytes to
develop in the ovaries of Rycnoscelus , Byrsotria, Blaberus craniifer ,
and B. giganteus; we therefore conclude that the inhibition of the
corpora allata during gestation, in these species at least, is dependent
upon nervous stimuli resulting from the presence of the egg case in
the uterus.

Engelmann (i960) concluded that in Leucophaea the inhibitory
influence of the ootheca may act on the last abdominal ganglion either
by nervous or chemical factors and that there was “no reason to
believe that the presence of an egg case in the brood sac is mechanically
recorded in the brain (Roth and Stay, 1959). The question is still
undecided.” Our conclusions in the 1959 paper were based on studies
of Rycnoscelus surinamensis and Blattella germanica. In the parthen-
ogenetic strain of Rycnoscelus there is no inhibition of corpora allata

206

Psyche

[December

in virgin females prior to the first pregnancy, and severance of the
nerve cord may affect the ability of the female to oviposit but has no
influence on the rate of maturation of the oocytes. There is no inhibi-
tory center in the last abdominal ganglion in this species before the
first oviposition. The insertion of wax into the uterus, after removal
of the ootheca, results in inhibition of the corpora allata, and indi-
cates that a chemical substance from uterine eggs is not necessary for
inhibition of corpora allata in Pycnoscelus. We interpret these results
to mean that pressure from the stretched uterus regulates the secretion
of the corpora allata. As suggested by Engelmann (1962) the inhibi-
tory center may be caudal to the site of the operation and “the brain
may act only as a way station for the transmission of nervous impulses.”

In Rhodnius prolixus the release of brain hormone was triggered by
the distension of the insect’s abdomen following a blood meal. Since
cutting the nerve cord eliminated this effect, Wigglesworth (1934)
inferred that the neurosecretory cells were influenced by nerve impul-
ses arising in abdominal proprioceptors. The two stretch receptors
found in each abdominal segment of Rhodnius adapt scarcely at all
and will continue to discharge as long as the abdomen is stretched
(Van der Kloot, 1961). In all of the false ovoviviparous cockroaches
the uterus becomes greatly distended as the eggs increase in size as a
result of water uptake and growth (Roth and Willis, 1955). It is pos-
sible that inhibition of the oocytes during pregnancy may be due to
pressure on abdominal stretch receptors as in Rhodnius. However, it
is also conceivable that there are mechanoreceptors in the uterus itself.
The present evidence indicates that the ovipositor is not involved in
transmitting the pressure stimulus from the ootheca in the uterus or
genital chamber of Pycnoscelus and Byrsotria ; similarly, the ovipositor
in Blattella germanica is not involved in corpora allata inhibition while
the female carries its egg case (Roth and Stay, 1962).

In Blattella which carries its ootheca externally, and in all cock-
roaches that incubate their eggs internally, the ootheca swells during
embryogenesis, particularly in the latter species (Roth and Willis,
1955, 1 955a, 1958). We (Roth and Stay, 1959, 1961, 1962)
have suggested that during pregnancy inhibition of the corpora allata
is due to nervous stimuli resulting from pressure of the ootheca. The
changing pressure stimulus resulting from the increase in size of the
ootheca would tend to prevent or retard adaptation of the receptors
involved so that the corpora allata are inhibited during the entire
(except in Diploptera and some Nauphoeta) gestation period. How-
ever, in virgins of Blattella germanica (Roth and Stay, 1962)
Blaberus craniifer, Byrsotria fumigata, and Pycnoscelus surinamensis

1962]

Roth and Stay — Cockroaches

207

(mated bisexual strain females whose uterine eggs do not develop)
where the ootheca does not increase markedly in size because the eggs
remain undeveloped, inhibition of the corpora allata, resulting from
the presence of the ootheca, ceases, and consequently the oocytes
develop in spite of the presence of the egg case; it seems that because
of the constant, more or less unchanging pressure stimulus resulting
from an ootheca that is not increasing in size, pressure receptors (or
the central nervous system) become adapted and nervous inhibition
of the corpora allata ceases.

ABSTRACT

The effect of mating on oocyte development and oviposition in
Pycnoscelus Surinam ensis , Byrsotria fumigata , Blaberus craniifer, Bla-
berus gigantms, Nauphoeta cinerea and Leucophaea tnaderaej all
cockroaches that incubate their eggs internally, was investigated. In
Diploptera punctata , the majority of females require mating for
maturation of the oocytes. In Pycnoscelus mating is unnecessary for
egg maturation. Between these two extremes are species which show
varying degrees of dependence on external mating stimuli for over-
coming inhibition or for stimulating corpora allata. Various species
also show different degrees of dependence on mating for normal form-
ation and retraction of the ootheca into the uterus.

The extent to which cockroaches depend upon food intake for stimu-
lation of the corpora allata also varies. The species may be arranged in
a series showing complete dependence to complete independence upon
food for oocyte development.

Experiments to determine the nature of inhibition of the corpora
allata during pregnancy indicate that inhibition is due to nervous
stimuli resulting from pressure of the growing eggs in the uterus.

References Cited

Barth, R. H. Jr.

1961. Hormonal control of sex attractant production in the Cuban
cockroach. Science 133:1598-9.

1962. The endocrine control of mating behavior in the cockroach Byrso-
tria fumigata (Guerin). Gen. and Comp. Endocrinology 2:53-69,

Engelmann, F.

1957. Die Steuerung der Ovarfunktion bei der ovoviviparen Schabe
Leucophaea maderae (Fabr.) Jour. Ins. Physiol. 1:257-78.

1957a. Bau und Funktion des weiblichen Geschlechtsapparates bei der
ovoviviparen Schabe Leucophaea maderae (Fabr.) und einige
Beobachtungen uber die Entwicklung. Biol. Zentr. 76 :722-40.

1959. The control of reproduction in Diploptera punctata (Blattaria).
Biol. Bull. 116:406-19.

208

Psyche

[December

1960. Mechanisms controlling reproduction in two viviparous cock-
roaches (Blattaria). Ann. New York Acad. Sci. 89:516-36.

1960a. Hormonal control of mating behavior in an insect. Experientia
16:69-70.

1962. Further experiments on the regulation of the sexual cycle in
females of Leucophaea maderae (Blattaria). Gen. and Comp.
Endocrinology 2:183-92.

Johansson, A. S.

1955. The relationship between corpora allata and reproductive organs
in starved female Leucophaea maderae (Blattaria). Biol. Bull.
108 :40-4.

Roth, L. M., and Barbara Stay

1959. Control of oocyte development in cockroaches. Science 130:271-2.

1961. Oocyte development in Diploptera punctata (Eschscholtz) (Blat-
taria). Journ. Ins. Physiol. 7:186-202.

1962. Oocyte development in Blattella germanica (Linn.) and Blattella
vaga Hebard (Blattaria). Ann. Ent. Soc. Amer. 5 5:633-42.

Roth, L. M., and E. R. Willis

1954. The reproduction of cockroaches. Smithson. Misc. Coll. 122:1-49.

1955. Water content of cockroach eggs during embryogenesis in relation
to oviposition behavior. Jour. Exp. Zool. 128:489-510.

1955a. Intra-uterine nutrition of the “beetle-roach” Diploptera dytiscoides
(Serv.) during embryogenesis, with notes on its biology in the
laboratory (Blattaria: Diplopteridae) . Psyche 26:55-68.

1956. Parthenogenesis in cockroaches. Ann. Ent. Soc. Amer. 49:195-204.
1958. An analysis of oviparity and viviparity in the Blattaria. Trans.

Amer. Ent. Soc. 83:221-38.

1961. A study of bisexual and parthenogenetic strains of Pycnoscelus
surinamensis (Blattaria: Epilamprinae) . Ann. Ent. Soc. Amer.
54:12-25.

SCHARRER, B.

1946. The relationship between corpora allata and reproductive organs
in adult Leucophaea maderae (Orthoptera) . Endocrinology 3 8 :46-
55.

Van der Kloot, W. G.

1961. Insect metamorphosis and its endocrine control. Amer. Zool. 1 :
3-9.

WlGGLESWORTH, V. B.

1934. The physiology of ecdysis in Rhodnius prolixus (Hemiptera). II.
Factors controlling moulting and ‘metamorphosis’. Quart. Jour.
Micr. Sci. 77:191-222.

MORE AMERICAN SPIDERS OF THE GENUS
CHRYSSO (ARANEAE, THERIDIIDAE)*

By Herbert W. Levi

Museum of Comparative Zoology, Harvard University

In a previous paper (Levi, 1955) I revised the genus Chrysso , which
I thought at the time to be a compact group of closely related species
quite distinct from Theridion , and all coming from America. How-
ever, soon after publishing a description of the genus Arctachaea for
only two species (Levi, 1957), I discovered species intermediate
between the new genus and Chrysso.

Though species bridging genera have been found in various groups
of theridiids, it was a surprise to see the description of Argyria venusta
Yaginuma, 1957 (Figs. 3-5) from Japan, a species close to Chrysso
and having similar habits, but less specialized in genitalic structure.
Soon several other species were examined that are best placed in
Chrysso unless we want numerous monotypic genera. One of these
species is Argyrodes nigra O. P. -Cambridge (Figs. 1,2) from Ceylon
(with genitalia superficially resembling those of Theridion punctospar-
sum Emerton) though it has the palpus still less specialized than does
Chrysso venusta (Yaginuma). Allied to Chrysso nigra (O. P. -Cam-
bridge) is the American Chrysso nigrosternum Keyserling (Fig. 13).
Several of the species now placed in Chrysso have the cymbium modi-
fied [e.g. Chrysso perplexa (Keyserling), Fig. 18], and bridge the
gap to the two species previously placed in Arctachaea. Further, a
male of Chrysso cainbridgei (Petrunkevitch) (Figs. 43-47) bridges
the gap between the latter groups and the species tentatively placed
in Meotipa. Meotipa is a genus of doubtful validity; the male of the
type species remains unknown. The three species placed in Meotipa
differed from Chrysso mainly by their longer legs and by a somewhat
different shape of abdomen.

There is still some doubt that the groups considered in this paper
are monophyletic. Certainly all are related to Theridion, from which
they can be separated by the modified shape of the abdomen and by
the more specialized palpus. The specialization of the palpus, how-
ever, may go in different directions. Most species resemble the T .
frondeum group. A distally truncate cymbium, otherwise rare in
theridiids, is common in Chrysso and probably indicates relationship.
Another unusual character, grooves on the sides of the abdomen, is
found in some of the species here considered to belong to Chrysso and

* Manuscript received by the editor April 18, 1962.

209

210

Psyche

[December

may be further evidence of common origin. In addition, several
species have teeth on the posterior margin of the chelicerae.

I would like to thank Dr. W. J. Gertsch for the loan of collections
from the American Museum of Natural History (AMNH) ; Dr. A.
M. Chickering for large collections from Panama and Jamaica now
in the Museum of Comparative Zoology; for additional collections
Mrs. D. L. Frizzell (Dr. H. Exline) of Rolla, Missouri; Mrs. Friz-
zell and Dr. E. S. Ross for specimens from the California Academy
of Sciences (CAS) ; Dr. O. Kraus of the Senckenberg Museum,
Frankfurt (SMF) ; and Dr. R. V. Chamberlin for a specimen of the
University of Utah collection (UU) ; Dr. E. Dresco, Paris, for
making drawings of Meotipa species; Prof. M. Vachon of the Museum
National d’Histoire Naturelle, Paris (MNHN) ; Prof. G. C. Varley
of the Hope Department of Entomology, Oxford, and Dr. G. O.
Evans, Mr. E. Browning, and Mr. K. Hyatt of the British Museum,
Natural History for their hospitality while visiting their institutions.
A. Riedel and J. Pro'szynski of the Polish Academy of Sciences, W ar-
saw (PAS) ; Dr. L. Brundin, Riksmuseum, Stockholm, Mr. T.
Yaginuma, Osaka, and Dr. J. V. Scorza, Caracas loaned valuable
specimens. A National Science Foundation grant (G-4317) made
possible a trip to Europe to examine types and the National Institute
of Health grant (E-1944) facilitated completion of the study and
publication of the manuscript.

Chrysso O. P. -Cambridge

Chrysso O. P.-Cambridge, 1882, Proc. Zool. Soc. London, p. 429. Type species
by original designation Chrysso alhomaculata O. P.-Cambridge.

Meotipa Simon, 1892, Histoire Naturelles des Araignees, 1: 519. Type species
by original designation Meotipa picturata Simon.

Physcoa Thorell, 1895, Descriptive Catalog of the Spiders of Burma, p. 83.

Type species by original designation Physcoa scintillans Thorell.

Argyria Yaginuma, 1957, Acta Arachnologica, 15: 11. Type species by origin-
al designation Argyria venusta Yaginuma. Homonym of Argyr.a Hueb-
ner, 1818, and Argyria Robineau-Desvoidy, 1863.

Argyroaster Yaginuma, 1958, Acta Arachnologica, 15: 37. New name for
Argyria Yaginuma preoccupied.

Arctachaea Levi, 1958, Psyche, 64: 102. Type species by original designation
Arctachaea pelyx Levi.

Diagnosis. Chrysso differs from Theridion by a hump or posterior
dorsal extension of the abdomen ; from A chaearanea by having a much
more complicated palpus containing a radix; from Argyrodes and
Spintharus by lacking a colulus (Levi and Levi, 1962) .

Comments. 1 he types of Chrysso nigriceps Keyserling and Helvibis
sulcata Keyserling have been examined since publishing my paper in

1962]

Levi — Chrysso

21

1 957. The specimens given those names were correctly identified.

The following species have recently been misplaced in Chrysso:
Chrysso maronica Caporiacco, 1954 (Comm. Pontificia Acad. Sci. 16:
74) is Achaearanea hirta (Taczanowski) . NEW SYNONYMY. C*
pentagona Caporiacco, 1954, ibid., 16: 75 belongs to Achaearanea.

While many species of Chrysso are strikingly colored, the coloration
usually cannot be used for determination or as a key character. As in
most theridiids, it is variable, e.g. Chrysso compressa (Keyserling) has
a striking black sternum and abdomen with a black venter in Brazil,
but the venter is yellow in Peruvian specimens. Uyemura (1957) has
observed rapid color change in Chrysso venusta (Yaginuma) when
disturbed. Probably other species of Chrysso can also change color.

Species of Chrysso are absent from the United States and Canada
except C. albomaculata and C. clementinae (Map) found in the south-
eastern states and the two species previously placed in Arctachaea
found from Wyoming to Alaska. None are known from Chile or
southern Argentina. Most species are found in the hotter parts of
America, where the spider fauna is least known, and our knowledge of
the species and their distribution is fragmentary.

Key to the American species of Chrysso

1 a. Mesal tip of male cymbium with teeth (1957, figs. 11-13), fe-
male chelicerae toothless; Wyoming to Alaska 2

ib. Cymbium without teeth; female chelicerae usually with teeth;

southeastern United States to South America 3

2a. Mesal corner of cymbium projecting (1957, figs. 12, 13) ; female

abdomen higher than long C. pelyx (Levi)

2b. Mesal corner of cymbium rounded (1957, fig. 11); female
abdomen longer than high .... C. nordica (Chamberlin and Ivie)

3a.

3b.

4a.

4b.
5 a.

5b.

6a.

Males 4

Females 25

Thread-shaped portion of embolus originating in center of bulb,
looping towards outside and base before continuing straight

towards distal end (1955, figs. 1-5) 5

Embolus otherwise 1 1

Width of area enclosed by embolus loop almost equals width of

basal portion of embolus (1955, fig. 15) ; Huanuco, Peru

C. niariae Levi

Width of area enclosed by embolus loop at least twice width of

basal portion of embolus thread (1955, figs. 7, 9, 13) 6

Median apophysis with a large thorn ( 1955, fig. 13) ; Amazon,
Peru C. sulcata (Keyserling)

212

Psyche

[December

6b. Median apophysis without thorn 7

7a. A thorn on ectal side of palpal bulb ( 1 955, figs. 7, 9) 8

7b. No thorn on ectal side of bulb 9

8a. Radix with evenly rounded swelling ; ectal thorn large (1955,

fig. 7) ; Panama C. vail crisis Levi

8b. Radix with round boss; ectal thorn small (1955, fig* 9) >* Pana-
ma, Peru C. diplosticha Chamberlin and Ivie

9a. Radix with a swelling; conductor almost lanceolate in ventral
view (1955, fig. 11) ; coloration as in 1955 fig. 12; Panama ....

C. indicifer Chamberlin and Ivie

9b. Radix without swelling; conductor hook-shaped (1955, fig.

1-6) ; coloration otherwise 10

10a. Radix with a mesal fold, (1955, figs. 1-4); abdomen usually

with spots; North Carolina to Brazil

C. albomaculata O. P.-Cambridge

10b. Radix without fold (1955, figs. 5, 6); abdomen without dis-
crete spots; Panama to Argentina C. vexabilis Keyserling

1 1 a. Prominent ventral embolus biforked (Figs. 8, 13) 12

11b. Embolus otherwise 13

12a. Embolus long, reaching base of bulb (Fig. 13) ; southern Brazil

C. nigrosternum Keyserling

12b. Embolus short, covering distal two-thirds of bulb (Fig. 8) ;

Minas Gerais C. gounellti sp. n.

13a. Distal end of cymbium truncate 14

13b. Distal end of cymbium rounded 15

14a. Ventral view as in Fig. 18; southern Barzil

C. per pie xa Keyserling

14b. Ventral view as in Fig. 28; Venezuela C. simoni sp. 11.

15a. With long thread-shaped embolus of even diameter (Fig. 23) ;

Costa Rica to Trinidad C. questona sp. n.

15b. Embolus otherwise 16

16a. Embolus corkscrew-shaped, on distal end of bulb (Figs. 89, 94)

: 17

1 6b. Embolus otherwise 18

17a. Embolus larger than remaining portion of bulb (Fig. 89)
abdomen higher than long; Panama to Ecuador .... C. silva sp. n
17b. Embolus one-third length of remaining portion of bulb (Fig

95) ; abdomen longer than high; Costa Rica, Panama

C. volcanensis sp. n

1 8a. Embolus with distal portion hook-shaped (Fig. 47) ; Mexico to

Venezuela C. cambridgei (Petrunkevitch)

18b. Embolus otherwise 19

1962]

Levi — Chrysso

19a.

19b.

20 a.

20b.

2 1 a.

2ib.

22a.

22b.

23a.

23b.

24a.

24b.

25a.

25b.

26a.

26b.

27a.

27b.

28a.

28b.

29a.

29b.

30a.

30b.

3ia.

3ib.

213

Two transparent sickle-shaped structures on venter of bulb

(Fig. 42) ; Colombia to Bolivia C. vittatula (Roewer)

Palpal bulb without sickle-shaped structures 20

Conductor prominent and with distal hook (Fig. 74) ; cosmo-

tropical C. clementinae ( Petrunkevitch)

Conductor otherwise 2 1

Embolus a ventral bottle-shaped structure (Fig. 69) ; Peru to

Brazil C. compressci ( Keyserling )

Embolus otherwise 22

Median apophysis not apparent in ventral view (Fig. 30) ;

southern Brazil C. drops sp. n.

Median apophysis visible in ventral view 23

Median apophysis in distal half of bulb, palpus lightly sclero-

tized (Fig. 79) ; Panama C. melba sp. n.

Median apophysis mostly on proximal half of bulb; palpus

heavily sclerotized 24

Palpus as in Figure 36, 37; Colombia to Bolivia

C. ecuadorensis Levi

Palpus as in Figure 51 ; Venezuela

C. barro mac had 0 i Caporiacco

Epigynum with openings in coiled borders of fossa (Fig. 78) ;

Costa Rica to Trinidad C. quest ona sp. n.

Epigynum otherwise 26

Ducts showing through epigynal wall long and coiled, openings

indistinct (Figs. 82, 84) 27

Epigynum otherwise 28

Ducts loosely coiled in large loops (Fig. 83) ; Panama

G. ale cula sp. n.

Ducts tightly coiled (Fig. 85) ; Trinidad C. arima sp. n.

Epigynum with a central black mark (Fig. 45) ; sternum, venter
black; Mexico to Venezuela .... C. cambridgei (Petrunkevitch)
Epigynum otherwise; sternum and venter usually not black

(except C. compressa, C. nigrosternum ) 29

Openings of epigynum widely separated (Fig. 88) ; Panama to

Ecuador C. silva sp. n.

Epigynum otherwise 30

Epigynum with a light, oval swelling, bordered all around (Figs.

34, 35, 40) Colombia to Bolivia 31

Epigynum otherwise 32

A dark mark anterior to swelling (Figs. 34, 35)

C. ecuadorensis Lev i

A dark mark posterior to swelling (Fig. 40)

C. vittatula (Roewer)

214

Psyche

[December

32a. Epigynum with a light swelling bordered anterior and sides

(Fig. 50) ; Venezuela C. barromachadoi Caporiacco

32b. Epigynum otherwise 33

33a. Epigynum with openings near posterior margin in a depression,

ducts loop anterior (Fig. 54) ; southern Brazil

C. ribeirao sp. n.

33b. Epigynum otherwise 34

34a. Epigynum with a central fossa 35

34b. Epigynum without fossa 41

35a. Two openings in center of fossa (Fig. 93) ; Costa Rica, Panama

G.volcanensis sp. n.

35b. Epigynum otherwise 36

36a. A black circular mark on each lateral end of fossa (Fig. 73) ;

cosmopolitan C. clementinae (Petrunkevitch)

36b. Epigynum otherwise 37

37a. Fossa far anterior, usually bordered on sides only (Fig. 27) ;

Venezuela C. simoni sp. n.

37b. Fossa in center or posterior, bordered anterior or posterior as

well as sides 38

38a. Fossa in a dark spot; ducts visible posterior to fossa (Figs. 16,

17) ; Brazil, Argentina C. rubrovittata (Keyserling)

38b. Epigynum otherwise 39

39a. Narrow ducts leading from sides into fossa (Figs. 57, 62) .... 40
39b. Ducts wider and leading from anterior into fossa (Figs. 64, 66,

67), Peru to Brazil C. cornpressa (Keyserling)

40a. Fossa bordered on sides and posterior (Fig. 57) ; southern

Brazil C. pulchra (Keyserling)

40b. Fossa bordered on sides and anterior (Fig. 62) ; Minas Gerais ..

C. caraca sp. n.

41a. Epigynal openings in a slit anterior to a dark area (Fig. 60) ;

ducts simple (Fig. 59) ; San Fernandez Isl.

C. backstromi (Berland)

41b. Epigynal openings otherwise; usually with internal sacs 42

42a. Epigynum a central transverse connected dark mark as in Fig-
ure 21 ; Brazil C. ant 0 mo sp. n.

42b. Epigynum otherwise 43

43a. Epigynum with two central dark curved marks (Figs. 10, 12);

southern Brazil C. nigrosternum (Keyserling)

43b. Epigynum otherwise 44

44a. Epigynum with an anterior dark mark and light posterior as in

1 95 5 > hg- 33i head region blackened; Colombia, Ecuador

C. nigriceps Keyserling

44b. Epigynum otherwise, coloration usually otherwise 45

1962]

Levi — Chrysso

2i5

45a.

45b.

46a.

46b.

47a.

47b.
48 a.

48b.

49a.

49b.

50a.

50b.

Epigynum with two lines, diverging posterior (i955> fig- 31)

Panama to Peru C. diplosticha Chamberlin and Ivie

Epigynum otherwise 46

Epigynum with a transverse dark mark posterior to two spots

( 1955, fig. 29) ; Panama C. vail e nsis Levi

Epigynum otherwise 47

Epigynum with a transverse line anterior to two spots (i955>

fig- 37) ,* southern Brazil C. sicki Levi

Epigynum otherwise 48

Epigynum with a pair of longitudinal lines in light area posterior
to transverse dark mark (1955, fig. 35) dorsum of abdomen
with two black long lines which fuse at posterior end ; Panama
C. indicifer Chamberlin and Ivie

Epigynum otherwise 49

Carapace dark brown ; abdomen with a longitudinal dark band

( 1955, fig. 20) ; Peru C. huanuco Levi

Coloration otherwise 5°

Abdomen without discrete white spots; Panama to Argentina

C. vexabilis Keyserling

Abdomen usually with white spots; North Carolina to Brazil
C. albomaculata O. P. -Cambridge

Chrysso albomaculata O. P. -Cambridge

Argyrodes elegans Taczanowski, 1872, Horae Soc. Ent. Rossicae, 9: 118, pi. 5,
fig. 11, $. Female syntypes from Uassa, French Guiana [now Rio Uaga,
Amapa, Brazil] in the Polish Academy of Sciences, Warsaw, examined.
NEW SYNONYMY. Not T heridion elegans Blackwall, 1862.

Chrysso albomaculata O. P. -Cambridge, 1882, Proc. Zool. Soc. London, p. 429,
fig. 6, $, $. Male, female syntypes from the Amazon in the Hope
Department of Entomology, Oxford, examined. — Levi, 1955, Jour. New
York Ent. Soc., 68: 61, figs. 1-4, 18-19, 25-27, 9, $ .

Chrysso elegans, — Keyserling, 1884, Die Spinnen Amerikas, Theridiidae 2
(1): 151.

Theridion elegans, — Simon, 1903, Histoire Naturelle des Araignees, 2: 990.
Theridion emendatum Roewer, 1942, Katalog der Araneae 1 : 429. New name
for Argyrodes elegans Taczanowski. NEW SYNONYMY.

Distribution. Southeastern United States, Mexico, West Indies,
northeastern South America to Bahia, Brazil.

Additional records. Lesser Antilles. Grenada: St. George’s (N. L.
H. Krauss, AMNH). Trinidad: Mt. St. Benedict (J. G. Myers).
Venezuela. Carabobo: San Esteban, 1888 (E. Simon, MNHN).
Grand Elang. Dist. Fed.: hacienda Corosal, N of Mt. Silla, 1888
(E. Simon, MNHN). British Guiana. Georgetown (A. M. Nadler,
AMNH). Surinam. Paramaribo. Brazil. Ceara: Sierra Com-

2l6

Psyche

[December

munaty (E. Gounelle, MNHN). Pernambuco : Recife (SMF; A.
M. Nadler, AMNH). Bahia: Salvador (A. M. Nadler, AMNH) ;
Terra Nova (MNHN).

Chrysso vexabilis Keyserling

Chrysso vexabilis Keyserling, 1884, Die Spinnen Amerikas, Theridiidae, 2(1) :
155, pi. 7, fig. 96, 2, $. Syntypes from N. Granada [now Ecuador,
Colombia, Panama, Venezuela] in the British Museum, Natural History,
examined. — Levi, 1955, Jour. New York Ent. Soc., 63: 63, figs. 5, 6, 23,
24, 2, $.

Steatoda rubicunda F. P.-Cambridge, 1902, Biologia Centrali-Americana,
Araneidea, 2: 386, pi. 36, fig. 21, $. Male type from Bugaba, Panama
in the British Museum, Natural History, examined. (Not Theridion
rubicunda Nicolet, 1849.) NEW SYNONYMY.

Theridion rubicundulum Roewer, 1942, Katalog der Araneae, 1: 498. New
name for T heridion rubicunda (F. P.-Cambridge). NEW SYNONYMY.

Examination of Keyserling’s type indicated that this species had
been correctly determined. The internal female genitalia of Argentine
specimens like the ones from Panama, have the posterior portion of
the sacs of the internal genitalia ending in narrow extensions (Levi,
1 955 > fig- 23) quite different from the broadly rounded sacs of C.
albomaculata. Unlike C. albomaculata it has no white spots on abdo-
men.

Distribution. Panama to Argentina.

Additional records. Venezuela. Dist. Fed.: hacienda Corosal, $,
cf (E. Simon, MNHN) . Ecuador. Guayas: Milagro, $, cf (H. E.,
D. L. Frizzell); Rio Chimbo, cf (Edwards, MNHN). El Oro:
Arenillas (E. L. Moore). Brazil. Bahia: Salvador, 9, (E. Goldi,
MNHN); Rio Salobro, 9 (E. Gounelle, MNHN, doubtful det.).
Argentina. Cordoba: Cosquin, 9-

Chrysso diplosticha Chamberlin and Ivie

Chrysso diplosticha Chamberlin and Ivie, 1936, Bull. Univ. Utah, biol. ser.,
3(5): 36, fig. 83, 2. Female type from Barro Colorado Island, Panama
Canal Zone in the University of Utah Collection. — Levi, 1955, Jour.
New York Ent. Soc., 63 : 65, figs. 9, 30, 31, 2 , $ .

Distribution. Nicaragua, Panama, Trinidad and Dept. Huanuco,
Peru.

Additional records. Nicaragua: Musawas, Waspuc River, cf (B.
Malkin, AMNH). Lesser Antilles. Trinidad: Simla near Arima,
cf (A. M. Nadler, AMNH).

Chrysso indicifer Chamberlin and Ivie

Chrysso indicifer Chamberlin and Ivie, 1936, Bull. Univ. Utah, biol. ser..,
3(5): 36, figs. 82, 96, 2 . Female type from Barro Colorado Island in the

1962]

Levi — Chrysso

217

University of Utah collection. — Levi, 1955, Jour. New York Ent. Soc.,
6 3 : 66, figs. 11, 12, 17, 34, 35, $ , $.

Distribution. Panama to Peru.

Additional records. V enezuela. Dist. Fed. Caracas, (E. Simon,
MNHN). Carabobo: La Cumbre, S. of San Esteban, (E. Simon,
MNHN). Ecuador. Guayas: Milagro, (H. E., D. L. Frizzell);
Rio Chimbo, (MNHN). El Oro: Arenillas, (E. L. Moore) ; Pasaje,
(R. Walls). Peru. Huanuco: Monzon Valley, Tingo Maria, $, cf
(E. I. Schlinger, E. S. Ross, CAS).

Chrysso gounellei sp. n.

Figures 6-8

Type. Male from Caraca, Minas Gerais, Brazil (E. Gounelle),
in the Museum National d’Histoire Naturelle, Paris (no. 9487).
The species is named after E. Gounelle who collected for E. Simon.

Description. Carapace orange, eye region black. Sternum black.
Legs yellowish to gray. Abdomen white with two black spots above
spinnerets (Fig. 7), sometimes also with a pair of anterior spots.
Carapace low and flat. Anterior median eyes larger than others, one
and one-third their diameter apart, three-quarters from laterals.
Posterior median eyes one and one-quarter their diameter apart, one
and one-half from laterals. Chclicerae with three teeth on anterior
margin, the middle one smallest, five sharp teeth posterior (Fig. 6).
Abdomen longer than wide with indistinct grooves on sides (Fig. 7).
Total length 1.9 mm. Carapace 1.0 mm long, 0.8 mm wide. First
femur, 1.1 mm; patella and tibia, 1.1 mm; metatarsus, 0.9 mm; tarsus,
0.5 mm. Second patella and tibia, 0.9 mm; third, 0.6 mm; fourth,
0.9 mm.

Diagnosis. The two prongs of the embolus (Fig. 8) of C. gounellei
are shorter than those of the related C. nigrosternum (Keyserling) .
Record. 6 paratypes collected with holotype.

Chrysso nigrosternum Keyserling
Figures 9-13

Theridium rotundum Keyserling, 1891, Die Spinnen Amerikas, Brasilianische
Spinnen, 3: 191, pi. 6, fig. 137, $. Female type from N. Freiburg [Nova
Friburgo, Est. Guanabara], Brazil in the British Museum, Natural His-
tory, examined. NEW SYNONYMY.

Chrysso nigrosternum Keyserling, 1891, Die Spinnen Amerikas, Brasilianische
Spinnen, 3: 206, pi. 7, figs. 148, $, $. Female, male syntypes from
Taquara do Mundo Novo, [Rio Grande do Sul], Brazil in the British
Museum, Natural History, examined. The female syntype is in poor
condition.

2 I 8

Psyche

[December

Note. The similarity of my illustrations of the female types of the
two names was not noted until some time after the examination*
Keyserling’s descriptions, except for the color of the abdomen, are also
similar. Although rotundum has page priority, the name Chrysso
nigrosternum Keyserling is used here, because the male palpus is very
distinct while the epigyna are not. Figures n, 12 were made from
the type of T. rotundum , Figures 9, 10, 13 from those of C. nigro-
sternum.

Chrysso rubrovittata (Keyserling), new combination
Figures 14-17

Faiditus rubrovittatus Keyserling, 1884, Die Spinnen Amerikas, Theridiidae,
2(1): 159, pi. 7, fig. 98, $. Female type without locality in the Hope
Department of Entomology at Oxford University, examined. It probably
came from South America.

Faiditus lacteovittatus Mello-Leitao, 1945, Rev. Museo La Plata, nueva serie,
sec. Zool., 4: 233, fig. 7. Female type from Puerto Victoria, Missiones,
Argentina in the La Plata Museum, examined. NEW SYNONYMY.

Description. Female from Pernambuco. Carapace, sternum, legs
yellow. Abdomen grayish white with a dorsal, longitudinal white
band and some white pigment spots (Fig. 14). Eyes subequal in size.
Anterior median eyes one and one-half diameters apart, their radius
from laterals. Posterior median eyes one and one-half diameters apart,
their radius from laterals. Chelicerae with two teeth on anterior
margin. Abdomen without lateral grooves (Fig. 14). Epigynum with
central, sclerotized opening having a posterior lip (Figs. 16, 17).
Total length 3.9 mm. Carapace 1.11 mm long, 1.02 mm wide. First
femur 2.10 mm; patella and tibia 1.95 mm; metatarsus 1.62 mm;
tarsus 0.78 mm. Second patella and tibia 1.17 mm; third 0.75 mm;
fourth 1.30 mm.

Explanation of Plate 14

Figs. 1, 2. Chrysso nigra (O. P. -Cambridge) (Ceylon). 1. Left palpus. 2.
Epigynum cleared.

Figs. 3-5. C. venusta (Yaginuma) (Japan). 3. Female genitalia, dorsal
view. 4. Epigynum. 5. Palpus.

Figs. 6-8. C. gounellei sp. n. 6. Male left chelicera, posterior view. 7.
Abdomen, dorsal view. 8. Palpus.

Figs. 9-13. C. nigrosternum Keyserling. 9, 11. Female genitalia, dorsal
view. 10, 12. Epigynum. 13. Palpus.

Figs. 14-17. C. rubrovittata (Keyserling). 14. Female abdomen, lateral
view. 15. Female genitalia, dorsal view. 16, 17. Epigynum.

Fig. 18. C. perplexa Keyserling. Palpus.

Figs. 19-21. C. antonio sp. n. 19. Female abdomen, lateral view. 20. Female
genitalia, dorsal view. 21. Epigynum.

Figs. 22, 23. C. questona sp. n. palpus. 22. Mesal view. 23. Ventral view.

Figs. 24-28. C. simoni sp. n. 24. Female abdomen, lateral view. 25. Female
genitalia, dorsal view. 26. Epigynum cleared. 27. Epigynum. 28. Palpus.

Psyche, 1962

Vol. 69, Plate 14

220

Psyche

[December

Figures 15, 16 were made from the type of Faiditus rubrovittatus.

Distribution. Brazil, Argentina.

Record. Brazil. Pernambuco : Recife, 9 (SMF) ; Minas Gerais:
Caraga, ? (E. Gounelle, MNHN).

Chrysso perplexa Keyserling
Figure 18

Chrysso perplexa Keyserling, 1886, Die Spinnen Amerikas, Theridiidae, 2(2) :
242, pi. 20, fig. 296, $. Male type from Blumenau, [Santa Catarina],
Brazil in the British Museum, Natural History, examined. Figure 18 was
prepared from the type.

Record. Brazil. Santa Catarina: Nova Teutonia, lat 270 1 V S,
long 520 23' W, cf (F. Plaumann, SMF) .

Chrysso antonio sp. n

Figures 19-21

Type. Female from Sao Antonio de Barro [Condeuba, Bahia],
Brazil (E. Gounelle), in the Museum National d’Histoire Naturelle,
Paris (no. 11518). The specific name is a noun in apposition after the
type locality.

Description. Carapace dark brown, dusky on sides. Chelicerae,
sternum black. Legs yellowish. Abdomen with dorsal white patch
and five white spots on sides on gray background a darker, indistinct
gray band behind dorsal white spot continuing to a black posterior
tip (Fig. 19). Eyes subequal in size. Anterior median eyes one diame-
ter apart, their radius from laterals. Posterior eyes their diameter
apart. Chelicerae with two long teeth on anterior margin. Abdomen
without grooves (Fig. 19). Total length 2.5 mm. Carapace 0.85 mm
long, 0.65 mm wide. First femur, 1.20 mm; patella and tibia, 0.98

Explanation of Plate 15

Figs. 29-30. Chrysso arops sp. n. 29. Male abdomen, lateral view. 30. Left
palpus.

Figs. 31-37. C. ecuadorensis Levi. 31, 32. Female abdomen, lateral view.
33. Female genitalia, dorsal view. 34, 35. Epigynum. 36, 37. Palpus. 36. Mesal
view. 37. Ventral view.

Figs. 38-42. C. Fittatula (Roewer). 38. Female abdomen, lateral view.
39. Female genitalia, dorsal view. 40. Epigynum. 41, 42. Palpus. 41. Mesal
view. 42. Ventral view.

Figs. 43-47. C. cambridgei (Petrunkevitch) . 43. Female, lateral view. 44.
Female genitalia, dorsal view. 45. Epigynum. 46. Palpus expanded, mesal
view (cymbium removed; C, conductor; E, embolus; M, median apophysis;
R, radix). 47. Palpus.

Figs. 48-51. C. barromachadoi Caporiacco. 48. Abdomen of juvenile,
lateral view. 49. Female genitalia, dorsal view. 50. Epigynum. 51. Palpus.

Psyche, 1962

Vol. 69, Plate 15

222

Psyche

[December

mm ; metatarsus, 0.78 mm ; tarsus, 0.50 mm. Second patella and tibia,
0.71 mm; third, 0.52 mm; fourth, 0.91 mm.

Diagnosis. The epigynum, which has a dark patch in a light trans-
versely folded area (Fig. 21), separates this species from C. alboma-
culata. The sacs of the internal genitalia (Fig. 20) are more complex
in shape than in related species.

Ghrysso questona sp. n.

Figures 22, 23, 76-78

Type. Female from El Valle, Panama, July 1936 (A. M. Chicker-
ing), in the Museum of Comparative Zoology. The specific name is
an arbitrary combination of letters.

Description. Carapace, sternum, legs whitish. Legs with black
rings on middle and distal ends of segments. Abdomen whitish with
some black patches (Fig. 76). Chelicerae with two anterior teeth.
Anterior median eyes one and one-fourth diameters apart, three-
fourths from laterals. Posterior median eyes slightly more than a diam-
eter apart, one diameter from laterals. Eyes of male smaller and
slightly farther apart. Legs very long. Total length of female 4.5 mm.
Carapace 1.3 mm long, 1.0 mm wide. First femur, 4.9 mm; patella
and tibia, 4.5 mm; metatarsus, 4.9 mm; tarsus 1.4 mm. Second patella
and tibia, 2.3 mm; third, 1.2 mm; fourth, 2.0 mm. Total length of
male 3.0 mm. Carapace, 1.6 mm long. Second patella and tibia, 2.7
mm; third, 1.3 mm; fourth, 2.3 mm.

Diagnosis. The coiled rim of the epigynum (Fig. 78), the coiled
ducts (Fig. 77) and the long embolus and narrow cymbium (Figs. 22,
23) distinguish this species from C. volcanensis.

Records. Costa Rica. San Isidro del General, 600-1200 m, $ (D.
Rounds). Panama: Chiriqui Prov., 1938, $ (UU). Trinidad:
Simla, 29, 30 Dec. 1954, ?; 26 Feb. 1954, $ (A. M. Nadler,
AMNH).

Ghrysso simoni sp. n.

Figures 24-28

Type. Male from Caracas, Venezuela, December 1887 to February
1888 (E. Simon), in the Museum National d’Histoire Naturelle,
Paris (no. 16392). The species is named after the great araneologist
E. Simon.

Description. Carapace yellow-brown, slightly darker in cephalic
region. Sternum, legs yellow-brown. Abdomen gray with variable
white, darker gray or black patches. Some specimens have a median

1962]

Levi — Chrysso

223

dorsal white longitudinal band, in others the band is almost black
(Fig. 24). Eyes subequal in size. Anterior median eyes about one and
two-thirds diameter apart, three-quarters from laterals. Posterior
median eyes less than two diameters apart, one from laterals. Anterior
median eyes of male slightly farther apart than in female. Chelicerae
with two teeth on anterior margin, none on posterior. Abdomen
longer than wide or high, with posterior tip of female overhanging
(Fig. 24) but posterior tip of male more blunt. Total length of female
3.1 mm. Carapace 1.1 mm long, 1.0 mm wide. First femur, 2.2 mm;
patella and tibia, 2.0 mm; metatarsus, 1.9 mm; tarsus, 0.8 mm.
Second patella and tibia 1.3 mm ; third 0.9 mm ; fourth 1.4 mm. Total
length of male 2.5 mm. Carapace 1.1 mm long, 0.9 mm wide. First
femur, 2.2 mm ; patella and tibia, 2.0 mm ; metatarsus, 1.9 mm ; tarsus,
0.8 mm. Second patella and tibia, 1.3 mm; third, 0.8 mm; fourth,
1.3 mm.

Diagnosis. The palpus of the male (Fig. 28) is quite similar to that
of C. perplexa Keyserling but can be separated from the latter by the
shape and length of the embolus, the base of which lies against the
alveolus of the cymbium. The internal female genitalia have a lateral
•duct loop (Figs. 25, 26) unlike that of the related C. rubrovittata
(Keyserling). The internal genitalia are asymetrical in the specimens
examined.

Records. 3 cf and 3 ? paratypes from type collection.

Chrysso barromachadoi Caporiacco
Figures 48-51

Chrysso barromachadoi Caporiacco, 1955, Acta Zool. Venezuelica, 1 : 337.
Juvenile type from Rancho Grande, Aragua, Venezuela, in the Museum
at Caracas, examined.

Description. Carapace, sternum dark brown. Legs yellow. Abdo-
men gray to black with white patches on sides of female (Fig. 48)
but lacking in male. Anterior median eyes slightly smaller than others,
one and one-half diameters apart, two from laterals. Posterior median
eyes one diameter apart one and one-half from laterals. Anterior eyes
are slightly closer in male. Abdomen with lateral grooves; female
abdomen extension, two-thirds length of abdomen between spinnerets
and pedicel. Male abdomen blunt and short. This species can be
diagnosed by the genitalia (Figs. 49-51).

Figure 48 was made from the juvenile type specimen.

Records. Venezuela. Aragua: Rancho Grande, March 1945; 20
Dec. 1954 (A. M. Nadler, AMNH) ; Choroni, 9 March 1959, (A.

224

Psyche

[December

M. Nadler, AMNH) ; Carabobo: Le Cumbre, S of San Esteban,
1888 (E. Simon, MNHN).

Chrysso cambridgei ( Petrunkevitch) , new combination
Figures 43-47

Achaea vittata O. P.-Cambridge, 1894, Biologia Centrali-Americana, Aranei-
dea, 1: 130, pi. 17, fig. 11, 2. Female type from Teapa, [Tabasco,
Mexico] in the British Museum, Natural History, examined. Not Argy-
rodes vittata Keyserling, T heridion vittatum Nicolet.

Thu'aitesia vittata, — O. P.-Cambridge, 1896, op. cit., p. 197.

Thwaitesia lepida O. P.-Cambridge, 1896, op. cit., p. 197, pi. 25, fig. 5, 2.
Female type from Teapa, [Tabasco, Mexico] in the British Museum,
Natural History, examined.

Chrysso splcndida Banks, — 1898, Proc. California Acad. Sci., (3) 1: 237, pi.
14, fig. 13, 2. Female syntype from Tepic, Nayarit, Mexico in the
Museum of Comparative Zoology, examined.

T heridion vittatum, — F. P.-Cambridge, 1902, Biologia Centrali-Americana,
Araneidea, 2: 391, pi. 37, figs. 6, 7, 2, $ .

Theridion splendidum, — F. P.-Cambridge, 1902, op. cit., p. 391.

Theridion cambridgei Petrunkevitch, 1911, Bull. Amer. Mus. Nat. Hist., 29:
192. New name for Achaea vittata thought preoccupied by T heridion
vittatum Nicolet and for Thnvaitesia lepida thought preoccupied by
Theridion lepidum Walckenaer. Reimoser, 1939, Ann. Naturhist. Museum
Wien, 50: 346. Roewer, 1943, Katalog der Araneae, 1: 490. Kraus, 1955,
Abhandl. Senckenbergischen Naturf. Gesell. 493: 17.

T heridion splendens Roewer, 1942, op. cit. p. 498. New name for Chrysso
splendida thought preoccupied by Theridion splendidum Taczanowski.
Conopistha acuminata Schenkel, 1953, Verh. Naturf. Gesell. Basel, 64: 12, fig.
12, 2. Two female syntypes from El Pozon, Venezuela in the Natur-
historisches Museum, Basel, examined. NEW SYNONYMY.

Comments. F. P.-Cambridge first suspected that Bank’s splendida
is a synonym of vittata. The epigynum, a shallow black depression
(Fig. 45) sometimes has an anterior lip. The only male examined was
caught in December at Barro Colorado Island by Dr. A. M. Chick-
ering.

Distribution. Mexico to Venezuela.

Records. Mexico. Veracruz: 16 km S of San Jose del Carmen
(L. I. Davis, AMNH). Tabasco: 3 km NE of Comalcalco (G.
Ekholm). Guatemala: Variedades, 300 m (C., P. Vaurie, AMNH).
El Salvador. (Kraus, 1955). Costa Rica. (Reimoser, 1938). Pana-
ma: El Valle. Panama Canal Zone: Barro Colorado Island (sev.
collections) .

Chrysso vittatula (Roewer), new combination
Figures 38-42

Argyrodes vittatus Keyserling, 1884, Die Spinnen Amerikas, Theridiidae,
2(1): 191, pi. 9, fig. 114, 2. Female syntypes from St. Fe de Bogota
[Bogota, Colombia] and female, male syntypes from Pumamarca,

1962]

Levi — Chrysso

225

[Junin], Peru in the British Museum, Natural History, examined.
Homonym of A. vittatus Bradley, 1877.

Argyrodes vittatula Roewer, 1942, Katalog der Araneae, 1 : 439. New name
for A. vittatus Keyserling.

Description. Carapace, sternum dark brown, legs yellow. Abdomen
with a longitudinal dorsal black band, series of white patches on sides
and venter partly black. Eyes subequal in size. Anterior median eyes
one and one-half diameters apart, almost two from laterals. Posterior
median eyes one diameter apart, one and one-half from laterals. Cheli-
cerae with two teeth on anterior margin. Total length of female 6.7
mm. Carapace 1.6 mm long, 1.2 mm wide. First femur 3.5 mm long;
patella and tibia. 3.4 mm long; metatarsus 3.0 mm long; tarsus 1.1
mm. Second patella and tibia 1.8 mm; third 1.2 mm; fourth 2.3 mm.

It is uncertain if females have been correctly separated from C.
ecuadorensis. Females of C. vittatula have a dark spot posterior in the
light area of the epigynum (Fig. 40). No other differences were
noted. The male palpi, however, are very distinct (Figs. 41, 42).
Only the availability of larger collections will permit finding charac-
ters separating the females. Illustrations were made from the syntypes.

Distribution. Colombia to Bolivia.

Records. Peru. Junin: Pumamarca, c? (PAS). Cuzco: San
Miguel, 9 (H. W. Foote). Bolivia: Tunari-Massiv, 4000 m, 1948,
$ (Zischka, AMNH).

Chrysso ecuadorensis Levi
Figures 31-37

Chrysso ecuadorensis Levi, “1955” (1957) Jour. New York Ent. Soc. 63: 66,
figs. 16, 38, 39, $. Female type from Runtun Trail, Banos, [Tungurahua],
Ecuador in the American Museum of Natural History.

Description. Carapace, sternum red-brown. Legs yellow with ends
of first tibiae and all tarsi darker to brown. Abdomen with white
patches on sides and above spinnerets and with some black pigment
in a median dorsal longitudinal line and above spinnerets (Figs. 31,
32). Posterior median eyes slightly larger than other eyes. Anterior
median eyes one and one-half diameters apart, two from laterals in
female, slightly more than two in male. Posterior median eyes one
diameter apart, one and three-quarters from laterals in female ; in male
one and one-quarter diameters apart, two and one-half from laterals.
Chelicerae with two teeth on anterior margin, apparently none pos-
terior. Total length of female from Bolivia 5.5 mm. Carapace 1.7
mm long, 1.3 mm wide. First femur, 3.9 mm; patella and tibia, 3.6
mm; metatarsus, 3.6 mm; tarsus, 1.2 mm. Second patella and tibia,
1.9 mm; third, 1.3 mm; fourth, 2.5 mm. Total length of male from

226

Psyche

[December

Bolivia 4.0 mm. Carapace 1.7 mm long, 1.4 mm wide. First femur,
3.7 mm; patella and tibia, 3.5 mm; metatarsus, 4.2 mm; tarsus, 1.2
mm. Second patella and tibia, 2.1 mm; third, 1.4 mm; fourth, 2.3
mm.

Diagnosis. It is not certain whether females have been correctly
separated from C. vittatula Roewer. The light area of the epigynum
has a dark spot anterior (Figs. 34, 35). Only large collections of
accompanied females will make the separation of the species possible.
Distribution. Colombia to Bolivia.

Records. Colombia. Cauca: 43 km S Popayan, March 1955, ?
(E. I. Schlinger, E. S. Ross, CAS). Ecuador. Los Rios: Macuchi,
copper mining camp inland from Rio Palenque, March, 1943 9 (H.
E. Frizzell). Tungurahua: Mt. Tungurahua, 1900-2000 m, Apr.
1939, 9 ( W. C. Macintyre) ; Banos, 1850-2000 m, Apr. 1939, 9 (W.
C. Macintyre, AMNH) . Bolivia. Chaco, 2000 m, 9, c? (MNHN).

Chrysso compressa (Keyserling) , new combination
Figures 63-70

Achaea compressa Keyserling, 1882, Die Spinnen Amerikas. Theridiidae,
2(1): 107, pi. 5, fig. 69, 9. Female type from Amable Maria,

[Junin], Peru in the Polish Academy of Sciences, Warsaw.

Theridion conspersa, — Simon, 1894, Histoire Naturelle des Araignees, 1: 535
(misspelling) .

Theridion compressum, — Roewer, 1942, Katalog der Araneae, 1 : 491.

Comments. A specimen, presumably collected with the type, in the
British Museum, Natural History, was examined. The abdomen
resembles C. cambridgei , it has slight grooves on sides; a black band
covers dorsum.

Description. Carapace yellow-white, yellowish in head region.
Sternum yellow in Peruvian specimens, black in Brazilian. Legs yel-
low-white, distal ends of most leg segments darker yellow with denser,
darker hairs. Abdomen with median dorsal colorless area and with
white pigment on each side. Brazilian specimens have genital area

Explanation of Plate 16

Figs. 52-54. Chrysso ribeirao sp. n. 52. Female abdomen, lateral view. 53.
Female genitalia, dorsal view. 54. Epigynum.

Figs. 55-57. C. pulchra (Keyserling). 55. Female abdomen, lateral view.
56. Female genitalia, dorsal view. 57. Epigynum.

Figs. 58-60. C. backstromi (Berland). 58. Female. 59. Female genitalia,
dorsal view. 60. Epigynum.

Figs. 61, 62. C. caraca sp. n. 61. Female genitalia, dorsal view. 62. Epigy-
num.

Figs. 63-70. C. compressa (Keyserling). 63, 65. Female genitalia, dorsal
view. 64, 66, 67. Epigynum. 68. Female abdomen, lateral view. 69, 70. Left
palpus.

Psyche, 1962

Vol. 69, Plate 16

Levi — Chrysso

228

Psyche

[December

black and a black spot anterior to spinnerets. Anterior median eyes
slightly larger than others or subequal, their diameters apart, and one-
third from laterals. Posterior median eyes slightly oval, their shorter
diameter apart, and one and one-half from laterals. Chelicerae with
two teeth on anterior margin. Abdomen usually without grooves.
Total length of female from Peru 6.1 mm. Carapace 1.8 mm long,
1-4 mm wide. First femur 5.4 mm ; patella and tibia 5.4 mm ; metatar-
sus 5.5; tarsus 1.8 mm. Second patella and tibia 3.0 mm; third 1.8
mm; fourth 3.2 mm. Total length of male from Brazil 4.1 mm.
Carapace 1.7 mm long. First femur 6.1 mm; patella and tibia 6.1
mm; metatarsus 3.6 mm; tarsus 1.8 mm. Second patella and tibia 3.3
mm; third 1.8 mm; fourth 3.3 mm.

The shape of the epigynum fossa is variable (Figs. 64, 66, 67).
Figures 63, 64 were prepared from the types.

Distribution. Peru to southeastern Brazil.

Records. Peru. San Martin: Mishqui-yacu, 1600 m, 20 km NE
of Moyobamba, Aug. 1947, $ (F. Woytkowski, AMNH). Hudnuco:
Monzon Valley, Tingo Maria, Oct. 19, 1954, $, cf (E. I. Schlinger,
E. S. Ross, CAS). Brazil. Minas Gerais: Matosinhos, 9 (MNHN).
Guanabara: Teresopolis, 9001000 m, March, 1946, 9 (H. Sick,
AMNH). Sao Paulo: Sao Paulo, Jan. i960, cf (A. M. Nadler,
AMNH) ; Jabaquara, Cidade Sao Paulo, 700-800 m, Dec. 1945, 9>
cf (H. Sick, AMNH). Santa Catarina : Nueva Teutonia, lat 270
1 1' S, long 520 23' W, 9> cf (F. Plaumann, SMF) .

Chrysso arops sp. n.

Figures 29, 30

Type. Male from Teresopolis, 900-1000 m alt., Est. Guanabara,
Brazil (H. Sick), in the American Museum of Natural History. The
specific name is an arbitrary combination of letters.

Description. Carapace with clypeus, labium, maxillae orange. Ster-
num black, except anterior portion. Legs orange to gray. Abdomen
orange-white with several black spots (Fig. 29). Eyes subequal in
size. Anterior median eyes one diameter apart, their radius from
laterals. Posterior median eyes one and one-fourth diameters apart,
one and one-half from laterals. Chelicerae with one tooth on the
anterior margin. Abdomen as illustrated with a hump above spinnerets
but without grooves (Fig. 29). Total length 2.4 mm. Carapace 0.98
mm long, 0.93 mm wide. First femur 2.58 mm; patella and tibia 2.50
mm ; metatarsus 2.34 mm. Fourth patella and tibia 1.28 mm.

Diagnosis. This species probably belongs to Chrysso , it is not cer-

1962]

Levi — Chrysso

229

tain whether the palpus (Fig. 30) has a radix. The palpal structure
separates C. arops from other Chrysso.

Chrysso ribeirao sp. n.

Figures 52-54

Type. Female from Ribeirao Pires, 700-800 m alt., Est. Sao Paulo,
Brazil, Dec. 1945 (H. Sick), in the American Museum of Natural
History. The specific name is a noun in apposition after the type
locality.

Description. Carapace yellow-white with some gray around pos-
terior margin. Sternum all black in most specimens, in some only
partially. Legs yellow-white with a black band on distal end of first
femora and first tibiae. Abdomen with black and white areas and a
median dorsal white band (Fig. 52). Eyes subequal in size. Anterior
median eyes their diameter apart, almost touching laterals. Posterior
median eyes their diameter apart, two-thirds from laterals. Abdomen
without grooves (Fig. 52). Total length 2.7 mm. Carapace 0.89
mm long, 0.77 mm wide. First patella and tibia 1.8 mm; second 1.04
mm; third 0.74 mm. fourth 1.22 mm.

Diagnosis. The curved connecting ducts opening near the posterior
rim (Figs. 53, 54) separates this species from others. The distance
between the opening and the rim is variable.

Records. Brazil. Guanabara: Teresopolis, March 1946, 3 9 para-
types (H. Sick, AMNH).

Chrysso pulchra (Keyserling) , new combination
Figures 55-57

Achaea signata Keyserling, 1891, Die Spinnen Amerikas, Brasilianische
Spinnen, 3: 199, pi. 7, fig. 1+3, 9. Female type from Alto da Serra do
Novo Friburgo [Nova Friburgo, Est. Guanabara], Brazil: in the British
Museum, Natural History, examined. NEW SYNONYMY.

Achaea pulchra Keyserling, 1891, op', cit., p. 200, pi. 7, fig. 144, 9. Female
type from Alto da Serra do Novo Friburgo [Novo Friburgo, Est. Guana-
bara], Brazil in the British Museum, Natural History, examined.

Theridion pulchrum, — Roewer, 1942, Katalog der Araneae, 1: 497.

T heridion signatellum Roewer, 1942, op. cit., p. 498. New name for signata,
thought to be preoccupied by Theridion signata Walckenaer, 1805. NEW
SYNONYMY.

Comments. Although signata has page priority, it will help nomen-
clatural stability to use the namei pulchra , since Theridion signata is
preoccupied and some authors might consider Chrysso and Theridion
synonymous. The types of signata and pulchra differ only in the color-
ation of the abdomen, a variable character. Chrysso pulchra has the

230

Psyche

[December

dorsum of the abdomen black, the sides white, and the venter yellow-
ish without pigment (Fig. 55). The carapace and palpi of this species
are black. The illustrations were prepared from the types of Achaea
pulchra.

Ghrysso caraca sp. n.

Figures 61, 62

Type. Female from Caraca, Minas Gerais, Brazil (E. Gounelle),
in the Museum National d’Histoire Naturelle, Paris (no. 8881).
The specific name is a noun in apposition after the type locality.

Description. Carapace yellow. Sternum yellow, gray or black.
Legs yellow, sometimes with irregular black marks. Abdomen with a
median longitudinal dorsal white band and white patches on sides; a
black spot on each side of posterior tip and sometimes a black patch
on each side. Carapace fairly high. Anterior median eyes slightly
smaller than others, their diameter apart, one-quarter from laterals.
Posterior median eyes one diameter apart, their radius from laterals.
Chelicerae with two teeth on anterior margin, probably none posterior.
Abdomen barely longer than high. Total length 2.9 mm. Carapace
0.98 mm long, 0.79 mm wide. First femur, 1.7 1 mm; patella and
tibia, 1.69 mm; metatarsus, 1.43 mm; tarsus, 0.62 mm. Second
patella and tibia, 1.00mm; third, 0.59 mm; fourth, 1.12 mm.

Diagnosis. The opening of the epigynum is bordered by an anterior
lip (Fig. 62) unlike C. pulchra ; the shorter, straighter ducts (Fig.
61 ) separate this species from C. ribeirao.

Records. 3 paratypes collected with type.

Chrysso backstromi (Berland), new combination
Figures 58-60

Theridion backstromi Berland, 1924, The Natural History of Juan Fernandez
and Easter Island, 3 : 426, figs. 4-6, $ . Female type from Juan Fernandez
Island, [Chile] in the Naturhistoriska Riksmuseum, Stockholm, examined.

Description. Yellow-white with white and black markings (Fig.
58). Legs yellow-white with black rings on ends of femora and tibiae
of fourth leg. Anterior median eyes smaller than others, a little more
than one diameter apart, their radius from laterals. Posterior eyes
two-thirds their diameter apart. Eyes appearing white and relatively
large. Chelicerae without teeth. Abdomen with a hump (Fig. 58).
Total length 2.5 mm. Carapace, 1.1 mm long, 1.0 mm wide. First
femur, 2.2 mm ; patella and tibia, 2.2 mm ; metatarsus, 2.0 mm ; tarsus,
0.9 mm. Second patella and tibia, 1.3 mm; third, 0.9 mm; fourth 1.5
mm.

1962]

Levi — Chrysso

231

Chrysso clementinae (Petrunkevitch), new combination
Figures 71-75; Map 1

Meotipa clementinae Petrunkevitch, 1930, Trans. Connecticut Acad. Sci., 30.
212, fig. 61, $. Female type from Bayamon, Puerto Rico, in Petrunkevitch
personal collection. Schmidt, 1956, Zool. Anz., 15: 240. [Not Bryant,
1940, Bull. Mus. Comp. Zool., 86 : 314, figs. 79, 80, $ ].

Description. Color white, with narrow reddish line in thoracic
region, some scattered black spots on venter of legs and white pigment
spots and some scattered small spots on dorsum of abdomen. Eyes
subequal in size. Anterior median eyes a little more than one diameter

Map. 1. Distribution of Chrysso clementinae (Petrunkevitch).

apart, almost touching laterals. Posterior eyes separated by about one
diameter. Abdomen often with several strong feather shaped ( ?
movable) spines near tip (Fig. 71). Epigynum with two dark spots
in a lightly sclerotized area (Fig. 73). Male palpus with a short
embolus and large conductor (Figs. 74, 75). Total length of a female
from Puerto Rico 2.3 mm. Carapace 0.85 mm long, 0.78 mm wide.
First femur, 2.45 mm; patella and tibia, 2.31 mm; metatarsus, 1.5 1
mm; tarsus, 0.61 mm. Second patella and tibia, 1.30 mm; third, 0.8 1
mm; fourth, 1.55 mm. Total length of a male from Venezuela 2.0
mm. Carapace 1.00 mm long, 0.78 mm wide. First femur 3.60 mm;
patella and tibia, 3.35 mm; metatarsus, 3.45 mm; tarsus, 0.91 mm.
Second patella and tibia, 2.05 mm; third, 1.13 mm; fourth, 1.96 mm.

232

Psyche

[December

Natural History. This species is found on the underside of leaves
of shrubs and small trees. Schmidt (1956) reports that it is common-
ly brought with bananas to Europe.

Distribution. Probably cosmotropical ; the absence of closely related
species in America might suggest that it has been introduced.
(Schmidt, 1956, records having obtained specimens with fruit from
Guatemala, Colombia, French West Africa and Belgian Congo) .

Records. Florida. Alachua Co.: Gainesville (H. K. Wallace,
Hubbell, AMNH). Dade Co.: Kendall (A. M. Nadler, AMNH).
Highlands Co.; 3 mi. S of Lk. Istokpoga (A. M. Nadler, AMNH).
Seminole Co.: near Geneva (W. J. Gertsch, AMNH). Mexico.
V eracruz. 30 km S of Jesus Carranza; Lake Catemaco, 300 m (C.
P. Vaurie, AMNH). Panama Canal Zone. Pedro Miguel; Corozal;
Barro Colorado Island; Summit; France Field (all A. M. Chicker-
ing). Cuba. Vega Alta, Santa Clara (P. Bermudez) ; Soledad, Las
Villas (P. J. Darlington). Jamaica. Mona, St. Andrew Par.; Gas-
tonbury, Trelawny Par.; Rio Cobre Gorge, St. Catherine Par.;
Ewarton, St. Catherine Par. (all A. M. Chickering) ; Kinloss (A. M.
Nadler, AMNH). Dominican Republic, near Ciudad Trujillo (A.
M. Nadler, AMNH); Boca Chica (A. M. Nadler, AMNH).
Puerto Rico. Rio Piedras (A. M. Nadler, AMNH). Venezuela.
Aragua: Maracay; Rancho Grande (A. M. Nadler, AMNH). Dto.
Federal: Caracas (A. M. Nadler, AMNH ) . British Guiana. Sauri-
Wau River near Rio Tacutae (W. G. Hassler, AMNH). Surinam:
Paramaribo (A. M. Nadler, AMNH). French Guiana. Cayenne
(A. M. Nadler, AMNH). Ecuador. Pichincha: 35 km NW of
Santo Domingo de los Colorados (A. M. Nadler, AMNH). Tun-

Explanation of Plate 1 7

Figs. 71-75. Chrysso clementinae (Petrunkevitch) . 71. Female, lateral

view. 72. Female genitalia, dorsal view. 73. Epigynum. 74. Left palpus. 75.
Palpus expanded, submesal view.

Figs. 76-78. C. questona sp. n. 76. Female abdomen, lateral view. 77.
Female genitalia, dorsal view. 78. Epigynum.

Fig. 79. C. melba sp. n., palpus.

Figs. 80,81. C. vesiculosa (Simon) (Philippines) , Epigynum.

Figs. 82, 83. C. alecula sp. n. 82. Female genitalia, dorsal view. 83.
Epigynum.

Figs. 84, 85. C. arima sp. n. 84. Female genitalia, dorsal view. 85. Epigy-
num.

Figs. 86-90. C. silva sp. n. 86. Female. 87. Female genitalia, dorsal view.
88. Epigynum. 89. Palpus. 90. Palpus expanded, subventral view.

Figs. 91-95. C. volcanensis sp. n. 91. Female. 92. Female genitalia, dorsal
view. 93. Epigynum. 94. Palpus expanded, cymbium removed, dorsal view.
95. Palpus. (C, conductor; E, embolus; M, median apophysis; P, paracym-
bium; R, radix; Y, cymbium).

Psyche, 1962

Vol. 69, Plate 17

234

Psyche

[December

gurahua: Banos (H. E. Frizzell) ; Punapi (H. E., D. L. Frizzell).
Guayas: Milagro (H. E., D. L. Frizzell). Peru. “San Martin”
(J. C. Pallister). Pium: Bellavista, Prov. Jean, (D. L. Frizzell).
Brazil. Maranhao : Sao Luis (A. M. Nadler, AMNH). Pernam-
buco: Recife (A. M. Nadler, AMNH). Espirito Santo: Santa
Teresa (A. M. Nadler, AMNH). Sao Paulo: Sao Paulo (A. M.
Nadler, AMNH) . Paraguay. San Luis [?].

Ghrysso melba sp. n.

Figure 79

Type. Male from Summit, Panama Canal Zone, July 21-29, 1950
(A. M. Chickering) in the Museum of Comparative Zoology. The
specific name is an arbitrary combination of letters.

Description. Yellow-white with a dusky median band on carapace,
ends of femora, on patella, and distal ends of tibiae and metatarsi.
Dorsum of abdomen with two rows of about ten dusky bars. Eyes
subequal in size; anterior median eyes, one and one-third diameters
apart, their radius from laterals. Posterior median eyes one and one-
half diameters apart, a little more than one from laterals. Chelicerae
with two teeth on anterior margin. Abdomen longer than wide or
high; highest above spinnerets. Total length 2.2 mm. Carapace 1.0
mm long, 0.9 mm wide. First femur, 3.5 mm; patella and tibia, 3.5
mm; metatarsus, 3.7 mm; tarsus, 1.2 mm. Second patella and tibia,
2.0 mm ; third 0.9 mm ; fourth, 1.7 mm.

Diagnosis. The palpal structure (Fig. 79) separates this species
from C. clementinae.

Ghrysso alecula sp. n.

Figures 82, 83

Type. Female from El Valle, Panama, July 1936 (A. M. Chicker-
ing) in the Museum of Comparative Zoology. The specific name is
an arbitrary combination of letters.

Description. Yellow-white with a dusky median band on carapace,
small dark spots on venter of legs and white pigment spots and some
dusky marks on dorsum of abdomen. Eyes subequal in size. Anterior
median eyes their diameter apart, one-fourth from laterals. Posterior
median eyes one-half diameter apart, one-fourth from laterals. Shape
of abdomen like that of C. silva (Fig. 86). Total length of female,
1.6 mm. Carapace, 0.68 mm long 0.65 mm wide. First femur, 1.71
mm; patella and tibia, 1.61 mm; metatarsus, 1.43 mm; tarsus, 0.50

1962] Levi — Chrysso 235

mm. Second patella and tibia, 1.06 mm; third, 0.71 mm; fourth,
1. 1 7 mm.

Diagnosis. The long loosely coiled ducts of the genitalia (Figs. 82,
83) separate this species from C. arima.

Records. Panama: El Valle, July 1936, $ paratypes (A. M.
Chickering) .

Chrysso arima sp. n.

Figures 84, 85

Type. Female from Simla, near Arima, Trinidad, 28 Dec. 1954
(A. M. Nadler) in the American Museum of Natural History. The
specific name is a noun in apposition after the type locality.

Description. Yellow-white with white pigment spots on dorsum of
abdomen. Eyes subequal in size. Anterior median eyes separated by
one diameter, one-fourth from laterals. Posterior eyes separated by
one diameter. Total length of female 1.6 mm. Carapace, 0.63 mm
long, O.52 mm wide. First femur, 1.42 mm; patella and tibia, 1.28
mm; metatarsus, 0.98 mm; tarsus, 0.50 mm. Second patella and
tibia, 0.80 mm ; third, 0.53 mm ; fourth, 0.98 mm.

Diagnosis. The narrow, tightly coiled ducts of the genitalia and
the median opening (Figs. 84, 85) separate this species from C. alecula.

Record. Trinidad. Simla near Arima, 9 paratype (A. M. Nadler).

Chrysso silva sp. n.

Figures 86-90

Type. Male from Forest Reserve, Panama Canal Zone, 25-28 July
1954 (A. M. Chickering) in the Museum of Comparative Zoology.
The specific name is an arbitrary combination of letters.

Description. Yellow-white with a median longitudinal dusky band
on carapace; abdomen with white pigment patches on dorsum, dusky
on sides. Anterior median eyes subequal or slightly larger than others,
three-quarters diameter apart, almost touching laterals. Posterior eyes
separated by one-third their diameter. Abdomen much higher than
long in female (Fig. 86). Total length of a female, 1.6 mm. Carapace
0.65 mm long, 0.59 mm wide. First femur, 1.56 mm; patella and
tibia, 1.43 mm ; metatarsus, 1.30 mm ; tarsus, 0.45 mm. Second patella
and tibia, 0.88 mm; fourth, 0.97 mm. Total length of male 1.4 mm.
Carapace, 0.66 mm long, 0.66 mm wide. First femur, 1.70 mm ; patel-
la and tibia, 1.75 mm; metatarsus, 1.53 mm; tarsus, 0.52 mm. Second
patella and tibia, 1.09 mm ; third, 0.63 mm ; fourth, 0.98 mm.

236

Psyche

[December

Diagnosis. The larger diameter and shortness of the female ducts
(Figs. 87, 88) separate this species from C. alecula and C. arima. The
large coiled embolus of the palpus (Fig. 89) is distinctive.

Natural History. This species is found by beating vegetation.

Records. Panama. Porto Bello, $ (A. M. Chickering). Panama
Canal Zone. Forest Reserve, cf paratype (A. M. Chickering) ; Sum-
mit, $ paratype (A. M. Chickering, A. M. Nadler, AMNH) ; Barro
Colorado Island, $, cf paratypes (A. M. Chickering, A. M. Nadler,
AMNH) ; Madden Dam (A. M. Chickering) ; Ft. Randolph (A.
M. Chickering). Ecuador. El Oro: Rio JuDanes, Pasaje, Oct. 23,
1942, cf (R. Walls).

Chrysso volcanensis sp. n.

Figures 91-95

Thwaitesia vittata, — Banks, 1929, Bull. Mus. Comp. Zook, 69: 86. Not C.
vittata O. P. -Cambridge.

Type. Male from El Volcan, Panama, Aug. 9-14, 1950 (A. M.
Chickering) in the Museum of Comparative Zoology. The species is
named after the type locality.

Description. Carapace, sternum, legs yellow-white. Legs with dark
bands on middle and ends of segments. Abdomen yellow-white with
white pigment areas on dorsum, sometimes with small black spots (Fig.
91). Anterior median eyes one and one-half diameters apart, their
radius from laterals in female ; one diameter apart, one-third diameter
from laterals in male. Posterior median eyes a little more than one
diameter apart, one from laterals. Total length of females 3. 5-4.5 mm.
One female measured 3. 7 mm total length. Carapace, 1.2 mm long,
0.8 mm wide. First femur, 5.0 mm; patella and tibia, 5.0 mm; meta-
tarsus, 5.4 mm; tarsus, 1.4 mm. Second patella and tibia, 2.7 mm;
third, 1.3 mm ; fourth, 2.6 mm. Total length of a male 2.5 mm. Cara-
pace 1.2 mm long, 1.0 mm wide. First femur, 5.9 mm; patella and
tibia, 6.0 mm; metatarsus, 7.5 mm; tarsus, 1.7 mm. Second patella
and tibia, 3.6 mm ; third, 1.4 mm ; fourth, 2.6 mm.

Diagnosis. The shorter ducts and embolus (Figs. 92-95) distinguish
this species from C. questona. The epigynum (Fig. 93) has an indis-
tinct shallow depression of variable shape. The small palpal embolus
(Figs. 94, 95) separate the species from C. silva.

Records. Costa Rica: Santa Maria Dota (Tristan). Panama. El
Volcan, 9, cf paratypes (A. M. Chickering).

Literature Cited

Levi, Herbert W.

“1955” (1957). The spider genera Chrysso and Tidarren in America.

Jour. New York Ent. Soc., 63 : 59-81.

1962]

Levi — Chrysso

237

“1957” (1958). Spiders of the new genus Arctachaea. Psyche, 64: 102-
106.

Levi, Herbert W. and Lorna R. Levi

1962. The genera of the spider family Theridiidae. Bull. Mus. Comp.
Zool. 127(1) : 1-71.

Uyemura, T.

1957. Colour change in two species of Japanese spiders. Acta Arach-
nologica 15: 1-10.

Yaginuma, T.

1957. Two new conopisthine spiders from Japan, ibid. 15:11-16.

THE NEOTROPICAL SPECIES OF THE ANT GENUS
STRUMIGENYS FR. SMITH: SYNOPSIS AND KEYS
TO THE SPECIES1

By William L. Brown, Jr.

Department of Entomology, Cornell University

Introduction

The New World Strumigenys have been revised through a series of
twelve papers bearing the general foretitle, “The Neotropical species
of the ant genus Strumigenys Fr. Smith,” plus several articles by Dr.
W. W. Kempf and by myself, beginning with my “Preliminary generic
revision of the higher Dacetini” (Brown, 1948). It now seems appro-
priate to offer a unifying synopsis of the New World species of the
genus, along with keys for identification and some general remarks.

Species Synopsis of New World Strumigenys

The synopsis below includes the names, each with author and date
of publication, plus citation of the principal references in the Brown
or Kempf papers already mentioned, which are listed in the section of
“References” at the end of this article. These papers contain refer-
ences to original descriptive and distributional material for each species,
but I have included in the synopsis new or supplementary information
wherever it seemed useful to do so. The species are listed by groups
in order of apparent relationship, as closely as it is possible to place
them in a purely linear order. The probable relationships within the
genus in the New World are discussed at the end of the synopsis. It
will be noticed that the group placement of some species differs from
that of the previous parts published. The present grouping represents
a reconsideration of all of the New World species taken together.

i Group of mandibularis

I. Strumigenys mandibularis Fr. Smith, i860
Brown, 1953b: 53-55, worker, synonymy.

Frederick Smith confused two species under this name; one of these
was later described as S. prospiciens by Emery. In order to fix these
names unambiguously according to present usage, I hereby designate
as lectotype of S. mandibularis the worker in the British Museum
(Natural History), which was called “holotype” in my 1953 paper.

Published) with the aid of a grant from the Grace Griswold Fund of the
Department of Entomology, Cornell University.

Manuscript received by the editor January 25, 1962.

238

1962]

Brown — Strumigenys

239

Although this specimen is labeled as “type,” Smith never designated
a type in print, and at least some of his original specimens exist else-
where (e.g., in the Munich Museum).

Distribution : Amazon Basin ; known only from the type series.
Synonym: S. batesi Forel.

2. Strumigenys godmani Forel, 1899

Brown, 1953b: 55-56, worker, female, variation.

Biology: Lives in wet forest. The nest I found in Panama was in
a small rotten log in cloud forest.

New records: Panama: Progreso, Chiriqui Prov. (F. M. Gaige
leg.) ; Cerro Campana, about 950 m altitude, Panama Prov. (W. L.
Brown, Jr. leg.).

Distribution: Costa Rica, Panama, British Guiana.

Synonym : S. ferox Weber.

3. Strumigenys planeti Brown, 1953

Brown, 1953b: 57-59, worker, female, variation, distribution.

Biology: Apparently a rain forest species. Weber (1952) reports
a nest taken in a wet mossy log in a cacao plantation on Trinidad.

New record: Peru: Monson Valley, Tingo Maria, winged female:
>( E. I. Schlinger and E. S. Ross leg.) .

Distribution : Trinidad, Amazon Basin to Bolivia and Peru.

4. Strumigenys smithii Forel, 1886

Brown, 1953c: 104-107, worker, variation, distribution, biology.

Biology: Nests in rotten logs, rotten twigs or, on St. Vincent, rare-
ly in sod. Primarily a forest species.

New records: Colombia: Loma Larga, Sierra Santa Marta (F. M.
Gaige leg.). Panama: Cerro Campana, 800 m, Province of Panama
(G. B. Fairchild and W. L. Brown, Jr. leg.) .

Distribution: Costa Rica south to Santa Catarina, Brazil, and
Amazonian Bolivia; St. Vincent, B. W. I.

Synonym : S. smithi var. inaequalis Emery.

5. Strumigenys hemidisca Brown, 1953 (Fig. 22)

Brown, 1953c: 107-108, worker.

Biology: The type series came from orchid plants intercepted in
U. S. plant quarantine, and so were probably nesting amid the epiphy-
tes in trees.

Distribution : Venezuela ; known only from the type series.

6. Strumigenys prospiciens Emery, 1906
Brown, 1953c: 108-110, worker, female, distribution.

240

Psyche

[December

Distribution : Amazon Basin south to Bolivia and to northern
Argentina : Misiones.

7. Strumigenys biolleyi Forel, 1908 (Fig. 28)

Brown, 1953c: 101-104, worker female, variation, distribution.

Biology: A forest species, nesting mainly in rotten logs.

New records : Ecuador : 10 miles north of Manglar Alto, Guaymas
(E. I. Schlinger and E. S. Ross leg.). Panama: Cerro Campana,
Panama Prov., 800 m altitude (W. L. Brown, Jr. leg.).

Distribution: Southern Mexico (Chiapas) south through Central
America to Ecuador.

Synonyms: S. tridens Weber, S. luctuosa Menozzi.

8. Strumigenys saliens Mayr, 1887

Brown, 1954b: 55-57, worker, female, distribution, biology.

Biology: Nests in rotten logs and branches lying on the floor of
forest.

Distribution: Southeastern Brazil and northeastern Argentina:

Misiones.

Synonyms: S. saliens var. procera Emery and var. angusticeps Forel.

9. Strumigenys borgmeieri Prown, 1954
Brown, 1954b: 57-59, worker.

Distribution : Brazil : Pernambuco ; known only from the holotype.

10. Strumigenys trinidadensisVJ heeler, 1922 (Figs. 14,23)

Brown, 1954b: 59-62, worker, male, distribution.

New record: Esmeralda, Ecuador (J. Foerster leg.).

Distribution: Trinidad, northeastern Brazil, Ecuador, Amazonian
Bolivia; probably widespread in the interior of South America.

11. Strumigenys sanctipauli Kempf, 1958 (Fig. 24)

Kempf, 1958b: 556-559, figs. 1-4, worker.

Distribution : Brazil : Serra do Mar, Sao Paulo State ; known only
from the holotype.

12. Strumigenys sublonga Brown, 1958
Brown, 1958a: 221-222, fig. 1C, D, worker, female.

Distribution : Bolivia : Lower Rio Madidi ; known only from type
series.

13. Strumigenys rehi Forel, 1907
Brown, 1958a: 222-223, worker.

Biology: This species was taken from orchid plants arriving at
Hamburg, Germany, a circumstance agreeing with the large eyes of
the worker to indicate an arboreal habitat.

1962]

Brown — Strumigenys

241

Distribution : Amazon Basin ; exact type locality unknown.

14. Strumigenys cordovensis Mayr, 1887 (Figs. 25, 26, 27)

Brown, 1958a: 218-220, fig. IB, E, F, G, worker, variation, distribution.

Distribution : Southern Mexico to Trinidad and the Guianas.

15. Strumigenys mokensis Forel, 1905

Brown, 1958a: 221, raised from variety to provisional species rank.

This is a very doubtful form, most likely a synonym of cordovensis.
The whereabouts of the type is unknown. The species is not included
in the key.

Distribution : La Moka, Venezuela, type locality.

16. Strumigenys dolichognatha Weber, 1934
Brown, 1958a: 223-224, fig. 1A, worker.

Distribution: British Guiana: Kartabo; known only from the type
series.

Group of cultriger

17. Strumigenys cultriger Mayr, 1887 (Fig. 9)

Brown, 1957: 97-99, worker.

New record: Xaxim, Santa Catarina (F. Plaumann leg.).
Distribution : Southeastern Brazil.

18. Strumigenys deltisquama^YOwn, 1957
Brown, 1957: 99-101, fig. la; b, worker.

Distribution : Panama Canal Zone: Barro Colorado Island ; known
from types only.

Group of tococae

19. Strumigenys tococae Wheeler. 1929
Brown, 1957: 101-102, fig. lc, worker.

Biology: The types were taken from an abundant population inhab-
iting the foliar sacs of Tococa formicaria, a tall myrmecophytic shrub,
in the outskirts of Bffem. From this circumstance and the large size
of the eyes, S. tococae is judged to be an arboreal or subarboreal special-
ist.

Distribution : Belem, Brazil, at the mouth of the Amazon ; known
only from the type series.

20. Strumigenys fairchildi Brown, 1961
Brown, 1961: 60-61, worker.

This species, described from a single worker, is very close to S.
tococae , but differs markedly in gastric sculpture and pilosity. It is-

242

Psyche

[December

not known whether S. fairchildi lives in plant cavities, but it does seem
likely that it is a subarboreal forager.

Distribution : Panama : Cerro Campana, Panama Province ca.
800 m altitude ; known only from the holotype.

Figure 1. Strumigenys India, worker from Veracruz, dorsal full-face view
of head showing fringing pilosity only.

Group of ludia

21. Strumigenys longispinosa Brown, 1958
Brown, 1958b: 123-126, figs. 1, 2, worker.

Biology : Nests in the soil of tropical forest.

Distribution : Panama.

22. Strumigenys marginiventris Santschi, 1931
Brown, 1958b: 126-128, fig. 3, worker, female.

Biology: Nests in the soil, often in paths or other other openings,
in rain forest or plantations, and the workers forage over the open
ground among leaves or herbs by day as well as night. Common on
Barro Colorado Island.

New records: Palmar, Puntarenas Dept., Costa Rica, in soil of
banana plantation, several collections (E. O. Wilson leg.).

Distribution : Golfo Dolce region of Costa Rica to northern Colom-
bia.

23. Strumigenys ludia Mann, 1922 (Figs. 1, 5)

Brown, 1954a: 194-196, worker, female.

Biology: S. ludia has been investigated at length in the field by

1962]

Brown — Strumigenys

243

Wilson and in the artificial nest by Wilson and Brown, and the details
will be published elsewhere. S. India is a forest species and usually
nests in rotten branches or twigs lying on the forest floor. The food is
chiefly entomobryoid Collembola caught alive in the manner usual for
the genus.

New records: Mexico: Ridge between Antiguo Morelos and Nue-
vo Morelos (E. S. Ross leg.). Pueblo Nuevo, near Tetzonapa, Vera-
cruz (E. O. Wilson leg.). Costa Rica: Abaca Plantation, Bataan (C.
H. Batchelder).

Distribution : Southern Mexico to Costa Rica.

Synonym : S. ludia subsp. tenuis Weber.

Group of hindenburgi

24. Strumigenys hindenburgi For el, 1915 (Fig. 8)

Brown, 1961: 61-64, worker, pseudogyne, distribution.

Distribution : Northern Argentina extending into southeastern Bra-
zil.

25. Strumigenys lanuginosa'W\\zz\zr, 1905 (Fig. 4)

Brown, 1961: 61-63, worker, female, distribution.

Distribution: Southern Mexico, Panama; Bahamas, where prob-
ably introduced.

26. Strumigenys ogloblini Santschi, 1936
Brown, 1958c: 136-137, fig. lb, worker, female.

Distribution: Northern Argentina, probably also in southern

Brazil.

Group of elongata

27. Strumigenys precava Brown, 1954 (Fig. 7)

Brown, 1954a: 196-200, worker, female.

Biology: I found this species rather common on Barro Colorado
Island in the Panama Canal Zone, nesting in red- or chocolate-rotten
logs. One nest found was very large, containing several hundred — -
perhaps a thousand or more — workers. Workers were seen carrying
a mycetophilid larva and a termite nymph into this nest as it was being
opened, and a captive colony fed on a wide variety of small arthropods,
including entomobryoid collembolans.

New record: Panama: Cerro Campana, Panama Province, about
800 m altitude, in a small rotten log in a cloud forest ravine, with
winged females, Jan. 16, i960 (G. B. Fairchild and W. L. Brown,
Jr- leg.).

244

Psyche

[December

Figures 2-6. Strumigenys spp., workers. Figure 2, S. lacacoca, paratype,
dorsal full-face view of head, showing fringing pilosity only. Figure 3, S.
nevermanni, same. Figure 4, S. lanuginosa, same. Figure 5. S. ludia, Vera-
cruz, side view of posterior alitrunk, nodes and anterior part of gaster.
Figure 6, S. lacacoca, paratype, same. Roughly to same scale.

1962]

Brown — Strumigenys

245

Distribution : Panama, British Guiana, Amazonian Bolivia ; prob-
ably widespread in hylaean South America.

28. Strumigenys elongata Roger, 1863

Brown, 1954a: 189-192, worker, female, male, variation, synonymy, distribu-
tion, biology.

Biology : This species is definitely a collembolan feeder, common in
the leaf litter of tropical forest. It seems to tolerate drier as well as
wet forest types.

New records: Mexico: Pueblo Nuevo and El Palmar, near Tet-
zonapa, Veracruz (E. O. Wilson leg.). Ocosingo Valley, Chiapas (C.
and M. Goodnight and L. Stannard leg.).

Distribution : Southern Mexico to Bolivia and southeastern Brazil.
Synonyms: S. imitator Mayr, S. elongata subsp. nicaraguensis
Weber.

29. Strumigenys consanii Brown, 1954
Brown, 1954a: 192-194, worker.

A larger, more robust relative of elongata with smooth and shining
postpeticlar disc.

Distribution: Costa Rica: La Palma, near San Jose, 1500 m alti-
tude ; known only from the type series.

Group of emeryi

30. Strumigenys emeryi Mann, 1922

Brown, 1959a: 97-99, worker, variation, distribution.

Distribution : Honduras, southern Mexico.

31. Strumigenys boneti Brown, 1959 (Fig. 12)

Brown, 1959a: 103-104, worker.

Distribution : Southern Mexico.

32. Strumigenys nevermmni Brown, 1959 (Fig. 3)

Brown, 1959a: 99-100, worker, female.

Distribution: Costa Rica: Hondura, 1050 m altitude; known only
from types.

33. Strumigenys micretes Brown, 1959 (Figs. 13, 19)

Brown, 1959a: 100-101, worker. Brown, 196: 58-60, variation, distribution.

As mentioned in the note in couplet 21 of the key (below), this
species and S. lacacoca may actually represent different populations of
the same species.

Biology : A species of rain forest and cloud forest.

34. Strumigenys lacacoca Brown, 1959 (Figs. 2, 6)

246

Psyche

[December

Brown, 1959a: 101-102, worker. Brown, 196: 58-60, worker variation, distri-
bution.

Distribution: Central Panama.

Group of silvestrii

35. Strumigenys silvestrii Emery, 1905 (Fig. 18)

Brown, 1959c: 25-28, fig. 1, worker, female, synonymy, variation, distribution.

Distribution: Northern Argentina, southern Brazil; also in Cuba
and Louisiana, U. S. A., where probably introduced by commerce.
Synonym : S. caribbeci Weber.

36. Strumigenys carinithorax Borgmeier, 1934
Brown, 1959c: 29-30, worker, female, male.

Distribution : Dutch Guiana : vicinity of Paramaribo.

37. Strumigenys schmalzi Emery, 1905
Brown, 1959c: 28-29, worker.

Distribution : Southeastern Brazil.

38. S tru m igenys perparva Brown, 1958
Brown, 1958c: 133-135, fig. la, worker, female.

Distribution: Trinidad and the Guianas to Sao Paulo; probably
interior Brazil.

Group of louisianae

39. Strumigenys mixta Brown, 1953 (Figs. 15, 21)

Brown, 1953a: 4-5, worker.

Biology: One of the two original series was taken in orchid plants
at quarantine, so the species may be arboreal or subarboreal.

Distribution: Guatemala; known only from the types (two locali-
ties) .

40. Strumigenys louisianae Roger, 1863

Brown, 1953a: 2-3, description of synonymous S. clasmospongia, worker.
Brown, 1953d: 28-31, figs. 1, 3, worker, variation, synonymy, distribution.
Brown, 1961: 64-68, geographical variation, synonymy.

Biology: The feeding habits of this species have been studied in
some detail by Wilson (1950, 1954) and by myself. The food con-
sists of a variety of small arthropods found in and on the soil cover
and caught by the workers with their trap-like jaws. The preferred
prey are entomobryoid and symphypleonan Collembola; poduroid
collembolans are not taken.

Distribution : Widespread in the Americas from Virginia and
Tennessee south at least to the Tucuman area of Argentina; north-

1962]

Brown — Strumigenys

247

ward in Mexico to sheltered canyons and cultivated areas of southern
Arizona; greater Antilles (except Jamaica). Unaccountably absent
from certain well-collected areas within this range, such as parts of the
Canal Zone, Trinidad and British Guiana, although plentiful in Costa
Rica and at least some localities in Colombia. This species tolerates
much drier conditions and will live in plantations and other cultivated
situations, so perhaps it is found mostly in habitats outside the primary
forest in the central parts of its range. Its range and ecological ampli-
tude are greater than those of any other New World Strumigenys.

Synonyms: S . unidentata Mayr, S. unispinulosa Emery, S. uni-
spinulosa var. longicornis Emery, S. fusca Emery, S. louisianae var.
obscuriventris Wheeler, 5. bruchi Forel, S. infidelis Santschi, S. eggersi
Var. cubaensis Mann, S. louisianae subspp. laticephala M. R. Smith,
soledadensis Weber, guatemalensis Weber, and costaricensis Weber,
S. clasmospongia Brown. The long list of synonyms reflects in part
the rather extreme variation shown by this species on the South Ameri-
can continent. More peripheral populations (North and Central
America, West Indies, Argentina) tend to be more uniform both
within and among themselves.

41. Strumigenys producta Brown, 1953
Brown, 1953a: 3-4, worker.

This species is a larger, long-mandibulate version of S. louisianae.
In view of the extensive variation now known for the latter species in
South America, it would not be surprising to find that S. producta is
just an extreme local variant of S. louisianae.

Distribution: Basin of the Rio Beni, Bolivia; known only from the
types.

Group of connectens

42. Strumigenys connectens Kempf, 1958 (Fig. 11)

Kempf, 1958a: 59-64, figs. 1-3, worker, variation.

Biology: The paratype series was taken in orchid plants in U. S.
quarantine, so the species is presumably arboreal.

Distribution : The species is known from two localities, both in
Colombia.

43. Strumigenys laevipleura Kempf, 1958
Kempf, 1958a: 64-65, figs. 5-7, worker.

Biology: Like S. connectens, this species was also taken from an
orchid shipment, and it may therefore be arboreal in habits.

Distribution : Known only from the type series from Colombia,
apparently from the vicinity of Medellin.

248

Psyche

[December

44. Strumigenys xenognatha Kempf, 1958
Kempf, 1958a: 65-66, fig. 4, female.

Biology: The holotype female, a unique, was taken from orchid
plants and bears the same data as the S. laevipleura types, from which
it differs too widely to be their queen. Perhaps it is a social parasite of
S. laevipleura.

Distribution : Colombia.

45. Strumigenys subedentata NLayr, 1887

Brown, 1960: 48-50, figs. 7, 9, worker, female, male, variation, distribution,
biology.

Biology : This species nests in small colonies in the soil or soil cover
in mesic tropical forest and feeds chiefly on entomobryoid Collembola.

Distribution: Southern Mexico south to southeastern Brazil; Trin-
idad ; probably v/idespread in interior South America.

Synonyms: S. tristani Menozzi, S. clavata Weber.

Figures 7-8. Strumigenys spp., workers, dorsal full-face view of head.
Figure 7, S. precava from Panama Canal Zone, showing fringing pilosity
only. Figure 8, S. hindenburgi from Tucuman, Argentina. Not to same scale.

46. Strumigenys trieces Brown, i960
Brown, 1960: 50-51, fig. 8, worker.

Distribution : Costa Rica ; known only from the holotype.

Group of gundlachi

(

1962]

Brown — Strumigenys

249

47. Strumigenys denticulata Mayr, 1887
Brown, 1960: 47-48, fig. 3, worker, female.

Biology: Occurs in both primary and second-growth forest, in leaf
litter ; epiphytes and in termite nests.

Distribution: Trinidad and the Guianas south to southeastern
Brazil ; probably occurs widely in interior South America as well.

48. Strumigenys jamaicensis Brown, 1959

Brown, 1959b: 6, worker. Brown, 1960: 45-46, fig. 4, worker.

Distribution : Mountains of Jamaica.

49. Strumigenys gundlachi (Roger, 1862)

Brown, 1960: 40-45, figs. 1, 5, worker, female, synonymy, distribution, biology.

In addition to the characters cited in the key, it may be mentioned
that fully-colored S. gundlachi workers and females are usually darker
in color (brownish-red to dark brown) than are those of S. eggersi
(ferruginous yellow).

Biology: S. gundlachi feeds chiefly if not entirely on entomobryoid
and sminthuroid Collembola, which it catches by employing a rela-
tively inactive “ambush” type of hunting, but if the prey struggles
after being struck, it may be lifted off the ground and stung in the
manner of other Strumigenys. In many parts of the Caribbean coun-
tries, this is a very abundant ant in the leaf litter of tropical forest,
thickets and plantations, and it tolerates a wide variety of ecological
conditions.

Distribution : Central America and southern Mexico, southern

Florida, West Indies to Trinidad.

Synonyms: S. eggersi varieties vincentensis Forel, banillensis Sant-
schi, isthmica Santschi and herlesei Weber; S. eggersi subsp. infuscaia
Weber, and S. bierigi Santschi.

50. Strumigenys eggersi Emery, 1890 (Figs. 10, 20)

Brown, 1960: 46-47, figs. 2, 6, worker, female, variation, distribution, biology.

Biology: Found in forests, thickets, gardens, etc. Almost certainly
a collembolan feeder.

Distribution: Trinidad and the Guianas to southeastern Brazil and
Amazonian Bolivia. Widespread (possibly by recent introduction) in
the West Indies; southern Florida; southern Mexico.

Group of rogeri

51. Strumigenys rogeri Emery, 1890 (Figs. 16, 17, 29)

Brown, 1954, Bull. Mus. Comp. Zool., 112: 20-23, worker, female, feeding
habits.

Although S. rogeri was first described from West Indian material, I

250

Psyche

[December

showed in 1954 that it is a tramp belonging to a characteristically
African species-group, and itself probably West African in origin.

Distribution: Widespread in the West Indies, from Cuba to Trini-
dad; British Guiana; West Africa; Hawaii, Tahiti, Fiji, Micronesia;
greenhouses in England and Scotland; apparently spreading rapidly
through commerce.

Synonyms : S. incisa Godfrey, S. sulfurea Santschi.

Phylogeny of the New World Strumigenys

I belong to the school that believes that since Darwin phylogenetic
reasoning is inseparably a part of the taxonomic ordering of any group.
The work of the more outspoken “aphyletic taxonomists” shows that
they have not escaped the influence of evolutionary thinking, either,
when it comes to revising a species-group or genus or family. Phylo-
genetic thinking is usually more or less implicit in the grouping of
species, as I have grouped the New World species (above). In Figure
30 I have shown my best guess as to how the species groups are related
one to the others. This diagram should not be taken too seriously,
because Strumigenys species are very likely to be convergent from
different groups, and the convergence may be very close and may
involve several to many characters.

The most serious problem in Strumigenys is the question of direction
of evolution ; in other words, which species or groups are primitive, and
which derived? One can look to the other two faunas of the genus
(Ethiopian-Malagasy and Indo-Australian) , but these give little help
at present. I used to think, for no very good reason, I suppose, that
certain species with large, ruggedly modelled heads and heavy, more or
less closely approximate mandibles \chyzeri group of Melanesia,

Explanation of Plate 18

Figures 9-29. Strumigenys spp., workers. Figures 9-12 and 14-16 show left
mandibles in dorsal view; Figure 13 is a dorsal enlarged view of the apices
of both mandibles. Figure 9, S. cultriger. Figure 10, S. eggersi. Figure 11, S.
connectens, paratype. Figure 12, S. boneti, paratype. Figure 13, S. sp. near
micretes from Boquete, Panama — one of several variants from this locality.
Figure 14, S. trinidadensis. Figure 15, S. mixta, paratype. Figure 16, S. rogeri.
Figures 17-21 are end-on views of the apical fork of the mandibles, much
enlarged. Figure 17, S. rogeri. Figure 18, S. silvestrii. Figure 19, S. micretes,
paratype. Figure 20, S. eggersi. Figure 21, S. mixta, paratype. Figures 22-28,
lateral view of propodeal lamella. Figure 22, S. hemidisca, holotype. Figure
23, S. trinidadensis, paratype. Figure 24, S. sanctipauli, holotype after Kempf.
Figure 25, S. cordovensis. Figures 26, 27, same, showing extremes of variation
in different individuals; the pattern of Figure 27 is common in southern
Mexico. Figure 28, S. biolleyi. Figure 29, S. rogeri, left side of head near eye
as seen from dorsal full-face view, to show “detached” eye.

Brown — Strumigenys

252

Psyche

[December

grandidieri Forel of Madagascar, precava of the present study (Fig.
7)], were primitive types within the genus, but now it seems to me
that the opposite is true. S. loriae Emery (of the chyzeri group) and
S. precava are viewed as derivative species with secondarily broadened
prey specificity, and it is predicted that S. grandidieri will also event-
ually be found to feed on a wide range of small arthropods instead of
the usual Strumigenys diet consisting mainly of collembolans. The
powerful head and mandibles of these species are probably an adapta-
tion to prey less fragile than Collembola.

Mandibular armament is probably the best character to use for
determining direction of descent within Strumigenys. More primitive
dacetine genera ( Acanthognathus , Orectognathus , Microdaceton) have
strumigenite mandibles with three long teeth in the apical fork ; often
the most dorsal of the three is also displaced slightly basad. In cases
where such displacement has taken place, we have what in the genus
Strumigenys would be called an apical fork (with two teeth) plus a
preapical tooth. This is the condition found, with greater or lesser
modification, in most Indo-Australian members of the genus as well
as several New World species. In the African group, the species
judged to be the more primitive ones have two preapical teeth, and
derivative species mostly are smaller in size and tend to lose one or
both distal preapical teeth. Quite a few of the New World forms,
most notably those of the mandibularis group, have two well-developed
preapical teeth on each mandible. In other New World forms, chiefly
among smaller species, one or both of these teeth are present in greatly
reduced form — in fact, in form so greatly reduced as to suggest that
they serve no present function in holding struggling prey. It seems
more likely to me that such feeble denticles represent vestiges of
larger, functional teeth rather than the reverse, especially since so
many of the species, and particularly the smaller species, have them.
From this hint (which is no more than that) , I take it that in the New
World fauna of Strumigenys the mandibularis groups two large pre-
apical teeth represent the primitive condition. The extensive radiation
of undoubted mandibularis group species also speaks for a relatively
long-term existence of this armament pattern. I have accordingly
placed the mandibularis group at the base of my phyletic scheme (Fig.
30), despite the very good possibility that the earliest Strumigenys on
a world basis may have had but a single preapical tooth.

The mandibularis group shows what appears to be a clear double
morphocline. Beginning with a more “normal” or “average” species
such as S. smith'd, a string of species of increasing size and develop-
ment (width) of occipital lobes, concurrent with a shortening and

1962]

Brown — Strumigenys

253

thickening of the mandibles, leads through S. planeti and S. godmani
to S. mandibularis. In the other direction, we find a trend toward
lengthening of the mandibles through the series S. biolleyi, S. saliens,
S. sanctipauli and so on to the species near S. cordovensis , climaxed by
the remarkable S. dolichognatha , the mandibles of which are relatively
longer than in any other ant known to me. Side offshoots of the
mandibularis group are species such as S. borgmeieri and S. trinidaden-
sis; the greatly weakened proximal preapical tooth of the last species
shows the first stages of a trend that apparently led to groups such as
the hindenburgi and emeryi assemblages, and beyond these to the
elongata and silvestrii groups respectively. Species such as S. perparva
and S. ogloblini , both of which have a single preapical tooth on each
mandible, were previously grouped together, but now I think it more
likely that their similarities are due to convergence. Such highly
reduced species are doubtfully placed at best.

The emeryi group, especially S. emeryi itself, is linked to the lo'uisi-
anae group by the virtually perfect intermediate S. mixta. The louisi-
anae group leads to the connectens group and through this to the
gundlachi group. These last three groups all have two (or rarely
more) intercalary denticles between the main teeth of the apical fork.
The genus N eostruma represents a further development of the louisi-
anae groups connectens group-> gundlachi group trend or morpho-
cline.

The three remaining species groups, all small, appear to be derivable
directly from the mandibularis group: the tococae group by addition
of a second intercalary denticle in the apical fork, the cultriger group
by development of a mandibular lamella, and the ludia group by the
serial loss of mandibular teeth.

Identification of Species

This section is intended to provide materials with which any compe-
tent entomologist can hope to identify quickly and surely the Strumi-
genys species at present known from the New World. Of course,
there are certainly species remaining to be discovered in this hemi-
sphere, but I believe that we now know all or nearly all of the species
that are both widespread and reasonably common, and many of the
rare or local species as well.

Before discussing the species, though, it is necessary that we charac-
terize the genus Strumigenys well enough to recognize it in this hemi-
sphere. It will be enough to say that any New World ant with the
following combination of characters is a Strumigenys : Worker and

254

Psyche

[December

female — Exactly 6 antennal segments, of which the third and fourth
are very short and the first (scape) and sixth (apical) are very long
(Figs. 1-4) ; mandibles long and linear, straight or bowed, more than
1/3 as long as the head proper, with an apical fork of two prominent
teeth, other teeth absent to few, usually separated (Figs. 1-4, 7-16) ,*
occiput with a deep median posterior excision between two broad,
rounded lobes, head in front distinctly narrowed (Figs. 1-4) ; spongi-
form appendages, or at least their vestiges, present on petiole and
postpetiole (Figs. 5, 6) ; head and often most of alitrunk reticulate-
punctulate and opaque, rarely with superimposed rugulation. Males
are not dealt with here, since few of them are known, and they cannot
be separated as a group from a number of other dacetine genera.

Measurements, and the proportions derived from them, are very
important in dacetine taxonomy, so it is necessary to measure with a
high degree of accuracy. Measurements should be made to the nearest
hundredth of a millimeter at least. A stereomicroscope magnifying at
least 90X is required, plus a carefully calibrated reticule of the ocular
squared disc type having finer subdivisions in one or more of the
squares. The art of measuring dacetines is discussed at length else-
where (Brown, 1953d: 7-15), so I shall repeat here just the essentials
for use with this paper.

Head length (HL), maximum measureable length of head proper
as seen from dorsal full-face view, including all of clypeus and occipi-
tal lobes.

Head width ( HW) is the maximum width of the cranium measured
in the same view as for HL.

Mandible length (ML), exposed length of mandibles, including
apical teeth, measured in same view from which HL is obtained.

Weber’s length (WL), oblique length of alitrunk from side view,
measuring from base of anterior pronotal declivity to metasternal
extremity.

Total length (TL) of the body is the summed lengths of ML, HL,
WL plus the axial lengths of petiole, postpetiole and gaster measured
separately.

Cephalic index (Cl), head width expressed as a percentage of head
length, or HW/HL X 100.

Mandibulo-cephalic index (MI), ML/HL X 100.

In addition to the dichotomous key to the species, I have constructed
a table giving known ranges of values for the seven quantitative
characters most used in species-level taxonomy of the genus. This
table may be used either as a primary key or as a check on the deter-
minations made with the dichotomous key. Number of individuals

1962]

Brown — Strumigenys

255

Figure 30. Diagram to illustrate the possible phyletic relationships among
the species-groups of New World Strumigenys, based upon the hypothesis that
the group of S. mandibularis represents the primitive stock in this hemisphere.
S. rogvri, being African in origin, does not figure in this scheme.

256

Psyche

[December

and localities on which the measurements were based are given under
“Sample” so that the user can judge crudely how nearly the natural
variance may be represented by the range of values given.

Following the dichotomous key is a glossary of the most important
morphological terms used in species identification.

Table of the most valuable quantitative characters of the
Strumigenys species of the New World

The table is arranged in order of size as based chiefly on head
length. The measurements (in hundredths of millimeters) HL (head
length), ML (mandibular length) and WL (Weber’s diagonal length
of alitrunk), as well as the proportions Cl (cephalic index) and MI
(mandibulo-cephalic index), are explained in the preceding section.
ID indicates the number of small teeth or denticles lying between the
two main teeth of the apical fork of the mandible, and PT is the
number of teeth along the inner margin of the mandible proximad of
the dorsal apical tooth (not counting the basal lamella, which is usual-
ly hidden beneath the clypeal margin) ; these preapical teeth may be
large and spiniform or may be reduced to exceedingly minute denticles.
The sample indicates the number of specimens measured and, follow-
ing a dash, the number of separate localities represented by the speci-
mens.

Species

HL

Cl

ML

MI

WL

ID

PT

Sample

mandibularis

131

- — 402

' — '51

' — 38

—

1

2

1

godmani

106-120

87-93

51-55

46-48

109-122

1

2

8-2

sanctipauli

98

76

96

98

103

1

2

1

fairchildi

96

75

59

61

101

2

2

1

precava

87-101

71-79

50-56

54-61

81-97

1

1

92-7

sp. nr. micretes

86-90

75-76

58-61

67-68

92-99

1

0-2

25-1

saliens

81-95

69-77

50-60

60-65

82-97

1

1

38-9

dolichognatha

84-90

75-77

104-108

119-124

—

1

2

6-1

cultriger

85

77

51

60

90

1

2

1

planeti

80-89

80-86

45-50

52-57

80-90

1

2

39-5

tococae

80-85

75

44-47

55

80-87

2

2

8-1

trimdadensis

78-83

78-82

52-56

66-69

80-88

1

2

12-6

longispinosa

78-80

72-74

71-72

90-91

80-81

0

1

2-1

cordovensis

73-82

76-80

83-92

109-123

—

1

2

40-8

prospiciens

74-80

79-83

41-46

56-60

75-82

1

2

8-3

sublonga

74-79

76-80

72-75

96-98

73-79

1

2

7-1

rehi

76

76

76

100

80

1

2

1

borgmeieri

74

65

52

67

77

1

2

1

lacacoca

70-76

67-71

45-53

60-70

72-82

1

0-1

8-2

1962] Brown — Strumigenys 257

micretes

70-76

71-74

45-51

62-68

72-

1

2

18-2

smithii

67-76

80-87

36-42

52-61

70-78

1

2

26-6

biolleyi

60-81

78-85

35-51

58-66

-86

1

2

51-11

ludia

67-74

81-84

49-56

73-76

62-73

1

0

9-6

marginiventris

68-73

78-82

51-57

75-79

68-72

0

1

11-3

producta

65-70

78-82

45-48

69-70

69-74

2

1

15-1

consanii

64-66

78-80

41-42

63-64

76-80

0

0

5-1

hindenburgi

60-69

83-88

38-40

58-64

63-71

1

2

9-3

deltisquama

62-67

95-97

34

51-55

60-63

1

2

12-1

xenognatha $

65

79

41

63

84

2

2

1

connectens

60-65

81-86

33-37

53-56

64-73

2

3-4

7-2

nevermanni

61-63

77-80

36-38

58-60

67-

1

1

8-1

emeryi

59-64

80-82

38-40

61-64

65-69

1

2

10-4

laevipleura

58-62

79-82

33-35

56-57

65-66

2

3

3-1

hemidisca

60

82

39

65

64

1

2

3-1

rogeri

57-62

70-74

30-34

53-55

—

0

2

11-9

trieces

58

78

28

48

62

2+?

3

1

elongata

50-63

72-79

28-41

56-67

47-62

0

0

161-25

ogloblini

55-56

80-81

28-29

51-52

' — '55

1

1

2-1

louisianae

46-64

77-92

23-37

49-67

—

2

1

130-71

mixta

51-59

80-84

28-32

54-60

51-60

2

2

20-2

jamaicensis

52-56

81-82

40-42

75-77

55-57

2

6-8

12-2

boneti

50-56

80-82

28-30

50-60

50-57

1

1

4-3

subedentata

48-56

82-86

26-30

53-54

47-60

2

4-6

36-15

denticulata

42-48

77-80

31-39

74-83

42-49

2

5-9

11-6

gundlachi

40-49

79-86

24-32

59-66

40-50

2

4-9

36-20

silvestrii

42-47

77-81

24-26

55-60

41-50

1

2

11-4

eggersi

39-47

83-88

22-27

56-64

39-46

2

4-8

32-22

schmalzi

42

79

27

64

44

0

2

1

perparva

40

81

20

50

40

0

1

2-1

carinithorax

36

81-83

21-22

59-61

41

0

2

3-1

Dichotomous key to the known species of Strumigenys occurring in the
New World, based chiefly on the worker caste, but applying
to the females of most species as well

1. Apical fork of mandible without distinct intercalary teeth or

denticles (Fig. 17) 2.

Apical fork of mandible with a single intercalary tooth or denti-
cle, either separate or occurring as a spur on the inner side of the

ventral tooth (Figs. 18, 19) 9.

Apical fork of mandible with 2 (rarely 3-4) intercalary denticles
(Figs. 20, 21) 38.

2. Mandible without preapical teeth or denticles (Fig. 1) 3.

Mandible with 1 or 2 preapical teeth and/or denticles (Fig. 16)

4.

258 Psyche [December

3. Postpetiole large and convex, its dorsum smooth and shining;
larger, more robust species (Costa Rica) consanii Brown

Postpetiole small, its dorsum densely punctulate and opaque;
smaller, more slender species (s. Mexico to se. Brazil and Bolivia)

elongata Roger

4. First segment of gaster margined for its full length on each side
by a strong, raised dorsolateral carina (Costa Rica to Colombia)

marginiven.tr is Santschi

First gastric segment smoothly rounded dorsolaterally, without

raised margins apart from the basal costulae 5.

5. Larger species with very long mandibles; combined length of
head + mandibles > i.iomm (Panama) .... longispinosd Brown
Smaller species with mandibles not so long; combined length of
head + mandibles < i.iomm 6.

6. Compound eye anteriorly detached, i.e., bounded in front by a

narrow cleft or notch in the ventrolateral margin of the head
(Fig. 29) ; combined length of head + mandibles > 0.80 mm;
2 preapical teeth on each mandible, the distal being smallest
(Figs. 16, 17) (West Indies, Trinidad, British Guiana, intro-
duced from Africa) rogeri Emery

No preocular notch in ventrolateral border of head; combined
length of head + mandibles < 0.80 mm 7.

7. Each mandible with a single preapical tooth ; no minute denticle

near mandibular midlength (Trinidad to se. Brazil)

perparva Brown

In addition to the preapical tooth, each mandible bears a minute
denticle somewhere near the midlength of its inner margin 8.

8. Mandibles (MI 54-61), scape (L0.23 mm) and apical funicular
segment (L ca. 0.22 mm) shorter; promesonotum with a distinct

median longitudinal carina (Dutch Guiana)

carinithorax Borgmeier

Mandibles (MI >61), scape (L > 0.27 mm) and apical funi-
cular segment (L > 0.25 mm) longer; no distinct carina in the
middle of the promesonotum (se. Brazil) schmalzi Emery

9. Mandible with no preapical teeth, or with a single preapical
tooth or denticle, or with a preapical tooth or denticle plus another

minute denticle proximal to it (Figs. 1-4, 7, 8, 12, 13) 10.

Mandible with 2 well-developed preapical teeth (Figs. 9, 14).

22.

to. Mandible without preapical teeth or denticles (Fig. 2) 11.

Mandible with a preapical tooth or denticle, or both (Figs. 3, 4,

7, 8, 12, 13, 15) 12.

11. Petiole claviform, the node only feebly differentiated from its

1962]

Brown — Striunigenys

259

anterior peduncle; gastric hairs mostly stiff, remiform (i.e., with
broadened apices) (Fig. 5); head broader (Cl > 79; Fig. 1)

(Nicaragua to s. Mexico) ludia Mann

Petiolar node with a steep anterior face, set off from its anterior
peduncle; gastric hairs long, finely flagelliform (Fig. 6) ; head
narrower (Cl < 78; Fig. 2) lacacoca; go to 2 1

12. Large hairs of gastric dorsum remiform (i.e., oar- or paddle-
shaped at apex) ; smaller species 13-

Large hairs of gastric dorsum fine, long, flagelliform 15.

13. First gastirc tergite reticulate-punctulate and opaque; preapical
armament of mandible reduced to a single minute denticle situ-
ated somewhat distal to the midlength of the inner border, but

remote from the apex (Fig. 12) (s. Mexico) boneti Brown

First gastric tergite smooth and shining beyond the basal belt of
longitudinal costulae ; preapical armament of mandible consisting
of a distinct tooth, with or without an additional minute denticle
near mandibular midlength 14.

14. Dorsal borders of antennal serobes broad, lamellose; preapical
armament of mandible consisting of a single strong tooth (n.

Argentina) ogloblini Santschi

Dorsal scrobe borders merely narrowly cariniform; preapical
mandibular armament consisting of a tooth near the apex plus
an additional minute denticle near the midlength (n. Argentina
to s. Brazil; also Cuba and Louisiana, where probably intro-
duced) silvestrii Emery

15. Ventral ends of propodeal lamellae at most rounded or bluntly

angulate, not dentiform 16.

Propodeal teeth large and acute, matched on each side below by
a (metasternal) tooth of nearly the same size and shape arising

from the ventral end of the infradental lamella (Fig. 23)

trinidadensis ; go to 26.

16. Long fine flagelliform hairs on nodes of petiole and postpetiole
and on gastric dorsum very numerous, too many to count, and
evidently always > 16 + 20 + 50, or > 86 total (partially
denuded specimens or those with hairs plastered down can of

course be deceptive) 1 7.

Long flagelliform hairs of nodes and gastric dorsum much fewer,
at most about 8 + 10 + 34, or about 52 hairs total 18.

17. Dorsal scrobe border on each side produced as a narrow but
distinctly lamellar margin ; inner mandibular border with a
minute denticle near the apical third, in addition to the preapical
tooth ; basal gastric costulae short, coarse, remainder of first

26o

Psyche

[December

tergite smooth and shining (Fig. 8; Argentina, se. Brazil)

hindenburgi Forel

Dorsal scrobe borders merely narrowly cariniform, not lamellate ;
mandibles without preapical dentical proximad of preapical tooth ;
basal costulae of gastric dorsum extended over basal third or
more of first tergite as fine, sericeous-opaque striolation (Fig. 4;
C. America to s. Mexico; Bahamas) lanuginosa Wheeler

18. Head with a strong concavity anterior to each eye, and thus
appearing constricted in full-face view (Fig. 7) ; humeral tuber-
cles large and produced (mandibles broad, contiguous or nearly
so when closed, each with a single short, broad preapical tooth;

Fig. 7) (Panama; hylaean S. America to Bolivia)

precava Brown

Head parallel-sided or gently tapered in front of eyes, without
marked preocular concavities; humeral tubercles or angles small,
not produced (mandibles slender, not contiguous at full closure,
preapical dentition diverse, but not as above) 19.

19. Smaller species, with mandibles < 0.42 mm long; head broader

(Cl > 75); ; _ 20.

Larger species, with longer mandibles (ML > 0.42 mm) ; head
narrow (Cl 75 or less) 21.

20. Inner mandibular margin with a minute denticle near the apical

third in addition to the preapical tooth ; head distinctly longitudi-
nally rugulose (s. Mexico, Honduras) emeryi Mann

Mandible with a single preapical tooth and no additional denticles
proximad of this (Fig. 3) ; head at most weakly and indistinctly
rugulose above (Costa Rica) nevermanni Brown

21. Mandible with a small preapical tooth or denticle and, near it
proximally, an additional minute denticle (Fig. 13; Costa Rica,

Panama) rnicretes Brown

Mandible with no teeth or denticles, or with a single minute

preapical denticle (Fig. 2; Panama) lacacoca Brown

(A population from Boquete, Chiriqui Prov., Panama, has 1 or
2 preapical denticles on the inner mandibular border, and is thus
intermediate between rnicretes and lacacoca , but this population
is also distinctive in having the promesonotum coarsely longitudi-
nally striate. The specimens are also larger. Possibly rnicretes
lacacoca and the Boquete sample are simply local variants of a
single unusually variable species, or perhaps three distinct species
are represented here.)

22. Inner mandibular border extended as a straight-edged lamella
that terminates abruptly and subangularly at its distal end near

1962]

Brown — Strumigenys

261

the proximal preapical tooth (Fig, 9) 23.

Inner mandibular border without a lamellar extension, or, if a
lamella of sorts is present, its form is not as above (Fig. 14) .. 24.

23. Lamelliform margin of inner mandibular border ending near mid-
length of mandible (Fig. 9) ; propodeal teeth very small; gastric
dorsum predominantly smooth and shining, with vestiture of
abundant fine, short reclinate hairs (se. Brazil) .. cultriger Mayr
Lamelliform margin of inner mandibular border ending near
apical quarter of mandible; propodeal teeth large; first gastric
tergite predominantly reticulate-striate, opaque, with about 20

apically-broadened, short erect hairs (Panama)

deltisquama Brown

24. Large forms (head width > 0.85 mm) with massive head and

short, heavy mandibles (MI 50 or less) 25.

Smaller forms with narrower heads (head width under 0.85 mm)
and longer, more slender mandibles (MI > 50) 26.

25. First gastric tergite finely longitudinally striolate for most or all
of its length; head about as broad as, or broader than, long

(Amazon Basin) mandibularis Fr. Smith

Gastric dorsum smooth and shining, with only a narrow basal
band of reduced costulae; head slightly longer than broad
(Guiana to Costa Rica) godmani Forel

26. Gastric dorsum predominantly finely longitudinally striolate,

sericeous-opaque, with very abundant, fine, erect flagelliform
pilosity; proximal preapical mandibular tooth small (only about
half the length of the distal preapical) and situated toward the
mandibular midlength (Fig, 14; Trinidad to Bolivia and Ecua-
dor) trinidadensis Wheeler

Gastric dorsum with either sculpture or pilosity or both different
from the above ; proximal preapical tooth of mandible more than
half as long as distal preapical tooth and situated well beyond the
mandibular midlength 27.

27. Mandibles very nearly as long as, to distinctly longer than, the

head proper (MI > 90) 28.

Mandibles relatively shorter (MI < 75) 32.

28. Mandibles slightly > 1.00 mm long; distal preapical tooth closer

to proximal than to apical fork (British Guiana)

dolichognatha Weber

Mandibles < 1.00 mm long; distal preapical tooth closer to
apical fork than to proximal preapical tooth 29.

29. Mandibles > 0.80 mm long; longitudinal costulation of post-

petiolar disc absent or incomplete 30.

262

Psyche

[December

Mandibles < 0.80 mm long; longitudinal costulation on post-
petiolar disc complete from anterior to posterior border 31.

30. Size larger (HL 0.98 mm in holotype worker) ; infradental
lamella of propodeum low and cariniform, terminating below in
a small, obtuse ventral angle that is much smaller than the dorsal
tooth (Fig. 24) ; antennal scape straight to its base (se. Brazil)

sanctipauli Kempf

Size smaller (HL < 0.85 mm) ; infradental lamella high, termi-
nating below in a prominent tooth or angle which is subequal
to, or often larger than, the dorsal propodeal tooth (Figs. 25-27) ;
antennal scapes gently but distinctly curved in basal half (s.
Mexico to Trinidad and the Guianas) cordovensis Mayr

31. Pilosity of head, alitrunk and nodes rather abundant and con-
spicuous, narrow-spatulate ; eyes 0.09 mm in greatest diameter;

MI 1 00 db (Amazon Basin) rehi Forel

Pilosity less abundant and conspicuous, that of nodes and first
gastric segment sparse, fine and filiform; eyes 0.07-0.08 mm in
greatest diameter; MI 94-99 (Amazon Basin) .. sublonga Brown

32. Propodeal lamellae evenly rounded, without dorsal or ventral

angles or teeth (Fig. 22; Colombia) hemidisca Brown

Propodeal lamellae angulate or toothed above and/or below,
more or less as in Figs. 23, 24 or 28 33.

33. Propodeal lamellae without dorsal teeth or angles; ventral angle

present and prominent (Fig. 28 ; C. America s. to Ecuador)

biolleyi Forel

Propodeal lamellae angulate or toothed both above and below
(more or less as in Figs. 23-26) 34.

34. Mandibles longer and more slender (MI > 63) ; head narrower

(CK80) 35.

Mandibles shorter and more robust (MI < 63) ; head broader
(Cl 80 or more) 36.

35. Smaller (HL < 0.80 mm), more slender (Cl < 68) ; preapical
teeth small and crowded toward apical fork, occupying little or
no more than the apical 1/5 of the mandibular length (Brazil:

Pernambuco) borgmeieri Brown

Larger (HL 0.80 mm or more), not so slender (Cl > 68) ;
preapical teeth more widely spaced, occupying the apical 2/5,
more or less, of the mandibular length (se. Brazil, n. Argentina)
s aliens Mayr

36. Larger (HL 0.80 mm or more), with heavy, distinctly arcuate

mandibles planeti Brown

1962]

Brown — Strumigenys

263

Smaller (HL < 0.80 mm) ; mandibles narrower, not or only
indistinctly arcuate 37*

37. Basigastric costulae absent or extremely reduced, never much
longer than the space separating one from the next ; anterodorsal
face of petiolar node convex in both directions (Amazon Basin

to Bolivia) prospiciens Emery

Basigastric costulae fine but numerous, extending at least 1/8 the
length of gastric tergite I; anterodorsal face of petiolar node
obliquely depressed, nearly or quite plane (tropical S. and C.
America, St. Vincent I.) smithii Forel

38. Mandible with a single small preapical tooth; no additional pre-

apical teeth or denticles on inner border 39.

Mandible with 2 or more preapical teeth and/or denticles ... 40.

39. Larger form with long mandibles (ML > 0.42 mm; MI 68 or

more; see discussion, p. 247) (Bolivia) producta Brown

Smaller form with shorter mandibles (ML < 0.42 mm; MI <
68) (Tennessee and Arizona to n. Argentina and Bolivia, W.
Indies) louisianae Roger

40. Mandible with at most 2 preapical teeth and/or denticles (Fig.

15) 41-

Mandible with 3 or more preapical teeth and/or denticles (Figs.
10, 11) 44.

41. Mandible with 1 preapical tooth and a single additional minute
denticle near the apical third of the mandibular length (Fig. 15);
gastric dorsum predominantly reticulate-striate, opaque, with stiff

remiform erect hairs (Guatemala) mixta Brown

Mandible with two well-developed preapical teeth 42.

42. Smaller species, HL < 0.75 mm; erect hairs of gaster stiff, slight-

ly clavate or remiform (known from female only; possibly an
inquiline in nest of S. laevipleura; Colombia)

xenognatha Kempf

Larger forms, HL > 0.75 mm ; erect hairs of gaster few, strag-
gling fiagelliform 43.

43. Dorsum of basal gastric segment with longitudinal costulae only

at base, otherwise smooth and shining; short, thickened reclinate
ground hairs of gastric dorsum abundant and conspicuous (Ama-
zon Basin) tococae Wheeler

Dorsum of basal gastric segment longitudinally striolate for its
full length; reclinate ground hairs of gastric dorsum obsolete or
apparently so (Panama) fairchildi Brown

44. Postpetiolar node smooth and shining when clean ; mandible with
2 preapical teeth and a denticle (Colombia) .... laevipleura Kempf

264

Psyche

[December

Postpetiolar node densely reticulate-punctulate, opaque 45.

45. Preapical armament of mandible consists of 2 close-set preapical
teeth, of which the second is much the longer, followed closely

basad by 1 or 2 denticles (Fig. 11) (Colombia)

connectens Kempf

Preapical armament otherwise; consisting either of three small
subequal teeth, or of 4-9 minute teeth and/or denticles (Fig. 10)
46.

46. Antennal scape 0.33 mm or more long; larger, dark-colored

species (Jamaica) jamaicensis Brown

Antennal scape < 0.33 mm long 47.

47. Mandibles short and thick (MI < 56) ; robust species, worker

HL mostly > 0.48 mm 48.

Mandibles longer and slender (MI 56 or more) ; smaller species,
worker HL mostly 0.48 or less (Fig. 10) 49.

48. Mandible short (MI 48 in unique holotype), with exactly 3
small preapical teeth ; ground pilosity of head nearly or quite

obsolete ; pronotum markedly flattened (Costa Rica)

trieces Brown

Mandible relatively longer (MI ca. 53-54), with > 3 preapical
teeth and/or denticles; ground pilosity abundant and conspicuous
on head ; promesonotum strongly rounded, not depressed ( Mexico
to s. Brazil) subedentata Mayr

49. Mandibles very long and slender (MI > 70), bowed outward

(Trinidad to n. Argentina) denticulata Mayr

Mandibles not so long (MI < 70), their shafts approximately
straight (Fig. 10) 50.

50. Ventral spongiform appendages of postpetiole small but distinctly
developed (side view) ; gastric dorsum of worker predominantly
smooth and shining (when clean!), at most with a few basal
longitudinal costulae, but female gaster commonly shagreened

above (Caribbean countries) gundlachi (Roger)

Ventral spongiform appendages of postpetiole obsolete; gastric
dorsum of both worker and female with fine, mostly opaque

reticulation (Brazil, Bolivia, Caribbean countries)

eggersi Emery

Glossary

Alitrunk: The second tagma of the body in Hymenoptera, incor-
porating the thorax and the closely fused propodeum (first true abdom-
inal segment).

1962]

Brown — Strumigenys

265

Antennal scrobes: Broad longitudinal excavations or grooves, one
on each side of the head above the eye, for the reception of the folded
antennae.

Apical fork: The two large teeth at the extreme apex of the mandi-
ble, converging to form a U or V ; between them may occur one or
more intercalary denticles (Figs. 9-21).

Basal costulae (basigastric costulae) : Numerous raised longitudinal
lines of the integumental sculpture originating from the base of the
first gastric segment (tergite) and extending caudad for distances
varying with the species (Figs. 5, 6) .

Basal tooth (or lamella) : A process, usually digitiform or denti-
form, arising from the inner mandibular border at its base, and usually
covered by the clypeus when the mandibles are closed; not to be con-
fused with the preapical teeth.

Flagelliform hairs: Very long, slender, tapered setae, often wavy,
looped or otherwise contorted, i.e., whip-like (Figs. 4, 6).

Intercalary tooth (or denticle) : Abbreviated “ID,” a tooth (or
denticle) occurring between the main teeth of the apical fork of the
mandible, or as a spur on the inner side of one of the main teeth (Figs.
18-21 ) .

Preapical tooth (or denticle) : Abbreviated “PT,” a tooth (or
denticle), one or more of which occur along the inner mandibular
border proximal to the apical fork, but not at or beneath the clypeal
margin; not to be confused with the basal tooth, q. v. (Figs. 3, 4,
7-16).

Propodeal lamella: One of a pair of raised lobes or flanges guard-
ing the sides of the propodeal declivity, sometimes incorporating the
(dorsal) propodeal tooth and/or a ventral (metasternal or meta-
pleural) tooth or angle (Figs. 5, 6, 22-28).

Remiform hairs : Setae with a more or less oar-like form ( Fig. 5 ) .

Spongiform appendages : Lobes, flaps and collar-like strips of light-
colored spongy integumental material situated in definite, symmetrical
positions on the petiole, postpetiole and anteroventral face of the gaster
(Figs. 5, 6), and sometimes even on the alitrunk, in the higher dace-
tines and a few other ants. Their function is unknown.

266

Psyche

[December

INDEX TO NAMES OF STRUMIGENYS SPECIES OF
THE NEW WORLD AND THEIR SYNONYMS

(Names in italics are synonyms; page references are to the accompanying

article)

angusticeps — saliens, 240,
banillensis — gundlachi, 249
batesi — mandibularis, 239
berlesei — gundlachi, 249
bierigi — gundlachi, 249
biolleyi, 240, 253, 257, 262
boneti, 245, 257, 259
borgmeieri, 240, 253, 257, 262
bruchi — louisianae, 247
caribbea — silvestrii, 246
carinithorax, 246, 257, 258
clasmospongia — louisianae, 247
clavata — subedentata, 248
connectens, 247, 253, 257, 264
consanii, 245, 257, 258
cordovensis, 241, 253, 256, 262
costaricensis — louisianae, 247
cubaensis — louisianae, 247
cultriger, 241, 253, 256, 261
deltisquama, 241, 257, 261
denticulata, 249, 257, 264
dolichognatha, 241, 253, 256, 261
eggersi, 249, 257, 264
elongata, 245, 253, 257, 258
emeryi, 245, 253, 257, 260
fairchildi, 241, 242, 256, 263
ferox — godmani, 239
fusca — louisianae, 247
godmani, 239, 253, 256, 261
guatemalensis — louisianae, 247
gundlachi, 249, 253, 257, 264
hemidisca, 239, 257, 262
hindenburgi, 243, 253, 257, 260
imitator — elongata, 245
inaequalis — smithii, 239
incisa — rogeri, 250
infidelis — louisianae, 247
infuscata — gundlachi, 249
isthmica ~ gundlachi, 249
jamaicensis, 249, 257, 264
lacacoca, 245, 256, 259, 260
laevipleura, 247, 248, 257, 263
lanuginosa, 243, 260

laticephala — louisianae, 247
longicornis — louisianae, 247
longispinosa, 242, 256, 258
louisianae, 246, 247, 253, 257, 263
luctuosa — biolleyi, 240
ludia, 242, 243, 253, 257, 259
mandibularis, 238, 252, 256, 261
marginiventris, 242, 257, 258
micretes, 245, 256, 257, 260
mixta, 246, 253, 257, 263
mokensis, 241
nevermanni, 245, 257, 260
nicaraguensis — elongata, 245
obscuriventris — louisianae, 247
ogloblini, 243, 253, 257, 259
perparva, 246, 253, 257, 258
planed, 239, 253, 256, 262
precava, 243, 245, 252, 256, 260
procera — saliens, 240
producta, 247, 257, 263
prospiciens, 238, 239, 256, 263
rehi, 240, 241, 256, 262
rogeri, 249, 250, 257, 258
saliens, 240, 253, 256, 262
sanctipauli, 240, 253, 256, 262
schmalzi, 246, 257, 258
silvestrii, 246, 253, 257, 259
smithii, 239, 252, 257, 263
soledadensis — louisianae 247
subedentata, 248, 257, 264
sublonga, 240, 256, 262
sulfurea — rogeri, 250
tenuis — ludia, 243
tococae, 241, 253, 256, 263
tridens — biolleyi, 240
trieces, 248, 257, 264
trinidadensis, 240, 253, 256, 259, 261
tristani — subedentata 248
unidentata — louisianae 247
unispinulosa — louisianae 247
vincentensis — gundlachi 249
xenognatha, 248, 257, 263

1962]

Brown — Strumigenys

267

References

Brown, W. L., Jr.

1948. A preliminary generic revision of the higher Dacetini (Hymenop-
tera: Formicidae). Trans. Amer. Ent. Soc., 74: 101-129.

1953a. Three new ants related to Strumigenys louisianae. Psyche, 60: 1-5.

1953b. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of mandibularis Fr. Smith. Jour. N. Y. Ent. Soc., 61: 53-59.

1953c. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of smithii Forel. Jour. N. Y. Ent. Soc., 61 : 101-110.

1953d. Revisionary studies in the ant tribe Dacetini. Amer. Midi. Nat..
50: 1-137, ill.

1954a. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of elongata Roger. Jour. N. Y. Ent. Soc., 61: 189-200, 1953.

1954b. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of saliens Mayr. Jour. N. Y. Ent. Soc., 62: 55-62.

1957. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of cultriger Mayr and S. tococae Wheeler. Jour. N. Y. Ent.
Soc., 63: 97-102, 1955.

1958a. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of cordovensis Mayr. Stud. Ent., Petropolis, Brazil, (n.s.)
1 : 217-224.

1958b. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of marginiventris Santschi. Jour. N. Y. Ent. Soc., 65: 123-
128, 1957.

1958c. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of ogloblini Santschi. Jour. N. Y. Ent. Soc., 65 : 133-137,
1957.

1959a. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of emeryi Mann. Ent. News, 70: 97-104.

1959b. Some new species of dacetine ants. Brev. Mus. Comp. Zool., 108:
1-11.

1959c. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of silvestrii Emery. Stud. Ent., Petropolis, Brazil, (n.s.) 2:
25-30.

1960. The neotropical species of the ant genus Strumigenys Fr. Smith:
Group of gundlachi (Roger). Psyche 66: 37-52, 1959.

1961. The neotropical species of the ant genus Strumigenys Fr. Smith:
Miscellaneous concluding studies. Psyche, 68 : 58-69.

Kempf, W. W.

1958a. Three new ants of the genus Strumigenys from Colombia. Rev.
Brasil. Ent., 8 : 59-68.

1958b. The ants of the tribe Dacetini in the State of Sao Paulo, Brazil,
with the description of a new species of Strumigenys (Hymenop-
tera: Formicidae). Stud. Ent., Petropolis, Brazil, (n. s.) 1 : 553-560.

Weber, N. A.

1952. Biological notes on Dacetini (Hymenoptera, Formicidae). Amer.
Mus. Novit., 15 54: 1-7.

Wilson, E. O.

1950. Notes on the food habits of Strumigenys louisianae Roger (Hymen-
optera: Formicidae). Bull. Brooklyn Ent. Soc., 45: 85-86.

1954. The ecology of some North American dacetine ants. Ann. Ent.
Soc. Amer., 46: 479-495, 1953.

THE EGG-SAC IN THE IDENTIFICATION OF SPECIES
OF LATRODECTUS (BLACK-WIDOW SPIDERS)1

By J. W. Abalos

Institute) Nacional de Microbiologia,

Santiago del Estero, Argentina

The genus Latrodectus comprises a few species very hard to classify.
The geographical distribution of some spreads across several contin-
ents, making the problem even more complex and, at the same time,
the synonymy richer. The difficulty lies in the lack of taxonomic
characters which allow true differentiation among the species. Levi
(J959)2 used for species identification, among other devices, the male
genitalia as well as the female; thanks to this the problem of system-
atics of so ill-treated a group begins to be clearer.

In the course of the research we are carrying out on Latrodectus
in Santiago del Estero (north-central region of Argentina) we have
recognized the presence of five different species living in the same area.
We were able to separate two of them according to their morphological
characters ; L. geometricus C. L. Koch, a domestic species we identified
by its well-defined color and genitalic characters, among them the coils
of the embolus of the male palpus, coils corresponding to those in the
connecting ducts of the female. The other species, temporarily named
Latrodectus No. i in order not to add further confusion of names,
resembles L. curacaviensis (Muller) in the genitalic characters pointed
out by Levi : the embolus of the male palpus and the connecting ducts
of the female with two coils. The three remaining species correspond
to what we call “L. mactans complex” with their three coiled embolus
and female ducts. The morphological characters are not enough to
separate these species. The width of the red spots on the abdomen
guides the identification but is not reliable. The three of them live in
the same area and ecologic environment. The only elements that allow
us at present to separate these species in an accurate way are the size,
shape and texture of the egg-sacs. We have temporarily called these
species: Latrodectus No. 2, Latrodectus No. 3, and Latrodectus No. 4.
However, many names are available for the species,

Latrodectus No. 2: The egg-sac is white when just formed, but
usually turns yellowish in the course of days; it is spheroidal, slightly

Research under the sponsorship of Consejo Nacional de Investigaciones
Cientificas y Tecnicas of Argentina.

2Levi, H. W. 1959. The spider genus Latrodectus (Araneae, Theridiidae)
Trans. Amer. Micros. Soc., 78(1) :7-43.

Manuscript received by the editor October 31, 1962.

268

Psyche, 1962 Vol. 69, Plate 19

Abalos — Latrodectus

Figs. 1-6. Egg-sacs of Latrodectus. Fig. 1. Latrodectus geometricus C. L.
Koch. Fig. 2. Latrodectus No. 1. Fig. 3. Latrodectus No. 2. Fig. 4. Latrodectus
No. 3. Figs. 5-6. Latrodectus No. 4.

270

A halos — Latrodectus

[1962

pyriform and its most frequent size is about io mm. The sac is formed
by threads 8 microns in diameter. They form a tissue, not very dense,
but tight, that constitutes a thin cover, soft to pressure of the fingers.
On the surface of the egg-sac we can observe numerous little web
pompons that give a characteristic granulous aspect (Fig. 3). These
little pompons are made by the spider with the hind legs; they are not
firmly joined to the surface of the egg-sac and they usually fall off
with handling.

Latrodectus No. 3 : The egg-sac of this spider generally has the
same shape as the one mentioned above, even though it sometimes
changes, up to the point of presenting the aspect of an elongated pear ;
its size varies, but it is always bigger than the former species, reaching
21 mm in its biggest diameter. The sac wall is thicker and is formed
by threads of a diameter of about 16 microns. These threads form a
tight tissue, with a parchment texture that makes the egg-sac resistant
to pressure. The egg-sac completely lacks pompons or any other orna-
ment on its surface (Fig. 4).

Latrodectus No. 4: The egg-sac is white, pyriform, and its common
size is about 10 mm. The sac-wall is thin, with threads of about 8
microns in diameter forming a less compact tissue. On this cover the
spider weaves a coat about 3 mm thick, of loose tissue, that gives; the
egg-sac the appearance of a spherical woolen tuft (Figs. 5, 6).

We can also observe that Latrodectus No. 1 differs from the other
species in its egg-sac. Its size is always smaller than in any of the
former species, about 7 mm, although it sometimes reaches 10 mm; its
shape is spherical pyriform. Its color is white, often turning greyish
in the course of days. The cover is thin with threads of about 8 mic-
rons in diameter, forming a very tight tissue of smooth surface, with
a paper texture, and completely lacking exterior ornaments (Fig. 2).
The egg-sac is fairly resistant to finger pressure.

The egg-sac of L. geometrious . many times figured, is very charac-
teristic: its color is whitish when just formed, turning yellowish in the
course of days. It is spherical; its size about 10 mm in diameter; it is
covered with conic protuberances that give it the curious aspect of a
war-mace (Fig. 1).

We are thankful to Dr. Herbert W. Levi, for his generosity and
advice on the publication of this note and to Emilio Clementi for
taking the photographs.

PSYCHE

INDEX TO VOL. 69, 1962

INDEX TO AUTHORS

Abalos J. W. The Egg-sac in the Identification of Species of Latrodectus
(Black-Widow Spiders). 268

Brady, A. R. The Spider Genus Sosippus in North America, Mexico, and
Central America (Araneae, Lycosidae). 129

Brown, W . L., Jr. A New Ant of the Genus Amblyopone from Panama. 73
A New Ant of the Genus Epitritus from South of the

Sahara. 77

The Neotropical Species of the Ant Genus Strumigenys
Fr. Smith: Synopsis and Keys to the Species. 238

Bush, G. L. The Cytotaxonomy of the Larvae of Some Mexican Fruit Flies
in the Genus Anastrepha (Tephritidae, Diptera). 87

Carpenter, F. M. A Permian Megasecopteron from Texas. 37

Chapin, E. A. Pseudoscymnus, a New Genus of Asiatic Scymnini (Coleoptera:
Coccinellidae) . 50

Crabill, R. E., Jr. A New Damothus and a Key to the North American
Dignathodontid Genera (Chilopoda: Geophilomorapha : Dignathodonti-
dae). 81

Darlington, P. J., Jr. Australian Carabid Beetles XI. Some Tachys. 117

Eisner, T. and G. M. Happ. The Infrabuccal Pocket of a Formicine Ant:
a Social Filtration Device. 107

Eisner, T. and F. C. Kafatos. Defense Mechanisms of Arthropods. X. A
Pheromone Promoting Aggregation in an Aposematic Distasteful Insect.
53

Ghent, R. L. and N. E. Gary. A Chemical Alarm Releaser in Honey Bee
Stings (Apis mellifera L.) . 1

Kmgsolver, J. M. Notes on Fossil Cleoninae (Coleoptera: Curculionidae) . 47

Levi, H. tV. More American Spiders of the Genus Chrysso (Araneae, Theri-
diidae). 209

The Spider Genera Steatoda and Enoplognatha in America
(Araneae T heridiidae) . 11

Lindroth, C. H. The Male of Platypatrobus lacustris Darlington (Coleoptera
Carabidae). 7

Roth, L. M. and Barbara Stay. A Comparative Study of Oocyte Development
in False Ovoviviparous Cockroaches. 165

Sharov, A. G. Redescription of Lithophotina floccosa Cock. (Manteodea)
with Some Notes on the Manteod Wing Venation. 102

Slater, J. A. A Remarkable New Genus of Lygaeidae from Sumatra (Hemip-
tera: Heteroptera) . 42

Wilson, E. O. The Trinidad Cave Ant Erebomyrma ( — Spelaeomyrmex)
urichi (Wheeler), with a Comment on Cavernicolous Ants in General. 62

271

INDEX TO SUBJECTS

All new genera, new species and new names are printed in capital type.

A chemical alarm releaser in honey
bee stings (Apis mellifera L.) 1
A comparative study of oocyte de-
velopment in false ovoviviparous
cockroaches, J65
ACTINOHYMEN RUSSELLI, 37
Alarm releaser in honey bee stings, 1
Amblyopone tropicalis, 73
Anastrepha aphelocentema, 95
Anastrepha distincta, 93
Anastrepha f rater cuius , 93
Anastrepha ludens, 91
Anastrepha mombinpraeoptans, 93
Anastrepha serpentina, 95
A nastrepha spatulata, 94
Anastrepha striata, 94
Anastrepha zuelaniae, 91
A new ant of the genus Amblyopone
from Panama, 73

A new ant of the genus Epitritus
from south of the Sahara, 77
A new Damothus and a key to the
North American dignathodontid
genera (Chilopoda: Geophilomor-
pha : Dignathodontidae) , 81
Ants, 62, 73, 77, 107, 238
A Permian megasecopteron from
Texas, 37
Apis mellifera, 1

A remarkable new genus of Ly-
gaeidae from Sumatra (Hemip-
tera), 42

Araneae, 11, 129, 209
Asiatic Scymnini, 50
Australian carabid bettles XI. Some
Tachys, 117
Bardohymenidae, 37
Bees, 1

Beetles, 7, 50, 117, 147
Black-widow spiders, 268
Blattaria, 165

Camponotus pennsylvanicus , 108
Carabidae, 7, 117
Cavernicolous ants, 62
Chaeteesidae, 102
Chilopoda, 81
Chrysso albomaculata, 215
Chrysso alecula, 234
Chrysso antonio, 220
Chrysso arima, 235
Chrysso arops, 228
Geophilomorpha, 81

Chrysso backstromi, 230
Chrysso barromachadoi , 223
Chrysso cambridgei, 224
Chrysso caraca, 230
Chrysso clemintinae, 231
Chrysso compressa, 226
Chrysso diplosticha, 216
Chrysso ccuadorensis , 225
Chrysso gounellei, 217
Chrysso indicifer, 216
Chrysso melba, 234
Chrysso nigrosternum, 217
Chrysso perplexa, 220
Chrysso pulchra, 229
Chrysso questona, 222
Chrysso ribeirao, 229
Chrysso rubrovittata, 218
Chrysso silva, 235
Chrysso simoni, 222
Chrysso vexabilis, 216
Chrysso vittatula, 224
Chrysso volcanensis, 236
Cleoninae, 47
Coccinellidae, 50
Cockroaches, 165
Coleoptera, 7, 47, 50, 117
Curculionidae, 47

Cytotaxonomy of the larvae of some
Mexican fruit flies in the genus
Anastrepha (Tephritidae, Dip-
tera), 87

Damothus alastus, 82
Defense mechanisms of Arthropods.
X. A pheromone promoting aggre-
gation in an aposematic distasteful
insect, 53

Dignathodontidae, 81
Diptera, 87

Egg-sac in the identification of
species of Latrodectus (black-
widow spiders), 268
Enoplogiiatha juninensis, 17
Enoploganatha maricopa, 15
Enoplognatha marmorata, 15
Enoplognatha peruviana, 16
Enoplognatha puno, 17
Enoplognatha selma, 15
Enoplognatha tecta, 15
Enoplognatha zapfei, 19
Epitritus laticeps, 77
Erebomyrma urichi, 62
Formicine ants, 107
Fruit flies, 87

272

Hemiptera, 42
Heteroptera. 42

Hymenoptera, 1, 62, 73, 77, 107, 238
Infrabuccal pocket of a formicine
ant: a social filtration device, 107
Latrodectus, 268
Lithophotina floccosa, 102
Lycosidae, 129
Lycus loripes, 54
Lygaeidae, 42
M ACELLORIS INCRASSATUS, 43
Male of Platypatrobus lacusiris
Darlington (Coleoptera: Carabi-

dae), 7

Manteodea, 102
Megasecoptera, 37
Mexican fruit flies, 87
More American spiders of the genus
Chrysso (Araneae, Theridiidae) ,
209

Neotropical species of the ant genus
Strumigenys Fr. Smith: synopsis
and keys to the species, 238
Notes on fossil Cleoniniae (Coleop-
tera: Curculionidae) , 47
Oocyte development, 165
Orthoptera, 102
Pheromone, 53
Platypatrobus lacustris, 7
pseudoscymnus, a new genus of
Asiatic Scymnini (Coleoptera: Coc-
cinellidae), 50

Redescription of Lithophotina floc-
cosa Cock. (Manteodea) with
some notes on the manteod wing
venation, 102
Scymnini, 50

Sosippus agalenoides, 146
Sosippus calif ornicus, 139
Sosippus floridanus, 151
Sosippus mexicanus, 145

Sosippus michoacanus, 148
Sosippus mimus, 156
Sosippus plutonus, 150
Sosippus texanus, 160
Spider genera Steatoda and Eno-
plognatha in America (Araneae,
Theridiidae), 11

Spider genus Sosippus in North
America, Mexico, and Central
America (Araneae, Lycosidae),
129

Steatoda ancorata, 34
Steatoda andina, 26
Steatoda chinchipe, 32
Steatoda diamantina, 31
Steatoda erigoniformis , 25
Steatoda grossa, 26
Steatoda iheringi, 31
Steatoda marta, 28
Steatoda moesta, 25
Steatoda porteri, 33
Steatoda sabulosa, 30
Steatoda triangulosa, 25
Steatoda variipes, 33
Strumigenys , 238, 267
Tachys bolellus, 127
Tachys bolus, 125
Tachys ectromioides, 124
Tachys fitzroyi, 123
Tachys hobariti, 118
Tachys kingi, 121
Tachys lutus, 120
Tachys murrumbidg ensis , 122
Tephritidae, 87
Theridiidae, 11, 209
Trinidad cave ant Erebo myrma
( — Splaeomyrmex) urichi (Whee-
ler), with a comment on cav-
ernicolous ants in general, 62

273

CAMBRIDGE ENTOMOLOGICAL CLUB

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The illustration of a spider on the front cover of this issue of
Psyche was reproduced from an original drawing made by J. H.
Emerton and contained in the Museum of Comparative Zoology.
First published in the Transactions of the Connecticut Academy of
Arts and Sciences (1909, vol. 14, plate 7, fig. 5), it shows the dis-
playing posture of the male of Habronattus viridipes (Hentz).

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