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ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
INDEX TO
VOLUME Ill
SEPTEMBER 1921-1923 OCTOBER
NUMBERS 1-23 INCLUSIVE
PUB LPs. B.D Buy fH E St 8s Sa! Bd ie! ae
Rit 2OOLOCGCTCAL: PARK, NEW YORK
L9O:2-5
%
2
+ Gon) ~ = . 4
P j . = Fate tn ering: S =i { Z
7 ial el Fe Eanes at eet a. Le . ,
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
‘ STATION IN BRITISH GUIANA
VOLUME III. NUMBERS 1 AND 2
a OBJECTS OF THE TROPICAL RESEARCH
STATION
By HENRY FAIRFIELD OSBORN,
President of the New York Zoological Society
2. CONTRIBUTIONS OF THE TROPICAL
‘ RESEARCH STATION, 1916 TO 1921 ~
By WILLIAM BEEBE,
Director of the Tropical Research Station and Honorary Curator of Birds
PUBLISHED BY THE OV. TBs
THE .ZOOLOGICAL PARK, NEW YORK
SEPTEMBER, 1921
New York Zonlogiral Society
General Office: 111 Broadway, New York City A
Officers
President, HENRY FAIRFIELD OsBoRN;
Vice-Presidents, MADISON GRANT and FRANK K. STURGIS;
Secretary, Chairman, Exec. Committee, MADISON GRANT;
Treasurer, PERCY R. PYNE.
Board of Managers
Glass nf 1922
Percy R. PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DODGE,
C. LEDYARD BLAIR, EMERSON McMILLIN, ANTHONY R.
KUSER, WATSON B. DICKERMAN, MORTIMER L.
SCHIFF, FREDERIC C. WALCOTT, BEEKMAN
WINTHROP, GEORGE C. CLARK,
W. REDMOND CROSS.
Glass nf 1923
HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F.
DIETERICH, GEORGE F. BAKER, WM. PIERSON HAMILTON,
ROBERT 8S. BREWSTER, EDWARD S. HARKNESS,
WILLIAM B. OSGooD FIELD, A. BARTON
HEPBURN, WILLIAM WOODWARD,
EDWIN THORNE, PERCY A.
- ROCKEFELLER.
Glass of 1924 ae
MADISON GRANT, WILLIAM WHITE NILES, HENRY A. C. ve
FRANK K. ‘STURGIS, GEORGE J. GouLp, OGDEN MILLS, :
LEWIS RUTHERFURD _ Morris, ARCHER M.
HUNTINGTON, GEORGE D. PRATT, T. COLE-
MAN DUPONT, HENRY D. WHITON,
EDWARD HATCH, JR.
Scientific Staff .
WILLIAM T. HORNADAY, Director of the Zoological Park;
CHARLES H. TOWNSEND, Director of the Aquarium;
RAYMOND L. DITMARS, Curator of Reptiles;
WILLIAM BEEBE, Honorary Curator of Birds and Director of the
Tropical Research Station;
LEE 8S. CRANDALL, Curator of Birds;
GEORGE 8. HUNTINGTON, Prosector;
GEORGE A. MACCALLUM, Pathologist;
W. REID BLAIR, Veterinarian;
ELWIN R. SANBORN, Photographer and Editor.
Editorial Comunittee
HENRY FAIRFIELD OSBORN, Chasrviine |
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND.
Corrected to September, 1921 ;
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 1
(Tropical Research Station Contribution Number 94)
1. OBJECTS OF THE TROPICAL RESEARCH
STATION
By HENRY FAIRFIELD OSBORN,
President of the New York Zoological Society
Elebini S Ee ED. « BY A EEA SOOM fT ee ¥.
Mite sOObLOGciCAL -~PARK, ~NEW YORK
SEPTEMBER, 1921
OAVLUVM LV NOILVIS HOUVASAY AHL T ‘Old
Volume III, Number 1
OBJECTS OF THE TROPICAL RESEARCH
STATION
By HENRY FAIRFIELD OSBORN.
The main object of the Tropical Research Station from the
beginning has been the observation of living organisms in their
natural environment. To make this intensive and inclusive of
the entire fauna, a single region has been selected and the fauna
studied as a whole and in detail so far as possible through the
expert knowledge of members of the staff and of the several
experts who have been invited to work at the laboratory from
time to time.
The work of the Station, therefore, is unique in two respects:
First, its very intensive character; second, its restriction to a
single locality on the border of a great zoological region, namely,
the eastern tropical forests of the Amazonian basin. In many
biological aspects it is new, in others it is the sequel to the long,
distinguished period of exploration of this general region, begin-
ning as early as 1812. The six best known explorers of eastern
South America have been the following:
Charles Waterton, four voyages from 1812 to 1824, chiefly
in Guiana.
-Charles Darwin, 1832-1835, from Bahia around the entire
South American coast to the Galapagos.
Alfred Russel Wallace, 1848-1852, the Amazon and Rio
Negro.
Henry Walter Bates, 1848-1859, the Amazon basin from
Para to Peru.
Thomas Belt, 1868, Nicaragua.
W. H. Hudson, born in Argentina seventy-odd years ago,
and left for England about 1890.
4 Zoologica: N. Y. Zoological Society [Iiis2
To Waterton we owe the pioneer review of the life of this
wonderful forest region; to Darwin, Wallace, Bates, and Belt
the intimate relations of animals and plants with each other and
with their environment, color and form adaptation, and struggle
for existence; to Darwin and Hudson especially the complex chain
of relations which connect the whole series of organisms together.
In the work of the Station it has been found absolutely
necessary to lay a secure foundation in systematic zoology. For
this purpose efforts are being made to complete and round out the
faunal lists of various systematic workers in this region.
These lists will give a common language to the distinctive
research feature of this Station, which, as remarked above, is
intensive biologic observation in one region, in fact, in one local-
ity, as distinguished from the observations of those who have
covered and are covering the whole biologic field of South
America.
The area chosen by our Honorary Curator of Birds, William
Beebe, when he founded the Station in 1916, is the eastern edge
of the tropical rain-forest of South America, which extends
unbroken across the greater part of the continent. The fauna
and flora are in general uniform with those of the entire Ama-
zonian region. The locality at Kartabo, Bartica District, British
Guiana, the point of junction of the Mazaruni and Cuyuni rivers,
demonstrated in the first season its exceptional advantages as
the site for a permanent station. Within ten minutes walk are
sandy and rocky beaches, mangroves, grassland, swamp, and
high jungle, each with a growth of life peculiar to itself. Free
exposure to the trade winds, the absence of flies and mosquitos,
invariably cool nights, excellent buildings assigned by the govern-
ment—all these features contribute to the wide range of life and
the unbroken health of the scientific staff. The work of the year
1916 was so full of promise that Mr. Beebe, then Curator of
birds of the New York Zoological Park, was promoted to the
rank of Director of the Tropical Research Station and given
entire charge both of the choice of the personnel and of the super-
vision of the scientific work.
The Station is now entering its sixth year. Owing to the
difficulty of transportation at the time of the war, there was a
1921] ‘Osborn: Research Station Objects 5
lapse during 1917, but work was resumed in 1918, continued in
1919, and the present VOLUME III of ZOOLOGICA opens the con-
tributions of the year 1920-1921, which has proved to be the most
productive of all.
The staff of specialists, artists and investigators has included
in the course of the past six years the following persons:
WILLIAM BreBeE, Director, Columbia University and
New York Zoological Society
General Evolutionary Problems in Ornithology and
Ecology
IRVING W. BAILEY Harvard University
Relations of Ants to Certain Plants
T. DONALD CARTER, Collector, New York Zoological
Society
ISABEL COOPER, Artist Bryn Mawr College
ALFRED EMERSON Cornell University
Life Histories of Kartabo Termites
GERTRUDE EMERSON University of Chicago
Anthropology
WINIFRED J. EMERSON, Artist Cornell University
J: F. M. FLoyD University of Glasgow
Parasites of Vertebrates
W. T. M. FORBES Cornell University
Organs of Hearing in Lepidoptera
H. GIFFORD University of Nebraska
Comparative Opthalmology
G. I. HARTLEY Cornell University
Relationships of Certain Non-oscine Birds
RACHEL HARTLEY, Artist New York
PAUL G. HOWES Bruce Museum
Studies in Hymenoptera
GEORGE W. HUNTER Carlton College
General Biology
BRITISH
SCALE 58°742'W.
5 MILES
FIG, 2. LOCATION OF THE TROPICAL RESEARCH STATION OF THE
NEW YORK ZOOLOGICAL SOCIETY
The circle represents a radius of six miles.
1921] Osborn: Research Station Objects a
CLIFFORD POPE University of Virginia
Life Histories of Kartabo Fish
ALBERT M. REESE University of West Virginia
Embryology of Crocodiles and Investigation of
Microscopical Beach Life
MABEL SATTERLEE ~ Columbia University
Coloration of Ameiva and Painting Optical Fundi
T. V. SMOLUCHA New Jersey
Photography and Pen-and-Ink Drawing
ANNA TAYLOR South Carolina
Botanical Paintings
JOHN TEE-VAN New York Zoological Society
Ecology of Certain Lepidoptera
Won. MorTON WHEELER Harvard University
Ants of Kartabo
C. A. Woop Leland Stanford University
Optical Fundi of Birds and Other Vertebrates
Ninety-three contributions have already been published from
the Tropical Research Station, of which a complete annotated list
is given in ZOOLOGICA Il, No. 2. Of these eighty-nine are scientific
papers and magazine articles, while four are included in the fol-
lowing bound volumes:
TROPICAL WILD LIFE IN BRITISH GUIANA, by Beebe, Hartley
and Howes.
INSECT BEHAVIOR, by Paul G. Howes.
JUNGLE PEACE, by William Beebe.
EDGE OF THE JUNGLE, by William Beebe.
These published observations of the Station are so broad in
scope that only a few salient features can be noted in this Intro-
duction. They extend from color changes and adaptations to
anatomical and physiological characters of the archaic forms of
life, like the hoatzin, as well as of the most highly modernized
and specialized forms. The colors of living amphibians and rep-
tiles are almost an untouched field, since all modern systematic
8 Zoologica: N. Y. Zoological Society (TIT; 2
zoology and description of these phyla have been founded on
alcoholic specimens, in which the colors are either modified or
lost altogether. It is only in the feathers of birds and in the coats
of mammals that the natural color hues can be preserved. The
paintings made directly from life by the artists of the Station,
Miss Cooper and Miss Satterlee, will be published in a series of
plate volumes accompanying the text volumes of ZOOLOGICA. The
opportunity of studying the faunal and floral complex and the
independent and interrelated adaptations in all grades of life,
both in the vertical and in the horizontal life zones, opens up
vistas for future research extending over many years. The ver-
tical division of the fauna and flora into distinctive life zones,
extending from the tree summits to the sub-soil, is a biologic
contribution of first importance.
The Station was honored during the year 1920-1921 by the
presence of Dr. Wm. Morton Wheeler, who makes an extremely
important contribution to entomology in the present volume of
ZOOLOGICA through his article on A Study of Some Social Beetles
in British Guiana and of Their Relations to the Ant-plant Tachi-
galia.
Of the work of the Staff of the Research Station the follow-
ing may be mentioned as having been accomplished to date:
Life History Notes on 445 species of birds, by Beebe
and Hartley.
Life History Notes on 106 species of sea and
amphibians, by Beebe.
One thousand five hundred and thirty-two photo-
graphic negatives, by Beebe, Howes, Tee-Van and
Smolucha.
Ten thousand: feet of moving picture film, by
Tee-Van.
Collection of 340 water color drawings by Isabel
Cooper.
Collection of types of 50 new species of termites.
Four hundred transparent preparations of embryos
and tongues,. etc.
1921] Osborn: Research Station Objects 2
Four hundred skeletons of mammals and birds.
Nests and eggs of 132 species of birds, many new to
science.
Materials for a monograph on the syrinx and the
voice of tropical birds.
Collection of 75,000 insects.
Collection of 776 bird skins.
Collection of 110 embryos of birds.
Materials for the study of the optical fundi of birds.
Monographic work on Trogons.
COOPERATION WITH THE ZOOLOGICAL PARK, THE AQUARIUM, THE
AMERICAN MUSEUM OF NATURAL HISTORY, AND
OTHER INSTITUTIONS
Besides the research work carried on at the Station there
are three general lines of cooperation with other institutions.
First, living organisms collected for the New York Zoological
Park and the New York Aquarium, among which the most inter-
esting forms are the following:
BIRDS
3 Cocks-of-the-Rock
Hawk-headed Parrot
Imperial Amazon Parrot
White-necked Rails
Bat Falcons
imThurn’s Blackbird
Etc.
MAMMALS
Silky Anteater
Tayra
2-toed Sloth
3-toed Sloth
Spotted Cavy
Red Howling Monkey
Jaguarondi
Wild Dog
Ete.
Rupicola rupicola (Linné)
Deroptyus accipitrinus accipitrinus (Linné)
Amazona imperialis (Richm.)
Porzana albicollis Vieill.
Falco rufigularis rufigularis Daud.
Agelaius imthurni Sclater
Cyclopes didactylus didactylus (Linné)
Tayra barbara barbara (Linné)
Choloepus didactylus (Linné)
Bradypus tridactylus Linné
Agouti paca paca (Linné)
Alouatta seniculus macconnelli Elliot
Herpailurus jaguarondi unicolor (Traill)
Cerdocyon thous thous (Linné)
Zoologica: N.
REPTILES
8-foot Bushmaster
Iguanas
White Amphisbena
Black and White Amphisbena
5 Crocodiles
Ete.
AMPHIBIANS
Marine Toads
Five-fingered Jungle Frog
Sharp-nosed Toad
Harlequin Frogs
Etc.
FIsH
Electric Eels
Marbled Eel
Perai
Y. Zoological Society
Lachesis mutus (Linné)
Iguana iguana (Linné)
Amphisbena alba Linné
Amphisbena fuliginosa Linné
Caiman sclerops (Schneid.)
Bufo marinus (Linné)
Leptodactylus pentadactylus (Laur.)
Bufo typhonius (Linné)
Dendrobates sp.
Electrophorus electricus (Linné)
Symbranchus marmoratus Bloch.
Pygocentrus niger (Schomb.)
[Tet
Ete.
Second, for the American Museum of Natural History, there
has ‘been brought together a collection of 485 mammals be-
longing to numerous species, preserved with their skins, skulls
and skeletons. Many of these mammals are of especial interest
because of the fact that most of the South American types of
the great Swedish naturalist Linnaeus were brought originally
from this region of the continent. Alcoholic collections of several
hundred reptiles, amphibians and fish have been made, preserved,
labeled, and shipped to the American museum.
Third, of especial significance is the collection of photo-
graphic and botanical material which, together with the actual
specimens themselves, has been gathered and furnished to the
Museum for large groups of Red Howling Monkeys, Alouatta
seniculus macconnelli Elliot, and Hoatzin, Opisthocomus hoazin
(P: .L. 8. Mull.)
Other institutions have been aided as follows: (1) Speci-
mens have been supplied to the Embryological Laboratories of
the Carnegie Institution at Johns Hopkins University, for study
of the embryology of the Red Howling Monkey; (2) numerous
electric eels have been captured and sent to Dr. Ulric Dahl-
1921] Osborn: Research Station Objects 11
gren at Princeton University, for investigation of the electric
organs of these animals; (3) a large collection of birds in alcohol
has been made for Dr. C. A. Wood, for future study, at Leland
Stanford University, of the various structures of the eye.
ZOOLOGICAL SOCIETY, New York, July 12, 1921.
‘OqdvyWey 78 uo4wS yorvasoy oy} woayz ysvo Suryoory
YAATY INNUVZVW AHL *s “Dla
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 2
(Tropical Research Station Contribution Number 95)
2. CONTRIBUTIONS OF THE TROPICAL
RESEARCH STATION, 1916 TO 1921
By WILLIAM BEEBE,
Director of the Tropical Research Station and Honorary Curator of Birds
Pee LIS H ED BY A StOeG ie i ¥
wer AOOLOGICAL PARK, NEW YORK
SEPTEMBER, 1921
Volume III, Number 2
CONTRIBUTIONS
OF THE
TROPICAL RESEARCH STATION
OF THE
NEW YORK ZOOLOGICAL SOCIETY
By WILLIAM BEEBE.
From the date of the establishment of the Tropical Research
Station in January, 1916, to September, 1921, there have been
published ninety-three contributions. Of these, eighty-nine are
scientific papers or magazine articles and four bound volumes.
To correlate, in convenient form, these contributions with the
more important facts set forth in them, I have prepared the fol-
lowing list:
1916
Contribution Number
1. ESTABLISHMENT OF THE TROPICAL RESEARCH STATION
WILLIAM BEEBE
Bull. Zool. Soc. XIX, No. 4, July, p. 1369
Hints for the formation of a tropical station, and photographs
of jungle and laboratory.
2. NOOSING A BUSHMASTER
WILLIAM BEEBE
Ibid, p. 1372
Method of capturing an eight-foot venomous snake alive and
unharmed. Lachesis mutus (Linné)
3. THE COCK-OF-THE-ROCK
LEE S. CRANDALL
Ibid, p. 1375
Habits of the bird, wild and in captivity. Rupicola rupicola
(Linné).
4. NESTLING HOATZINS AT HOME
WILLIAM BEEBE
Bull. Zool. Soc. XIX, No. 5, Sept., p. 13893
Thorough account of the activities and psychology of hoatzin
chicks, photographs of the chicks climbing and in the nest,
adult on nest, nest and eggs, and general environment. Opis-
thocomus hoazin (P.L.S. Mull.)
16 Zoologica: N. Y. Zoological Society [Ilis2
5. PICTURES FROM THE TROPICAL RESEARCH STATION
WILLIAM BEEBE and PAUL G. HOWES
Ibid, p. 1400
Photographs of
House Bats, Vampyrus sp.,
Beesa Monkey, Pithecia pithecia (Linné)
Aracari Toucan, Pteroglossus aracari atricollis (P.L.S. Mull.)
and Akawai Indian Benab.
6. WASPS AT THE TROPICAL RESEARCH STATION
PAUL G. HOWES
Ibid, p. 1412
Brief account of wasp life at the Station. Photographs of the
insects and their nests.
7. NOTES ON THE PERAI
G. INNESS HARTLEY
Bull. Zool. Soc. XIX, No. 6, p. 1428
Description, habits and methods of capture, photographs of
teeth and entire fish, Serrasalmo niger (Schomb.).
8. A JUNGLE-BOUND RESEARCH STATION
WILLIAM BEEBE and PAUL G. HOWES
New York Tribune, Photogravure Section, December 24th, p. 3
Photographs of
Trumpeter Chicks, Psophia crepitans Linné
Rhinoceros Beetle Grub, Megasoma actacon Linné
House Bats, Hemiderma p. perspicillatum (Linné)
Beesa Monkey, Pithecia pithecia (Linné) ,
Young Aracari Toucan, Pteroglossus aracari atricollis
(P.L.S. Mul.)
Young Hoatzins, Opisthocomus hoazin (P.L.S. Mull.)
Perai, Serrosalmo niger (Schomb.)
Giant Tree Frog, Hyla maxima (Laur.).
9. ANNUAL REPORT TO THE ZOOLOGICAL SOCIETY
Annual Report, New York Zool. Soc., 1916, p.,113
Résumé of first season’s work at the Station, with personal
contributions; photographs of laboratory.
LOLT
10. THE ALLIGATORS OF GEORGETOWN
WILLIAM BEEBE
Bull. Zool. Soc. XX, No. 1, p. 1437
Life history, breeding habits, method of capture, variation of
young, photographs, drawings of young, Caiman sclerops
(Schneid.).
1921]
ce.
12.
13.
15.
16.
Beebe: Research Station Contributions ki
INTERESTING BIRD’S NESTS FROM SOUTH AMERICA
WILLIAM BEEBE and PAUL G. HowWES
Ibid., p. 1458
Folio of photographs of Bird’s Nests from the Station.
Cayenne Hermit Hummingbird, Phoethornis superciliosus su-
perciliosus Linné
Guiana Pygmy Flycatcher, Tyranniscus acer (Salv. and God.)
Oily Flycatcher, Pipramorpha oleaginea oleaginea (Licht.).
THE POMEROON TRAIL
WILLIAM BEEBE
Atlantic Monthly, January
Note on distribution of opossums, reptiles, insects and seeds
by floating logs and currents; account of a road newly built
from the sea through a tropical jungle; bird life of rice-fields.
A NATURALIST’S TROPICAL LABORATORY
THEODORE ROOSEVELT
Scribner’s, January, p. 46.
Detailed account of the founding, the operation and the envi-
ronment of the Research Station, and of jungle life as observed
on walks. Photographs of jungle, Indian huts, bats, Phyllos-
tomus h. hastatus (Pallas), Nestling Parrots, Pionus fuscus
(P.L.S. Mull.), and Mouse Opossum, Marmosa murina murina
(Linné).
TROPICAL WILD LIFE IN BRITISH GUIANA
WILLIAM BEEBE, G. INNESS HARTLEY and PAUL G. HowWEs, with
introduction by THEODORE ROOSEVELT
Published by New York Zool. Soe.
General and detailed account of the first year’s work of the
Station. 1438 photographs by William Beebe and Paul G.
Howes. This volume contains contributions 15 to 50, which
are reviewed under their respective numbers.
INTRODUCTION TO TROPICAL WILD LIFE IN BRITISH
GUIANA
THEODORE ROOSEVELT
Introduction, p. ix.
Character of the work accomplished at a static research sta-
tion in the tropics.
ESTABLISHMENT OF THE TROPICAL RESEARCH STATION
WILLIAM BEEBE
Chap. I,..p.. 23
Detailed account of the conception and establishment of the
Research Station at Kalacoon, on the Mazaruni River, British
Guiana. Map and photographs of the Station.
18
WE
18.
iis
20.
21.
22.
Zoologica: N. Y. Zoological Society [iitS2
HISTORICAL BARTICA
WILLIAM BEEBE
Chap. II, p. 31
History of three hundred years of occupation by the Spanish
and Dutch.
THE NATURALISTS OF BARTICA DISTRICT
WILLIAM BEEBE
Chap. III, p. 38
Brief account of the work of Schomburgk, Hilhouse, Appun,
Lloyd, imThurn, Whitely and McConnell.
THE GENERAL FIELD OF WORK
WILLIAM BEEBE
Chap. IV, p. 43
General surroundings and geology; map and photographs of
the country.
THE OPEN CLEARING AND SECOND-GROWTH
WILLIAM BEEBE
Chap. V, p. 51
Succession of plant communities consequent upon clearing
primitive jungle and a subsequent four years’ second growth.
Birds inhabiting the various areas.
THE JUNGLE AND ITS LIFE
WILLIAM BEEBE
Chap. VI, p. 69
Temperature and lack of dangers in jungle; seasons; vertical
life zones of the jungle, with detailed distribution of birds and
mammals.
THE BIRD LIFE OF BARTICA DISTRICT
WILLIAM BEEBE
Chap. VII, p. 91
Comparison of temperate with tropical avifauna; daily and
seasonal migrations; roosting habits; northern migrants;
classification of birds as to sociability, voice, coloration, food,
friends, enemies; relations to the Indians; legal protection of
wild birds in British Guiana; collecting for the Zoological
Park.
23. LIST OF THE BIRDS OF BARTICA DISTRICT
WILLIAM BEEBE
Chap. VIII, p. 127
Classified list of 850 species of birds. By a proof-reading error
on the part of one of the authors, the 22 starred species were
made to read as new to the Colony of British Guiana, instead
1921]
24.
25.
26.
27.
28.
Beebe: Research Station Contributions 19
of new to Brabourne and Chubb’s List of the Birds of South
America, the error being more absurd because some are among
the most abundant species. This slip was deservedly corrected
in a detailed article by Thomas E. Penard, (Auk, Vol. 36, No.
Zepealts)
AKAWAI INDIAN AND COLONIAL NAMES OF BIRDS AND
MAMMALS
WILLIAM BEEBE
Chap. IX, p. 188
Akawai names of 143 species of birds and 41 species of mam-
mals, taken orally, and after reéonfirmation, transcribed.
METHODS OF RESEARCH
WILLIAM BEEBE
Chap. X, p. 147
Methods of shooting, trapping, or otherwise collecting and
observing tropical organisms.
FURTHER NOTES ON THE LIFE HISTORY OF HOATZINS
WILLIAM BEEBE
Chap. XI, p. 155
Life History Notes of the Hoatzin, Opisthocomus hoazin
(P.L.S. Mull.), in addition to those presented in “Ecology of
the Hoatzin,” Zoologica, Vol. I, No. 2, 1909, pp. 45-66. Photo-
graphs of environment, birds, nests and eggs.
THE HOMES OF TOUCANS
WILLIAM BEEBE
Chap. XII, p. 1838
Detailed account of discovery of breeding habits of five species
of Toucans, Red-billed Toucan, Ramphastos monilis Muller,
Sulphur-and-white-breasted Toucan, Ramphastos vitellinus
Licht., Black-necked Aracari, Pteroglossus aracari atricollis
(P.L.S. Mull.), Green Aracari, Pteroglossus viridis (Linné),
and the Guiana Toucanet, Selenidera culik (Wagler). Record
and description of eggs of Ramphastos monilis, and of young
Pteroglossus aracari atricollis. Photographs of nesting sites,
eggs, young; diagrams of heel-pads of Black-necked Aracari.
ORNITHOLOGICAL DISCOVERIES
WILLIAM BEEBE
Chap. XIII, p. 213
Detailed description of nests and eggs of seventeen species of
tropical birds which “have not heretofore been described, or
are almost unknown.” (For critical discussion of this paper,
see Thomas E. Penard, Auk, Vol. 36, No. 2, p. 221).
20
29.
30.
ol.
32.
Zoologica: N. Y. Zoological Society [lites
Talpacoti Ground Dove Chaemepelia talpacoti (Temm. and
Knip.), Red Mountain Dove Oreopelia montana (Linné),
White-necked Crake, Porzana albicollis (Vieill.),. Cayenne
Crake, Creciscus viridis viridis (P.L.S. Mull.), Dusky Night-
hawk, Caprimulgus nigrescens Cab., Guiana Tyrantlet,
Tyranniscus acer (Salv. and God.), Oily Flycatcher, Pipra-
morpha oleaginea oleaginea (Licht.), Varied Flycatcher, Hm-
pidonomus varius varius (Vieill.), Cinereous Bushbird, Tham-
nomanes glaucus Cab., Rufus-fronted Antcatcher, Anoplops
rufigula rufigula (Bodd.), Quadrille Bird, Leucolepia musica ~
musica (Bodd.), Orange-headed Manakin, Pipra aureola
aureola (Linné), Brown-breasted Pygmy Grosbeak, Oryzo-
borus angolensis brevirostris (Berlepsch), Chestnut-bellied
Seedeater, Sporophila castaneiventris Cab., Black-headed
Seedeater, Sporophila bouvronides (Less.), Blue Honey
Creeper, Cyanerpes cyaneus cyaneus (Linné), and the Moriche
Oriole, Icterus chrysocephalus (Linné). Photographs of nests.
YOUNG GREY-BACKED TRUMPETERS
WILLIAM BEEBE
Chapter XIV, p. 247
Detailed description of appearance, molt and habits of young
Trumpeter, Psophia crepitans Linné, with photographs and
colored plate.
THE WAYS OF TINAMOU
WILLIAM BEEBE
Chap. XV, p. 253
Life histories, nests, eggs and general account of Guiana
Great Tinamou, Tinamus major major (Gmel.), Pileated
Tinamou, Crypturus soui soui (Hermann), and the Variegated
Tinamou, Crypturus variegatus variegatus (Gmel.), with
photographs.
WILD LIFE NEAR KALACOON
WILLIAM BEEBE
Chap. XVI, p. 271
Detailed account of awakening of tropical wild life from 5.30
to 6.45 A.M., on mornings of March 26, May 16 and July 2nd,
1916. Log of occurrences, broods, migrations and nesting
seasons of tropical organisms, from June 15th to August 16th.
THE ALLIGATORS OF GEORGETOWN
WILLIAM BEEBE
Chap. XVII, p. 283
Detailed account of life history, breeding habits, eggs, young,
variation in young, photographs and diagrams of young.
Caiman sclerops (Schneid.).
1921]
33.
34.
35.
36.
37.
Beebe: Research Station Contributions ad
NOTES ON THE DEVELOPMENT OF THE JACANA
G. INNESS HARTLEY
Chap. XVIII, p. 293
General development, pterylosis, head of embryo, toes and
claws, spurs, relationships of growing leg and wing to body.
With diagrams of wing growth, of legs and wings compared
to body and of ontogenetic variations. Photograph of spurs.
Jacana jacana jacana (Linné).
NOTES ON THE DEVELOPMENT OF THE SMOOTH-
BILLED ANI
G. INNESS HARTLEY
Chap. XIX, p. 307
Detailed description of pterylosis of embryo and adult, wing
pads, development of wing, bones of leg, and development of
bill. With diagrams, charts, and photograph of embryo.
Crotophaga ani (Gmel.).
NOTES ON A FEW EMBRYOS
G. INNESS HARTLEY
Chap. XX, p. 321
Pterylosis and external characters of embryos of Caprimulgus
nigrescens Cab., Pitangus sulphuratus sulphuratus (Linné),
and Empidonomus varius varius (Vieill.).
NESTING HABITS OF THE GREY-BREASTED MARTIN
G. INNESS HARTLEY
Chap. XXI,p. 328
Breeding season, courtship, nest-building, eggs, young, food,
development of flight, and instincts of young birds. Photo-
graph of young. Progne chalybea chalybea (Gmel.).
PRELIMINARY NOTES ON THE DEVELOPMENT OF THE
WING
G. INNESS HARTLEY
Chap. XXII, p. 342
Development of the wing and pinion in relation to habits of
Opisthocomus hoazin (P. L. S. Mull.), Psittacula passerina
(Linné), Psophia crepitans Linné, Butorides striata (Linné),
Pteroglossus aracari atricollis (P.L.S. Mull.), Galeoscoptes
carolinensis (Linné), Cacicus cela cela (Linné), Pitangus sul-
phuratus sulphuratus (Linné), and Progne chalybea chalybea
(Gmel.). Development of the pinion of the following species:
Opisthocomus hoazin (P. L. S. Mull.), Psophia crepitans
Linné, Pitangus sulphuratus sulphuratus (Juinné), Pteroglos-
sus aracari atricollis (P. L. S. Mull.), Galeoscoptes carolinensis
' (Linné), Progne chalybea chalybea (Gmel.).
|
{
22 Zoologica: N. Y. Zoological Society [Tits
38. NOTES ON THE PERAI FISH
G. INNESS HARTLEY
Chap. XXIII, p. 342
‘ A general account of this dangerous fish elaborated from con-
tribution number 7.
39. THE BEES AND WASPS OF BARTICA
PAUL G. HOWES
Chap. XXIV, p. 371
General account, relation of rainfall to nesting, difficulties in
rearing larvae.
40. TWO POTTER WASPS
PauL G. HOWES
Chap. XXV, p. 376
General account, nests, food of young, egg, larva, cocoon, pupa-
tion. Red Potter Wasp Eumenes canaliculata, and Buff
Eumenes, Humenes, sp. Colored plates of egg, larva, pupa
and imago. Photographs of wasp on nest, nests, and of larva
within the nests.
41. LARVAL SACRIFICE
PAauL G. HOWES
Chap. XXVI, p. 386
Life history, nest, eggs, discussion of transformation from
larva to pupa and imago. Colored plate of egg, larva, pupa
and imago, Figure 125—9, 10, 11 and 12. Photographs of
pupa, and of transformation from larva to pupa. Podium
rufipes (Fabr.).
42. THE BLACK REED WASP
PauL G. Howes
Chap. XXVII, p. 394
Nests, nesting, nest provisions, egg, larva, pupa, method of
depositing eggs in chambers of nest. Colored plate of egg,
larva, pupa and imago, Figure 125—13, 14, 15 and 16. Pho-
tographs of nests. Trypoxylon cinereohirtum Cam.
43. THE WHITE-FOOTED WASP
PauL G. HowEs
Chap. XXVIII, p. 401
General account, method of laying eggs, larval food, nests,
eggs, larva, cocoon, pupa, emergence. Colored plate of egg,
larva, pupa and imago, Figure 133—1, 2, 3 and 4. Photograph
of cocoon. Trypoxylon leucotrichium Rohmer. ,
1921]
44,
45.
46.
47.
48.
49.
50.
Beebe: Research Station Contributions 23
THE FOREST SHELL WASP
PAUL G. HOWES
Chap. XXIX, p. 407
General account, nests, egg, larval food, larva, artificial feed-
ing of larva, pupation, emergence. Colored plate of egg,
larva, pupa and imago, Figure 133—5, 6, 7 and 8. Photo-
graphs of nests and larvae. Zethusculus hamatus Zav.
THE ONE-BANDED DAUBER
PauL G. Howes
Chap. XXX, p. 413
General account, methods of constructing nests, discussion of
sense of direction, larval food, egg, larva, pupa, cocoon.
Colored plate, Figure 133—9,.10, 11 and 12. Photographs of
nest, wasp at nest and newly emerged specimen. Sceliphron
fistulare Dahlb.
THE BLUE HUNTRESS
PauL G. HOWES
Chap. XXXI, p. 413
General account, nest, larval food, egg, larvae, pupa, emergence
from cocoon. Colored plate, egg, larva, pupa and imago,
Figure 133—13, 14, 15 and 16. Photographs of wasp at nest,
cocoon. Chlorion neotropicus Kohl.
PARALYZED PROVENDER
PAUL G. HOWES
Chap. XXXII, p. 436
General account of the actions of wasps in paralyzing spiders
and insects. Forms of paralysis produced by wasps.
CONTROLLED PUPATION
PauL G. HOWES
Chap. XXXIII, p. 443
General account of larva of Trypetid flies of genus Spilographa.
Transformation of larva to pupa. Photographs of larva.
NOTES FROM THE HINTERLAND OF GUIANA
WALTER G. WHITE
Chap. XXXIV, p. 453
General account of the country and the animals found along
the upper Rupununni River.
INDIAN CHARMS
JAMES RODWAY
Chap. XXXV, p. 488
Charms and superstitions of the Guiana Indians, with colored
plate of plant leaves used as charms.
24
51.
52.
53.
54.
55.
56.
57.
Zoologica: N. Y. Zoological Society [iirs2
A HUNT FOR HOATZINS
WILLIAM BEEBE
Atlantic Monthly, February.
Detailed account of the general environment of hoatzins.
Opisthocomus hoazin (P. L. S. Mull.).
WITH ARMY ANTS SOMEWHERE IN THE JUNGLE
WILLIAM BEEBE
Atlantic Monthly, April
Method of attack, securing and carrying food, and general
account of the offensive activities of Hciton burchelli West-
wood.
A WILDERNESS LABORATORY
WILLIAM BEEBE
Atlantic Monthly, May
Method of founding and operating a jungle laboratory with
description of the environment, opportunities for research,
servants, etc.
COLONEL ROOSEVELT AND THE TROPICAL RESEARCH
STATION
WILLIAM BEEBE
Timehri (3), IV, June, p. 27
Brief narrative of the part which Colonel Roosevelt played in
the conception and development of the Station.
JUNGLE NIGHT
WILLIAM BEEBE
Atlantic Monthly, July
Activities of organisms on moonlit nights in the Guiana jungle.
ANNUAL REPORT OF THE TROPICAL RESEARCH STA-
TION
WILLIAM BEEBE
Annual Report of the New York Zool. Soc. 1917, p. 99
Activities of the Station staff during the year 1917.
1918
A SILKY EATER OF ANTS
WILLIAM BEEBE
Bull. Zool. Soc. XXI, No. 1, January, p. 1561
General account of the Silky Anteater, Cyclopes didactylus
didactylus (Linné), with photographs and drawings.
1921]
58.
59.
60.
61.
62.
63.
64.
Beebe: Research Station Contributions 25
LABEL-MAKING IN THE FIELD
WILLIAM BEEBE
Ibid., p. 1574
Short account of simple photographic method of making field
labels of small size.
A SECOND IMPERIAL PARROT
WILLIAM BEEBE
Ibid., p. 1578
General account of the second specimen of this rare parrot
brought to the Zoological Park, Amazona imperialis (Rich.).
INSECT TYRANTS
WILLIAM BEEBE
Bull. Zool. Soc. XXI, No. 5, p. 1670
General account of the methods of attack, and methods of
transportation used by the army ant, Eciton burchelli West-
wood. With two drawings.
ANTS COLLECTED IN BRITISH GUIANA BY MR. WILLIAM
BEEBE
WILLIAM MoRTON WHEELER
Jour. N. Y. Ento. Soc., Vol. XXVI, No. 1, March, p. 23
Account of a small collection of ants taken from twenty square
feet of bushes at Penal Settlement, Bartica District. The
collection comprises 42 forms and the following new species:
Camponotus (Myrmobrachys) beebei Wheeler.
Crematogaster ornatipilis Wheeler.
ISLANDS
WILLIAM BEEBE
Jour. Am. Mus. Nat. Hist., Vol. XVIII, No. 6, p. 453
Notes on the West Indies, from St. Thomas to Barbados.
CONVICT TRAIL
' WILLIAM BEEBE
Atlantic Monthly, September
The cutting of a trail through second growth jungle and the
subsequent effect on flora and fauna; social sleeping habits of
Ithomiid and Heliconid butterflies.
SEA WRACK
WILLIAM BEEBE
Atlantic Monthly, October.
Study of the pelagic life and sargossa weed organisms between
New York and British Guiana.
26 Zoologica: N. Y. Zoological Society Babee
65. JUNGLE PEACE
WILLIAM BEEBE
8vo., New York, Henry Holt & Co. Illustrated by the author.
Collection in book form of contributions 12, 51, 52, 53, 55,
62, 63 and 64.
66. BEES FROM BRITISH GUIANA
T. D. A. COCKERELL
Bull. Amer. Mus. Nat. Hist., Vol. XXXVIII, Art. XX, p. 685
Account of a small collection of bees from Bartica District,
with key to all the forms in the collection, and descriptions
of the following new species and varieties:
Euglossa decorata Smith, var. ruficauda Cockerell
Euglossa ignita Smith, var. chlorosoma Cockerell
‘ Epicharis maculata var. barticana Cockerell
Rhathymus beebei Cockerell
Augochlora callichlorura Cockerell
Florilegus barticanus Cockerell
Included in this paper is a description of a new species from
French Guiana in the author’s collection:
Augochlora maroniana Cockerell.
67. ANNUAL REPORT OF THE TROPICAL RESEARCH STATION
WILLIAM BEEBE
Annual Report N. Y. Zool. Soc., 1918, p. 84
Activities of the Station during 1918.
1919
68. REVIEW OF “TROPICAL WILD LIFE IN BRITISH GUIANA”
GEORGE W. HUNTER
Bull. Zool. Soc., XXII, No. 1, p. 21
Critical and detailed review of the work of the Tropical Re-
search Station as narrated in “Tropical Wild Life in British
Guiana.”
69. HAMMOCK NIGHTS
WILLIAM BEEBE
Atlantic Monthly, February
Night life of the jungle as observed from a hammock.
70. HIGHER VERTEBRATES OF BRITISH GUIANA, LIST OF
AMPHIBIA, REPTILIA AND MAMMALIA
WILLIAM BEEBE
Zoologica, Vol. II, No. 7; p. 205
Check list of 52 amphibians, 112 reptiles and 119 mammals
of British Guiana of which almost half occur at the Station.
1921} Beebe: Research Station Contributions 27
(See in connection with this paper, “Comments on a Recent
Check-list,” by Thomas Barbour, American Naturalist, Vol.
LIV, page 285.)
71. BIRDS OF BARTICA DISTRICT
WILLIAM BEEBE
Zoologica, Vol. II, No. 8, p. 229
Check list of 75 birds new to the Bartica District. This list
is a supplement to the list found in Contribution 23.
72. LIZARDS OF THE GENUS AMEIVA
WILLIAM BEEBE
Zoologica, Vol. II, No. 9, p. 235
A brief, critical study of the pattern and color of Amewa
lizards, with a discussion of Barbour and Noble’s paper, “A
Revision of the Lizards of the Genus Ameiva,’ Bull. Mus.
Comp. Zool., Harvard, LIX, No. 6, 1915, pp. 417-479.
73. THE TROPICAL RESEARCH STATION
WILLIAM BEEBE
Bull. Zool. Soc., XXII, No. 4, p. 274
Brief account of the re-establishment of the Tropical Research
Station at its new home, Kartabo, with short account of some
of the forms of life at the Station.
74. THE TERMITES OF KARTABO
ALFRED EMERSON
Ibid., p. 75
Notes on termite nests and general termite ecology.
75. A HOME TOWN OF THE ARMY ANTS
WILLIAM BEEBE
Atlantic Monthly, October
Detailed account of the temporary nest and the intricate nest
activities of a colony of Eciton burchelli Westwood.
76. INSECT BEHAVIOR
PauL G. HOWES
Richard G. Badger, Boston, Mass.
Chapters II to IX, inclusive, are reprints of contributions 41
to 48 from “Tropical Wild Life.”
77. ANNUAL REPORT OF THE TROPICAL RESEARCH STATION
WILLIAM BEEBE
Annual Report, N. Y. Zool. Soc. 1919, p. 115.
Activities of the Station for the year 1919.
28
78.
79.
80.
81.
82.
83.
84.
Zoologica: N. Y. Zoological Society i ae
1920
A JUNGLE CLEARING
WILLIAM BEEBE
Atlantic Monthly, January
Comparison of superficial aspects of tropical with temperate
fauna and flora; biocoenose of a weed, parrakeets, cotingas
and toucans.
THE LURE OF KARTABO
WILLIAM BEEBE
Atlantic Monthly, February
The founding of the Station at its permanent site, and de-
scription of the wild life in the immediate vicinity.
TROPICAL TADPOLES
JOHN TEE-VAN
Bulle Zool. soc:, Vole SO No: de san, p10
Brief account of tadpoles and tadpole habits at Kartabo.
FOURTH YEAR OF THE NEOTROPICAL RESEARCH STA-
TION
HENRY FAIRFIELD OSBORN
Science, June 11, 1920, pages 585-587
Résumé of the foundation, objects and achievements of the.
Station.
ANNUAL REPORT OF THE TROPICAL RESEARCH STATION
WILLIAM BEEBE
Annual Report, N. Y. Zool. Soc. 1920, p. 111.
Activities of the Station during 1920.
1921
A TROPIC GARDEN
WILLIAM BEERE
Atlantic Monthly, February
Habits of manatees, Trichechus manatus Linné, jacanas,
Jacana jacana jacana (Linné), mongoose, Mungos mungo
(Gmelin), ‘and herons.
GUINEVERE THE MYSTERIOUS
WILLIAM BEEBE
Atlantic Monthly, March
Phyllomedusa bicolor (Bodd.), its environment, metamorphosis
and habits.
1921] Beebe: Research Station Contributions 29
85. A NEW CASE OF PARABIOSIS AND THE “ANT GARDENS”
OF BRITISH GUIANA
WILLIAM MorTON WHEELER
Ecology, Vol. II, No. 2, April, 1921, p. 89
Résumé of the known cases of myrmicine parabiosis. Records
of the parabiosis of Camponotus and Crematogaster in 80 per
cent of the ant gardens found near the Station; detailed ac-
count of these gardens.
86. THE BAY OF BUTTERFLIES
WILLIAM BEEBE
Harpers Magazine, April
Habits of the Giant Singing Catfish, Doras granulosus Valen-
ciennes; the lLong-armed Beetle, Acrocinus longimanus
(Linné), and general life of the tidal area of the Mazaruni
shore; migration and social habits of five species of Catopsilia.
87. A JUNGLE BEACH
WILLIAM BEEBE
Atlantic Monthly, May
The ecological results, floral and faunal, of the falling of a
single tree into the water; organisms living in the tide-washed
roots of trees.
88. OBSERVATIONS ON ARMY ANTS IN BRITISH GUIANA
WILLIAM MorTON WHEELER
Proc. Am. Acad. Arts and Sci., Vol. 56, No. 8, June 1921, p. 291
Notes on the life histories of twelve species of army-ants,
three of which are new to science; description of the female of
Eciton burchelli Westwood, and the males of that species
and of EHeiton pilosum F. Smith. Descriptions of the follow-
ing new species and varieties: Eciton (Acamatus) an-
gustinode Emery subsp. emersoni Wheeler, Eciton (Labidus)
praedator F. Smith var. guianense Wheeler, Hciton (Acamatus )
pilosum F. Smith var. beebei Wheeler, and Cheliomyrmex
megalcnyx Wheeler. With many photographs and drawings.
89. THE GARDENS OF THE JUNGLE
WILLIAM BEEPE
House and Garden Magazine, July
An account of the plants raised by the Akawai Indians of
Guiana.
90. THE ATTAS—A JUNGLE LABOR UNION
WILLIAM BEEBE
Atlantie Monthly, July
Atta cephalotes Fab. Foundation of the nest by the queen,
leaf cutting activities, the trails, functions of the minims in
the field, and abnormal actions of the maxims.
30 Zoologica: N. Y. Zoological Society [III; 2
91. THE ATTAS AT HOME
WILLIAM BEEBE
Atta cephalotes Fab. The nest, its appearance, environment,
visitors, parasitic and otherwise, reactions in defense, fungus
‘gardens, habits of the parasitic cockroach Attaphila, marriage
flight of the males and females, founding of a new colony.
92. SEQUELS
WILLIAM BEEBE
Atlantic Monthly, October
Instrumental sounds made by Tapping Wasp, Synoeca irina
Spinola; voice of Trogonurus curucui curucui (Linné); re-
markable exhibition of instinct by Hciton burchelli Westwood,
an entire colony being confined to a one-hundred yard circle
for several days.
93. EDGE OF THE JUNGLE
WILLIAM BEEBE
8vo. Henry Holt & Co., New York, October
Collection in book form of contributions Nos. 69, 75, 78, 79,
83, 84, 86, 87, 89, 90, 91, 92.
APPENDIX
Before the establishment of the Tropical Research Station
there were published in the first and second volumes of ZOOLOGICA
a number of articles by William Beebe, then Curator of Birds,
dealing with Neotropical zoology. In order to correlate all the
past work of the Society in this direction a list of titles is
presented :
A CONTRIBUTION TO THE ECOLOGY OF THE ADULT HOATZIN
WILLIAM BEEBE
Zoologica, I, No. 2, 1909, p. 45
History, names, distribution, general appearance, parasites, food,
nests, eggs, enemies and odors. Field notes on this species made
in British Guiana and Venezuela. Methods of photographing.
Bibliography, map of distribution and photographs of the adults
in their natural surroundings. Opisthocomus hoazin (P. L. S.
Mull.).
AN ORNITHOLOGICAL RECONNAISSANCE OF NORTHEAST-
ERN VENEZUELA
WILLIAM BEEBE
Zoologica, I, No. 3, 1909, p. 67
1921]
Beebe: Research Station Contributions aL
General character of the pure mangrove forest, the mainland
forest and the pitch lakes. Annotated list of the birds observed,
and ecological conclusions concerning the birds of the Orinoco
region. Notes on the voice of many species of birds. Descrip-
tions of the nests and eggs of the following species: Great Blue
Tinamou, Tinamus tao Temm., Venezuelan Rufous-tailed Jaca-
mar, Galbula ruficauda Cuv., Yellow-backed Cassique, Caccicus
persicus Linné, and the eggs and young of the Yellow-fronted
Amazon Parrot, Amazona ochrocephala (Gmel.).
NEW SPECIES OF INSECTS COLLECTED BY C. WILLIAM
BEEBE IN SOUTH AMERICA
Zoologica, I, No. 4, 1910, p. 118
Descriptions of the following new genera and species of insects
collected by William Beebe in British Guiana and Venezuela:
MALLOPHAGA
Colpocephalum armiferum Kellogg—host Opisthocomus hoazin
(P. L. S. Mull.) ; Lipeurus absitus Kellogg—same host. De-
scribed by Vernon L. Kellogg, Stanford University, California.
ORTHOPTERA
Stagmomantis hoorie Caudell. Described from male and fe-
male specimens from near the Hoorie gold mine on Hoorie Creek,
a tributary of the Barama River, British Guiana, by A. N.
Caudell, U. S. National Museum, Washington, D. C.
LEPIDOPTERA
The following new species and genera were described by Har-
rison G. Dyar, U. S. National Museum, Washington, D. C.:
Hylesia indurata, Zatrephes cardytera, genus Zaevius, Zaevius
calocore, genus Thyonaea, Thyonaea dremma, Illice biota, Neo-
phaenis aedemon, Emarginea empyra, Hadena niphetodes, Cap-
nodes albicosta, Thermesia dorsilinea, Claphe laudissima, Rifar-
gia phanerostigma, Eois costalis, the female of Racheolopha
nivetacta, Acropteryx opulenta, Saccopleura lycealis, genus
Dichocrocopsis, Dichocrocopsis maculiferalis, Ischnurges bicolor-
alis, genus Hositea, Hositea gynaecia, genus Incarcha, Incarcha
aporalis, Macalla pallidomedia, Paracraga amianta, Minacragides
arnacis, Trosia nigripes, and Hemipecten cleptes. All of the
above insects are described from specimens captured at Hoorie,
British Guiana.
RACKET FORMATION IN TAIL FEATHERS OF THE MOTMOTS
WILLIAM BEEBE
Zoologica I, No. 5, 1910, p. 141
Methods and means of denudation of the barbs of the central
rectrices of the Motmots, Momotus, Eumomotus sp., to form the
characteristic racket.
Zoologica: N. Y. Zoological Society iit
NOTES ON THE ONTOGENY OF THE WHITE IBIS, GUIRA ALBA
WILLIAM BEEBE
Zoologica I, No. 12, 1914, p. 241
Breeding habits, eggs, young, food of young, development and
annual changes in coloration and form of the White Ibis, Guira
alba (Linné). With a colored plate.
SPECIALIZATION OF TAIL DOWN IN CERTAIN DUCKS
WILLIAM BEEBE and LEE S. CRANDALL
Zoologica I, 18, 1914, p. 247
Account of retention of the caudal down by attachment to the
growing juvenile rectrices in Aix sponsa Linné, Hrismatura
jamaicensis (Gmel.), and Merganetta columbiana Des Murs.
NOTES ON THE BIRDS OF PARA, BRAZIL
WILLIAM BEEBE
Zoologica II, No. 8, 1916, p. 55
Account of a visit to Para, Brazil, with discussion of the region,
the general ecology, and of the visit of seventy-six species of
birds to a single tree. Notes on some of the invertebrates of the
region. Notes on the molt of some Para birds. Annotated list
of the birds observed.
FAUNA OF FOUR SQUARE FEET OF JUNGLE
WILLIAM BEEBE
Zoologica, II, No. 4, 1916, p. 107
An investigation of four square feet of debris taken from the
jungle floor near Para, Brazil, and carefully searched for minute
forms of life. Accounts of the many interesting invertebrates
found within the debris. Illustrations of the following new
ants, Glamyromyrmex beebei Wheeler, and Blepharidatta brasi-
liensis Wheeler. .
Also belonging to this series is the following volume, dealing
with the account of a trip into the British Guiana jungle in the
year 1909.
OUR SEARCH FOR A WILDERNESS
M. B. & C. W. BEEBE
8vo., 408 pages, illustrated. Henry Holt & Co., New York, 1910
Pages 111 to 398 deal entirely with British Guiana and the animal
life found there.
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ZOOLOGICA
“SCIENTIFIC CONTRIBUTIONS OF THE
EW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBERS 3-11
. A STUDY OF SOME SOCIAL BEETLES IN BRITISH
~ GUIANA AND OF THEIR RELATIONS TO THE ANT-
~ PLANT TACHIGALIA. By WILLIAM M. WHEELER
THE TACHIGALIA ANTS. By WILLIAM M. WHEELER .
HABITS OF CUCUJIDAE. By WILLIAM M. WHEELER
NEW COLEOPTERA. By E. A. SCHWARZ
and H. S. BARBER
LARVAE AND PUPAE OF SOCIAL BEETLES.
By ADAM GIEDE BOvING
_ A NEW DIADIPLOSIS. | By E. P. FELT
. A NEW BLEPYRUS. -_-By C. T. Brues
TWO TACHIGALIA MEMBRACIDS. By HERBERT OSBORN
aeane | NEW ENTOMOBRYA. By J. W. FoLsom
PUBLISHED Oey. T H Bee o- Cie tT ¥
ZOOLOGICAL PARK, NEW YORK
_ DECEMBER 24, 1921
New York Zonlogival Soriviy
General Office: 111 Broadway, New York City ica
Officers
President, HENRY FAIRFIELD OSBORN;
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Class nf 1922
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C. LEDYARD BLAIR, EMERSON McMILLin, ANTHONY R,
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f WINTHROP, GEORGE C. CLARK,
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DIETERICH, GEORGE F, BAKER, WM. PIERSON HAMILTON, ~
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WILLIAM B. Oscoop FIELD, A. BARTON -
HEPBURN, WILLIAM WooDWarp,
EDWIN THORNE, PERCY A.
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Morris, ARCHER MM. HUNTINGTON, ' GEORGE D.
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Srientific Stat.
WILLIAM T. HORNADAY, Director of the Zoological Park: *
CHARLES H. TOWNSEND, Director of the Aquarium;
RAYMOND L. DITMARS, Curator of Reptiles; —
WILLIAM BEEBE, Honorary Curator of Birds and Director
Tropical Research Station; .
LEE 8. CRANDALL, Curator of Birds;
GEORGE S. HUNTINGTON, Prosector;
GEORGE A. MACCALLUM, Pathologist; _
W. REID BLAIR, Veterinarian:
ELWIN R. SANBORN, Photographer and Editor.
Enditorial Committee ‘
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WILLIAM T. HORNADAY, CHARLES Jee TOWNSEND .
Corrected to December, 1921
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 3
(Tropical Research Station Contribution Number 97)
A STUDY OF SOME SOCIAL BEETLES IN BRITISH
GUIANA AND OF THEIR RELATIONS TO
THE ANT-PLANT TACHIGALIA
By WILLIAM MorRTON WHEELER
ee et sHED BY THE SO Ci1lE Ty
mt ZOOLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
If organic evolution had stopped with insects it would
still have been a succession of achievements that angels
might desire to look into. The entomologist watches by
the most copious fountain of wonder in the world—a
well of surprises for eye and intellect—J. ARTHUR
THOMSON, “The System of Animate Nature,” p. 545.
Volume III, Number 3
A STUDY OF SOME SOCIAL BEETLES IN BRITISH
GUIANA AND OF THEIR RELATIONS TO
THE ANT-PLANT TACHIGALIA*
By WILLIAM MortToN WHEELER
INTRODUCTION.
The materials for the following paper were obtained during
July, August and September, 1920, while I was working with
Prof. I. W. Bailey on the myrmecophytes of British Guiana. We
had gone to the Tropical Laboratory of the New York Zoological
Society at Kartabo for the purpose of making an intensive study
of the relations of the ants to such plants as Cecropia, Triplaris,
Cordia and Tococa; Prof. Bailey’s investigations being primarily
concerned with the anatomical peculiarities of the plants, my own
with the identification and habits of their various ants. During
these investigations I encountered two species of beetles of such
unusual social habits that I was led to devote considerable atten-
tion to their behavior. They live in the fusiform enlargements
of the petioles of a singular tree, Tachigalia, which is also inhab-
ited by numerous species of ants. Both the beetles and the ants
cultivate coccids in the hollow petioles, and with the beetles and
coccids there are, moreover, several associated, parasitic or
syneeketic insects. And since there are also other insects, besides
those already mentioned, associated with the plant, we may
regard the latter as the focus of a very interesting and compli-
cated biocoenose.
The elements of this biocoenose are so numerous and hetero-
geneous that I have had to appeal to several specialists for assist-
ance in making identifications and writing descriptions. Dr. E.
A. Schwarz and Mr. H. S. Barber kindly studied and described
the social beetles and a little Coccinellid which feeds on their
coccids. Dr. Adam Boving made a fine study of the beetle larve;
*Contributions from the Entomological Laboratory of the Bussey In-
stitution, Harvard University. No. 188.
36 Introduction
Mr. Harold Morrison identified the coccids; the Hymenopterous
and Dipterous parasites of the latter were studied by Prof. C. T.
Brues and Dr. E. P. Felt. Dr. J. Bequaert, Prof. Herbert Osborn,
Prof. J. W. Folsom and Dr. R. V. Chamberlin identified a wasp,
two Membracids, a Collembolan and a Myriopod found living in
Tachigalia petioles; and Prof. Roland Thaxter found and iden-
tified a fungus growing on the surfaces of one of the species of
social beetles. I have added the reports of several of these inves-
tigators as appendices to the present paper, and wish to express
my great indebtedness to all of them for their generous assist-
ance and to Mr. Wm. Beebe for his many kindnesses to Prof.
Bailey and myself while we were at Kartabo and the pains he
has taken, since our return to Boston, to ascertain further facts
concerning the Tachigalia and other myrmecophytes and to
collect additional species of Formicidee. So many of the ants in-
habiting the Tachigalia prove to be new to science that I have had
to provide a special paper for their taxonomic descriptions
(Zoologica III, No. 4).
Although I have endeavored to acquire a comprehensive
knowledge of the various insects associated with the Tachigalia,
I am aware that my account must be very fragmentary. The
territory covered was limited and in other parts of British Guiana
the same tree undoubtedly harbors other insects. This is indi-
cated by the few published observations on the trees of the genus
Tachigalia and their insects in Brazil and other parts of South
America. Moreover, only the leaves and petioles and their
inhabitants were studied and there are probably many peculiar
insects that live only in the wood and seeds or merely visit the
flowers, which in some species, at least, are conspicuous and
sweet-scented. The seeds were found only after they had germi-
nated, and the trees showed no indications of flowering during
our stay in British Guiana. Our inference that they might bloom
during the winter months has not been confirmed, since up to the
time of this writing (March, 1921), Mr. Beebe has seen no
flowers on any of the trees which Prof. Bailey marked for obser-
vation. I trust, nevertheless, that my account of the Tachigalia
biocoenose will give such a picture of the astonishing complexity
and exuberance of the insect fauna of the Neotropical Region,
and of the fierce competition among these organisms on the one
Introduction 37
hand and of their intimate co-operation on the other, as to
stimulate some future investigator to complete my observations.
At any rate, the following pages may serve to direct the atten-
tion of our younger entomologists to one of the many wonderful,
almost untouched fields for investigation in tropical America.
BRITISH
BRITISH GUIANA ; GUIANA
EORGE TOWN
SCALE 58742'W.
5 MILES
LOCATION OF THE TROPICAL RESEARCH STATION OF THE
NEW YORK ZOOLOGICAL SOCIETY
The circle represents a radius of six miles.
THE TACHIGALIA.
The Leguminous myrmecophytes of the genus Tachigalia
comprise more than a dozen known species of trees and shrubs
belonging to the forest formation (hylea) of the Guianas and
the Amazon basin. The first species, paniculata, was described
by Aublet as long ago as 1775, and re-described by Tulasne in
1844. The former derived the generic name from the Carib
“tachigali,” “tachi” being the term employed by the natives of
the Guianas and Brazil for the stinging ants of the genus Pseu-
domyrma, which regularly inhabit the swollen petioles of the
species of Tachigalia and the hollow branches and trunk of the
various species of Triplaris. Spruce (1869) was also familiar
with several species of Tachigalia and their ant-inhabited petioles.
Owing to our not finding the trees at Kartabo in bloom, Prof.
Bailey and I have been unable as yet to ascertain their specific
name. They closely resemble, however, at least in their younger
stages, the species described by Harms (1906) as T. formicarum
and figured by Ule (1907), who discovered it in Brazil (Plate I).
I first found the Tachigalia on the Puruni Trail near the
Kartabo laboratory on July 22. The specimens were small and
slender, from a foot and a half to six or seven feet tall, with only
two or three to about a dozen leaves, and were growing in the
shade. The petioles were inhabited either by beetles or by ants
of the genera Azteca and Pseudomyrma. On the following day
Prof. Bailey found what we at first took to be a different Tachi-
galia on the left bank of the Cuyuni River. It was a small tree, 30
to 40 feet high, with denser foliage and very different leaves. Its
petioles were all inhabited by parts of a single huge colony of a
yellow Pseudomyrma. Later, on finding more material, we
became convinced that both trees were merely the juvenile and
adult forms of the same plant, the former growing in the shade,
the latter in the sun. Prof. Bailey found the seedlings in various
stages and was able to ascertain that the seed is a peculiar
40 - Zoologica: N. Y. Zoological Society [IlI;3
samara, not a pod as in most Leguminose, and that the plants
grow in loose colonies, the seedlings springing up in the shade
about the base of the parent tree, which rises to a height of
40 to 60 feet, with a crown of foliage at the summit of a very
slender trunk.
Further search revealed the fact that the Tachigalia is not
uncommon in many localities within a mile of the laboratory,
especially along the Puruni and Cuyuni Trails. It was also found
scattered through the beautiful primeval forest at Kalacoon and
Baracara on the right bank of the Mazaruni and in the jungle
behind the Penal Settlement on the opposite banks of the same
stream. Though the tree was often found in all the localities
mentioned, it is, nevertheless, rather sporadic compared with
many other components of the hylea. It may also be very local
in British Guiana and the adjacent countries. This is indicated
by the fact that there seemed to be no specimens of it in the
herbarium of the Botanical Garden at Georgetown nor in that
of the Botanical Garden at Port of Spain, Trinidad, and neither
the botanists of those institutions nor the chief of the forestry
department at Georgetown had ever seen the plant. Moreover,
the halfbreed Indian caretaker of the Kartabo laboratory, though
familiar with it and its ant-inhabitants, did not know its native
name, notwithstanding his remarkably accurate memory for the
aboriginal names of most other trees of the jungle. Thus we
were unable to ascertain even the generic name of the Tachigalia
till we had gone over the works on the South American flora in
the library of the Arnold Arboretum.
As Prof. Bailey will publish a detailed account of the
anatomical peculiarities of the tree, I may here confine myself
to a brief sketch of its appearance. For the purposes of this
study it will be advantageous to distinguish rather sharply
between the young, shade and large, sun forms, since their insect
inhabitants are, as a rule, very different. In the shade form
(Plate I) the Tachigalia has an extremely slender, straight
trunk, from about a foot to eight or twelve feet in height, with
long alternate, pinnate leaves coming off of it at right angles and
at such long intervals that even the smallest of the young plants
1921] Wheeler: Some Social Beetles 41
have only two or three, the largest hardly more than a dozen
leaves. These are dark green, smooth and somewhat shining, a
except at the base, where it forms a fusiform swelling two to
four inches in length and a third to half an inch in diameter. The
longer, more distal portion of the petiole bears six to eight pairs
of broadly lanceolate leafiets, which are not drooping or pendant.
The stipules at the base of the petiole are small and inconspicu-
ous. The sun form (Fig. 4) is much more vigorous and has
numerous branches at or near the summit of the long, slender,
foot or less in length. The petiole (Fig. 5a) is very slender
smooth, gray-barked trunk. The leaflets are pendant, more
crowded, much coarser, brighter green, with more rugose surfaces
and the petiole is thicker, with a large, three-cornered basal swell-
ing more gradually continued into the leaflet-bearing portion
(Fig. 5b). The cavity of the petiole also extends nearly through-
out its length instead of being confined to the basal, swollen
portion, and there are also cavities in the branches near each
leaf. The stipules are large and conspicuous and palmately
multifid. Forms of the plant intermediate between the shade
and sun forms were rarely seen. The specific identity of the two
forms was proved by finding young shoots of the typical shade
form growing from the trunk of a large sun tree in an abandoned
cassava patch at points where the wood had been cut back by
the natives.
THE TACHIGALIA BIOCOENOSE.
The interrelationships of the various organisms constituting
that portion of the Tachigalia biocoenose with which this paper
deals, i. e., the numerous insects with their parasites and satel-
lites, that infest the young shoots and leaves, are represented in
the accompanying diagram (Plate II). It will be seen that the
portion of the plant which forms what may be called the center
of the biocoenose is the leaf-petiole, and it is, of course, the
peculiar structure of this organ that determines the specific
relations of the various insects to the host-plant and is, there-
fore, the key to an analysis and understanding of the whole living
complex. As Prof. Bailey will publish a detailed account of the
42 Zoologica: N. Y. Zoological Society [III ;3
histological composition of the petiole, I here confine myself to a
general description, dwelling only on the more salient features.
The petioles of the shade-plants are very long and slender
and their enlargement is fusiform and restricted to the base,
proximal to the leafiets. In cross-section (Pl. IV) the enlarge-
ment is nearly circular, with one surface, corresponding to the
dorsal surface of the petiole, flattened. In the sun-plants the
basal thickening passes gradually into the leaflet-bearing portion
of the petiole and the cross-section (Pl. III, figs. 3 and 4) is
distinctly triangular, one of the sides being dorsal, the two others
forming together the inverted roof-shaped ventral portion, so
that the petiole might be described as flattened dorsally and
carinate ventrally. The dorsal surface is distinctly winged, or
alate on each side. In the very young leaves the petiolar enlarge-
ment is solid, its interior being filled with juicy pith (PI. III,
fig. 3). This soon dries up, however, and is converted into a
flocculent or fibrillar, cinnamon brown substance, lying loosely
in a large cavity, the walls of which are lined by a thin layer
of the same reddish tissue. The same kind of tissue, the cells
of which are filled with a homogeneous amber-colored substance,
is also continually forming in four longitudinal strands or rays
in the dorsal wall of the petiole (Pl. IV, fig. 1). Later a few
similar strands make their appearance also in the ventrolateral
walls. Since the amber-colored substance which characterizes
this tissue evidently has a high nutritive value, I shall call it the
nutritive parenchyma. In a few petioles in this stage I found
from four to six small Curculionid larve feeding on the loose
material and reducing it to a red, powdery frass. Unfortu-
nately these larve could not be reared, so that their specific iden-
tity is unknown. They are not, however, necessary agents in the
preparation of the petioles for their future occupants.
As soon as the petiole has reached the stage just described
it is evident that any small insect, sufficiently enterprisng to bore
a hole in its wall, would find comfortable lodgings, which could
easily be rendered even more comfortable by tossing the dried
remnants of the pith out of the entrance or by compacting them
in the narrow ends of the spindle-shaped cavity. And if the
insect were a vegetable feeder it might find also an abundant
food-supply in the remains of the pith still forming the lining
‘ tn ~~ Snag
: ‘
- =- SS = + i ?
el = eee x a SS \
~ ae ae? ere oem ho ee _ ‘ . z ~~ a PT
me ah i i 3 .<.— . ety . SS LS ST
FIG. 4. BRANCH OF AN ADULT (SUN) TACHIGALIA
The leaflets have been removed from two of the petioles.
Photograph by John Tee-Van.
FIG. 5. BASES OF LEAF-PETIOLE OF TACHIGALIA SP.
a, of young, shade tree; b, of large, sun tree, both nearly 44 natural size. Pieces of
the older petiole and adjacent trunk have been cut out to show the cavity
inhabited by Pseudomyrma.
1921] Wheeler: Some Social Beetles 45
of the cavity and especially in the rays of nutritive parenchyma.
That the very exuberant neotropical fauna comprises a consid-
erable number of such enterprising insects, the following obser-
vations will show.
It will conduce to clearness if we divide the insects which
take possession of the petiolar enlargements into two series
(Plate II). The first series comprises a small number of ants,
social beetles and coccids which utilize all the advantages afforded
by the petioles, 2. e., use them not only as dwellings for them-
selves and their young but also as sources of food. Among these
three groups of insects the coccids occupy a peculiar and impor-
tant position. They are present in considerable numbers and of
all sizes in all petioles inhabited by established colonies of the
beetles and ants. Specimens of the coccids from petioles of both
young and adult trees were submitted to Mr. Harold Morrison,
who pronounces them all, without exception, to belong to a single
species of mealy-bug, Pseudococcus bromelizx Bouché (Fig. 9),
a well-known form which has been taken in widely scattered
tropical and subtropical localities (India, Zanzibar, Brazil,
Florida and even in the hot-houses of Massachusetts) and on a
variety of plants (mulberry, Canna, Hibiscus, pineapple). As
these insects are, of course, quite unable to make openings in the
_ walls of the petioles, they either wander into the cavities from
the surface of the plant, after the ants and beetles have entered,
or they are carried in by the former insects. Once inside the
petioles the coccids attach themselves to the longitudinal strips
of nutritive parenchyma in the walls, insert their beaks and find
an abundant supply of sap. The ants can then absorb the saccha-
rine excrement (“honey dew’’) of the coccids and thus vicariously
feed on the plant. The beetles, as we shall see, not only utilize
this same source of liquid nutriment but also feed on the solid
nutritive parenchyma of the petiolar walls.
The other series, which comprises many more forms, merely
use the petioles as nesting, dwelling or hiding places. They enter
petioles that have been previously occupied and abandoned, and
behave in all respects like the insects which live in old oak-galls
in our northern woods. They are in fact merely tenants, or
inquilines, or what German zoologists call ‘“‘“Raumparasiten.” No
coccids are found in the petioles inhabited by these insects, which
46 Zoologica: N. Y. Zoological Society [TI1;3
may be divided into two groups, miscellaneous arthropods and
ants. The former are probably very numerous but I have notes
only on the following five:
1. In one abandoned petiole I found that a female katydid
had enlarged the opening with her jaws, had thrust her ovi-
positor through it and had deposited her flattened eggs in the
cavity.
2. On another occasion I found a petiole of a young plant
inhabited by termites (Nasutitermes sp.) They had built an
earthen gallery along the stem from the ground to an opening
in the petiole and were living in the cavity.
3. At the Penal Settlement, near Bartica, I found in one
old petiole the mature and pigmented pupa of a solitary wasp,
which Dr. J. Bequaert has identified as Podiwm ruficrus Fabr.
The mother wasp had stored the petiole with spiders, laid an egg
on them and sealed up the entrance.
4. Another petiole, found near the laboratory, contained
a small, red female centipede, identified by Dr. R. V. Chamber-
lin as Otostigmus limbatus Meinert, previously known from
Brazil and Paraguay. She was coiled about her small, white,
recently hatched young.
5. Some of the old and abandoned petioles on young trees
were occasionally seen to be inhabited by spiders, especially by
Attids.
‘Inasmuch as the ants comprise the more numerous and more
important insects associated with the Tachigalia they may be
considered in somewhat greater detail, and since their relations
to the plant are of five different kinds (Plate II) they may be
most conveniently grouped under as many captions.
1. Defoliators. Only one species, Atta cephalotes L., the com-
mon leaf-cutting ant of Central America and northern South
America, was actually proved to defoliate young Tachigalias at
a time when many of their petioles were not inhabited by other
ants or inhabited only by recently fecundated queens of Pseu-
domyrma that had not yet produced their colonies of belligerent,
stinging workers. In the jungle behind the Penal Settlement
I observed a few such trees which had been either completely
’
1921] Wheeler: Some Social Beetles AT
defoliated or had had large semicircular pieces bitten out of their
leaflets by Atta workers. I have no doubt that these ants would
carefully refrain from thus injuring larger Tachigalias in the
possession of well-developed Pseudomyrma colonies. In all prob-
ability the smaller leaf-cutters of the genus Acromyrmex, not
uncommon in the same locality, may also occasionally visit and
defoliate the young trees, but this was not actually observed.
2. Attendants of Homoptera. The terminal shoots of young
plants are often infested with a small brown Membracid in all
stages (Endoastus (?) productus Osborn) and peculiar flat Mem-
bracid nymphs (probably belonging to the genus Microcentrus,
according to Osborn) the piercing mouthparts of which leave on
the surfaces of the petioles permanent scars that may perhaps
serve later as convenient points for the beetles and queen ants
to bore into the enlargements. At least four species of ants
were taken in attendance on these Membracids: Camponotus
femoratus Fabr., Crematogaster limata parabiotica Forel, Ecta-
tomma tuberculatum Oliv. and Dolichoderus attelaboides Fabr.
The first two were the most frequently encountered and belong
to another interesting biocoenose, that of the ‘“ant-gardens,”
which I have described in a recent paper (1921), the last two
were of more sporadic occurrence. A few notes on their habits
are recorded in Zoologica, Vol. III, No. 4.
3. Inquilines. These comprise no less than sixteen different
forms, representing thirteen species, belonging to the genera
Neoponera, Leptothorax, Crematogaster, Allomerus, Solenopsis,
Pheidole, Camponotus and Brachymyrmex. Most of them are
small and all are timid and nonaggressive species which never
keep coccids, and are more frequently found in other situations,
especially in the dead twigs and branches of various shrubs and
trees. And most or all of them merely take possession of peti-
oles that have been previously perforated, inhabited and aban-
doned by other ants or by the social beetles. The inquiline ants,
moreover, are confined to the young Tachigalias growing in the
shade. Although the number of them I have collected is con-
siderable and although they represent very diverse genera and
even subfamilies, it is certain that further search, especially in
other localities, will greatly increase the number, for the petiolar
cavities of Tachigalia are sufficiently commodious to accomodate
48 Zoologica: N. Y. Zoological Society [Ties
at least the young colonies of nearly all of the twig-inhabiting
ants of British Guiana and the list of these is a long one, com-
prising many small species of Cryptocerus, Procryptocerus, Pseu-
domyrma, Leptothorax, Crematogaster, Azteca, Tapinoma,
Myrmelachista, Camponotus, ete.
4. Thief-ants. The only species belonging to this category
is the tiny Solenopsis altinodis Forel, (Fig. 15). which is closely
related to a series of ‘“‘lestobiotic” Solenopsis species that nest in
the walls of ant-or termite-nests and prey on their brood. It
does not live in the Tachigalia petioles but enters those inhabited
by the social beetles, when their entrances happen to be un-
guarded, and destroys their larve. In all probability it also
attacks small, defenceless colonies of the inquiline ants and
devours their brood. I infer that it nests in the ground from
the fact that it often appeared suddenly in considerable numbers
during the night and exterminated the colonies of the Pseudo-
myrmas in vigorous Tachigalia and Triplaris branches that had
been left on a table in the yard of the laboratory. I have also
seen it wandering about on the laboratory tables indoors, over
the foliage of the undergrowth in the dark jungle or feeding
on the pulp of injured fruits of the water cocoa (Pachira aqua-
tica Aubl.) growing along the river banks.
5. Obligatory Ants. I would thus designate the ants that
are definitively attached to the Tachigalia as their host-tree.
There are only four species: Pseudomyrma damnosa sp. nov.,
(Fig. 18), Ps. maligna sp. nov. (Fig. 14), with its two varieties, -
cholerica and crucians, Azteca foveiceps sp. nov. (Fig. 16), and
A. traili Emery. The last is doubtfully included for reasons to
be given below. The recently fertilized queens of these ants
perforate and enter the petioles of young Tachigalias growing in
the shade, close the openings behind them with particles gnawed
from the walls and eventually produce their broods in the cavi-
ties. Occasionally queens of the two species of Psewdomyrma
or of these and A. foveiceps may be found, each in a petiole of
the same plant and even of one with only a few leaves. That
the founding of the colonies may be frought with many dangers
is shown by the fact that dead queens are often found shut up
in the petioles precisely as in the case of the Azteca queens in
the internodes of young Cecropias. Some of the unfortunate in-
1921] Wheeler: Some Social Beetles 49
sects undoubtedly succumb to hunger or the attacks of fungi,
but in many cases they seem to be killed by alien ants or by the
beetles boring into the same petioles. If the queens survive,
however, and produce their broods of workers, the latter open
from the inside the entrances made by the queens and eventually
take possession of the whole plant. Since all the petioles of
larger trees are invariably inhabited by a single flourishing
colony of a single species or variety, we must suppose that the
offspring of different queens on opening their respective petioles
either fight for the possession of the plant or, if they belong
to the same species or variety, unite to form a single polycladic
colony. The fact that the petioles of large trees contain several
dedlated fertile females of Ps. damnosa or maligna would seem
to indicate that the eventual climax colony, as it may be called,
is established by alliance of several broods rather than by the
survival of the offspring of a single queen. Furthermore, the
climax colony seems to be far too populous to represent the off-
spring of a single mother.
No sooner are the petioles opened by the young broods of
workers than the coccids enter or are carried in by the ants and
attach themselves to the areas of nutritive parenchyma which
furnish optimum conditions for feeding and growth. Here they
can multiply and be cared for by the ants which undoubtedly
find in them a most welcome source of food. As the tree grows
and puts forth new leaves, each young petiole as soon as it has
reached the proper size is perforated by a detachment of ants and
coccids. The later leaves, as I have stated, have cavities extend-
ing nearly the entire length of the petioles and adjacent to their
insertions there are also cavities in the branches, which are like-
wise entered and occupied by the insects. The largest leaves have
petioles with such broad cavities at their bases that the ants
find it necessary to make chambers in them by building carton
partitions. The materials for the carton are gnawed from the
walls. Azteca foveiceps builds a much more elaborate system of
partitions than the Pseudomyrmas. ‘This is not surprising, be-
cause the Aztecas are nearly all wonderful experts in carton
construction.
When the climax stage of the Tachigalia biocoenose has been
reached, it has been reduced to only three organisms, the plant,
50 Zoologica: N. Y. Zoological Society [III;3
the coccids and Azteca fovetceps or more frequently one of the
two species of Pseudomyrma or one of the varieties of Ps.
maligna. Merely rapping on the tree then reveals the situation,
for the angry workers rush out of the petioles, cover the leaflets,
trunk and branches and can be readily identified by their size and
color, Ps. damnosa being yellow, A. foveiceps and Ps. maligna var.
erucians black; maligna s. str. and its var. cholerica both black
with yellow markings, but differing in size. Of A. traili only a
single very young colony was seen, and this was inhabiting a
petiole of a small Tachigalia. There were coccids in the cavity
of the petiole, however, and both for this reason and because
the ant is known to be associated with other myrmecophytes, I
have included it among the obligates in the diagram. It is, in
fact, conceivable that in some localities in British Guiana traili
may be the dominant species and replace foveiceps in the climax
stage of the biocoenose.
I have noticed unmistakable indications of two other insects
on young Tachigalias, both attacking the foliage. One was a
caterpillar of some kind, which had gnawed the leafiets of a few
plants and had disappeared, so that I was unable to secure a
specimen for identification. The other was an Itonidid gall which
occurred in great numbers on a single plant on both the upper
and lower surfaces of the leaflets. This gall was 2-4 mm. long,
erect and cylindrical, with a conical tip, and somewhat resembled
the galls produced by Cecidomyia caryxcola Osten Sacken on the
leaves of our northern hickories. The smaller galls were unde-
veloped and contained no larve but each of the larger ones had
a round hole at the tip and contained in the base of its tubular
cavity a cylindrical cocoon enclosing the pupa of a small parasitic
Hymenopteron, evidently a Eulophid. Some of the specimens
were fully pigmented and nearly ready to hatch, others had been
killed by an Entomophthora-like fungus. Probably the mother
of the parasite had made the round hole in the tip of the gail
in order to enter and oviposit on its maker.
It will thus be seen that the Tachigalia biocoenose comprises
at least 51 different organisms, the host plant and 50 organisms
associated primarily with its leaves and terminal shoots or secon-
darily with the organisms thus associated.
1921] Wheeler: Some Social Beetles 51
THE SOCIAL BEETLES.
A. COCCIDOTROPHUS SOCIALIS SCHWARZ AND BARBER.
After a careful taxonomic study of the beetles which I found
in the Tachigalia petioles, Messrs. Schwarz and Barber conclude
that they belong not only to perfectly distinct species but to
different genera of the family Silvanide. While still at Kartabo
I convinced myself of their specific difference, but unfortunately
not till it was too late to make a detailed study of the differences
in their behavior. While the species described in Zoologica III,
No. 5 as Coccidotrophus socialis was so abundant that I obtained
nearly four hundred of its colonies, Hunausibius wheeleri was so
scarce that I encountered it in less than a dozen petioles. For some
time I took it to be merely a depauperate variety or aberration
of the Coccidotrophus and therefore gave it little attention.
Finally, when I had come to appreciate its distinct specific status,
my stay at the laboratory was drawing to a close and I could
secure only a few of its colonies. My account, therefore, relates
almost entirely to Coccidotrophus and my notes on Eunausibius
yield only a few rather summary remarks which may be most
conveniently relegated to a brief separate caption (p. 88).
The Coccidotrophus (Plate III, fig. 2; Plate VI, figs. 1-5)
was taken only near Kartabo and only in the petioles of young
Tachigalias growing in the shade along the Puruni, Hacka and
Cuyuni trails, within a mile of the Tropical Laboratory. Although
the tree was not uncommon in the jungle behind H. M. Penal
Settlement and along the trails in the primeval forest near
Kalacoon and Baracara, I found no traces of the beetle in these
localities. It would seem, therefore, to be rather local. Its body
is flattenéd-cylindrical, long, slender and parallel-sided, with
short, stout legs, the femora, especially the fore pair, being con-
spicuously thickened. It varies in size from 3.5 to 4.5 mm. and
when fully mature is deep castaneous brown to almost black, with
more reddish appendages. Specimens recently emerged from the
pupa, however, are pale reddish testaceous, with more yellowish
elytra. The surface of the body is shining and glabrous, the
head and thorax punctate, the elytra punctate-striate. The head
bears a singular and significant resemblance to that of certain
small ants of the genus Cryptocerus. The antenne are short,
52 Zoologica: N. Y. Zoological Society [III ;3
compactly jointed and distinctly clubbed at the tip. The female
is, perhaps, somewhat smaller than the male, but I have been
unable to detect any other external differences between the sexes,
and Schwarz and Barber in their more careful study of the struc-
tural peculiarities of the species, have been no more successful.
For further details the reader is referred to their taxonomic
description in Zoologica III, No. 6.
The eggs (Fig. 11) are pure white, regularly elliptical, 0.55
long and 0.25 mm. broad, with rather thick, leathery, smooth
and shining chorion. The larva is described and figured in great
detail in Dr. Boéving’s article (Zoologica III, No. 7, Plates VII
and VIII). In life it is whitish and beautifully translucent so that
the straight alimentary tract and its contents can be distinctly
seen, but as maturity approaches the body becomes more opaque
and milk-white, owing to a considerable increase in the fat-body.
The pupa is even more opaque, but just before eclosion becomes
yellowish or testaceous, with pigmented eyes. Both beetle and
larva are very active and alert, the latter at all times, the former
when hungry or disturbed. At other times the beetle is rather
sluggish or quiescent, but shows no tendency to “feign death”
when handled. Though its wings are well-developed (Plate VI,
fig. 5) I have never seen it attempt to fly. The pupa (Plate
IX, figs. 19-21) is also very active when stimulated, wriggling
its abdomen from side to side, but is unable to move about.
The beetles enter petioles which have either not been pre-
viously perforated by other insects or those which have been
occupied for some time and have then been abandoned by ants
or other beetles or have been occupied by queens of Pseudomyrma
or Azteca that have died before they could produce their broods
of workers. The beetles either enter as a single pair or one
beetle enters and is very soon joined by an individual of the
opposite sex. I have been unable to decide which of these
methods is followed or whether the opening in the petiole is
made by the male or the female or by either indifferently. Cer-
tainly the great majority of colonies in their first, or incipient
stage consist only of a male and a female beetle. On the rare
occasions when only a single beetle was found in a petiole, the
other may have escaped or eludéd my attention while I was
cutting into the cavity. Twice I have actually seen both beetles
FIG. 6. SIX SUCCESSIVE STAGES IN THE CONSTRUCTION OF THE COCOON
BY THE FULL-GROWN LARVA OF COCCIDOTROPHUS SOCIALIS.
See text for explanation.
1921] Wheeler: Some Social Beetles 55
in a cavity diligently cleaning it up for occupancy. Such house-
cleaning is necessary both in previously unoccupied and in pre-
viously occupied petioles since in the former the particles of fibril-
lar or powdery pith, which partially fill the cavity, and in the
latter the dead bodies of ants, beetles, ete. must be removed. The
entrance opening gnawed in the wall is transversely elliptical
and just large enough to admit the slender bodies of the beetles.
It is most frequently made in the lateral wall some distance
from the narrow ends of the petiolar swelling. The petiole
occasionally has several openings, sometimes as many as five to
seven. I believe that each of these must be the work of the
founders of one of the colonies which have successively occupied
the same cavity. In other words, the pair of beetles seems not to
utilize the openings of previous occupants for the purpose of
entering the petiole but insists on making an opening of its
own, so that the considerable number of orifices occasionally
noticed in an old petiole represent the number of colonies of
beetles or potential colonies of ants that have from time to time
taken up their abode in its cavity.
The beetles accomplish the removal of the loose pith or the
remains of previous tenants by pushing this refuse into the
pointed ends of the cavity with their flattened heads, much as a
slovenly servant might tidy a room by sweeping things under
the furniture or into closets. Smaller particles of pith are
sometimes thrown out of the entrance but the decomposed and
more or less disarticulated bodies of queen ants and beetles are
too voluminous to be disposed of in this manner so that they can
only be packed away compactly into the ends of the cavity. This
behavior brings the insects into contact with the outermost layer
of pith still adhering to the ligneous walls of the cavity and the
strips of nutritive parenchyma laden with amber-colored sub-
stance. (Plate IV, fig. 1, Plate III, fig. 5). That-this tissue
actually constitutes the food of the beetles is proved not only
by finding it in their intestines but also by actually observing
their feeding activities. Very soon, however, young coccids
begin to enter the petiole through the opening made by the
beetles and take up their positions on the walls of the cavity and
preferably along these very strips of nutritive tissue which, as
the beetles feed, become gradually deepened into grooves. The
56 Zoologica: N. Y. Zoological Society [III;3
coccids station themselves in a row in each groove, with the long
axes of their bodies parallel with the long axis of the petiole.
Since both beetles and coccids center their feeding activities
on the tissue forming the floor of these grooves it is important
that they shall be kept clean so that the parenchyma, which is
continually proliferating, can be easily reached by both species
of insects. (Plate III, fig. 5). Hence the beetles carefully
deposit their feces, or frass, which has a chocolate brown color,
on the areas between the grooves. As time goes on the accumu-
lations of frass acquire the form of more or less longitudinal
ridges projecting into the petiolar cavity (Plate IV, figs. 2-3 and
Plate V). In many petioles these ridges are strikingly regular,
in others more vermiculate, interrupted or anastomosing. In
old petioles inhabited by old colonies of Coccidotrophus the inter-
ior of the petiolar cavity presents the appearance of the figures
on Plate V. In addition to these frass ridges the beetles also
build a more or less circular wall of the same substance around
the entrance, so that the latter is converted into a short tube, in
which one of the beetles often stations itself on guard for
hours at a time, with the long axis of its body at right angles
to the long axis of the petiole and its flattened head exactly
filling the elliptical orifice. (Fig. 11). From time to time the
beetle may project its antenne out into the air and wave them
about, so that the petiole from the outside suggests the nest
of one of the smaller species of Cryptocerus, with a soldier or
worker ant on guard at the orifice.
The female beetle begins to lay her eggs either before or
after the entrance of the coccids. They are deposited singly
along the edges of the frass ridges and evidently at intervals of
several hours or even days, for dissection of the beetles shows
that only a few eggs mature at a time in the ovaries. They -
are glued ‘to the wall of the petiole rather firmly and always
with their long axis parallel with its long axis and that of the
food-grooves and frass ridges (Fig. 11). As I have not wit-
nessed oviposition I have been unable to determine the length
of the embryonic period. The eggs hatch, of course, at intervals,
so that the larve vary greatly in size, some being very small
and evidently just hatched, others a third or half-grown or
actually full-grown and ready to pupate. Like the beetles, the
1921] Wheeler: Some Social Beetles 57
FIG. 7. ORYZAEPHILUS SURINAMENSIS lL.
Beetle, pupa and larva, showing the teeth on the sides of the pronotum
of the beetle and tubercles of the pupa in which they are formed.
Courtesy of the Federal Bureau of Entomology.
larve feed on the nutritive parenchyma. Its amber colored
cells in the intestines may be distinctly seen through the clear
integument and body cavity of the larve. The colony, which is
now in its second stage consists of the pair of parent .beetles,
about one or two dozen larve, mostly immature and in most
cases of about the same number of young or half-grown coccids.
When mature the larve make brown cocoons and pupate
in them, as will be described in detail below. These are formed
singly and the beetles emerging from them remain in the petiole
with their parents and larval brothers and sisters, mate and
produce eggs and larve in turn, thus leading to the third or
climax stage of the colony, which may eventually consist of
several dozen beetles of both sexes and numerous larve and
pupe in all stages of development. The coccids also increase
in number, so that the cavity of the petiole sometimes becomes
so crowded that its inmates must find their movements greatly
impeded. In the meantime the old and exhausted beetles gradu-
ally die off and their bodies are consigned to the refuse accumu-
58 Zoologica: N. Y. Zoological Society [TII;3
a
i
:
i
4
i
i
\
\
i
{
'
/
ade ee
FIG. 8.
a, pronotum of pupa of Eunausibius wheeleri showing lateral tubercles; b, to f, pronota
of five pupz of Coccidotrophus socialis, showing vestigial lateral tubercles and
their variation.
lations, or kitchen-middens in the pointed ends of the cavity.
When this crowded condition is reached, beetles begin to leave
the colony either singly or in pairs, seek and enter other petioles
of the same or other Tachigalias and thus establish new colonies.
In order to study the stages of colony development just
described as well as those that supervene, I found it necessary to
split each petiole longitudinally and to place its two halves, with
their cavities turned outward in a slender vial or test-tube plug-
ged with cotton. The cotton could be pushed in till it held the
two halves firmly against the inner surface of the glass. Through
the latter the behavior of the beetles could then be studied with
the pocket-lens (magnifying 10-20 diameters) or the binocular
dissecting microscope. Splitting the petiole, of course, so greatly
disturbs the insects that many of them at once escape into the
tube. Moreover, a certain number are killed or injured by the
1921] Wheeler: Some Social Beetles 59
knife-blade. But all the uninjured soon return to the two half-
cavities and remain in them. At first I carefully kept the tubes
in the dark, but I soon found that they could be left in the
diffuse day-light on my table without disturbing the activities
of the insects. It was necessary, however, to keep them in a
horizontal position, like that of the petiole on the living plant,
for when they were placed upright, gravity seriously interfered
with the activities of the beetles and especially of the larve,
causing them to drop and accumulate at the lower ends of the
cavities or of the tubes. Of course, this position did not inter-
fere with the coccids which remained attached by their sucking
mouthparts to the nutritive parenchyma. Colonies of Coccido-
trophus can be kept in tubes and under close observation for
a week to ten days but by the fifth or sixth day the petioles are
apt to become so dry even during the rainy season that the
beetles, larve and coccids become demoralized. The modifica-
tions of behavior thus induced will be considered in the sequel.
There is, perhaps, nothing very remarkable in the fact that
both the beetles and their larve feed on the nutritive paren-
chyma of the Tachigalia, since other Silvanide, e.g., certain spe-
cies of Oryzephilus, Silvanus, Cathartus and Nausibius are
known to be vegetarian, but the fact that both the imaginal and
larval Coccidotrophus actually solicit and imbibe the saccharine
excrement (“honey dew’’) of the coccids, is so unusual and start-
ling that it will be advisable to give a more detailed account of
these insects and of their treatment by the beetles.
The adult female Pseudococcus bromelix (Fig. 9) measures
nearly 3 mm. in length and is broadly and regularly elliptical,
evenly convex dorsally, flattened ventrally and of a pinkish flesh-
color or pale dull red, but the body is so completely covered
with snow-white wax as to be scarcely visible in healthy speci-
mens. The wax is secreted in a thin, even, mealy layer over
the dorsal surface but around the periphery of the body as a
regular fringe of stiff, blunt pencils which are longest on the
posterior segments, somewhat shorter on the anterior border
of the head and still shorter along the sides. Large specimens
of the insect are less numerous in the petioles than the smaller
or recently hatched individuals, many of which scarcely exceed
.5 mm. in length. These are reddish because they have not yet
60 Zoologica: N. Y. Zoological Society (TI1;3
FIG. 9. PSEUDOCOCCUS BROMELIAE BOUCHE.
Sketch of an adult living female with intact covering and
peripheral pencils of wax.
secreted an appreciable quantity of wax from their dorsal in-
tegument, and lack the peripheral fringe of snow-white pen-
cils. I have not seen the males. Young individuals are rather
active, not infrequently moving about in the petiole, but the
older ones remain stationary in the food grooves till the peticle
begins to dry up, when they take to wandering about aimlessly
in search of more succulent pastures.
In specimens of the Psewdococcus treated with caustic potash
and stained according to the method recommended by Mac-
Gillivray (1921, Chapter II), the details of the integument are
clearly visible (Fig. 10). Only a few of the structures are of
interest in connection with the following behavioristic account,
such as the anus, which is on the ventral surface of the flat-
ee
P+. tes > cee ges
1921] Wheeler: Some Social Beetles 61
tened posterior border of the body, and, on the dorsal surface,
two pairs of peculiar organs which have the form of transverse,
mouth-like slits with thick lips. One pair of these organs is
situated near the posterior corners of the head, the other between
the sixth and seventh abdominal segments. Coccidologists have
long been familiar with these organs in certain genera of mealy-
bugs of the subfamily Eriococcine, and have called them “eye-
like glands” “cicatrices,” ‘‘osteoliform or labiate fovee,’” or
“dorsal ostioles.” Sule called them ‘‘adipopugnatorische Organe’”’
and MacGillivray has recently dubbed the two pairs ‘‘cephalabize”’
and “caudalabie”’ respectively, terms so barbarous that they
make one’s flesh creep. I shall call them anterior and posterior
ostioles. Berlese regarded them as apodemes, or invaginations
of the integument for the insertion of muscles. Comstock, New-
stead, Sule and MacGillivray regard them as glands. In 1882
Comstock stated that he had “observed in Dactylopius a pair of
openings on the dorsal side of the sixth abdominal segment,
which are evidently homologous with the honey tubes of Aphi-
dide. A female mealy-bug was gently rubbed near the caudal
end of the body, when suddenly there appeared two drops of a
clear fluid, resembling in appearance the honey-dew of plant-
lice. This experiment was repeated many times and with many
specimens. Mr. Pergande assures me that he has observed a
similar excretion from a pair of openings on the cephalic mar-
gin of the first thoracic segment.” Comstock was, of course,
under the erroneous impression that the honey-dew of aphids
is a secretion of the cornicles instead of being the excrement of
the insects and therefore extruded from the anus. According
to MacGillivray: “There can frequently be observed on living
specimens a small globule of a clear fluid over the mouth of each
labia, more frequently the caudalabie than the cephalabiz, so
that they are probably also glandular in structure as suggested
by Comstock. For, as he suggested, when the specimens are
stroked with a pencil or dissecting needle, the insect will hump
up its back and extrude a globule of liquid. The insect is unable
to repeat this operation until the pocket is again filled with the
clear fluid. Specimens have been observed to extrude globules
from all four labize at the same time. The labize undoubtedly
have a glandular function which is probably of later origin than
their earlier function, a parademe for the attachment of muscles.”’
62 Zoologica: N. Y. Zoological Society [pTHS 3S
FIG. 10. PSEUDOCOCCUS BROMELIAE BOUCHE.
Adult female treated with caustic potash to show the open-
ings of the anterior and posterior ostioles, anal orifice, etc.
The specimen is mounted dorsal surface uppermost.
Sule (1909) made a histological and physiological study of
the anterior and posterior ostioles in Pseudococcus farinosus De
Geer. When the female of this insect was stroked with a brush,
each of the ostioles suddenly emitted a droplet of orange-yellow
liquid, which partly adhered to the brush and partly rolled off
from the wax-powdered dorsal surface. The liquid was found to
consist of cells and a few blood-corpuscles. Sule concludes that
the organs are repugnatorial, and that their secretion is employed
like that of the cornicles of aphids for gluing up the appendages
of insect enemies. His account of the function of the secretion
is by no means convincing, since it might also be regarded as an
exudate, derived directly from the fat-body, like the exudates
produced by various termitophiles and myrmecophiles (Cf.
Wheeler 1918), and hence employed for allurement instead of
repulsion.
Returning to a consideration of the Coccidotrophus in the
Tachigalia petioles, we find that the beetle often remains motion-
less for hours at a time, in a food-groove, which just fits its long
slender body. If at other times, when it is moving about, it
1921] Wheeler: Some Social Beetles 63
chances upon a coccid, it stops suddenly and seems at once
to become more alert or excited, for as soon as its clubbed anten-
nz touch the dorsal surface of the insect, their beat, hitherto
leisurely and exploratory, becomes greatly accelerated. With
each beat, each antenna rapidly describes a minute transverse
ellipse on the surface of the coccid, and the beats of the two
appendages seem not to be quite synchronous. At the same
time the beetle, with a much slower rhythm, rocks its body for-
ward and backward by bending its legs, while the mobile arti-
culations between the head and prothorax and between the pro-
thorax and mesothorax enable it to cover more of the coccid’s
dorsum and to keep the antennal clubs in contact with its
rounded surface. While engaged in this perfomance the beetle
resembles an expert pianist moving his hands from side to side
over the key-board, or a masseur with his hands in soft gloves,
massaging a patient. The beetle undoubtedly distinguishes a
large coccid’s posterior from its anterior end, since it lavishes
most attention on the former. It seems, however, to be quite
as interested in the medium-sized or smallest coccids and will
spend just as much time in stroking them. The time devoted
to the-performance in any particular case seems to vary directly
with the beetle’s appetite or thirst. A beetle may thus spend
ten, twenty or even forty or more minutes massaging a single
coccid, with occasional short pauses. After a coccid in the
proper condition has been stroked in this manner for a few
moments it may slowly turn up its wax-penciled posterior seg-
ments and discharge from the anal orifice a perfectly limpid
droplet of sweet excrement, which the beetle at once greedily
swallows. The coccid then flattens down its posterior segments
and the beetle resumes its massage. The coccid may thus con-
tribute a droplet every few minutes or it may remain inert and
unresponsive. An ant confronted with such a situation would
take the hint and at once look up another coccid, but the beetle
stubbornly keeps on and may work for an hour or more with-
out receiving another drink. Usually, however, some of the
larve or one of the other beetles of the colony intervene and the
scene may change, as described in a later paragraph.
That the antennez of the beetle are beautifully adapted for
stroking the coccids is apparent at a glance (Plate III, fig. 2,
64 Zoologica: N. Y. Zoological Society [III ;3
Plate VI, fig. 1). Their compact structure and clubbed extremi-
ties recall the antenne of many myrmecophilous beetles or of
certain ants, for many of these insects, of course, use their
antenne in soliciting liquid food from one another. The basal
joint of the Coccidotrophus antenna is even elongated to form a
crude scape, although the remaining joints do not form an angle
with it as in ants. The relations of the beetle to the coccids,
moreover, are physiologically similar to those of symphilic beetles
to the host ants that feed them with regurgitated liquids, and
the Coccidotrophus like the symphiles has a short, broad tongue
and short labial palpi. Wasmann (1896) and Escherich (1902)
have dealt with these antennal and labial adaptations in detail,
pointing out that the tongue in symphilic beetles becomes short,
broad and spoon-like and that the palpi, especially those of the
labium, become shorter and have a reduced number of joints.
Precisely this condition is seen in the labium of Coccidotrophus
‘as shown in the figure of Schwarz and Barber (PI. VI, fig. 1).
The greater development of the maxillary palpi indicates that
they may occasionally function like the antenne in soliciting
honey-dew.
Coccidotrophus larve of all stages, from those just hatched
and less than a millimeter in length, to those almost four milli-
meters long and nearly ready to pupate, likewise solicit and
obtain food from the coccids by stroking them with the antenne.
The small beetle larve show no preference for small coccids
since just hatched larve are often seen on the backs of adult
female coccids, feverishly stroking their waxen surfaces and
full grown larve may often devote themselves to coccids smaller
than their heads. The movements of the larva’s antenne, though
similar to the antennal strokes of the beetles, cover a smaller
portion of the coccid but the larve reinforce the titillation by
a simultaneous use of the maxille and maxillary palpi. The
larva is almost or quite as persistent as the beetle and drinks
up the periodic globules of honey-dew with quite as much gusto.
Both beetles and larve, however, stroke the dorsal surfaces of
the coccids so gently that their waxen bloom is neither removed
nor diminished even after the most prolonged solicitations.
In connection with the behavior of the larva, Dr. Béving’s - |
figures of its mouthparts and antenne are very interesting and
io
ra
i
i
+
Pr.
by oe
i
TIT
I ET a i EIT I SO ENS
LEE I TPIS EE 3 EOS EI To
ee eae ere
Bin
FIG. 11. ENLARGED DRAWING OF A PART OF THE WALL OF A TACHI-
GALIA PETIOLE INHABITED BY COCCIDOTROPHUS SOCIALIS.
Showing the food grooves and frass ridges, the entrance with its wall, the eggs,
an intact and broken cocoon of the Coccidotrophus and two cocoons of Blepyrus
tachigaliae, one of them after the eclosion of the parasite.
1921] Wheeler: Some Social Beetles 67
suggestive. It will be noticed that his Figs. 1, 2, 5, 8, 16, Plates
VII and VIII etc. show that the tongue is large and flat like a
spatula or ladle and well-adapted for receiving the globules of
coccid excrement. The labial palpi are very small and 2-jointed,
but the maxilla is large, with extensive stipital articulation and
a large lacinia tipped with a claw-like tooth and fringed with
stiff hairs along the medial border. The maxillary ‘palpi are
long and 3-jointed and the large articular membranes of the
separate joints suggest great mobility. The antenne are un-
usually interesting (Plate VII, figs. 1, 2, 5, 7). Though they
consist of only three joints, the second is greatly elongated and
distinctly drum-stick-shaped, the apical joint being much reduced
to form merely a sensory cap for the second joint. Now this
drum-stick-type of antenna is precisely the one found in a long
series of symphilic ant-guests of the Coleopterous family Clavi-
geride (Fustiger, Rhynchoclaviger, Adranes, etc.), which use
their antenne for soliciting liquid food from their hosts. Dr.
Boving’s figures of other larval Silvanids and of genera belong-
ing to closely allied genera of the Cucujid complex, namely
Cathartus (Plate VIII, fig. 12), Nawsibius (Plate VIII, fig. 18),
Dryocera (Plate IX, fig. 33), Telephanus (Plate X, figs. 34, 37)
and Scalidia (Plate X, fig. 39), show very different conditions.
Thus we may say that the antenne and maxille of the Coccido-
trophus larva are specially adapted to their active réle of solicit-
ing and the labium to its passive, spoon-like réle of receiving the
liquid excreta of the Pseudococcus.
The question naturally arises as to the function of the
anterior and posterior ostioles, which, as I have shown, are
highly developed in Ps. bromelix, and the probability of their
secreting substances that may be ingested by the beetles. Un-
fortunately I did not know of the existence of the ostioles while I
was at Kartabo. My attention was called to them by Prof. Mac
Gillivray several months after my return. I feel very confident,
nevertheless, that these organs in Ps. bromelie cannot have the
function ascribed to them by Sulc in Ps. farinosus. I have so care-
fully watched the coccids of all ages that I could not have over-
looked the emission of orange-yellow droplets from the ostioles
or of any sticky liquid that would glue up the appendages of
small insects. The smallest beetle larve are so delicate that
68 Zoologica: N. Y. Zoological Society [TII;3
they are at once immobilized or impeded in their movements
when they happen to run or fall into a minute drop of water, and
if the Ps. bromelix were at all hostile to the beetles or their
larve in the manner described by Sulc, their presence would
not only be a nuisance but a serious menace to the colonies. The
further fact that the coccids in a petiole are frequently decimated
or even exterminated by several small predatory insects (vide
infra p. 78) is also unfavorable to Sulc’s contention.
I am, of course, willing to admit that the ostioles may be
glands or exudate organs which secrete substances that may be
ingested by the beetles and their larve, but the closest observa-
tion of which I was capable showed that the onty liquid visibly
imbibed by the Coccidotrophus was the saccharine excrement, or
honey-dew. If the secretion of the ostioles is a liquid, it must
be emitted in droplets too minute to be visible under a Zeiss lens
magnifying 20 diameters, or it must be a volatile substance like
that secreted by the peculiar tubular organs which occur on
the eighth abdominal segment of many ant-attended Lycaenid
caterpillars and have been described by de Niceville (1890),
Thomann (1901) and others.t That the ostioles of Ps. bromeliz
may actually emit some substance attractive to the beetles and
their larve is indicated by their often very prolonged stroking
of the dorsal, lateral or anterior portion of a coccid. That they
prefer to stroke its terminal abdominal segments may be due
to the fact that that region bears both the anus and the poster-
ior pair of ostioles.
The attraction of the mealy-bugs, whether due solely to their
ability to excrete honey-dew or because they can also secrete some
delicious exudate or fascinating aroma, is so great, that in popu-
lous colonies a single coccid may often become the center of a
circle of actively competing beetles and larve of various sizes.
This is not so apparent in colonies that have just been collected
and placed in glass tubes, because the petiole still contains a cer-
tain amount of sap and the coccids are able to excrete normally,
but after the colonies have been kept for several days or a
* Conf. also the paper of Newcomer ee who figures a section of one
of these organs in Lycaena piasus (Pl. 2, Fig. 3). Although he believes
that the tubular organs are not glandular, the structural details certainly
seem to support Thomann’s contention.
1921] Wheeler: Some Social Beetles 69
week -and the supply of water in the plant tissues has consider-
ably diminished, the excretions of the coccids are less frequent
and copious, so that the beetles and larve become more and
more thirsty and therefore more desperate and exacting. Thus
by the very simple device of keeping the colonies for some time
in tubes, it is possible greatly to exaggerate the attentions of
the beetles and their larve to the coccids and to witness certain
peculiarities of behavior which are less obvious in recently col-
lected colonies.
When a number of thirsty beetles and larve surround a
large coccid, all stroking different parts of its dorsal surface
and periphery, only the individual that happens to be stationed
at its posterior end is able to secure any honey-dew. The beetles
and larve all keep at work, however, till the antenne of two of
them happen to meet. Then the larger individual stops stroking
for a moment and butts its competitor with the side of its head. If
the group is formed by a single beetle and several larve, the beetle
being the stronger, soon pushes any larva with which it may come
in contact away, but the latter usually at once returns and re-
sumes its stroking till contact with the beetle again occurs and
the butting is repeated. When several larve of different sizes
have preempted a coccid, the largest treats the others in the same
manner. This behavior is so suggestive of that of a number of
pigs eating out of the same trough that one can hardly doubt that
something more than a mere reflex is involved in the butting. The
larger beetle’s or larva’s indefatigable perseverance in butting
is only equalled by the pertinacity with which the butted indivi-
dual keeps returning and resuming its stroking movements. I
can illustrate this best by transcribing a few observations from
my note-book.
August 10, a beetle was seen standing over the posterior end
of a large coccid and stroking it busily. At short intervals the
coccid raised its anal segments and discharged a minute limpid
globule of honey-dew, which was at once avidly seized and
swallowed by the beetle. During 11 minutes the coccid raised
its tail 25 times and the globule could be distinctly seen on
most occasions, as the beetle paused suddenly in its manipula-
tions and moved its labium and palpi each time a globule was
imbibed. Sometimes the beetle would pause for a moment,
70 Zoologica: N. Y. Zoological Society [III;3
before proceeding with its titillation and remain with its‘head
flattened down, even when the coccid failed to move its anal seg-
ments. Once a large larva came up and endeavored to get some
of the excretion but was promptly butted away, and once another
beetle was treated in the same manner. After the 25 feedings
the beetle moved away and another beetle came up and received
from the same coccid four globules in less than two minutes.
August 12, I observed a beetle (No. 1), which was red and
therefore immature, soliciting from a nearly full-grown coccid
in a petiole collected a few days previously and already beginning
to become dry. The beetle stroked the coccid for 15 minutes,
during which time the latter produced only five droplets of
honey-dew at intervals of two to five minutes. Then a mature,
dark colored beetle (No. 2) came up and began to stroke the
anterior end of the coccid, gradually moving back over it. When-
ever the beetles met they butted each other with their heads or
even locked mandibles for an instant and then returned to their
former position and occupation. Beetle No. 1 worked for an-
other 15 minutes without a reward. The coccid then rotated
180 deg. on its dorsoventral axis so that its anal end was now
presented to beetle No. 2, and inserted its beak into another part
of the nutritive parenchyma. The beetle at once became more
alert and accelerated the beat of its antenne. During the suc-
ceeding eight minutes it received seven globules of honey-dew
in quick succession, probably as a result of the coccid’s change
of pasture. Throughout this period beetle No. 1 kept titillating
the coccid’s side, pausing now and then for a few seconds, and
after 40 minutes from the time I began the observation, moved
away. Beetle No. 2 continued to stroke the coccid for some
time, but I did not follow its behavior further.
In another colony at 8 P. M. on the same day I noticed a
nearly mature beetle (No 3) vigorously stroking the hind end
of a small coccid, while its sides were being stroked simultane-
ously by two just-hatched larve (A and B). From time to time
other beetles and older larve came up and joined the party.
Beetle No. 3 continually butted the newcomers away and they
at last rather reluctantly departed, leaving the original trio in
possession of the coccid. Every few seconds the beetle gave
1921] Wheeler: Some Social Beetles “1
one of the larve a shove with its head, but the tiny creature
instantly returned and went on with its stroking. At 8.14 the
beetle gave A such a vigorous knock that it stayed away from
the coccid for some time. B, however, kept returning so pertin-
aceously that the beetle twice seized it in its mandibles for an
instant and then dropped it. The larva was uninjured, however,
and at once returned and went to work. Then the periodic but-
ting continued till 8.35 when larva A returned and went to work
with B on the side of the coccid. One or the other was butted
away by the beetle every few seconds till 9 P. M. During the
entire hour the coccid remained stationary and unresponsive,
never once raising its caudal segments nor emitting a droplet.
All this time the beetle had remained in the same spot and had
butted every beetle or larva with which its antennze had come
in contact. The beetles soon left after receiving a few knocks
but the little larve A and B, which seemed to be famished,
persisted for a whole hour side by side, except for the 20
minutes during which A was absent. Larva B must have been
struck by the beetle more than a hundred times. Finally the lat-
ter’s patience seemed to be exhausted; it seized first A and then
B in its mandibles, carried the latter three millimeters away
from the coccid and hurled it to one side. Larva A returned, but
B had fallen out of the petiolar cavity onto the moist wall of
the glass tube, adhered and was unable to leave the surface. The
beetle now left the coccid and another very mature beetle (No.
4) took its place. It permitted larva A to stroke the posterior
end of the coccid without molestation, but beetle No. 3 soon
bustled up from the opposite direction, locked mandibles with
beetle No. 4 and pushed it away. During the scrimmage the
coccid suddenly raised its caudal end and discharged a droplet
which was eagerly inbibed by the larva, at length rewarded for
its incredible pertinacity and the innumerable knocks it had
received. Then beetle 3 and larva A, now in undisturbed posses-
sion of the coccid but in the reversed position, the former being
at the anterior, the latter at the caudal end of the coccid, con-
tinued their stroking, interrupted every few seconds by the
butting of the beetle and the temporary withdrawal of the
larva. This went on till 9.20. By that time my eyes which had
been following the performance under the lens for an hour and
twenty minutes were so fatigued that I had to desist from further
12 Zoologica: N. Y. Zoological Society [III ;3
' observation, just after the beetle had tossed the larva to a dis-
tance of about four millimeters from the coccid by an unusually
well-aimed blow.
Scenes of this description were so frequently enacted that
they could be readily observed in almost any of the colonies after
they had been kept for several days and the beetles, larve and
ecoccids had all grown very thirsty. In such colonies the bodies
of the coccids and larve become visibly attenuated and some-
what shrivelled as a result of the loss of water from the tis-
sues of the Tachigalia petioles. All the insects now become rest-
less. The beetles leave the petioles, wander about on the walls
of the tubes and finally collect about the plugs of cotton in an
endeavor to escape to the outside. The coccids, too, withdraw
their beaks from the parenchyma in the floors of the food-
grooves and wander aimlessly about, vainly seeking more favor-
able pastures. But before this stage of demoralization is reached,
both the beetles and the larve become cannibalistic and one
may often see them, singly or in groups voraciously devouring
partly dismembered larve or immature beetles. Within a few
days all the larve and immature beetles are consumed, but the
coccids, immune from attack, still wander about till they die
of starvation.
I believe that Coccidotrophus is rarely or never cannibal-
istic under normal conditions. It is, as already stated, almost
impossible to split a freshly gathered Tachigalia petiole con-
taining one of the beetle colonies, without cutting some of the
insects iu two, and such disabled individuals are soon devoured
by their fellows, but both in such cases and in the cannibalism
that supervenes in dried petioles, I believe that thirst or the need
of water and not a veritable carnivorous instinct, such as seems
to be manifested by some species of Cucujid beetles and their
iarve, must be regarded as the true explanation. I am con-
firmed in this view by Heins’ recent investigations (1920) on
meal-worms (Tenebrio molitor). He found that when the larve
of this beetle are reared in dry meal as many as 24.2% of
them may be devoured by their fellows, but that if wet slices .
of rusk or of vegetables are placed in the breeding jars the
mortality from cannibalism is reduced to 7.5%. In this connec-
tion Bodine’s observations (1921) on grasshoppers are also of
1921] Wheeler: Some Social Beetles 73
interest. He finds that during starvation, the loss of water in
these insects is always greater than that in body weight or in
the solids. “This shows that starvation in the grasshopper
results in a rapid loss in water which has a decidedly quick and
fatal effect.”
I have been unable to ascertain the length of the larval per-
iod or the number of larval moults of Coccidotrophus. As no
exuvie were found in the petioles it would seem that they
must be devoured either by the beetles or by the larve them-
selves. The food of the larvee, as we have seen, consists of the
amber-colored nutritive parenchyma and of the sweet excreta of
the coccids, the former evidently supplying the proteids, the
latter the sugar and most of the water. So concentrated a diet
should be very favorable for growth and probably the whole lar-
val period at tropical temperatures occupies only two or three
weeks. The fat-body, however, does not seem to become very
voluminous till the last larval instar when the segments of
the body become more convex and puffed out with the accumula-
tions of adipocytes. Yet this condition, which immediately pre-
cedes pupation, does not tend greatly to inhibit the activities of
the larva.
When a petiole containing a colony in what I have called the
third stage is opened, one or more cocoons are invariably found
in the cavity (Plate V). They are oblong-elliptical structures,
5-6 mm. long and 2-3 mm. broad and seem to consist of the
same chocolate brown substance as the frass-ridges. Their
walls are of uneven thickness, with smooth inner and rough-
ened outer surfaces, and are easily fractured. These cocoons
do not lie loosely in the cavity but are attached to some flattened
surface of the wall where the lumen of the petiole is rather
broad, i.e., away from the pointed ends, and always have their
long axes parallel with the long axis of the cavity. They are
sometimes single but more frequently occur in pairs or in
groups of three or four. When in pairs, the two cocoons lie
abreast of each other, when in threes or fours, the third and
fourth cocoon are often built on top of a basal pair. Such groups
of cocoons are so voluminous that they obstruct the lumen of
the petiole and leave only a narrow passage for the beetles to
move between the more roomy spaces at either end of them.
74 Zoologica: N. Y. Zoological Society [Ili:3
One naturally infers that either the larve must make the
cocoons of frass or the beetles must envelop the pupz with this
material, but observation shows that both inferences are incor-
rect. The larva does, indeed, build the cocoon, but utilizes nei-
ther the frass nor the materials of old, abandoned cocoons in its
construction. I have not seen the earliest stages in the process
but it is evident that the larva selects a flat surface and begins
to build a wall around an elliptical area, which thus becomes
the fioor of the cocoon. Little material is added to the wall
at the end of the ellipse compared with the sides, where the
material is built up as a pair of folds like those shown in Fig. 16a.
I have seen several cocoons that had been abandoned in this or
a somewhat more advanced stage, but on two occasions I was
able to observe the completion of the structure from a stage like
the one figured. Since in both cases the insects behaved in
essentially the same manner I shall describe only one of them.
The larva was first seen working inside the cocoon in the
stage of Fig. 6a, but it soon came out, wandered away to a
distance of a few millimeters and, after careful search, bit off a
minute particle of the living tissue of the petiolar wall, avoiding
any frass-covered surface, returned, entered the cocoon at one
end (left hand side of Fig. 6a), carefully masticated the particle
with its maxille, while mixing it with saliva, applied it to the
border of one of the folds, pressed it into place, crept out of
the other end of the cocoon and went in search of another parti-
cle. Then it returned, entered the cocoon as before and repeated
the building process. Excursions were made every few minutes
and within a radius of 8 to 10 millimeters from the cocoon. The
particles, which were selected with the greatest care and often
after what seemed like some hesitation, were very minute and
greenish when first bitten off but had become brown (by a pro-
cess of oxydation?) by the time they had been incorporated in
the walls of the cocoon. The particles were applied now to
one fold of the wall, now to the other so that the edges became
rather irregular (Fig. 60), but as most of the particles were
added to the middle of the folds, they began to approach each
other. Still, their growth was very slow, owing to the minute
size of the particles and the time consumed in their selection.
The larva labored incessantly, making trip after trip and choos-
1921] Wheeler: Some Social Beetles 75
ing every particle with the same diligence and avoiding the re-
mains of empty cocoons in the immediate vicinity, although
their materials, one would suppose, might have been easily ap-
propriated and quickly built into the cocoon under construction.
The two folds or side-walls slowly approached as the work pro-
gressed and eventually fused with each other, the larva always
entering at the same end, applying the particle to one of the
edges from the inside and leaving by the opening at the opposite
end. Then it set itself to building the walls around this latter
opening, which grew smaller and smaller (Fig. 6c), till the larva
could no longer squeeze through it and was compelled when
about to leave the cocoon to turn back on itself, bending its
body in a loop with the two limbs in contact, and crawl out of
the opening by which it had entered. This feat seemed to be
accomplished with considerable effort but had to be performed
after each particle had been built into the wall of the cocoon.
Eventually the small opening was closed and the cocoon had
only a single large elliptical orifice at one end (Fig. 6d). The
larva now began to contract this orifice, but after a time, as it
grew smaller, the insect on returning, no longer entered the
cocoon and reversed its body in order to apply the particles to
the edge of the orifice, but merely thrust its head and a few of
its anterior segments into the cocoon and left the remainder
of its body outside. At such times it used as a support or ful-
crum a structure which I had not seen used at any previous
stage of larval life, namely, the proleg which terminates the
conical tenth abdominal segment. This structure is described by
Dr. Boving (Zoologica III, No. 7) and clearly shown in his figures
(Pl. VII, figs 1 and 2). When the size of the orifice had been
reduced till the larva could only just squeeze through it, the
insect entered the cocoon, reversed its position and continued
building along the edges of the orifice with particles scraped
from the inner surface of the structure. The orifice thus soon
grew too small to permit the egress of the builder. (Fig. 6e).
Then the imprisoned creature slowly closed the opening and the
cocoon was completed (Fig. 6f).
A few days after the larva has thus immured itself, it sheds
its cuticle and becomes a pupa which lies loosely in the cavity of
the cocoon and has the appearance of Dr. Boving’s Plate IX, figs.
76 Zoologica: N. Y. Zoological Society [III;3
19-21. Owing to the minuteness of the particles used in building
the cocoon, the care with which they are chosen and the many
trips necessary to secure them, the time consumed in completing
the structure is considerable. The earliest stage figured (Fig. 6a)
was first seen at 8 P. M., July 25. By 6 A. M. the following
morning the cocoon was in the stage shown in b. By 12.30 P.
M. the small opening had been closed and the large opening was
being contracted (c). At 6:30 P. M. a small opening remained
(e), and the cocoon was completed an hour later (f). As the
first stage must have been the work of the greater part of a day,
the structure was probably begun not later than 7 A. M. on
July 25th. At least 36 hours of continuous labor, requiring
hundreds of trips back and forth between the cocoon and ex-
posed patches of living parenchyma on the petiolar wall, were
therefore consumed in completing the cocoon. The second larva
observed in the act of building its cocoon was even slower, since
the latter was first seen on the evening of July 26 in a stage
corresponding to Fig. 6a and was not entirely completed till after
10 A. M., July 29th. There was nothing to indicate that the
first larva rested during the whole period of cocoon construc-
tion. While it was working in the manner described, it was
occasionally annoyed by some young or half-grown larva enter-
ing the cocoon and using it as a hiding place while its architect
was away gathering building materials. It was interesting to
see the latter on its return oust the intruder, which scampered
away with comical alacrity. When the cocoon is completed it
is rather smooth externally but may later become rough through
the beetles’ plastering their frass over its surface. This cer-
tainly strengthens the walls of the structure.
I endeavored to keep the two pupz enclosed in the cocoons
which I had seen built, in order to determine the length of the
pupal period, but both died when the petioles dried out. The
pupal period as inferred from other cases covers, at least, seven
days. The callow emerging beetle under norma} conditions
gnaws a round or elliptical opening at one end of the cocoon
and joins the other members of the colony. At first it is yellow-
ish white and etiolated, with the legs, dorsal surface of head
and prothorax and a transverse band on each segment of the
venter pale red. It runs about very actively, nevertheless, and
:
:
.
1921] Wheeler: Some Social Beetles cr
in the course of several days gradually takes on the deep chest-
nut brown color of the mature insect. But before this stage is
reached, and while the male and female are still of a bright red
or pale chestnut red color, they mate. On two of the three occa-
sions on which I witnessed copulation both the male and the
female were immature. The third couple, observed August
10, consisted of an immature female and a mature male. It
would seem, therefore, that mating occurs not only among im-
mature beetles of the same generation soon after they leave
the cocoons, but that old males of a preceding generation at
least occasionally fecundate the recently emerged females.
The observation of August 10 is here transcribed from
my note-book. The female was’ distinctiy immature but uni-
formly red, i.e. no longer in the white, callow stage, and dis-
tinctly smaller and more slender than the male. The latter was
certainly very nearly or quite mature. When first noticed the
female was eating the parenchyma of the petiolar wall. The
male mounted her back and remained for 18 minutes, clasping
her sides with his legs and occasionally attempting to insert his
aedoeagus. Now and then he rubbed her occiput from side to
side with his mandibles and antenne and sometimes shifted his
position very much to one side. The antennal movements were
precisely like those employed in titillating the coccids. The
female kept on feeding and pressed the tip of her abdomen
against the wall of the petiole so that the male was unable to
introduce his aedoeagus. He then dismounted and ran away
but soon returned and attempted to mount and grasp the female
again, but she was unwilling, slipt out from under him and
escaped. He permitted her to go only a very short distance,
however, before he again seized her just as she had stumbled on
a coccid and had begun to stroke its posterior segments. While
the male was strenuously endeavoring to copulate she continued
to stroke the coccid and on this occasion kept the tip of her abdo-
men tightly pressed upward against the tips of her elytra. Then
the male again dismounted and left her and she and another
immature beetle turned their attention to a partly eaten larva
which they proceeded to devour. In a moment the male, appar-
ently in a high state of excitement, returned, mounted the
female and this time succeeded in introducing his edoeagus by
78 Zoologica: N. Y. Zoological Society [IlI;3
forcing the tip of the female’s abdomen downward and away
from the tips of her elytra. During coitus, which lasted a little
more than six minutes, the female continued to partake of the
larva, but the male remained motionless.
B. THE ENEMIES OF THE BEETLES AND COCCIDS AND THE DECAY
OF THE COLONIES.
Attention has been called to the decay of the Coccidotro-
phus colonies when the flow of sap to the petioles is artificially
cut off by severing the latter from the plant and both coccids
and beetles are deprived of their nourishment. But even under
the natural conditions of the jungle the colonies are doomed ‘to
decay, though from very different causes. As has been shown,
Coccidotrophus lives only in the petioles of young Tachigalias
growing in the shade and in these plants inhabits the petioles
only till they are taken over by the obligate ants of the genera
Pseudomyrma and Azteca. The beetles are not permitted, so to
speak, to occupy their apartments after the rightful owners of
the plant have become sufficiently numerous and: aggressive to
eject them. Sometimes this may occur even in rather young
plants four to six feet in height. Still the period during which
the beetles may be allowed to inhabit the young Tachigalias
must cover several months. I infer this from the fact that dur-
ing the more than two months of my stay at Kartabo I saw little
change in the plants, which grew very slowly notwithstanding
the almost daily, drenching rains, and their growth is probably
almost nil during the dry season. Since the leaves are persist-
ent, at least during the rainy months, there is ample time for
the development of the beetle colonies, even if the growth of the
larval broods and the coccids requires more than the three weeks
above suggested. Throughout the latter part of July, August
and the first half of September colonies were found in all stages,
from those represented by a single pair of beetles to those com-
prising numerous beetles, larve, pupe and herds of coccids. It
would seem, therefore, that although the individual colony may
live for only a few months, new colonies must be formed contin-
ually, at least during the rainy season, by emigration of pairs of
young beetles from old colonies to other plants which have
1921] Wheeler: Some Social Beetles 79
attained a sufficient size, i.e. have produced at least three or four
fully developed leaves.
Still neither the beetles nor the coccids are permitted to
live in perfect security till the obligate ants take possession of
their quarters. Where competition among insects is so very
keen as it is in the Neotropical jungle it is not surprising to find
that several predators and parasites are continually gaining
access to the petioles and decimating or even completely destroy-
ing their occupants. The greatest enemy of the beetles is the
small thief-ant Solenopsis altinodis Forel (See p. 48 and Zoologica
III, No. 4, p. 154), and the coccids have at least three formidable
enemies. All of these insects enter the petiole through the open-
ings made by the beetles and must therefore elude their watch-
fulness. We should expect the beetles to keep one of their num-
ber constantly on guard at the entrance, but they are neither suffi-
ciently constant in this behavior nor sufficiently discriminating
to keep out all intruders. When the petioles are taken into the
laboratory the beetles are often seen to remain for hours with
their heads in the entrances and their bodies at right angles to
the longitudinal axis of the cavity, and even after the petioles
have been split longitudinally and placed in vials the insects still
exhibit this behavior, though it is now absurdly futile, since their
domicile is wide open. But not infrequently even the single
opening of a petiole may remain unguarded for long periods,
and when the petiole has several openings some of them are
apt to have no sentinels, so that predators and parasites small
enough to pass the narrow orifices, have no great difficulty in
gaining access to the colony. Moreover, a beetle that is guard-
ing an opening may fight off certain intruders but back away
and allow others to enter. On several occasions I held a beetle
with its head to a guarded entrance. The sentinel at once
grasped the stranger’s head with its mandibles and pushed it
away. But when I placed a worker Solenopsis altinodis in the
same position, the beetle beat a hasty retreat and the ant climbed
into the petiole. From these experiments we may infer that
the beetles are more intent on keeping alien beetles of their own
species than dangerous pests like the Solenopsis out of their
nests. More probably some peculiar odor of the ant induces the
beetle to withdraw. Thus while it seems to be probable that alien
80 Zoologica: N. Y. Zoological Society [TII;3
beetles are often kept out of the colony, the fact that the beetles
or two or more colonies occupying different petioles, will, when
the latter are split open and placed in the same tube, mingle
without the slightest signs of hostility, would seem to show that
even strange beeties may occasionaily enter an unguarded colony
and become members of it in good standing. It has since
occurred to me that female beetles, at least, might be permitted
to pass the sentinels unchallenged. Unfortunately I failed to
dissect and determine the sex of the beetles with which I
experimented. |
The laxity of the beetles in guarding the entrances is, indeed,
amply proved by the presence in their nests of several species of
insects, some of which are harmless or indifferent while others
are injurious either to the beetles and their larve or to the
coccids. From analogy to the guests of ants, those of the former
category may be called ‘‘synoeketes,”’ or indifferently tolerated
guests, the latter ‘“‘synechthrans” (predators) and parasites. To
the synoeketes belong a Collembolan, a mite and a small Phorid
fly. The Collembolan is most frequently seen, especially in large
petioles containing small colonies of beetles and therefore allow-
ing ample space for its movements. It is a minute silver gray
species, which Prof. Folsom has described and figured as E’nto-
mobrya wheeleri (Zoologica III, No. 11), and occurs in droves of
individuals of all stages, running hither and thither over the walls
of the petiole, like certain species of the same genus (EH. myrmeco-
phila Reut. and dissimilis Mon.) and Cyphodeirus (C. albinos
Nicol), which are often abundant in the nests of ants. Like the
ants the Coccidotrophus pay no attention to these diminutive in-
sects and are probably not even aware of their existence. The
mites (Hypoaspis sp.) and Phorid flies (Aphiochxta scalaris)
were more rarely seen. They probably breed in the accumulations
of refuse at the ends of the petiolar cavity and may therefore
be regarded as scavengers, like the mites and Phorids which
occur in many ant-nests. é
Careful examination of the alimentary tract of the Coccido-
trophus would probably show that the beetle harbors a number
of entoparasites, at least certain bacteria, but I could not find the
time to make such an examination when fresh material was avail-
able and my alcoholic specimens are worthless for the purpose.
1921] Wheeler: Some Social Beetles 81
FIG. 12. HYPHOMYCETOUS PARASITE ON COCCI-
DOTROPAUS SOCIALIS.
From a drawing by Prof. Roland Thaxter.
It seemed probable, nevertheless, that the beetle, living as it
does in dark, moist cavities, might be infested with ectoparasitic
fungi, especially of the group Laboulbeniales. I therefore re-
quested Prof. R. Thaxter, the leading specialist in this group, to
examine a large number of the beetles. After carefully scrutin-
izing their external surfaces he reports that he found no Laboul-
beniales, but only a sterile Hyphomycete, growing on the elytron
of one of the specimens. Referring to his work on the similar
fungi of other insects he writes me as follows: “The fungus on
the Coccidotrophus probably belongs to the group spoken of at
the bottom of p. 237 in my first paper (1914), the most striking
form of which (Aposporella elegans Thaxter), found on the
wings of a small fly from the Kamerun, is figured on Plate III,
Fig. 30 of the second paper (1920). These fungi seem to produce
no spores and to reproduce by a kind of fragmentation; pieces
breaking off with little or no differentiation, and starting to grow
where they adhere. I have seen a considerable number of them
on a variety of tropical insects, and ran across one of them a
few days ago growing on a Laboulbenia from Kamerun. It
has seemed hardly desirable to give names to such nondescript
82 Zoologica: N. Y. Zoological Society (Tite
forms till it is quite certain that they have no differentiated type
of reproduction.” Prof. Thaxter kindly enclosed a drawing of
the Coccidotrophus Hyphomycete which is here reproduced as
Pip. 12.
The various organisms just described can, of course, have
little or no effect on the health of the beetle colonies. Their
decay and eventual extinction is due to ants destroying the
beetles and their larve or to predators or parasites destroying
the coccids on which they depend for an important part of their
diet. Colonies in all stages may be invaded by the tiny Solenop-
sis altinodis, which I believe to be the most persistent and deadly
enemy of the beetles. I have opened petioles containing both
beetles and Solenopsis workers, but no beetle larve or only frag-
ments of them, showing that the ants begin their depredations by
slaughtering the offspring of the beetles. In such petioles the
coccids remain uninjured, and the same is true of the pupe im-
mured in their cocoons. Many other petioles reveal conditions
from which the last stages in the history of the colony may be
inferred. In such petioles only dead beetles and a number of
dead Solenopsis are found, indicating that the ants, after destroy-
ing all the larve, attack the beetles and that they, in the ensuing
bitter conflict, often defend themselves with their powerful jaws
to such good purpose that they succeed in killing many of the
ants. But as more Solenopsis are probably continually entering
the petioles as auxiliaries the beetles finally succumb and the
colony is exterminated, with the exception of such pup# as may
be present in their protective cocoons. The beetles emerging
from these may, conceivably, after the Solenopsis invasion is
over, start a new colony in the same petiole or emigrate to other
petioles. If no pupz are present the petiole is sooner or later
entered by a pair of young migrant beetles, which pack the dead
bodies of the previous occupants, together with the ants they
have slain, into the ends of the cavity and establish a new colony.
In some of the colonies in the last stages of devastation above
described, I failed to find any coccids. They may have been either
eaten or carried away by the Solenopsis. Sometimes they remain
undisturbed, however, and may, perhaps, be taken over by the
beetles emerging from the cocoons or by any pair of young
beetles entering the petiole and establishing a new colony.
1921] Wheeler: Some Social Beetles 83
We can picture to ourselves the fierce battles which rage
in the petioles between the beetles and the Solenopsis workers,
probably mostly at night, for the Solenopsis is a nocturnal spec-
ies, and the precarious life of the beetles in parts of the jungle
where the ant is abundant. The beetles must live, in fact, like
the ancient Greeks, always in danger of invasion from the war-
like hords of barbarians. Yet even in quiet recesses of the
jungle, where the Solenopsis may happen to be rare or absent,
the attachment of the Coccidotrophus colonies to the Tachigalia
is sure to be severed as the plant grows and the workers of the
colonies started in one or more of the petioles by dedlated queens
of Azteca or Pseudomyrma have become sufficiently numerous to
take possession of every petiolar cavity and patrol the whole sur-
face of the plant. Perhaps some of the inquiline ants may occa-
sionally kill or oust the beetles, but as these ants merely occupy
a petiole here or there on the young trees, they cannot be regarded
as very serious enemies. Many of them, too, are small, timid
ants, which probably have their own battles to fight with the
insidious Solenopsis and are destined to be supplanted, like
the beetles, by the obligate Pseudomyrmas and Aztecas.
During the struggles between the beetles on the one hand and
the Solenopsis, Pseudomyrmas, and Aztecas on the other,
the poor coccids evidently play somewhat the same defenceless
role as the cattle in a country overrun by contending armies—
they merely change masters and are either eaten, or carried off
or permitted to remain and produce honey-dew for the victors.
But before any such change of masters occurs they are often
decimated or even exterminated by three enemies of their own,
which may be briefly described seriatim.
In a few of the beetle colonies I have seen a number of
larve of a very small Coccinellid beetle, described by Schwarz
and Barber as Scymnus xantholeucus (Zoologica III, No. 6).
These larve when full-grown resemble the larger coccids so
closely in size, form and color and are covered with such a similar
layer of snow-white wax, that I frequently overlooked them in my
living colonies and detected their presence only in the preserved
material after my return to the United States. They move slowly
about among the beetles and their larve and devour the coccids.
I am inclined to believe that by the time they are ready to pupate,
84 Zoologica: N. Y. Zoological Society [III;3
they have also devoured all the beetle larve since I found two
petioles each containing nothing but a pair of Scymnus pupe, a
few shrivelled remains of coccids and the kitchen-middens at the
ends of the cavity. The pupz# were attached to the wall by
their caudal ends and with their longitudinal axis parallel with
that of the petiole. They were of a waxy yellow color, with their
surface studded with short, blunt hairs. Several days later the
imaginal beetles emerged. They measured 1.7 mm. in length
and 1.2 mm. in width and were pale yellow, with the basal two-
thirds of the elytra, the meso- and metasternum and the median
third of the first and second abdominal segments fuscous.
A much more abundant enemy of the the coccids is a peculiar
predacious Itonidid (Cecidomyid) fly, which Dr. Felt has de-
scribed as Diadiplosis pseudococci (Zoologica III, No. 8). The
larvee of this insect are orange red and are often found in clusters
of as many as eight to a dozen around groups of coccids. The
whole mass is covered by a tough, dense, white web, or tent, which
is spun by the larve in such a way as to shut them and the coccids
off from the cavity of the petiole and therefore from contact
with the beetles or their larve. Thus secure from interference
the Diadiplosis larve devour the coccids at their leisure, attack-
ing them from the ventral side where their integument is thin-
est and free from wax. The coccids are eventually reduced to
their dorsal integument. When mature the Diadiplosis larva
pupates where it has been feeding, often in the midst of a group
of young or full-grown larve, and without orienting itself with
respect to the longitudinal axis of the petiole. Just before eclo-
sion the pupa forces its body, head foremost, through the silken
tent and projects into the cavity. The fly then emerges and prob-
ably either lays its eggs among any surviving coccids or emigrates
from the petiole and enters other beetle colonies. The adult fly
is a very delicate little midge measuring only 1.25 to 1.5 mm.
and of a reddish-brown color, with the abdomen red internally
and the sclerites somewhat infuscated.
Considerable interest attaches to this insect, because, unlike
the great majority of Itonidids and the species noticed above (p.50),
it does not make plant galls but is predaceous. The genus Diadi-
plosis was originally established by Felt (1911a), for D. cocci Felt,
a species reared in the island of St. Vincent from larve preying
1921] Wheeler: Some Social Beetles 85
on the eggs of a coccid, Saissetia nigra Nietn., on stems of Sea
Island cotton. In another paper published during the same year
(1911b) he gives a list of 19 species of known zoophagous Amer-
ican Itonidids. The list includes species of Endaphis, Arthro-
cnodaxz and Mycodiplosis feeding on mites, an unidentified species
of Cecidomyia feeding on the eggs of Cicada septemdecim, sev-
eral species of Aphidoletes and Lestodiplosis preying on aphids
and of Lestodiplosis, Dentifibula, Diadiplosis, Coccidomyia, Ceci-
domyia, Lobodiplosis, Mycodiplosis and Dichrodiplosis preying
on various coccids. The Diadiplosis from British Guiana seems
to be closely related to the type of its genus. According to Kiis-
ter (1911), certain European Itonidid larve have been described
by Riibsamen (1899) and Kieffer (1902) as preying on the larve
of gall-makers of the same family.
Aimost as abundant as the Diadiplosis in the beetle colonies
is a Hymenopterous parasite of the Pseudococcus, namely Blep-
yrus tachigaliz Brues (Zoologica III, No. 9), a small Encyrtid of
the family Chalcidide. The white larva of this insect lives in the
coccid and grows with it, eventually becoming so voluminous that
the coccid’s body is very convex both dorsally and ventrally. The
coccid grows increasingly sluggish and inert and its wax-glands
cease to function so that its integument takes on a dull brown-
ish color and the wax-pencils disappear from its periphery. The
beetles and their larve are, of course, quite unaware of these pro-
found changes in their parasitized cattle and still continue to
stroke them, often for long periods, although there is no honey-
dew forthcoming as a reward for their efforts.
When full-grown the larval Blepyrus does not escape from
the coccid but remains within it and forms an amber-colored,
regularly elliptical cocoon about 2 mm. long and therefore very
nearly as large as the coccid, which is now reduced to a mere
skin enveloping the huge parasite. The cocoon seems to consist
of a hard, glassy substance, possibly a modified silk, and is cov-
ered except on its ventral side with small circular spots which
represent thinner, depressed areoles in its wall. Where these
areoles are lacking on the ventral side the wall is homogeneous
and distinctly thinner than elsewhere, but has a number of small
pointed projections which seem to pierce the ventral integument
of the coccid and to attach the cocoon rather firmly to the wall of
86 Zoologica: N. Y. Zoological Society [III ;3
the petiole. Since the parasitized coccid remains in the food-
groove with its body in the usual position the cocoon necessar-
ily has the same position and orientation, i.e. with its long axis
parallel with the long axis of the petiole. The cocoon gradually
grows darker, passing from amber-yellow to dark brown and by
the time it has reached this stage, the dead tissues of the coccid
enveloping it, except those on the ventral side, between the cocoon
and the wall of the petiole, disappear, leaving the lateral and dor-
sal surfaces of the cocoon fully exposed. I am inclined to believe
that the dead tissues of the coccid are eaten away by the beetles
or their larve, but as they are very soft and disintegrate easily,
they may perhaps be rubbed off merely by the attrition of the
insects as they move back and forth in the petiole. Two of the
denuded cocoons in the stage and with the orientation just des-
cribed are shown in Fig. 11.
When the completed Blepyrus cocoon is cleared in carboi-
xylol, mounted in balsam and examined as a transparent object,
the larva is found to have pupated within it, after extruding a
number of large meconial pellets, the undigested remains of all
the food it swallowed while it was living on the tissues of its
host. In most cases, at least, the head of the pupa is at the caudal
end of the coccid. The imago, when mature, cuts a large round
hole in the end of the cocoon (see upper part of Fig. 11) and
emerges as a short, thickset, broad-headed fly, only 1.5 mm. long,
with a metallic green face, a black, more or less bronzed body,
black and yellow antenne and legs and basally infuscated wings.
It is very active, and like other small Encyrtids skips about by
using the long saltatory spurs on its middle tibie. After mating
the female undoubtedly oviposits in the young coccids either in
the same or in some other beetle-inhabited petiole.
This parasite seems not to be nearly so serious a menace to
the Coccidotrophus colonies as the Scymnus and Diadiplosis, since
the infested coccids are probably able to supply the beetles and
their larve with honey-dew till both host and parasite are nearly
mature. Hence one often finds several infested coccids and
Blepyrus cocoons in petioles inhabited by flourishing beetle col-
onies. In one such colony I counted more than fifty cocoons and
a dozen large coccids swollen with parasites that were still in
the larval stage. 1
1921] Wheeler: Some Social Beetles | 87
I have failed to find more than one of the three species of
coccid enemies in a single petiole. Their combined action, if
they actually ever occur together, would, of course, not only
greatly hasten the extermination of the coccids, but would seri-
ously interfere with their own development. It may be noted
incidentally that none of these enemies occurs in the petioles of
the large Tachigalias inhabited by the obligate Pseudomyrmas
and Aztecas. In such plants the coccids are free from all preda-
tors and parasites and are not only more numerous but attain
a larger size than in the petioles of the small shade trees tenanted
by the Coccidotrophus. The ants are undoubtedly much more
efficient than the beetles in keeping small miscellaneous guests
and synoeketes out of their nests. This is particularly true of
the Pseudomyrmas. Although I have collected the entire colonies
of many of these ants on several different trips to the American
tropics, the only synoekete I have ever seen associated with them
was a Microdon larva described many years ago (1901) from
the nest of Pseudomyrma mexicana Roger. Even coccids are kept
and attended by only a few species of Pseudomyrma.
Before concluding my account of Coccidotrophus I may intro-
duce a few statistical data, which are probably valid only for
the particular time and locality of my observations. While at
the Tropical Labaratory I noted roughly the condition of the
contents of each of the Tachigalia petioles I opened on a particu-
lar day. On some days only a few petioles were opened and
the results are not worth transcribing. The following collection,
however, gives a more interesting picture owing to the number
of petioles examined:
August 9. Collected 253 petioles from young Tachigalias
114 to 7 ft. high growing along the Cuyuni Trail. Of these 37
or about 14% were either too young to have inhabitants or con-
tained solitary Pseudomyrma queens founding colonies or small
colonies of inquiline ants; 203 or about 86% either contained or
had contained beetle colonies. Of the latter number, 50 contained
incipient colonies, i.e., a single pair of beetles or more rarely sin-
gle beetles which had just entered the petioles and were busy
“cleaning house.” In one petiole one of the beetles of a pair was
guarding the entrance while the other was shovelling frass and
the remains of previous occupants with the top of its head into
88 Zoologica: N. Y. Zoological Society [111 ;3
the pointed ends of the cavity. Sixty-four of the beetle colonies
had larve and were in what I have called the second and third
stages. In nearly every case coccids were seen. Eighty-nine of
the colonies were either moribund or extinct. Solenopsis altinodis
workers either living or dead, were present, sometimes in con-
siderable numbers, in 35 of these petioles, and 10 of them still con-
tained large coccids that had been shut off by webs and were being
devoured or had been already devoured by Diadiplosis larve. In
one petiole two of the flies had emerged. In 6 of these petioles
the Solenopsis workers had destroyed the beetles and their larve
and were still running about. When I tore away the webs
covering the Diadiplosis larve the ants at once seized and killed
them. The webs serve, therefore, not only to protect the Itoni-
dids from the beetles and their larve, but also from the Solenop-
sis.
C. EUNAUSIBIUS WHEELERI SCHWARZ AND BARBER.
This beetle, though superficially very similar to Cocctdo-
trophus socialis, can be easily distinguished in all its postembry-
onic stages. The adult beetle (Plate III, fig. 1, Plate VI, figs. 6
to 10) is distinctly smaller, measuring only 3-3.5 mm., perman-
ently of a red color like the immature Coccidotrophus and there-
fore never deepening into the dark chestnut color of the latter.
The antennal clubs are larger and broader and much more dis-
tinctly marked off from the more proximal joints, the eyes are
much. larger, the anterior border of the front is much less
deeply emarginate, the femora are less incrassated and the pos-
terior pair has a small tooth on the flexor side. The surface of
the body is smoother, the punctation being less pronounced. The
larva is more slender, with the head and dorsal surface distinctly
gray, owing to a deposition of fine pigment granules in the integu-
ment. The pupa can be at once recognized by the presence of
four large, equidistant tubercles on each of the parallel lateral
borders of the pronotum (Fig. 8a, Plate IX, fig. 23). For many
of the less obvious differences between the various instars of
the two beetles the reader may be referred to the excellent des-
criptions and figures of Schwarz and Barber (Zoologica III, No.
6) and Boving (Zoologica III, No. 7).
1921] Wheeler: Some Social Beetles 89
The prominent tubercles on the sides of the pupal pronotum
of EHunausibius merit somewhat fuller consideration, because
they present a striking instance of the retention in an earlier
ontogenetic stage of a character which may be completely lost
in the adult. An examination of the common saw-toothed grain-
beetle, Oryzephilus (formerly Silvanus) surinamensis L., repre-
sented in Fig. 7, and other species of the same genus, shows
that the adult beetle has six acute teeth on each of the lateral
borders of the pronotum, corresponding to and arising within
as many large, blunt tubercles of the pupa. These structures
were long ago noticed by Coquerel (1849) and Perris
(1852) and by the former erroneously supposed to be por-
tions of some tracheal system peculiar to the pupa. In
Nausibius (N. clavicornis) the pronotum of the beetle bears
six obtuse teeth on each side. In other Silvanid genera, such
as Silvanus and Cathartus as well as in Eunausibius and
Coccidotrophus these teeth are either altogether absent in the
imago, or reduced to the first pair, which form the anterior
corners of the thorax. Hunausibius has well-developed teeth in
this position but in Coccidotrophus the anterior corners of the
pronotum are merely rectangular. It is therefore interesting to
find that the pupa of Hunausibius has four well-developed pairs
and a fifth vestigial pair of tubercles, that these tubercles
decrease in size anteroposteriorly, and that only the first pair
gives rise to teeth that persist in the adult. In Coccidotrophus
the reduction of the pupal tubercles is carried much further since
there are only small vestiges of the three anterior parts of Euna-
usibius, none of which gives rise to teeth in the imago. More-
over, the second and third pairs of pupal tubercles may be repre-
sented by only a single tubercle on one side of the pronotum, as
Boving observed (Zoologica III, No. 7). The tubercles are, in
fact, so evanescent that they have become very variable, like ves-
tigial organs in general. This is seen in Fig. 8—b to f, represent-
ing the prothoraces of five Coccidotrophus pupe selected from a
series of fifty specimens. We may safely conclude, therefore, first,
that Eunausibius and Coccidotrophus are derived from ancestors
which had a 12-toothed pronotum like the species of Oryzae-
philus; second that this condition disappeared first in the imago
and still tends to linger on in the pupa, and third, that the tuber-
cles have a tendency to disappear in sequence in a posteroanter-
90 Zoologica: N. Y. Zoological Society [III;3
ior direction. There can be little doubt that the dentation of
the sides of the thorax is a very ancient character not only in
the Silvanids but also in the Cucujids (as restricted by Boving),
since vestiges of the teeth can also be clearly seen in the imagines
of certain genera of the latter family (Cucujus, Brontes).
I have already alluded to the fact that Hunausibius colonies
are much rarer at Kartabo than those of Coccidotrophus though
both species may occur in the same localities and even in differ-
ent petioles of the same young Tachigalia. And not only are
all the instars of Hwnausibius smaller than those of Coccidotro-
phus but the colonies are also much less populous. The largest
I have seen consisted of less than a dozen beetles and not more
than two dozen larve. The habits, so far as I have been able
to observe them, are much like those of Coccidotrophus. The
Eunausibius also feed on the nutritive parenchyma in the walls
of the petiole but they do not dig long grooves in the tissue but
only narrow elongate pits, nor do they build up their frass in
parallel or vermiculate ridges but plaster it in a thin layer over
the walls of the petiole, so that the latter are smooth and even.
The elongate entrance to the petiole seems not to be provided
with a wall of frass. In one petiole I found that the pair of
parental beetles had entered through a large hole about 2.5 mm.
in diameter which had evidently been made by some larger insect.
The beetles had plugged the opening with frass, leaving a small
elliptical opening in the center just large enough to fit the head
of the beetle when acting as a sentinel. Coccids are found in
the elongated pits in the nutritive tissue but are few in number
and of small size, though the Hunausibius solicit and drink their
saccharine excretions in the same manner as Coccidotrophus. The
cocoons of Hunausibius, apart from their smaller size and some-
what more delicate walls, are very much like those of Coccido-
trophus and are, in all probability, constructed in the same man-
ner.
I have seen so few colonies of Hunausibius that I can give
no account of its enemies nor of those of its coccids. In all prob-
ability it is even less able than the more vigorous and more proli-
fic Coccidotrophus to withstand the insidious attacks of Solen-
opsis altinodis. The whole appearance of the beetle and its colon-
1921] Wheeler: Some Social Beetles 91
ies is that of a feeble, anemic and harried species on the verge
of extinction.
GENERAL CONSIDERATIONS
The behavior of the social Silvanids described in the pre-
ceding pages and the conditions under which they live are suf-
ficiently startling to stimulate reflection and a comparison with
other species of the same and allied families. Such comparison,
as an ethological method, has so often thrown light on what ap-
peared at first sight to be unique and incomprehensible instincts
and their settings that we may hope by resorting to it to trace
the peculiar conditions in Coccidotrophus and EHunausibius to
simpler and more general phenomena. Since both the setting,
or environment and the responses, or behavior of the beetles are
rather complicated it will be best to consider them separately and
to begin with the setting, i.e. with the Tachigalia biocoenose.
The general ethological concept of the ‘biocoenose’ was, of
course, more or less clearly recognized by many of the early zoolo-
' gists. Although the term seems to have been first used by Mo-
bius (1877), even Réaumur had an inkling of the value of study-
ing insects in association with their host plants. He says: “I
would that the observers who busy themselves with the history
of insects gave catalogues of those that feed on every plant.”
In the middle of the last century Perris (1852-1862) devoted
many years to the study of the insects associated with the mari-
time pine and the chestnut in France, and Kaltenbach (1874)
attempted to list all the phytophagous insects of Germany accord-
ing to their host plants. In the United States Packard’s volume
(1881) on the forest and shade-tree insects and Mrs. Dimmock’s
paper (1885) on the insects of the birch represent more modest
studies of the same kind. Perhaps none of our entomologists
has been more thoroughly convinced of the advantages of study-
ing insects and other animals as components of biocoenotic com-
plexes than Forbes. Forty years ago he expressed his general
- eonvictions on this subject in his paper on the food of fishes and
insects (1880) and he has returned to the subject in a recent
address (1915). His fine papers on the strawberry and maize
plants and their associated organisms (1884, 1894-1905) also
clearly illustrate the great value of biocoenotic investigations.
92 Zoologica: N. Y. Zoological Society [IlI;3
Picard (1919) has recently published an interesting paper on the
insects of the fig in Southern France.
As a mere record of the insects associated with a tropical
plant my study of the Tachigalia biocoenose is necessarily frag-
mentary, owing to the few weeks I could devote to it, but it ac-
quires considerable interest from the fact that the Tachigalia is a
myrmecophyte, or one of those plants which are supposed to be
peculiarly adapted structurally to association with battalions of
protecting ants. The only organs which can be cited, however, as
such an adaptation aré the fusiform enlargements of the petioles,
which undoubtedly furnish excellent lodgings for all the various
ants, both inquiline and obligate. The plant is utilized also as a
source of food by the obligate species through the instrumental-
ity of the coccids, which are kept in the petioles and draw their
food by preference from the strands of nutritive parenchyma.
The beetles also use the petioles as lodgings and not only utilize
the species of coccid as a copious source of sugar and water but
also feed directly on the tissues of the plant. The plant is there-
fore more completely exploited by the beetles than by the ants and
might be said to be more perfectly adapted to the former than to
the latter. But the question as to whether the peculiar structure
of the petiole is really an adaptation to either of these groups of
insects is one which I shall leave to the botanist. Prof. Bailey
will no doubt deal with it in connection with the same probler.
in the other South American myrmecophytes which he has inves-
tigated. That both the ants and the beetles have adapted them-
selves to the plant cannot be doubted and this adaptation, as I
have shown, is exhibited in three degrees, the inquiline ants
merely using the petiolar enlargements as lodgings, the obligate
ants as lodgings and through their herds of coccids as indirect
sources of food, and the beetles as lodgings and as both direct
and indirect sources of nutriment.
Of course, a particular biocoenose is not an isolated, per-
fectly self-contained association of organisms but shares some
of its components with other biocoenoses. Thus the Tachigalia
is part of a large association, or biocoenose of jungle trees grow-
ing under certain conditions of soil, humidity, light, tempera-
ture, ete. The Atta cephalotes, which occasionally defoliates the
young tree is the center of an elaborate biocoenose of its own,
1921] Wheeler: Some Social Beetles 93
comprising all the trees it habitually defoliates, its fungus gar-
dens, its myrmecophiles of the Blattid genus Atiaphila, the toads,
lizards and ant-eaters which feed on the foraging workers, the
Amphisbaenians which live in the penetralia of the huge nests,
etc. The two ants, Camponotus femoratus and Crematogaster
parabiotica, which attend Membracids on the young shoots of
the Tachigalia, are really characteristic members of the very
peculiar ‘ant-garden”’ biocoenose, which I have described in an-
other paper (1921), and the Dolichoderus attelaboides belongs
to still another biocoenose of which many Melastomaceous plants
and their Membracid parasites are important components.
A particular biocoenose must also, of course, have a phylo-
genetic history, i.e. we must conceive it to have been gradually
built up, integrated and organized in time from components
which detached themselves from other biocoenoses and attached
themselves as satellites to an organism which furnished more
congenial conditions of life. Owing to the basic nutritive inter-
dependance of animals and plants, a particularly favorable plant
usually constitutes the primary focus of a biocoenose. The vari-
ous parasites, scavengers and synoeketes, which live with the
insects that immediately depend on this plant merely use the
former as so many secondary or tertiary foci. Thus in the
Tachigalia biocoenose the primary focus is the young plant
and the center of the focus the leaf-petiole, the secondary focus
is represented by the coccids and the tertiary foci by the ants
and beetles to the extent that they attract predators, parasites
and scavengers.
It is permissible, perhaps, to reconstruct the phylogenetic
sequences of the various organisms that have become associated
to form the Tachigalia biocoenose. Not improbably the tree, like
many other trees of the Neotropical jungle, was originally peo-
pled throughout its life by a certain number of miscellaneous,
inquiline ants. Among these were several species of Pseudo-
myrma and Azteca, both large genera comprising numerous forms
which still habitually inhabit any available hollow twigs or peti-
oles of the most diverse trees and shrubs. Later the number
of these ants was reduced, through the advent of the coccids and
their definitive association with the Tachigalia, to a very few
species, the putative ancestors of the present Ps. damnosa and
94 Zoologica: N. Y. Zoological Society [III;3
maligna and A. foveiceps, because the coccids enabled them to
acquire very intimate trophic relations to the plant. The coc-
cids present an unsolved problem in this connection. It would
seem that the Tachigalia must be their true host-plant, and that
the various other plants on which they are known to live, are
subsequent, or secondary hosts, possibly acquired when the
natives of British Guiana and of the surrounding countries took
to making clearings in the jungle and growing in them various
introduced plants such as pine-apples, Hibiscus, etc. The truth of
this statement can, of course, be established only by further in-
vestigation of Pseudococcus bromelie throughout its range.
When the obligatory ants had thus acquired their definitive at-
tachment to the tree, the miscellaneous inquilines necessarily
became restricted to its youngest stages since they were no
longer able to compete with the obligates for the possession of
nesting sites on the adult plant. The Silvanid beetles were prob-
ably relatively late intruders which found that they could inha-
bit the young tree for a considerable period before the queens of
the obligate ants had succeeded in maturing their broods of
belligerent workers. At first the beetles merely used the petiolar
cavities as lodgings and fed on the nutritive parenchyma in their
walls, but later they discovered the coccids and learned how to
obtain their honey dew and came to depend more and more on
this saccharine nutriment. The various parasites, scavengers,
ete., which infest the beetle colonies and their droves of coccids
obviously represent still more recent accessions to the biocoenose.
The other insects, such as Atta cephalotes, the Membracids and
their attendant ants, the caterpillars and gall-flies of the leaves,
etc., may belong to the ancient miscellaneous fauna which origin-
ally attacked or frequented the Tachigalia in all its stages, when
it was quite as “unprotected” as the great majority of jungle
plants. ;
Turning now to a consideration of the beetles themselves,
it would seem to be desirable to review their activities in the
light of what is known concerning the other members of the
natural family to which they belong. Here, however, we en-
counter difficulties, for the family Cucujide (sensu lato) has been
more neglected by taxonomists and students of insect behavior
alike than any other family of equal size in the order Coleoptera.
1921] Wheeler: Some Social Beetles 95
As understood by Coleopterists the family Cucujide belongs to
the huge and very inadequately analyzed Clavicorn complex of
families, but its characters are so striking that they have arrested
the attention of some of the specialists. Thus Leconte and Horn
(1883) long ago remarked: ‘This family is evidently an antique
and synthetic type, which exhibits alliances with both Hetero-
mera and Rhynchophora more than any other Clavicorn fam-
ily.” And Handlirsch (1908) says: “The family Cucujide,
which Ganglbaur places in the midst of typically Clavicorn
forms, exhibits many primitive characters and at the same time
high specialization. I do not believe that their antennze can be
derived from those of the Clavicorn type, although the Cucujids
agree with this group in the number of their Malpighian tubules
(six). Perhaps the Cucujids branched off very near the base
from the Cantharid stem, but possibly, and I regard this as
more probable, they form an independent series.” In his phyle-
tic tree (opposite p. 1278), Handlirsch therefore depicts the
family as arising from the Protopolyphaga as far back as the
beginning of the Coenozoic. Several species of Silvanus and one
of Passandra are, in fact, known from the Baltic Amber (Lower
Oligocene Tertiary), and Wickham (1920) cites Laemophloeus
vestitus Scudder from the Green River Eocene and three species
of Lithocoryne and a Pediacus from the Miocene of Florissant.
That the family must be an old one is indicated also by the
fact that New Zealand possesses some 20 indigenous species of
Cucujide, distributed over 12 genera, mostly peculiar to the
islands, which are said to have been separated from Australia
during the Jurassic.
Kolbe (1910) is also of the opinion that the Cucujids are a
primitive group. He says: “The very lowly organized Cucu-
jids are not only in part characterized by a prothorax of very
primitive structure (as in the Adephaga) but primitively in-
serted (inframarginal) and primitively constructed (filiform or
moniliform) antenne.” Leng (1920) places the Cucujids in the
lower portion of the series of Clavicorn families, near the Rhizo-
phagide and Erotylide. In a brief study of the larve of Cucu-
jids, de Peyerimhoff (1902-’03) calls attention to their great
diversity and their resemblance on the one hand to the larve of
Cryptophagus among the Clavicorns and on the other to Pyro-
96 Zoologica: N. Y. Zoological Society [III;3
chroa among the Heteromera. Hamilton (1886) long ago noticed
the resemblance of the larval Cucujus clavipes to the Pyrochroid
Dendroides canadensis larva. As shown in his very valuable
paper on the larve of Coccidotrophus and other genera (Zoo-
logica III, No. 7) Béving divides the Cucujide auctorum into
four families, the Silvanide, Cucujide (sens. str.), Leemophleeideze
and Scalariide. The last of these he relegates to the group
Cleroidea, and states that they are closely connected with the
family Bothioderide of Craighead (1920).
Apart from several species of considerable economic im-
portance, the little that is known concerning the habits of these
four families of beetles is scattered through the literature. Such
data as I have been able to glean in regard to the European,
North American and cosmopolitan species have been brought to-
gether in condensed form in Zoologica III, No. 5. From these data
it will be seen that the Cucujids, taken as a whole, exhibit certain
tendencies which are not without significance in connection with
the peculiar behavior of Coccidotrophus and Eunausibius. If we
exclude the Scalariide, which Fiske (1905) has shown to be
parasitic in their larval stages—resembling in this respect the
Bothrioderide—we notice that the various genera and many of
the species of the remaining families show an extraordinary
diversity, one might say versatility of behavior. They occur in
a great variety of habitats such as stored human foods of vege-
table origin, under bark, in decaying wood, in the burrows of
bark-beetles, under dead leaves and rubbish, and feed on all
sorts of substances mainly of a vegetable and especially of a
concentrated or highly nutritious character. Many of the spe-
cies are scavengers, others are undoubtedly predaceous and prey
on the larve of other insects. The adult beetles seem to be
rather long-lived and usually, if not always, live gregariously
with their larve, all the active stages feeding on the same sub-
stances. The developmental period is certainly very brief in
some species, as e.g. in Cathartus advena, the whole life-cycle
of which, from the egg to the imago may require only three weeks
and in the saw-toothed grain beetle (Oryzzphilus surinamensis)
less than a month. As a rule both the beetles and the larve of
the vegetarian and detritivorous species are very tolerant of the
presence of other insects and actually seem to seek their compan-
1921] Wheeler: Some Social Beetles 97
ionship, especially when the food supply is abundant. Thus O.
surinamensis is often found living with the rice weevil (Cal-
andra oryzx), Cathartus advena with the Indian meal moth
(Plodia interpunctella) and O. mercator with a Tenebrionid
grain-beetle, Palorus subdepressus. Many of the species of
Lemophloeus constantly live in the burrows of Scolytid beetles
and feed on their dejecta. Owing to these peculiarities and espe-
cially to their very diverse and plastic feeding habits many of the
Silvanids and Lemophloeids have become cosmopolitan house-
hold pests capable of doing considerable damage to many of the
staple stored foods of our own species.
If with this general complex of behavioristic tendencies
exhibited by the European and North American Cucujids (sens.
lat.) we compare the activities of Coccidotrophus and Hunausi-
bius, we find that the latter while retaining many of the ancient
and primitive family traits nevertheless exhibit several of them
in a peculiar and highly specialized form. Thus the merely gre-
garious habits of the adults and larve of the northern and cos-
mopolitan Cucujids have become more definitely social in the
Tachigalia beetles, and their toleration of alien insects has
increased; the feeding of the ancient Cucujids on various vege-
table substances has become specialized to the point of concen-
tration on a particular tissue of a particular plant and both
adult beetles and larve have become coccidophilous. The con-
struction of the cocoon, too, exhibits peculiarities not found in
any other Cucujids. Owing to the unusual interest of these
various specializations they may be discussed at greater length
under separate captions.
1. SocIAL LIFE AMONG THE COLEOPTERA
If we regard as truly social only those insects in which the
parent or parents live with their offspring, protect them and
either feed them directly or prepare materials for their susten-
ance, there seem to be only three groups of beetles that meet these
requirements, namely, the Platypodide, the Scolytide (Ipide)
and the Passalide. The Platypodidze and that portion of the
family Scolytide, comprising, according to Hagedorn (1910) the
tribes Corythaline, Xyleborine and Spongiocerinz, with some
400 described species, mostly tropical, are commonly known as
98 Zoologica: N. Y. Zoological Society [III;3
ambrosia beetles, because like the Attiine ants of the Neotropical
Region and many Old World termites they cultivate fungi as
food for themselves and their larve. The remarkable social
organization and the food-fungi of these beetles have been stud-
ied by Eichhoff (1881), Hubbard (1897a, 1897b), Hopkins
(1898), Neger (1908a, 1908b, 1909, 1911), Schneider-Orelli
(191la, 1911b, 1912, 1913) and others, Hubbard’s account of
the habits of Platypus compositus of our Southern States is so
interesting that I quote it at length:
“These social instincts reach their highest development, ap-
parently in the genus Platypus. The species of this genus are
readily known by their very long cylindrical bodies, their promin-
ent head, flattened in front, the flattened and spur-tipped joint of
the front legs, and in the males the spine-like projections of the
wing cases behind. They are powerful excavators, generally
selecting the trunks of large trees and driving their galleries
deep into the heartwood... The female is frequently accompanied
by several males, and as they are savage fighters fierce sexual
contests take place, as a result of which the galleries are often
strewn with the fragments of the vanquished. The projecting
spines at the end of the wing cases are very effective weapons in
these fights. With their aid a beetle attacked in the rear can
make a good defence and frequently by a lucky stroke is able to
dislocate the outstretched neck of his enemy. The females pro-
duce from one hundred to two hundred elongate-oval pearly
white eggs, which they deposit in clusters of ten or twelve in the
galleries. The young require five or six weeks for their develop-
ment. They wander freely about in the passages and feed
in company upon the ambrosia which grows here and there upon
the walls. The chitinous ridges upon the thoracic segments, to-
gether with the row of tubercles upon the other segments, enable
the larva to move as rapidly through the galleries as if it were
possessed of well-formed legs. The mouthparts of the larva are
also provided with strong cutting mandibles, but the inner jaws
are not adapted to masticating hard food, such as particles of
wood. The older larve assist in excavating the galleries, but they
do not eat or swallow the wood. The larve of all ages are sur-
prisingly alert, active and intelligent. They exhibit curiosity
equally with the adults, and show evident regard for the eggs
1921] Wheeler: Some Social Beetles 99
and very tender young, which are scattered at random through
the passages, and might easily be destroyed by them in their
movements. If thrown into a panic, the young larve scurry
away with an undulating movement of their bodies, but the older
larvee will frequently stop at the nearest intersecting passageway
to let the small fry pass, and show fight to cover their retreat.
When full grown the larva excavates a cell, or chamber, into
which it retires to undergo its transformations. The pupa cells
are cut parallel with the grain of the wood and generally occur
in groups of eight to twelve along some of the deeper passages.
The older portions of the galleries are blackened by the long-
continued formation of the food fungus. In the ambrosia of
Platypus compositus the terminal cells are hemispherical, and
are borne in clusters upon branching stems.”
The habits of several genera of ambrosia beetles of the
family Scolytide were investigated by Hubbard, and one of our
species, Xyleborus xylographus, which has an extensive circum-
polar distribution and is common in the wood of fruit-trees, has
also been studied by EHichhoff (1881) and Hopkins (1898). I
quote the latter’s account of this insect which may serve as a
paradigm of the whole group:
“The fertilized females pass the winter in their brood cham-
bers and emerge in the spring (April and May, near Morgan-
town, W. Va.). They are then attracted to sickly, dying or
felled trees, in the living or moist dead wood of which they
prefer to excavate their brood galleries. A crevice or opening in
the bark, such as may be made by other insects, or, as I have
observed, those made by the yellow-bellied woodpecker, but more
commonly the edge of a wound, in a dead place on a living tree,
is selected as a favorite point of attack. Here a female will
commence the excavation of a mine, and after she has penetrated
the wood a short distance, another female (as I have observed)
will come to her assistance, one working at the excavation, while
the other guards the entrance and assists in expelling the borings.
The primary or main gallery is usually extended into the heart-
wood before eggs are deposited. When the primary gallery is
completed (according to Hubbard) a bed is provided on the
sides of the gallery for the propagation of the special species or
variety of ambrosia fungus which is to furnish food for the
100 Zoologica: N. Y. Zoological Society [Til ;3
future broods. The first set of eggs are few in number (five to
ten) and are placed without any protection on the sides near
the end of the main gallery, or in cavities or short branching
galleries, one-half to one inch from the end, where upon hatch-
ing, the young larve find a supply of ambrosial food. After the
first set of larve have attained considerable size, another set of
eggs are deposited, and so on at intervals until a large family
is reared, in which eggs, larve of all stages of development,
pups, and young and old adults are found crowded promiscu-
ously in leaf-like brood-chambers which are continually broad-
ened or extended by the adults and possibly by the larve, to make
room for the increase. It appears that the brood-chambers are
broadened and extended by the adults, and that the borings,
mixed with the fungus, are softened and furnish additional food
for the larve and young beetles.” At this point in his account
Hopkins introduces the following note: ‘In a brood-chamber
before me just cut from a nearby apple tree, I find a pupa minus
an abdomen. No predaceous enemies can be found, but two or
three half-grown larve are in such a position as to make the
circumstantial evidence quite plain that they are to blame for
the multilation. The remaining portion of the pupa is in a
normal condition, which would indicate that the attack had been
recent and when the victim was alive. This would also indicate
that the helpless pupz may furnish food for the larva in case
of a scarcity of ambrosia, or that*they may be thus disposed
of to prevent an overcrowded brood-chamber.”
The account of Xyleborus continues: “Mr. Hubbard records
the discovery of a death chamber, or a kind of catacomb, in
which the dead mother beetles and other dead friends or foes
of a large colony are consigned by the survivors. In some fresh
specimens of galleries before me I find the same thing, but it
appears that in addition to a resting place for the dead, it is also
utilized for the disposal of all objectional and refuse matter,
which owing to the crowded condition of the chamber, cannot
be conveniently expelled from the entrance. One of the males
found in this set of chambers was excavating a burrow in the
mass of material in the death or garbage chamber. Whether he
was excavating his own tomb, or simply providing bachelor
quarters, I cannot say. The proportion of males in this, as in
1921] Wheeler: Some Social Beetles 101
all other species of the genus Xyleborus, is remarkably small.
There are usually not more than three males in the largest
colonies, or groups of brood-chambers. It would appear from
observations made by Swiner and Eichhoff in Germany, and
the numerous colonies I have examined in this country that there
is, on an average, about one male to twenty females. The males
have no wings, therefore probably do not leave the brood-cham-
bers, but remain with the over-wintering colony until all have
emerged in the spring. They are then left to be smothered in
overabundant ambrosial food, or to the tender mercies of preda-
tory insect enemies which had previously been prevented from
entering the brood-chambers by one or more female sentinels
at the entrance. A few females may emerge from time to time
during the summer to start new colonies, but from the excessively
crowded condition of the brood-chambers during the fall and
winter months, it would appear that the older adults of the
broods excavate branching chambers in which new broods are
developed, and that in these old and new chambers they pass
the winter.”
The third group of social beetles comprise the large
Lamellicorns of the family Passalide, abundant in the tropics
of both hemispheres but represented in the United States by
only a single species, Passalus cornutus Fabr., which ranges as
far north as Massachusetts and Illinois. None of our Coleop-
terists seem to have taken the trouble to study the habits of this
common and conspicuous insect, so that it was left to Ohaus
(1899-1900, 1909) to discover the social behavior in certain
Brazilian species. He found that they live in rotten logs in
colonies, each consisting of an adult male and female with their
larve. The beetles excavate spacious galleries, comminuting
the wood and probably treating the particles with some digestive
enzyme, so that they can be eaten by the larve, which slowly
follow along the galleries just behind their tunneling parents.
Owing to the structure of their mouthparts the larve are quite
unable to break down the wood, and when removed from their
parents soon die. The beetles not only guard their greenish eggs
and diligently provide food for their larve, but also protect the
pupe and feed the imaginal young till their chitinous integument
is completely hardened. Ina former paper (Wheeler and Bailey,
102 Zoologica: N. Y. Zoological Society [Tits 2
1920) I have published an account of the stridulatory organs of
the larval and adult Passalus and have given reasons for believing
that all the members of colony are kept together by the shrill
sounds they are able to emit.
During the summer of 1920 while in Trinidad and British
Guiana my son Ralph and I made a few observations on severai
of the species of Passalus which are very common in rotten logs
throughout the jungle. Just under the bark the beetles make
large, flat cavities, which are later very often occupied by the
fungus-growing ants of the genera Apterostigma, Myrmicocrypta
and Cyphomyrmex, and evidently furnish just the right places
for their more or less globular gardens. In each of the Passalus
colonies examined during July and August there were only two
adult beetles, usually accompanied by a troop of larve varying
little in size and evidently belonging to a single brood. In one
log, however, we found a pair of the beetles guarding a batch of
about 40 large, olive-green, broadly elliptical eggs, some of which
had just hatched. The young larve closely resembled the older
individuals in the structure of the peculiarly modified paw-like
metathoracic legs, which are rubbed over the finely ridged middle
coxe during stridulation, but the hairs on the body were conspic-
uously longer and coarser. Our observations on the beetles and
their larve both in the field and in the laboratory, confirm the
statements of Ohaus.
The preceding account of the Platypodids, Scolytids and
Passalids will suffice to show that they have reached a more
advanced stage of social development than Coccidotrophus and
Eunausibius, though the latter exhibit certain interesting resem-
blances to such ambrosia beetles as Xyloterus. The two Silvanids
really represent a stage in social development intermediate
between that of the families mentioned and the merely gregarious
Silvanus, Oryzephilus, Nausibius etc. Although the colonies of
Coccidotrophus and Eunausibius are founded by pairs of parent
beetles and in the climax stage of their development may comprise
a considerable number of offspring in all stages of development, yet
the latter do not seem to be the recipients of any special care on
the part of the parents, unless we interpret as such the guarding
of the petiolar cavity and the deepening of the food-grooves which
would seem to render the nutritive parenchyma more accessible
ee »,
1921] Wheeler: Some Social Beetles 103
to the larve. And although all the members of the colony, beetles
and larve alike, seem to be very indifferent to one another, except
when they are competing for the honey-dew of the same coccid
or when larve occupy cocoons in process of construction by other
larve, yet under normal conditions there are no signs of hostility
on the part of the beetles and larve even when other individuals
are very annoying. Moreover, the use of the petiolar cavity as
a common domicile, with its kitchenmiddens and more or less
definite arrangement of the frass-ridges and wall about the
entrance, the droves of coccids and the definite orientation of the
eggs and cocoons, all show a much more socialized condition than
anything that has been hitherto observed in other Cucujids. I
believe, therefore, that I am justified in regarding the two
Tachigalia Silvanids as representing a fourth group of social
beetles, of a more primitive type than any of the three families
above considered and differing in the absence of any definite
preparation of larval food by the parents. No such preparation
is necessary, in fact, owing to the peculiar conditions under which
the Coccidotrophus and Eunausibius live, since both the young
and the adults feed on the same substances and these are
furnished by the plant and the coccids, which in turn feed on the
same specialized parenchyma as the beetles.
No doubt the toleration by the beetles and larve of such
different insects as the coccids, the Scymnus larve, the larval and
adult Diadiplosis, the Entomobrya, Aphiochaeta and probably
also of the adult Blepyrus, is due to the same causes as the tolera-
tion by so many ants and termites of numerous myrmecophiles
and termitophiles. Such guests, parasites and synoeketes can, of
course, manage to live only among insects which through long
association with individuals of their own species have come to
tolerate or even to seek the presence of insects belonging to alien,
or unrelated species.
2.—THE DEVELOPMENT OF THE FEEDING-HABITS OF
THE SOCIAL SILVANIDS.
oad
é There would seem to be little doubt that the primitive food
of Coccidotrophus and Hunausibius is the nutritive parenchyma
of the Tachigalia petioles. But this is a very specialized diet
104 Zoologica: N. Y. Zoological Society [TI ;3
compared with that of other Silvanide. Although such genera as
Silvanus, Oryzzphilus, Cathartus and Nausibius eat by prefer-
ence vegetable substances with high protein, starch or sugar con-
tent, none of these forms is known to devour the tissue of grow-
ing plants, and it is even doubtful whether White’s statement
(1872) that the Cucujid Dendrophagus crenatus feeds on the in-
ner bark of conifers, is correct (see Zoologica III, No. 5). The
Tachigalia beetles are primarily attracted by the tree and there
is every reason to regard this peculiar Leguminous plant as
their only host, so that in this respect, also, they are highly
specialized, for while some of the Silvanids, Lemophloeids and
Cucujids (sens. str.) prefer particular trees, they seem never-
theless to thrive equally well in trees of different species,
probably because they do not eat the living plant tissues
but merely require special moisture conditions or the presence
of certain -other insects. We must suppose, furthermore,
that the social Silvanids are primarily attracted to the Tachi-
galia by certain chemical substances in the petioles, a suppo-
sition which seems to offer the only satisfactory explanation, as
Picard (1919) has shown, for the selection of particular host
plants by particular insects. We should have to suppose also,
that the beetles can discriminate between the petiolar substances
of young shade—and older sun-trees, since they confine their
attentions to the former. This is not surprising when we consider
that some insects, e.g., certain Cynipid gall-flies exhibit even
more delicate powers of discrimination since, when ovipositing,
they seem to be able to distinguish between the viability of
different buds or leaves on the same branch.
An even more interesting problem is presented by the
coccidophily of Coccidotrophus and Eunausibius, for nothing like
it has been observed in any other beetles, and apart from the
ants, few insects are known to have developed the ability to
solicit honey-dew from any of the Homoptera. I find only the
two following cases in the literature, the first an observation by
Belt in his “Naturalist in Nicaragua” (1884, p. 228), on wasps
attending Membracids: “Similarly as, on the savannahs, I had
observed a wasp attending the honey-glands of the bull’s horn
acacia along with the ants, so at Santo Domingo another wasp,
belonging to quite a different genus (Nectarinia), attended some
1921] Wheeler: Some Social Beetles 105
of the clusters of frog-hoppers, and for the possession of others
a constant skirmishing was going on. The wasp stroked the
young hoppers, and sipped up the honey when it was exuded,
just like the ants. When an ant came up to a cluster of leaf-
hoppers attended by a wasp, the latter would not attempt to
grapple with its rival on the leaf, but would fly off and hover
over the ant; then when its little foe was well exposed, it would
dart at it and strike it to the ground. The action was so quick
that I could not determine whether it struck with its fore-feet or
its jaws, but I think it was with the feet. I often saw a wasp
trying to clear a leaf from ants that were already in full posses-
sion of a cluster of leaf-hoppers. It would sometimes have to
strike three or four times at an ant before it made it quit its
hold and fall. At other times one ant after the other would be
struck off with great celerity and ease, and I fancied that some
wasps were cleverer than others. In those cases where it
succeeded in clearing the leaf, it was never left long in peace.
Fresh relays of ants were continually arriving, and generally
tired the wasps out. It would never wait for an ant to get near
it, doubtless knowing well that if its little rival once fastened
on its leg, it would be a difficult matter to get rid of it again.
If a wasp first obtained possession, it was able to keep it; for the
first ants that came up were only pioneers, and by knocking
these off, it prevented them from returning and scenting the
trail to communicate the intelligence to others.”
The second case is more remarkable and refers to a Gerydine
Lyczenid butterfly of India, described by Bingham (1907, p. 287) :
“A remarkable habit in one member of the subfamily, viz.,
Allotinus horsfieldi, has been communicated to me by Colonel
H. J. Barrow, R. A. M.C. He writes: ‘I don’t know whether you
have observed the habits of a small plain butterfly which I caught
in Maymyo. I watched it often in the jungle, sometimes for an
hour at a time. It puzzled me at first to know why it took such
an immense time to settle. It would keep within one yard of a
spot and almost settle, twenty times perhaps, before it actually
did. Its legs are immensely long and I discovered why. It settles
over a mass of Aphides and then tickles them with its proboscis,
just as ants do with their antennze and seems to feed on their
exudations.’ The butterfly would settle over rather large ants
106 Zoologica: N. Y. Zoological Society [III;3
that were attending the aphids ‘and did not mind one or two
actually standing up and examining its legs to see who was
there. The ants did not attack it in any way.’ ”’
A comparison of the behavior of the insects considered in
the foregoing paragraphs is very instructive. The predilection
of ants for various Homoptera (aphids, coccids, membracids,
cercopids and psyllids) is well known. Though never observed
among the predatory Doryline and Cerapachyine and rare among
the Ponerine and Pseudomyrmine this predilection is, neverthe-
less, so prevalent among the higher subfamilies (Myrmicine,
Dolichoderine and Formicine) that it has not escaped the most
casual observer. When this type of behavior is highly developed,
as in our species of Lasius, the ants display not only an exquisite
deftness in stroking their trophobionts but also a decidedly pro-
prietary interest in them, most clearly evinced by building
peculiar carton or earthen shelters over them, aggressively
defending them from their foes, or even collecting them and
their eggs in the nests, distributing them over the surfaces of
suitable plants and conveying them to places of safety when the
colonies are disturbed. The whole performance is so elabo-
rately adaptive as to suggest on the part of the ants an intimate
acquaintance with the requirements and habits of their wards.
This is also indicated when the latter fail to respond to stroking,
for the ants do not wear themselves out by prolonged solicitation
after their cattle have discharged their honey-dew, but stand
around as if waiting for more of the saccharine liquid to
accumulate.
The wasp described by Belt and the butterfly described by
Bingham are really robbers, the former having learned to dispos-
sess the ants of their wards, at least temporarily, the latter to
overreach the ants and obtain the honey-dew by stealth. Though
very different, the relations of the beetles to their coccids are no
less extraordinary. The case is, indeed, so far as known, unique
among the Coleoptera. Unlike many species of ants, the beetles
have not yet learned to pick up the coccids and carry them about,
put merely accept them as an integral part of the normal environ-
ment or as members of the colony. The fact that the beetles
clearly recognize the signal of the prospective emission of the
honey-dew by the coecids, when they raise their caudal segments,
1921] Wheeler: Some Social Beetles 107
and the fact that they devote more attention to the posterior than
to the anterior end of the larger coccids, implies a delicate
discrimination, because both ends of the coccid’s body are so very
much alike. Furthermore, the beetle’s antennze and mouthparts
seem to be so clearly adapted to dealing with the coccids as to
indicate that the trophobiotic relations between the two species
have been in existence for a very long time. The beetle’s extraor-
dinary perseverence in stroking individual coccids after they
have been exhausted by repeated emissions of honey-dew might
be interpreted either as a very thorough and hence highly
adaptive method of exploiting the coccids, or as due to a very
imperfect discernment of their physiological peculiarities.
Obviously the most remarkable item of behavior in
Coccidotrophus and EHunausibius is the stroking of the coccids
by the larve of all stages, as well as by the adult beetles. No
one has even considered the possibility of a similar performance
by the larve of ants, wasps or butterflies, since it is difficult
to imagine creatures more unfitted for such behavior. Neverthe-
less, Mr. W. F. Fiske informs me that while he was investigating
certain injurious insects in British East Africa, he saw small
worker ants climbing a tree with their larve and holding them
to the posterior ends of aphids, so that they could feed on the
honey-dew voided in response to the antennal solicitations of their
nurses! Unfortunately no specimens of the ants were preserved,
and from Mr. Fiske’s description I am unable to determine even
the subfamily to which they belonged, but I have no reason to
doubt the statement of an entomologist so competent and so
keenly observant. I surmise that the ant must have been some
Myrmicine which is unable to feed its larve by regurgitation, as
otherwise such behavior would be superfluous.
So specialized a habit as the coccidophily of the two genera
of social Silvanids calls for some consideration of its possible
phylogenetic origin. Under existing conditions, the beetles either
find the coccids already established in petioles that have been
previously inhabited by other beetle colonies or by other insects,
or the coccids enter the young petioles just after they have been
perforated by the beetles,, for insects with sucking mouthparts
cannot, of course, gain access to the cavities in any other way.
That they migrate into the petioles as very young individuals is
108 Zoologica: N. Y. Zoological Society [IlI;3
certain. Later, after completing their growth, they are often
too bulky to escape through the entrances made by the beetles,
and as such imprisoned coccids contain eggs, they might be
supposed to breed in the petioles. I have never been able to find
either the males or the deposited eggs in the petioles, and as the
total number of coccids in a petiole is too small to indicate the
survival of many of the young, I suspect that the beetles, though
averse to devouring the young or mature coccids, nevertheless
consume many of the eggs. How the coccids manage in the first
place to reach the individual Tachigalia plant is a problem which
presents itself also in the case of any of the other often widely
distributed species of the family. That ants have much to do
with carrying certain species of coccids to their host-plants is
very probable. The only other active agents in such distribution
would seem to be birds or the wind.
Soon after they enter the petioles the coccids seek out the
strands of nutritive parenchyma, and sink their slender beaks
into the tissue. And as the beetles keep gnawing at the same
strands, grooves or narrow depressions are soon made in which
the coccids settle, one behind the other, in rows, with the long
axes of their bodies parallel with the long axis of the petiole.
Thus the coccids naturally and inevitably come to lie in the paths
of the feeding beetles, so that these can hardly avoid continual
contact with the waxy creatures and their excrement. Probably
at first the coccids simply voided their excrement in the grooves,
thus drenching the surface of the nutritive parenchyma, so that
the beetles found their bread spread with syrup. But this could
hardly be an unalloyed blessing, because a sticky liquid spread
on the walls of the petiole would almost certainly be injurious
or fatal to the eggs, pupze and younger larve of the beetles.. We
may therefore conjecture that the latter soon learned to stroke
the coccids and to swallow the honey-dew at its very source, and
that they have even acquired so keen an appetite for the liquid
that it has now become a very important if not an essential
constituent of their diet. The exploitation of such a constant
and energizing supply of syrup, moreover, would surely tend not
only to lengthen the original life-span of the adult beetles but
also to increase the number of their progeny and hence the size
and vigor of their colonies. This is suggested by the conditions
1921] Wheeler: Some Social Beetles 109
in certain ants, e. g., our yellow, hypogzie species of Lasius,
which are able to develop populous colonies mainly, if not exclu-
sively, on a diet of honey-dew derived from root-aphids and root-
coccids.
Inasmuch as the larve, from their very youngest stages,
no less than the adult beetles, continually stroke the coccids, the
question arises as to whether the habit was first acquired by the
larve, or by the beetles, or whether both instars developed it
simultaneously. An answer to this question might, perhaps, be
forthcoming if we could determine whether the larva or the
adult beetle shows the greater structural adaptation of the
antenne and mouthparts to dealing with the coccids. The larve
as I have shown, use both their antennze and maxille in the
process, the adults only the antenne, but although the beetles
are able to cover a greater area of the coccid’s surface with their
antennal clubs, the larger larve at least, by combining both pairs
of organs, can probably produce a stimulus no less intense and
effective. The larve are certainly more alert, restless and
inquisitive than the beetles and the mandibles in the youngest
stages seem to be very poorly adapted to feeding on the nutri-
tive parenchyma. It is therefore quite as probable that the habit
of stroking the coccids was first acquired by the young larve
and later continued in the adult as that it originally appeared in
the latter and was inherited in earlier and earlier ontogenetic
stages till it came to be manifested by the just-hatched larva
less than a millimeter in length. As shown by the observations
recorded on p. 70 such larve show an even greater avidity for _
the honey-dew than the adult beetles. Since the stroking of the
female beetle by the male during the courtship is precisely like
the stroking of the coccids, we might be tempted to conjecture
that the habit had arisen first in the adult as a modification of
the sexual appetite, but this is, perhaps, rather far-fetched.
38.—THE BUILDING OF THE COCOON.
The few data I have been able to gather concerning the
processes accompanying pupation in the beetles allied to Cocci-
dotrophus and Eunausibius are reproduced in Zoologica III, No. 5.
The pupating larvae of some Lemophloeids, Silvanids and Cucu-
110 Zoologica: N. Y. Zoological Society (iis
jids are described as attaching themselves by the last segment to
the substratum, and pushing back the larval skin to the tip of
the abdomen which remains fixed to the exuvium. In Oryze-
philus surinamensis the larva before pupating makes a rude
cocoon by agglutinating particles of food or detritus with an
oral secretion, but I have seen no circumstantial account of the
process which may be of considerable interest in connection with
the cocoon-building of Coccidotrophus. The Cucujus larva also
makes a rude cocoon (see p. 177).
The construction of a substantial cocoon by the larve of
the social Silvanids would seem to be necessary, because a nude
pupa, even if attached to the petiolar wall by its anal end, would
be exposed to injury by the numerous beetles and larve moving
about in the narrow cavity. But the way in which the cocoon is
constructed is, to say the least, very unusual. So far as known,
the larve of Coleoptera and other insects, when engaged in
making such structures, remain in situ and build the cocoon as
an envelope around the body, using for the purpose extraneous
particles of earth, detritus, wood or frass, or threads of silk
spun from the sericteries or more rarely a secretion of the
Malpighian tubules, as in ant-lions and certain weevils, as
described by Knab (1915a, 1915b) and others, or several secre-
tions as described by Boving for Donacia (1910). The Coccido-
trophus larva, however, laboriously collects minute particles of
living plant-tissue, mixes them with saliva and builds them up
in a very definite manner, repeatedly leaving the structure to
go afield for the purpose of collecting the necessary materials.
When the cocoon is all but completed the larva enters, and
becoming a voluntary prisoner, closes the aperture at the end with
materials scraped from the inner surfaces of the walls. I have
failed to find in the entomological literature any account of such
a method of cocoon-building, which in many particulars resembles
the nest-building of certain birds and rodents.
The only suggestion I can make in regard to the possible
origin of this behavior is that it may be derived in some way
from the beetle’s habit of building up its frass in more or less
regular ridges, or welts between the food-grooves or immediately
around the entrances to the petioles. I have been unable actually
1921] Wheeler: Some Social Beetles 111
to witness this performance as the beetles never exhibited it
after the petioles were cut open. I am inclined to believe that
the feces are not simply voided in the spaces between the food-
grooves but actually built up with the aid of the mouthparts.
This seems to be clearly indicated by the circular wall around
the entrances (Fig. 11). Perhaps the larve have quite as much
to do with the construction of the ridges as the beetles.
4.-CONCLUDING REMARKS ON THE BEHAVIOR OF THE
SOCIAL SILVANIDS
All modern observers of insects have been deeply impressed
by the highly mechanized character of their behavior, but it is
equally true that those who have most closely studied these
organisms both under natural and experimental conditions have
failed to find that the behavior of any one of them can be com-
pletely reduced to a rigid system of automatic or stereotyped
reactions. While the behavior of certain forms such as the
larval ant-lion, according to Doflein (1916) or the larval worm-
lion (Vermileo), as shown by my unpublished studies, seems to
consist almost entirely of a small number of reflexes, the behavior
of other insects, such as the solitary wasps, termites and social
Hymenoptera, often exhibits considerable plasticity, modifiability
or adaptability. Between these extremes we find the majority
of insects with a certain modicum of the latter type of behavior.
To this group we may assign the social Silvanids. The interpre-
tation of their various activities necessarily involves.some refer-
ence to the behavior of insects in general and the assumption
of a definite attitude towards certain intricate and much dis-
cussed questions. The limitations of space compel me, therefore,
either to leave the whole matter unconsidered or to treat it in a
very brief and sketchy manner. I prefer to adopt the latter
course.
The fashion which required one to explain as much as
possible of the behavior of an insect in terms of tropisms, or
taxes, and measured the value of one’s work by the success
achieved in the endeavor, seems to be rapidly passing. Thirty
years ago I followed the fashion with some enthusiasm, but
continued observation of the ways of insects has made me very
tM 3 Zoologica: N. Y. Zoological Society [rites
dubious in regard to the whole subject of the tropisms. My
present position concerning them is not essentially different
from that of Jennings (1904, 1906, 1909), von Buddenbrock
(1915), Claparede (1912, 1913), and others. I should, therefore,
interpret them as adaptive, secondarily developed reflexes and
not as unique, primitive elements in the genesis of instinctive
behavior.
There are, nevertheless, in the behavior of Coccidotrophus
certain phenomena, which some might be inclined to interpret
as tropisms, especially the reactions to contact, light and chemical
stimuli. The reader who has followed my account of the beetle
will have noticed the peculiar orientation of some of its stages
and of some of the associated insects with respect to the walls
of the long fusiform petiolar cavity which they inhabit. Thus
the eggs, cocoons and pupe of the beetle are always placed with
their long axes parallel with the long axis of the petiole, and
the coccids, while feeding, the cocoons of Blepyrus, which are
formed within the bodies of the coccids, and the nude pupez of
Scymnus assume the same orientation. The food grooves which
are excavated by the beetles and the frass-ridges which they
build, as well as the longer axis of the entrance are all longitu-
dinal. Moreover, the beetles spend much time lying in the food-
grooves with their narrow bodies longitudinally oriented and
with as much as possible of their surface in contact with the
fioor and walls of the grooves.
At first sight this striking series of orientations would seem
to be best described as tropistic, perhaps as due to some form ~
of thigmotropism, but it is evident that the only behavior which
might be legitimately regarded as such is that of the adult beetie
when resting or moving in the food grooves. The Collembolans
and the larve of Scymnus and Diadiplosis exhibit not the slightest
tendency to assume a similar orientation, and the Coccidotrophus
larve show no traces of it till they start their cocoons. Even
then, though they orient their cocoons, they assume a position
with their long axes parallel with the long axis of the petiole
only while actually adding particles to the walls and after they
have pupated. It is evident, furthermore, that nearly all the
orientations mentioned can be traced more or less directly to
peculiarities in the structure of the petiole, i.e., to the shape of
1921] Wheeler: Some Social Beetles 113
its cavity and the histological structure of its walls, and especially
to the singular arrangement of the nutritive parenchyma in long,
narrow strands. The beetles gnaw these out and thus form
grooves, which in turn orient the coccids and their internal para-
sites. The orientation of the frass ridges and beetle cocoons is
also determined by the grooves, and the position of the Coccido-
trophus eggs is very probably due to their being laid and attached
by the beetles while they are lying in the depressions between
the frass ridges. Thus the various orientations are merely so
many direct or indirect adaptations to the nutritive parenchyma
and the long narrow petiolar cavity. The latter clearly deter-
mines to some extent the longitudinal arrangement of the bulky
cocoons, just as a long Pullman car makes it advisable for us
to arrange the berths in a similar manner.
One orientation, that of the entrance, is not so easily
explained. In all the petioles I have examined, the long axis of
the entrance is parallel with the long axis of the petiole. It is
evident that it precisely fits the head of the beetle and that the
latter while gnawing it must stand on the outer surface of the
petiole at right angles to its long axis. When the surface is
longitudinally grooved, as is sometimes the case, the entrance can,
of course, have no other orientation, but often the surface is
quite smooth and it is difficult to see why it should be easier for
the insect to gnaw through the tissue lengthwise rather than
crosswise of the grain. It is also evident that while guarding
the entrance the beetle has its long axis at right angles to the
long axis of the petiole, but since a tubular wall is built around
the inside of the orifice (Fig.11) the insect’s reaction might be
regarded as thigmotropic, like its reactions to the walls of the
food grooves.
Nevertheless, I am convinced that the responses of the adult
beetle to contact stimuli may be more properly interpreted as
typical and highly adaptive reflexes. This is clearly indicated
by the structure of the insect. The long, parallel-sided, sub-
cylindrical form of the body and the shortness of the legs are
merely so many adaptations to living in narrow tubular cavities.
The same type of structure reappears as an independent develop-
ment in each of many different families of beetles, which live
in cylindrical cavities or burrows, e.g., the Scolytidz, Platypodi-
114 Zoologica: N. Y. Zoological Society (Tires
de, Brenthide, Bostrychide, Buprestide (Agrilus), Cleride,
Trogositide, Histeride (Teretrichus), Colydiide (Colydiwm),
Lyctide, Lucanide (Ceruchus), Elateride, Parandride, many
Cerambycide, Lymexylondide, etc. This type of body and one
more extremely flattened and adapted for living under bark
(Cucujus, Brontes), are very common among the Cucujide sens.
lat. The peculiar conformation of the front and of the mand-
ibles of Coccidotrophus and Hunausibius, so strikingly like that
of many ants (Cryptocerus, Cataulacus, Colobopsis, etc.) is, more-
over, a definite adaptation to guarding elliptical or circular en-
trances to solid-walled nesting cavities. It is interesting to note
also that the general shape of the body of Coccidotrophus and
Eunausibius reappears in several genera of ants which regularly
live in narrow plant cavities, e.g., Colobopsis, Simopone, Cylin-
dromyrmex, Metapone, Pachysima, Tetraponera and Pseudo-
myrma (see Plate III, figs. 1 and 2). Even the larve and pupe of
these ants have assumed a similar form. (See Wheeler, 1918, and
Wheeler and Bailey, 1920). We may say, therefore, that the
whole general bodily structure of the various insects I have
mentioned has been adaptively modified during their phyloge-
netic history and that such a modification can hardly be any-
thing but an expression of a concomitant adaptation of their
nervous system and reflexes.
Equally unsatisfactory from an ethological point of view
is the reference of other behavioristic peculiarities of the social
Silvanids to simple tropisms. Let us take as an example the
attraction to the Tachigalia. I agree with Picard (1919) in his
contention that phytophagous insects are attracted to their
respective host-plants by particular chemical substances in the
latter. Entomologists have always believed this but have usually
described the phenomena as due to “odor” or “taste.” To desig-
nate them as chemotropism really adds nothing to our knowledge
but a technical term. The ethological question remains: Why
is a particular insect species or sex attracted to a particular part
of a particular species of host plant, or, in the case of the social
Silvanids, why do they fly to and bore into the swellings of the
petioles of young individuals of a certain species of Tachigalia?
Undoubtedly the exquisite sense-organs in their antennal clubs
enable the beetles to detect certain very delicate effluvie emanat-
1921] Wheeler: Some Social Beetles 115
ing from the young Tachigalia as a whole or perhaps even from
the nutritive parenchyma in its petioles, but the attraction of
these odors is probably due to their having acquired a “meaning”
for the beetles, because the latter throughout their larval and
early imaginal stages fed on these very substances, and had, in
fact, long been familiar with the petiolar cavities, the coccids, etc.
The latter part of this statement also applies to the young queens
of the obligate Tachigalia ants of the genera Pseudomyrma and
Azteca and would account for the rather unusual attachment of
these insects to a definite host-tree. Hence in these cases organic
memory, or ““mneme,”’ or even individual memory yields a more
satisfactory explanation of the phenomena than a naked tropism.
A consideration of the responses to light leads to similar
results. The beetle colonies, as we have seen are “photophobic,”’
or live in the dark, and when the petioles are opened in the light
the insects are at first much agitated but soon settle down and
continue the regular routine of their existence even when the
pieces of petiole are kept exposed to artificial or diffuse day-light
in the laboratory. The adult beetles have well-developed eyes
and the larve have three pairs of small simple eyes on each side
of the head. (Plate VII, fig. 2, Plate VIII, fig. 10). And since a
certain amount of light enters the petiole through the entrance, at
least when it happens to be unguarded, it is probable that under
ordinary conditions the eyes are mainly useful in enabling the in-
sects and particularly the larve, to stay in the dark. There is noth-
ing to show that the young beetles, which leave the petioles to
establish new colonies, do so because they become positively
phototropic. The emigration may, perhaps, take place at night
and even if it occurs during the day the light in the parts of
the jungle where the young Tachigalias are growing, is very
subdued. I believe that the young beetles must emigrate either
because the space in the petiole has become too greatly reduced
by the growth of the colony or because the food-supply has for
the same reason become insufficient. In either case emigration
would be due to internal stimuli (‘‘physiological states’). These
may also be important factors in the swarming of ants, bees and
termites. We might, perhaps, even suppose that the guarding
of the entrance by the beetles is due to an abortive or inhibited
emigration impulse due to feebler or vaguer internal stimuli
116 Zoologica: N. Y. Zoological Society [III;3
comparable with those which sometimes impel a person who has
been sitting for hours in a dimly-lighted room, to stand at the
window or open door or to step out into the sunlight.
Thus while it is easy to interpret many of the activities of
the social Silvanids as adaptive reflexes it is difficult to assign to
such stimuli as light, contact, chemicals, etc., the leading role
which they have in the theories of Loeb and Bohn. Much of the
behavior of the beetles, such as their treatment of the coccids, the
building of the cocoon, mating, the guarding of the entrance,
etc., so obviously depends on internal or physiological states that,
in so far it has not become completely mechanized, we may more
properly regard it as made up of cyclical activities like those
recognized by Herrick (1910), Craig (1918) and others in birds.
Craig especially has shown how much of the behavior of birds
can be interpreted as cycles of appetence, or appetite and aver-
sion, which he defines as follows: ‘‘An appetite (or appetence,
if this term may be used with purely behavioristic meaning),
so far as externally observable, is a state of agitation which
continues so long as a certain stimulus, which may be called the
appeted stimulus, is absent. When the appeted stimulus is at
length received it stimulates a consummatory reaction, after
which the appetitive behavior ceases and is succeeded by a state
of relative rest. An aversion is a state of agitation which con-
tinues so long as a certain stimulus, referred to as the disturbing
stimulus, is present; but which ceases, being replaced by a state
of relative rest, when that stimulus has ceased to act on the
sense-organs.”
Rignano (1920) gives this same conception a more general
physiological formulation in the following passage: ‘Every
organism is a physiological system in a stationary condition and
tends to preserve this condition or to restore it as soon as it is
disturbed by any variation occurring within or without the
organism. This property constitutes the foundation and essence
of all “needs,” of all ‘‘desires,’’ of all the most important organic
“appetites.” All movements of approach or withdrawal, of
attack or flight, of taking or rejecting which animals make
are only so many direct or indirect consequences of this perfectly
general tendency of every stationary physiological condition to
remain constant. We shall soon see that this tendency in its
1921] Wheeler: Some Social Beetles 117
turn is only the direct result of the mnemic faculty characteristic
' of all living matter. This single physiological tendency of a
general kind, accordingly, is sufficient to give rise to a large num-
ber of the most diversified particular affective tendencies. Thus
every cause of disturbance will produce a corresponding ten-
dency to repulsion with special characteristics determined by
the kind of disturbance, by its strength, and by the measures
capable of avoiding the disturbing elements; and for every
incidental means of preserving or restoring the normal physio-
logical condition, there will be a quite definite corresponding
tendency such as “longing,” “desire,” “attraction” and so forth.
Even the instinct of self-preservation—when understood in the
usual narrow sense of “preservation of one’s own life’—is only
a particular derivative and direct consequence of this very gen-
eral tendency to preserve physiological invariability.” This ten-
dency, however, as Rignano remarks, is supplemented by another,
“for as soon as the previous stationary condition cannot be
restored by any means, that is by any movements or change ot
location, the organism disposes itself in a new stationary con-
dition consistent with its new external and internal environment.
In this way there originate a large number of new phenomena
called ‘adaptations’.”’
Of course, the contention that the appetites are fundamen-
tally important in animal behavior is not new. It is merely
astonishing that they have been so consistently ignored by many
modern observers. The role of appetency, or appetite, was set
forth with great acumen by the philosopher Fouillée, especially
in the third book of his “Evolutionnisme des Idées Forces’
(1920, first edition 1890’). The germ of the conception, however,
can be traced back to Reimarus (1798) and Leibnitz (Dwelshav-
ers, 1908, p. 181), to the “appetitus sensitivus” of the schoolmen
(see Maher, 1903, Chap. 10), and the doeEic and +6 doextixdv
of Aristotle (De Anim. 3, 10; Eth. Nic. 1, 18, 18). Recently the
behavioristic psychologists, psychopathologists and students of
the sympathetic nervous system and internal secretions, Mosso
(1899), Drever (1917), Smith and Guthrie (1921), Goddard
(1919), Kempf (1918), Crile (1915), Cannon (1915), and others,
have emphasized the importance of the appetites and others have
stressed their peculiarities in such terms as “‘tumescence” and
118 Zoologica: N. Y. Zoological Society [IlI;3
“detumescence” in sexual psychology and “enhancement,”
“relief” and “catharsis” in art. Kempf (1921) in his valuable °
contribution to psychoanalysis has used the term “craving” in-
stead of appetite, avoiding the “libido” of the Freudians which
embodies the same notion. In a recent volume Bertrand
Russell takes essentially the same view of the phenomena
of appetite but uses the word “desire.” Most of the authors
cited deal, of course, with man, but one can hardly over-
estimate the value of their work for the animal behaviorist and
entomologist. It is certain that insects have well-developed sym-
pathetic and glandular systems and that their alimentary and
sexual behavior presents quite as definite a picture of appetites
as does the corresponding behavior of the higher animals.
Fouillée believes that every appetition involves a rudimentary
cognition and that automatic behavior like that of the habits
and reflexes is merely lapsed appetition. If it could be shown
that the latter really can have this derivation and that such
ontogenetic mechanisms as habits can acquire representation in
the germ-plasm and hereditary transmission, we might be in a
position to give a consistent account of all animal behavior, and
one which would lead us to regard the reflexes and the tropisms
as ultimate, highly specialized end-stages instead of primitive,
elemental components of behavior.
“The Analysis of Mind,’ London and New York, Allen, Unwin &
Macmillan, 1921.
POSTSCRIPT.
Just as the final proof of this paper was being returned to
the printer Prof. I. W. Bailey received a letter from Col. David
Prain, Director of the Royal Botanical Gardens of Kew, with
the identification of the Tachigalia. It proves to be T. paniculata
Aublet. Col. Prain compared our specimens with Aublet’s type
in the British Museum Herbarium.
Among the various social beetles considered in the latter part
of my paper I should have included Phrenapates bennetti Kirby,
the habits of which were studied by Ohaus (1909) in Ecuador.
I shall have occasion to return to this insect in a future
publication.
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1921] Wheeler: Some Social Beetles 125
PIERCE, W. D.
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126 Zoologica: N. Y. Zoological Society [TII;3
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1920. The Feeding Habits of Pseudomyrmine and Other Ants. Trans.
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1872. Contributions Towards a Knowledge of the Life-histories of
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WIcKHAM. H. F.
1920. A Catalogue of the North American Coleoptera Described as
ew : Fossils. In Leng’s Catalogue (q. v.) pp. 349-365.
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PLATE III.
1. Eunausibius wheeleri Schwarz and Barber. X &.
Coceidotrophus socialis Schwarz and Barber. X 8.
Cross section of base of very young petiole of adult Tachigalia, showing pith
still in the cavity. Photograph by Prof. I. W. Bailey.
Cross section of base of large petiole of Tachigalia inhabited by Pseudomyrma
damnosa. The dark areas in the wall are nutritive parenchyma, which is not
disturbed by the ants but nourishes their coccids. A portion of one of the carton
partitions is shown on the left side. Photograph by Prof. I. W. Bailey.
Cross section through one of the strands of nutritive parenchyma showing the
cells with their homogeneous, amber-colored contents. Photograph by Prof.
I. W. Bailey.
Pig. 1.
Fig. 2.
Fig. 3.
PLATE IV.
Cross section of an as yet uninhabited petiolar swelling of a young Tachigalia,
showing the intact nutritive parenchyma, large pith cavity, and thin layer of
pith cells lining it. Photograph by Prof. I. W. Bailey.
Cross section of a petiolar swelling inhabited by Coccidotrophus socialis, show-
ing the gnawed nutritive parenchyma and the frass ridges. Photograph by
Prof. I. W. Bailey.
Section similar to that of Fig. 2 but showing three Coccidotrophus cocoons in
cross section. Photograph by Prof. I. W. Bailey.
PLATE V.
Halves of various Tachigalia petioles seen from the inside, showing the arrangements
of the cocoons, frass ridges, entrances, and in some cases also the beetles,
larve and coccids and a pupa (second figure from below).
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 4
(Tropical Research Station Contribution Number 98)
THE TACHIGALIA ANTS
By WILLIAM MORTON WHEELER
merPuBLISHED BY THE Sek.) ¥
eerie ZOOLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
Volume III, Number 4
THE TACHIGALIA ANTS
By WILLIAM MorTON WHEELER
The early botanists Aublet (1775) and Spruce (1869, 1908),
who first observed and collected Tachigalia in the forests of the
Guianas and adjacent regions mentioned ants as inhabiting the
swollen petioles but made no effort to have the species identified.
As we have seen, Aublet coined the name of the plant from
“tachi,” the general native-name for the stinging ants of the
genus Pseudomyrma. The white settlers of British Guiana call
them “long John ants,” but since the term “long John” is applied
to the trees of the genus Triplaris, the “palo santo” of the Latin
Americans, I have been unable to ascertain whether the name
of the tree is derived from that of the ant or vice versa, for the
tree is very tall and slender and the ants which inhabit its
cavities are long and narrow. “Tachi,” which is also used by
the natives of Brazil for various species of Pseudomyrma, would
seem to be the best name to introduce into the vernacular for all
the species of Pseudomyrma.
For more precise knowledge of the ant-fauna of Tachigalia
we are indebted to Ule (1907), who carefully collected the ants
of this and many other myrmecophytes and turned them over
to Forel to describe. In 1904 the eminent Swiss myrmecologist
published a comprehensive paper on Ule’s material, comprising
ants from various species of the following genera of myrmeco-
phytes: Tococa, Maieta, Pseudocatopsis, Triplaris, Sapium,
Cordia, Coussapoa, Duroia, Tachigalia, Platymischium, Pterocar-
pus, Pterocladon, Schwartzia and a peculiar Polypodium-like
fern. He described, however, only two ants from Tachigalia,
_Pseudomyrma latinoda Mayr subsp. tachigalix, from the petioles
and flower-bearing branches of Tachigalia formicarum Harms
(Plate I.), collected by Ule at Tarapoto, Amazonas, and Azteca
tachigaliz from the petioles of an undetermined Tachigalia from
Cerro de Escaler, in the mountains of Peru. Forel (1906) also
recorded the Ps. tachigalix from the upper Rio Purus, Amazonas
138 Zoologica: N. Y. Zoological Society [ALLIES 4!
(A. Goeldi and Huber). In 1912 he added Ps. latinoda var. endo-
phyta from the Rio Ariramba, near the Rio Trombetas, Ama-
zonas, basing the variety on specimens taken by A. Ducke in an
unidentified Tachigalia. Stitz in. 1912 records two additional
ants taken by Ule in Tachigatlia petioles: Ps. picta Stitz from Alto
Acre, Brazil, and Azteca brevicornis Mayr from Rio Branco,
Sierra de Maivasy, Brazil. I have been unable to find any other
records of Tachigalia ants in the literature. Strangely enough,
none of the five forms cited above occurs in the trees at Kartabo,
but instead, the following forms, 28 in number, half of them
belonging to new species, subspecies and varieties. I have no
doubt that I could have greatly increased the number of species,
had I been able to examine Tachigalias in other parts of British
Guiana, or even in the Bartica District, had I been able to pro-
jong my stay.
SUBFAMILY PONERINA.
1.—Ectatomma tuberculatum (Oliver).
This large, rather sluggish, yellow-brown ant, the well-
known “‘kelep,”’ which Dr. O. F. Cook several years ago attempted
to introduce into Texas from Guatemala for the purpose of prey-
ing on the cotton-boll weevil, was occasionally seen resting on the
terminal shoots of young Tachigalias along the Puruni trail at
Kartabo. It evidently feeds on the honey-dew of the Membracids,
but more frequently it visits other plants, and especially the con-
spicuous saucer-shaped nectaries on the petioles at the junction
of the leaflets of various species of Inga.
2.—Neoponera unidentata (Mayr).
A single dedlated female founding a colony in a Tachigala
petiole. This species nests normally in the hollow twigs of vari-
ous other trees.
3.—Neoponera crenata (Roger).
A single dedlated female founding a colony in a Tachigalia
petiole. This species nests in the hollow twigs of other trees.
lt is very common at Kartabo.
1921} Wheeler: Tachigalia Ants 139
SUBFAMILY PSEUDOMYRMIN A.
4.—Pseudomyrma damnosa sp. nov.
Ciice yale ac)
Worker. Length: 4.5-5 mm.
Head about one-quarter longer than broad, subrectangular,
slightly narrower in front than behind, with feebly and evenly
convex sides, and straight or very feebly convex pos-
terior border, somewhat flattened above, in profile about three-
fifths as high as long. Eyes rather small, only as long as their
distance from the anterior border of the head, feebly convex.
Mandibles stout, evenly convex, with two large apical and two
or three indistinct basal teeth. Clypeus small and very short,
convex but scarcely carinate in the middle, depressed on the sides,
its anterior border very feebly sinuate in the middle, more
strongly on each side. Frontal carine straight, subparallel,
closely approximated, extending back to a line joining the anter-
ior orbits. Antenne short and rather slender; scapes scarcely
incrassated apically, scarcely more than a third as long as the
head, their tips reaching only to the anterior third of the inner
orbits; first funicular joint longer than broad, constricted basally ;
joints 2-8 twice as broad as long, 9 and 10 somewhat longer in
proportion to their width; terminal joint as long as the two
preceding together. Thorax rather robust, shaped much as in
latinoda Mayr and arboris-sanctze Emery; pronotum with the
neck as long as broad, evenly rounded above, concave on the
sides but the dorsal surface not marginate laterally. Mesonotum
flattened, broader than long, its anterior border strongly arcuate,
its posterior border nearly straight. Mesoépinotal constriction
deep and abrupt. Epinotum longer than broad, a little broader
in front than behind, in profile with the base horizontal and on a
level with the pro- and mesonotum, distinctly longer than the slop-
ing declivity, into which it passes through an even curve. Petiole
from above subtriangular, about one and one-half times as long
as broad, with straight sides; in profile about half again as long
as high; the dorsal outline in profile concave anteriorly, the node
evenly rounded and convex above, abruptly truncated behind;
ventrally the surface is nearly straight, at its anterior end with
a strong, compressed, acuminate tooth, directed downward but
140 Zoologica: N. Y. Zoological Society [TiTs4
not backward. Postpetiole from above twice as broad as the
petiole, nearly twice as broad as long, very convex above, but
strongly narrowed and constricted anteriorly where it bears
on the ventral surface an acute downwardly directed projec-
tion, smaller than that on the petiole; middle of the ventral sur-
face strongly protuberant. Gaster elongate; sting long. Legs
rather slender.
Smooth and shining, especially the head, which is very
sparsely punctate; on the thorax the punctures are smaller and
somewhat denser, and the gaster is very finely and superficially
punctate and therefore appears somewhat less shining than the
head and pronotum. Mandibles at their tips very finely striate
and with coarse, elongate punctures; cheeks and pleure finely
and superficially punctate.
Hairs sparse, golden yellow, erect or suberect on all parts
of the body, rather short on the whole, longest on the petiole.
Pubescence fine, distinct on the gaster, postpetiole, petiole,
pleure, tarsi, tibiz, funiculi and gula, not concealing the surface
but rendering it somewhat less shining.
Brownish yellow; mandibles reddish brown, with black
teeth; ocellar region, sutures of thorax, petiole, postpetiole, a
patch on the center of each segment of the venter, the meso-
pleure, the femora, except their bases and tips and sometimes
portions of the tibizw, varying from pale to dark brown or black-
ish, clypeus and adjoining portions of head pale, clear yellow.
Female. Length: 7.5-9 mm.
Resembling the worker, but the head longer, with less con-
vex sides and somewhat larger and more convex eyes and ocelli.
Mandibles convex as in the worker, their external borders not
geniculate at the base. Thorax rather long, in the region of the
wing-insertions scarcely as broad as the head. Mesonotum rather
convex, as long as broad. Epinotum sloping, the base one and
one-half times as long as the declivity, which is even more abrupt
than in the worker. Petiole and postpetiole somewhat longer in
proportion to their width. Wings narrow and rather short,
only 6.5 mm. long.
1921] Wheeler: Tachigalia Ants 141
SC Caae.
FIG. 13. PSEUDOMYRMA DAMNOSA SP. NOV.
a, worker in lateral view; b, head of same from above; c, head of female from) above.
Head less shining, the fine punctures anteriorly and the
larger punctures on the vertex deeper and more conspicuous.
Gastric segments with large scattered punctures.
Hairs and pubescence much as in the worker but the pubes-
cence decidedly longer and more dilute, so that the gaster appears
more shining.
Color yellow, like the worker, but with more numerous and
more extensive spots. There is a brown spot on the middle of
the pronotum anteriorly, one on the mesonotum posteriorly, the
sides of the thorax have an irregular brown pattern, the petiole
is sometimes entirely brown, more often with the surface of
the node yellow; the ventral and anterodorsal portion of the
postpetiole, a broad band on each gastric segment above and a
large spot on each ventral sclerite, dark brown. All the femora
and the middle and hind tibiz, except their tips and bases, are also
dark brown. In some specimens the dorsal banding of the
gaster is more indistinct, in others the gaster may be described
as dark brown with yellow posterior borders to the segments, less
frequently the dark brown spot on the mesonotum becomes a
Y with its branches extending to the anterior border of the
sclerite. The scutellum and postscutellum are usually dark brown
throughout. Wings pale fuscous, iridescent, with dark brown
veins and pterostigma.
142 Zoologica: N. Y. Zoological Society [JI1; 4
Male. Length: 5.5-6 mm.
Head through the eyes as broad as long, narrowed behind,
with broadly concave posterior border; eyes and ocelli large;
cheeks very short; mandibles well developed, with two larger
apical and several indistinct basal teeth. Clypeus advanced and
convex in the middle, its anterior border deeply sinuate on each
side. Antenne short; scapes twice as long as broad; first funic-
ular joint as long as broad; joints 2-10 about twice as long as
broad, terminal joint longer. Thorax shaped much as in the
female, but the epinotum evenly rounded and sloping, without
distinct base and declivity. Petiole and postpetiole like those of
the worker but their ventral teeth are smaller. Gaster broad-
ened, or clavate at the tip. Legs slender.
Surface of body less shining and less distinctly punctate
than in the worker.
Pilosity and pubescence as in the female but less abundant.
Color of a duller brownish yellow; mandibles not darker;
posterior portion of head, large spots on pleure, a large spot at
the anterior end of the pronotum, lateral borders and a Y-shaped
spot on the mesonotum, the epinotum, except its disc, the petiole,
postpetiole and gastric segments, except the anterior and poster-
ior border of each segment, the femora and tibiz, except their
tips and bases and the anterior tibie, dark brown. Venter yellow-
ish, at least posteriorly; tips of genitalia infuscated. Wings col-
ored as in the female.
Described from numerous specimens taken at Kartabo and
the Penal Settlement, in the petioles of large specimens of
Tachigalia growing in the sun.
This species is very closely related to Ps. latinoda Mayr and
arboris-sanctz Emery. The worker differs from that of the
former, however, in its longer head, smaller and more flattened
eyes, more approximated frontal carine and smoother and more
shining surface. The var. nigrescens Mayr of latinoda ap-
proaches damnosa more closely, to judge from three workers
from Para received from Forel. The form described as var.
tachigalix Forel, of which I possess a couple of cotypes, is much
1921] Wheeler: Tachigalia Ants 143
more opaque, more brownish, with longer head and antenne,
larger eyes, more slender thorax, much more angular epinotum
and feebler pilosity. The var. endophyta Forel resembles dam-
nosa in coloration, but the description is very brief. In arboris-
sancte the body and eyes are larger, the antenne decidedly
longer, the petiolar node more strongly truncated behind, with
submarginate sides, the postpetiole is smaller and narrower, the
color uniformly brownish yellow. Ps. damnosa is also closely
related to dendroica Forel, but this form is larger, with a differ-
ent sculpture and color, a different petiole, etc. Stitz’s Ps. picta,
too, is an allied form but, to judge from his figure, has very
large eyes and a very different thorax, petiole and postpetiole.
It is certain, nevertheless, that all the forms mentioned—
damnosa, latinoda, arboris-sanctx, dendroica and picta—as well
as triplaridis Forel and the species maligna, described below,
constitute a peculiar group of very closely related tachis confined
to living myrmecophytes. One might regard the whole complex
as a single highly variable species, but in the present state of
our knowledge it is advisable not to indulge in too much “lump-
ing” in the genus Pseudomyrma.
5.—Pseudomyrma maligna sp. nov.
(Fig. 14 a-c)
Worker. Length: 4-4.6 mm.
Very similar to the preceding species but slightly smaller,
head somewhat broader and shorter, eyes distinctly smaller, less
elongate and more flattened; ocelli smaller and further apart;
mandibles more flattened with less curved external borders,
frontal carine less approximated, antennal scapes shorter and
more incrassated towards their tips. Clypeus sharply carinate,
depressed on the sides, its anterior border emarginate laterally,
produced in the middle as a short, transverse lobe, with feebly
concave margin and sharp corners. Thorax shorter and more
robust than in damnosa; pronotum broader, more convex on
the sides and above; mesoépinotal impression shorter but quite
as deep; epinotum decidedly shorter, the base from above sub-
hexagonal, broader behind than in front, and almost marginate
on the sides, more flattened, appearing straight in profile, the
declivity of the same length and forming with the base a more
144 Zoologica: N. Y. Zoological Society [(Tihs4
FIG. 14. PSEUDOMYRMA MALIGNA SP. NOV.
a, worker in lateral view; b, head of same from above; c, head of female.
distinct angle. Petiole shorter, only one and one-half times as
long as broad, the node more rounded and more abruptly trun-
cated behind, the spine at the anterior end of the ventral surface
longer, more curved and hook-like. Postpetiole more convex
above and at the sides, its anteroventral tooth less acute. Gaster
more slender; legs somewhat shorter.
Surface of body even smoother and more shining than in
damnosa; mandibles, head and thorax very smooth, with very
small, shallow, uniformly scattered punctures; pedicel and gaster
with even smaller and more indistinct punctures.
Hairs pale yellowish, short, uneven and similar to those of
damnosa but much less numerous. Pubescence shorter and more
dilute, so that it does not dim the shining surface of the gaster,
ionger and more distinct on the tibie and tarsi.
Deep castaneous; mandibles, clypeus, cheeks, antenne, pro-
notum usually, sides of epinotum, petiole and postpetiole, wholly
or dorsally, sides and bases of the gastric segments, legs, except
the middle portions of the femora of all the pairs and of the tibiz
of the middle and hind pairs, yellowish brown. In some speci-
mens the pronotum is also more or less castaneous above and
1921] Wheeler: Tachigalia Ants 145
in some the fore legs are brownish yellow throughout. There
is also considerable variation in the coloration of the petiole,
postpetiole and gaster. In some specimens the petiole is entirely
castaneous and only the node of the postpetiole is yellowish
brown, or the yellowish brown at the bases of the gastric seg-
ments may be much reduced. The whole epinotum may also be
castaneous in dark individuals.
Female. Length: 7.5-8.5 mm.
Head one and one fourth times as long as broad, subrectan-
gular, slightly narrower in front than behind, with rather
straight sides and feebly and broadly excised posterior border.
Eyes and ocelli larger and more convex than in the worker.
Mandibles short, with the external borders, abruptly geniculate
at the base, the upper surface flattened, their apical borders with
two sharp terminal and no basal teeth. Epinotum rounded, but
with distinct, subequal base and declivity. Petiole one and two-
third times as long as broad, the node somewhat depressed above,
its sides rather bluntly submarginate, its ventral tooth large;
postpetiole nearly as long as broad, less than twice as broad as
the petiole. Gaster elongate. Wings narrow and short (6 mm.).
Sculpture much as in the worker, but mandibles striatopunc-
tate and anterior portion of head more opaque and more densely
punctate.
Pilosity and pubescence as in the worker.
Black; mandibles castaneous red; antenne brownish yellow;
the scapes sometimes slightly infuscated in the middle; clypeus,
gula, pronotum and sutures of the thorax more or less castane-
ous; lateral borders of the ventral and dorsal sclerites of the
gaster, tarsi and tibie brownish yellow; the middle and hind tibie
infuscated in the middle of their extensor surfaces ; femora black-
ish, with brownish tips and bases. Wings grayish hyaline, iri-
descent, not infuscated, with very distinct blackish veins and
pterostigma.
Male. Length: 7.4-7.6 mm.
Head through the eyes as long as broad, eyes and ocelli
larger than in damnosa; antennal scapes and first funicular
joint, decidedly longer. Clypeus strongly carinate, slightly pro-
146. Zoologica: N. Y. Zoological Society [1it=4
jecting in the middle, its sides impressed, their anterior borders
bisinuate. Head behind the eyes narrowed but more convex
and rounded than in damnosa. Epinotum somewhat longer and
slightly more angular in profile, the ventral spine of the petiole
much larger, nearly as large as in the female, the terminal dorsal
segment of the thorax of a different shape, its posterior border
produced as a narrow blunt point in the middle.
Shining, like the worker; even the mandibles and head
very smooth and shining, the former striolate at the tips, punc-
tate at the base, the head more densely punctate anteriorly as in
the female.
Pilosity and pubescence much as in the worker.
Black; mandibles testaceous, infuscated at the base; antennz
tuscous, first funicular joint and tips and bases of scapes yellow-
ish. Thoracic and abdominal sutures brown. Legs yellowish
or pale brown, the femora, except their tips and bases, dark
brown, the extensor surfaces of the hind tibiz infuscated. Wings
as in the female.
Described from many specimens from Kartabo, nesting, like
the preceding species, in the petioles of large Tachigalias. Though
closely related to damnosa, arboris-sancte, etc., this species seems
to me to be specifically distinct. It also resembles triplaridis
Forel and another species, which is found in Triplaris surina-
mensis near Kartabo and will be described elsewhere. Among
my material there are also two varieties of maligna found nesting
in some of the smaller Tachigalias; namely:
6.—Pseudomyrma maligna var. cholerica var. nov.
Worker. Length: 3-3.5 mm.
Distinctly smaller than the typical form and similarly col-
ored, but the lighter parts are more extensive and often of a more
washed-out, sordid yellow, whereas the darker parts are more
piceous or blackish. The clypeus, cheeks, petiole and postpetiole
are pale yellow, as are also the thorax, except the base of the
epinotum and a fuscous cincture in the mesonotal region. The
gula and first gastric segment are apt to be paler than the
1921] Wheeler: Tachigalia Ants 147
upper surface of the head. Except in size, this variety is not
very sharply marked off from the type.
7.—Pseudomyrma maligna var. crucians var. nov.
Worker. Length: 3-3.5 mm.
Of the same size as the preceding variety but with a decided
tendency to melanism, the whole thorax, pedicel and gaster being
very dark brown or black in mature specimens. The middle of
the scapes is infuscated and there may be occasionally pale
markings on the node of the postpetiole and at the bases and
sides of the gastric segments.
SUBFAMILY MYRMICIN 2.
8.—Pheidole cramptoni Wheeler subsp. petiolicola subsp. nov.
Soldier. Length: 2.3-2.6 mm.
Decidedly smaller than the type, which measures 4 mm.,
and differing as follows: head with the anterior corners even
more decidedly turned outward, anterior gular teeth more slender
and more acute; eyes slightly larger and distinctly convex (flat
in the type) ; antenne shorter, the tips of the scapes reaching
the lateral borders of the head two-fifths of their length from the
anterior corners (nearly half their length in the type). Thorax
shorter and proportionally stouter; epinotal spines slightly
shorter; postpetiolar node hemispherical and evenly convex
above, without distinct anterior angles. Interrugal spaces on the
anterior portion of the head less distinctly reticulate, so that the
rugz seem to be sharper. Surface of the body smoother and
more shining. Pilosity quite as well developed as in the type.
Color more vivid but the dark and light areas distributed in the
same manner. Mandibles and clypeus red, with dark brown
borders; anterior two-thirds of head clear ivory yellow, with a
large elongate castaneous spot between the frontal carinz; pos-
terior two-thirds of head with a band extending forward on
each side to the eye; the thorax, petiole, postpetiole and gaster
rich castaneous, the first gastric segment, femora and pleure
somewhat paler; frontal carine, tibie and antenne reddish; tarsi
yellowish.
148 Zoologica: N. Y. Zoological Society [Tia
Worker. Length: 1.4-1.6 mm.
Slightly smaller than the worker of the typical form, some-
what darker brown, with distinctly shorter antenne, the scapes
reaching back beyond the border of the head only to a distance
equal to their greatest diameter. Head shorter, with more dis-
tinct posterior corners. Epinotal spines somewhat smaller.
Described from several specimens found with brood in a
young Tachigalia petiole near Kartabo. The types of the species
were collected by Dr. F. A. Lutz at Kaieteur, British Guiana.
9.—Pheidole tachigaliae sp. nov.
Soldier. Length: about 2 mm.
Head large, subrectangular, about one-quarter longer than
broad, as broad in front as behind, with rather straight, sub-
parallel sides and deeply, angularly excised posterior border,
vertex with a large and rather deep impression, without a median
groove between it and the frontal area where the head is uni-
formly convex. Gular teeth small and blunt. Eyes small and
moderately convex, only about their own length distant from the
clypeus. Frontal carine long, very far apart, curved, gradually
diverging behind towards the posterior corners of the head,
forming the inner borders of broad, shallow, but distinct scrobes
for the antennz. These scrobes and the frontal carine reach
nearly to the posterior corners of the head. Mandibles stout and
very convex, with a pair of blunt, apical teeth. Clypeus. flat,
ecarinate in the middle, its anterior border somewhat retuse,
feebly sinuous in the middle and on each side. Frontal area
rather large, triangular, shallow, with a small median pit.
Antenne slender, scapes curved, not incrassated or flattened but
slightly enlarged distally, reaching nearly to the middle of the
lateral borders of the head; antennal club somewhat shorter
than the remainder of the funiculus, its two basal joints sub-
equal, broader than long, together equal to the terminal joint;
first funicular joint large; joints 2-6 small and transverse.
Thorax very short, broad through the humeri which are promi-
nent and subangular; promesonotal suture distinct but not im-
pressed. In profile the pro- and mesonotum form together a
high hemispherical mass, the mesonotum descending abruptly
1921] Wheeler: Tachigalia Ants 149
behind to the pronounced mesoépinotal constriction. Epinotum
small and low, subcubvidal, broader than long, in profile with
subequal base and declivity, the former straight and horizontal
in profile, the latter very steep and concave, the spines very
small and acute, scarcely longer than broad at their bases,
directed backward and very slightly upward. Petiole slender,
pedunculate, the node at its posterior end abrupt, anteroposter-
iorly compressed, its border moderately sharp, transverse and
very feebly sinuate in the middle. Postpetiole slightly broader
than the petiolar node, transverse, with rounded sides. Gaster
oval, about as large as the head, convex above and below. Legs
moderately long, with thickened femora and tibize.
Shining ; mandibles and middle of clypeus smooth, the former
with coarse, scattered, piligerous punctures; antennal scrobes
reticulate; remainder of head longitudinally rugose, the ruge
rather straight and subparallel extending to the posterior cor-
ners, the interrugal spaces and posterior part of the head
rather loosely reticulately rugose. Pro- and mesonotum smooth
and shining above; pleure and epinotum more opaque, finely
reticulate or punctate, as are also the ventral portions of the
petiole and postpetiole; the nodes of the latter, the gaster and
legs smooth and shining.
Hairs pale, golden yellow, erect, of uneven length, moderately
abundant, rather uniformly covering the dorsal surface of the
body. Pubescence almost absent.
Reddish yellow; gaster and appendages paler yellow, the
former with a brown transverse band or cloud on the posterior
portion of each segment. Mandibles red, with black borders;
clypeal border also blackish. Impression on the vertex with a .
small, indistinct brown spot.
Worker. Length: 1-1.1 mm.
Head subrectangular, as long as broad, its sides rather
rounded, its posterior border feebly but distinctly emarginate in
the middle. Eyes rather large, flat, in front of the middle of the
head. Mandibles with oblique, indistinctly denticulate apical
borders. Frontal area obsolete. Clypeus rather convex in the
150 Zoologica: N. Y. Zoological Society [EF
middle, the anterior border transverse, entire. Antennal scapes
extending only very slightly beyond the posterior corners of
the head. Thorax shaped somewhat as in the soldier, but the
upper surface of the pro- and mesonotum more flattened. Epino-
tum longer than broad, with very minute, acute teeth. Petiole,
postpetiole, gaster and legs resembling those of the soldier.
Mandibles smooth and shining, sparsely and indistinctly
punctate. Clypeus, head, thorax, petiole and postpetiole sub-—
opaque, finely, densely and uniformly punctate; postpetiolar node
slightly shining, gaster and legs very smooth and shining.
Pilosity like that of the soldier.
Brownish yellow; mandibles, appendages and gaster paler,
yellow; the gaster without dark fascie.
Female (deadlated). Length: 2.5 mm.
Head subrectangular, a little longer than broad, a little nar-
rower in front than behind, with straight sides and posterior
border. Frontal carine and antennal scrobes as in the soldier.
Antenne longer, the scapes reaching to the posterior third of the
jateral borders of the head. Upper surface of head convex,
without vertical impression. Ocelli rather far apart. Thorax
elliptical, as broad as the head, the mesonotum and scutellum
flat above.. Epinotum very small and sloping, with small teeth.
’ Petiole and postpetiole much as in the soldier; gaster larger and
more elongate.
Sculpture, pilosity and color as in the soldier. There is a
black spot on the ocellar triangle and the brown fasciz on the
gaster are broader and darker. Wing insertions blackish.
Described from specimens from a single colony with brood,
taken August 23 at Kartabo from a petiole of a small Tachigalia.
This species is very peculiar on account of its small size
and the antennal carine of the soldier and worker. In the latter
particular it seems to approach Ph. scrobifera Emery of Costa
Rica, but this species is decidedly larger, with much longer
epinotal spines, quite different sculpture, etc.
1921] Wheeler: Tachigalia Ants 151
10.—Crematogaster (Orthocrema) limata F. Smith
var. palans Forel.
A single colony with two dedlated females, of what I take to
be this variety, was found August 6, nesting in a small Tachigalia
petiole at Kartabo. The workers are smaller than those of the
typical limata, possibly because the colony was young. The anten-
nal scapes are longer than in the type from Panama, but Forel
mentions specimens from Para Brazil, with longer antenne.
The female measures nearly 6 mm. and is very deep brown,
almost black, with the mandibles, legs and antennal clubs yellow-
ish brown, the femora and tibize somewhat infuscated in the
middle. Head subrectangular, broader than long, nearly as broad
in front as behind, with straight, subparallel cheeks as long as
the eyes. Antennal scapes extending a short distance beyond
the posterior corners of the head. Thorax a little more than
twice as long as broad; slightly narrower than the head, elongate
elliptical from above, the nearly vertical epinotum with a pair of
slender, pointed spines, longer than broad at their bases. Petiole
subelliptical, truncated behind, its posterior corners distinctly
angular; in profile twice as long as high, with nearly flat dorsal
surface and a small tooth at the anteroventral margin. Post-
petiole as broad as the petiole, somewhat depressed above, trun-
eated behind. Gaster rather large, convex on the sides and
above, pointed behind and emarginate anteriorly. Legs moder-
ately slender. Surface of body very smooth and shining. Mand-
ibles smooth, with a few small, scattered, piligerous punctures.
Cheeks, sides of front, sides and declivity of epinotum longitud-
inally striate; mesonotum -with coarse, scattered, piligerous
punctures, the punctures on the gaster quite as sparse, but
smaller. Hairs golden yellow, moderately abundant, erect, of un-
even length, most numerous on the head, well-developed on the
scapes and legs.
12.—-Crematogaster (Orthocrema) limata F. Smith subsp
ludio Forel.
Two colonies with deilated females, taken August 10 and
September 3 from petioles of young Tachigalias on the Cuyuni
trail at Kartabo. The workers agree well with Forel’s descrip-
152 Zoologica: N. Y. Zoological Society [III; 4
tion and with specimens taken elsewhere in British Guiana by
F. A. Lutz, F. M. Gaige and myself.
The female closely resembles that of the var. palans, but the
head is longer, fully as long as broad and more narrowed ante-
riorly; the epinotal spines are shorter and stouter, not longer
than broad at their bases, the posterior angles of the petiole are
less acute, the mandibles are darker, the legs paler and more
uniformly brownish yellow, the hairs on the body are whitish.
12.—Crematogaster (Orthocrema) limata F. Smith subsp.
parabiotica Forel.
This subspecies was not found nesting in the petioles but
attending Membracids on the shoots of young Tachigalias. As
I have shown in another paper (1921) it commonly lives in
parabiosis with Camponotus femoratus (vide infra p. 167) in
the peculiar “ant-gardens” attached to the trunks and branches
of trees in the moist jungle. Both species of ants forage together
and when found on Tachigalia the workers intermingle on the
shoots and are very friendly to one another and to the Membracids
but very pugnacious when interfered with.
13.—Crematogaster (Orthocrema) delitescens sp. nov.
Worker. Length 2-2.1 mm.
Head nearly circular, convex and rounded above, without
distinct posterior corners; eyes moderately convex, situated just
behind the middle of the sides, distant nearly twice their length
from the anterior border of the head. Mandibles narrow, convex,
their short apical borders with four small teeth. Clypeus convex
in the middle, its anterior border slightly reflected, broadly and
evenly rounded. Frontal carine short, subparallel. Antennz
slender ; scapes extending a little beyond the posterior border of
the head; funicular joints 2-8 as long as broad, ninth joint longer
than broad; club slender, decidedly shorter than the remainder
of the funiculus, its basal about two-thirds as long as its terminal
joint. Thorax short; pro- and mesonotum together subtriangular
from above, as broad as long; the pronotum flattened, marginate
on the sides, the mesonotum very short and narrow, declivous,
1921] Wheeler: Tachigalia Ants 153
strongly carinate on each side. Promesonotal suture feeble;
mesoépinotal suture distinct, but very short. Epinotum very
short, sloping between the spines, which are slender, straight and
acute, a little longer than the sides of the base, much shorter than
their distance apart and directed backward and slightly upward.
Petiole oblong, a little longer than broad, with parallel sides,
constricted at the anterior end, the posterior corners projecting
as minute tubercles. In profile it is about one and one-half times
as long as broad, with very feebly convex dorsal and ventral
surfaces and without a distinct anteroventral tooth. Postpetiole
small, broader than long, the node entire, anteroposteriorly com-
pressed. Gaster elongate triangular, tapering and pointed
posteriorly, concave dorsally, convex ventrally, the anterior
border of the first segment straight and transverse.
Shining: mandibles smooth, with a few small, scatterea
punctures. Head very smooth and shining, cheeks indistinctly
striolate-reticulate. Clypeus with two sharp, longitudinal ruge
on each side, smooth and shining in the middle. Pronotum with
a few longitudinal ruge on the disc, abbreviated behind; meso-
notum and epinotum subopaque, regularly and evenly reticulate,
except the most posterior portion of the latter, which is smooth
in the middle; pronotum on the sides, petiole and gaster smooth
and shining; postpetiole with short longitudinal furrows above.
Hairs whitish, long, sparse, rather blunt but not clavate
on the dorsal surface of the body; anterior surfaces of antennal
scapes with a row of long, rather flexuous hairs. On the legs
the hairs are very short, appressed and rather sparse.
Piceous brown; head and gaster darker; base of scape and
of funiculus paler; mandibles and tarsi brownish yellow.
Female (dedlated). Length 4.7 mm.
Head subrectangular, nearly as broad as long, and nearly
as broad in front as behind, with rounded posterior corners and
straight, parallel cheeks. Eyes at the middle of the sides,
elongate. Antennal scapes scarcely reaching to the posterior
corners of the head. Thorax from above elongate elliptical,
decidedly narrower than the head; mesonotum one and one-third
times as long as broad; epinotum abrupt, without base or
154 Zoologica: N. Y. Zoological Society [III ; 4
declivity, the spines of the worker represented by blunt angles
on the sides. Gaster elongate, with subparallel sides, the first
segment subrectangular, nearly as long as broad.
Smooth and shining, the epinotum between the angles reti-
culate, the node of the postpetiole smooth.
Hairs decidedly shorter than in the worker, more abundant
on the head and thorax than on the abdomen.
Color much like that of the worker, but the thorax as dark
as the head and gaster, except the anterior portion of the
pronotum, the prescutellum and epinotum which are more
castaneous.
Described from a female and several workers constituting a
single small colony found July 24 nesting in the petiole of a
young Tachigalia growing on the Cuyuni trail, near Kartabo.
This form approaches C. sumichrasti Mayr in many particu-
lars but is, I believe, quite distinct, especially in the shape of
the head, mesonotum and petiole, in the pilosity of the legs, color,
etc. It also recalls forms like limata Smith, brasiliensis Mayr
and lutzi Forel. The antennal club might be regarded as indis-
tinctly three-jointed. The female is very small compared with
the corresponding phase of the limata group. Mayr’s laevis is
also a closely allied form, but the petiole and median funicular
joints of this species are shorter, the pro- and epinotum smoother,
the color is paler, etc. I have three workers taken by Mr. F. M.
Gaige at Castries, St. Lucia, W. I., which agree very closely with
the types of delitescens.
14.—Solenopsis altinodis Forel
(Fig. 15, a-c).
Worker. Length 1.8-2.1 mm.
Head about one and one-sixth times as long as broad, sub-
rectangular, as broad in front as behind, with straight posterior
and very feebly convex sides. Eyes consisting of about a dozen
ommatidia, well in front of the median transverse diameter of
the head, and about four times their diameter from the anterior
corners. Mandibles with evenly convex external borders, the
1921} Wheeler: Tachigalia Ants 155
FIG. 15. SOLENOPSIS ALTINODIS FOREL, WORKER.
a, lateral view of body; b, head from above; c, petiole and post petiole from above.
apical borders oblique, with four distinct teeth. Clypeus project-
ing, with the usual pair of sharp carine. These do not end at
the border in sharp teeth, but in small, rather blunt projections.
Lateral denticles lacking. Frontal area distinct, impressed.
Antennal scapes reaching nearly to the posterior fourth of the
head. Funiculi about as long as the two-jointed club; joints 2-7
broader than long, 2-6 being fully twice as broad as long; basal
joint of club a little longer than broad, somewhat less than a
third as long as the terminal joint. Thorax slender; pronotum
with subrectangular humeri and straight subparallel sides, in
profile evenly and feebly convex above. Mesoépinotal constric-
tion short but pronounced; epinotum long, its base feebly convex,
indistinctly bidenticulate behind and nearly twice as long as the
156 Zoologica: N. Y. Zoological Society [1It;4
declivity which is abrupt and distinctly marginate on the sides.
Petiole in profile very large, with distinct peduncle and longer
abrupt node, which is much higher than the epinotum, higher
than long, in profile subrectangular, with vertical anterior and
posterior borders and subhorizontal superior outline rounded at
each end. The ventral surface is convex behind and concave in
front, with an angular projection but no distinct tooth near the
anterior end. Seen from above the petiole is twice as long as
broad, gradually enlarged behind, so that the node is seen to be
much compressed laterally. Postpetiole in profile much higher
than long, the summit of its node rounded, not attaining the
height of the petiole. From above the postpetiole is somewhat
broader than the petiole and slightly broader than long. Gaster
large, the anterior border of the first segment somewhat concave.
Sting long and powerful. Legs moderately long and slender.
Smooth and shining, with minute, scattered, indistinct,
piligerous punctures; cheeks and sides of front finely striate.
Hairs pale yellowish, bristly, of uneven length, moderately
abundant, erect or suberect, rather long on the body, shorter on
the antenne and legs.
Dull reddish brown; head and especially the gaster dark
brown; mandibles, clypeus, cheeks, antennze and legs yellow;
middle portions of scapes, femora and tibi# more or less
infuscated.
I have redescribed this species, originally described from
Venezuela and Trinidad, from many specimens taken at Kartabo.
It does not live in the petioles of Tachigalia but in the ground
and often enters those inhabited by the Coccidotrophus and
-Hunausibius and destroys their colonies (vide supra p. 82). I
was never able to secure the sexual forms and brood. The
worker is readily distinguished from the numerous other South
American species of Solenopsis by the peculiar, high, compressed
petiolar node, marginate and subbidentate epinotum, nearly
unarmed clypeus and unusual coloration.
1921] Wheeler: Tachigalia Ants 157
15.—Solenopsis helena Emery subsp. hermione subsp. nov.
Worker. Length 1.6-1.8 mm.
Differing from the typical helena of Chile in its somewhat
greater size (helena measures 1.3-1.5 mm.), in having the head
slightly shorter, with more rounded sides, joints 2-7 of the
funiculus very slightly longer as compared with their width, so
that the club is shorter in proportion to the remainder of the
funiculus. The epinotum is evenly convex, without indications
of an angle between the base and declivity, and the summit of
the petiolar node is not so thick in profile. The eyes, which are
not figured or described by Emery for the type, are very smal}
and consist of only four ommatidia.
Female. Length 3.3 mm.
Differing from the typical helena in color, being yellow,
like the worker, instead of pale brown, with the ocellar triangle,
a broad dorsal band and a large ventral spot on each gastric
segment dark brown. Each of these spots and bands is emargi-
nate posteriorly. In old, dedlated females the wing-insertions are
also dark brown. Wings gray, rather opaque, with distinct brown
veins and dark brown pterostigma.
Described from numerous specimens taken from several
colonies found in the petioles of young Tachigalias along the
Cuyuni trail at Kartabo. It nests also in the hollow twigs of
various other plants.
16.—Solenopsis helena subsp. ultrix subsp. nov.
Worker. Length 1.6-1.8 mm.
Of the same size as the preceding subspecies but differing
in having the head still broader in proportion to its length and
with more rounded sides and posterior corners, the posterior
margin straight or even slightly convex. The antennal scapes
are distinctly longer, attaining the posterior fourth of the dis-
tance between the eyes and posterior corners of the head. Color
pale piceous brown; tarsi and funiculi, except the clubs, pale
whitish yellow; gaster dark brown, with the anterior and
posterior borders of the segments yellowish.
158 Zoologica: N. Y. Zoological Society [III; 4
Female (dedlated). Length nearly 4 mm.
Distinctly larger than the female of hermione and of a
different color, being castaneous brown; mandibles and clypeus
pale reddish brown; antennze and legs pale yellow, with the
antennal clubs and scapes, femora and tibiz, except their bases
and tips, infuscated. Piligerous punctures on the head more
distinct than in hermione. Epinotum in profile scarcely angular,
but rounded as in that form and the typical helena.
Described from numerous workers and a single female
taken from two colonies inhabiting the petioles of young Tachi-
galias at Kartabo. I have attached this form to helena though
the shape of the head and the length of the antennal scapes are
different. The other characters, however, such as the shape of
the thorax and petiole, the sculpture and pilosity are so similar
that it can hardly be regarded as a distinct species.
17.—Leptothorax (Goniothorax) echinatinodis Forel subsp.
aculeatinodis Emery var. pleuriticus var. nov.
Worker. Length 2.2-2.5 mm.
Differing from the typical aculeatinodis in the following
particulars: Head very dark brown or piceous, like the gaster;
whole upper surface of the thorax ivory or brownish yellow, only
the mesopleure and sides of the epinotum dark brown. Petiole
pale, like. the thoracic dorsum; postpetiole brown, paler than the
gaster; scapes, base of funiculi, tarsi, tibiz and bases and tips
of femora pale ivory yellow, remainder of femora dark brown,
as are also the clubs of the antenne. Pilosity abundant, long,
erect, present also on the scapes and legs, the hairs being obtuse
but neither stout nor clavate. Ruge of the petiole somewhat
concentric. Tubercles on the petiole and postpetiole small and
blunt. Epinotal spines long, slender, curved downward and
directed upward, backward and outward, slightly shorter than
the base of the epinotum.
Female. Length about 3 mm.
Head smooth and shining only in the middle between the
frontal carine and imaginary lines continuing them to the
posterior border of the head; lateral to these lines the surface
1921] Wheeler: Tachigalia Ants 159
is coarsely, loosely and longitudinally rugose, with reticulate
interrugal spaces. Thorax subopaque, its sides obscurely longi-
tudinally rugose; mesonotum and scutellum longitudinally and
‘rather regularly rugulose, the former somewhat smooth
anteriorly. Epinotum punctate-rugulose, the declivity regularly
traversely rugulose. Pilosity as in the worker. The spines of
the epinotum are reduced to two stout teeth, as long as their width
at the base. Petiole and postpetiole nearly smooth, except for
their tubercles, which are decidedly smaller than in the worker.
Gaster elongate elliptical, transversely truncated anteriorly.
Brownish yellow; upper surface of head, except its posterior
corners, antennal clubs, scutellum, postscutellum, sometimes a
posteromedian spot on the mesonotum, femora, except their bases
and gaster, except the anterior corners of the first segment,
dark brown. Wings whitish hyaline, with very pale veins and
distinct brown pterostigma.
Male. Length 2.2 mm.
Head distinctly longer than broad, with distinct posterior
corners, the posterior orbits very near the median transverse
diameter. Ocelli small. Cheeks very short. Clypeus with
arcuate, projecting anterior border. Mandibles small, with
oblique 3- or 4-toothed apical borders. Antenne slender, 12-
jointed; scapes reaching nearly to the posterior corners of the
head; first funicular joint longer than broad, not broader than
the remaining joints, which are somewhat fusiform; fifth joint
long, representing two incompletely separated joints. Thorax
long and slender, humeri distinctly angular; mesonotum with
Mayrian furrows; epinotum low and sloping, moderately convex,
without distinct base and declivity. Petiole smooth and slender,
the node represented by a swelling in the middle, the sides with
feeble traces of the lateral tubercles of the worker and female;
postpetiole smooth, without tubercles, nearly as long as broad,
broader behind than in front and distinctly broader than the
petiole. Gaster and legs slender.
Head and thorax finely and densely punctate; vertex and
mesonotum smooth and shining, as are also the pedicel, gaster
and appendages.
160 Zoologica: N. Y. Zoological Society [III ; 4
Hairs whitish, short, less abundant and more delicate than
in the worker and female; the longer hairs on the femora and
tibiz reclinate.
Piceous brown; head darker and more blackish; mandibles,
antenne, legs, genitalia and incisures of gaster pale brownish
yellow. Wings as in the female.
Described from specimens belonging to a single colony
inhabiting the petiole of a young Tachigalia at Kartabo (Sept. ae
18.—Leptothorax (Goniothorax) umbratilis sp. nov.
Worker. Length 1.7-2 mm.
Head subrectangular, very slightly longer than broad and
slightly broader behind than in front, with nearly straight
posterior border and very feebly convex sides; the eyes moderately
convex, their posterior orbits at the median transverse diameter
of the head. Mandibles convex, their apical borders with five or
six small teeth. Clypeus distinctly concave in the middle, without
carine, the anterior border entire and transverse in the middle,
large but rounded and lobular. Mesoépinotal impression distinct
but shallow. Frontal groove absent. Antenne 11-jointed; scapes
reaching to within a distance equal to their greatest diameter
of the posterior corners of the head; second funicular joint as
iong as broad, joints 3-7 distinctly broader than long; two basal
joints of club subequal, longer than broad, together shorter than
the enlarged terminal joint. Thorax shaped much as in
echinatinodis, flattened above, marginate on the sides, with
bluntly acuminate anterior humeral angles; lateral mesonotal
angles rather acute, those at the base of the epinotum quite as
large but rounded and lobular. Mesoépinotal impression distinct
but shallow. Epinotal spines short, stout and curved, not longer
than their distance apart at the base, directed upward, backward
and outward, much shorter than the base of the epinotum.
Petiole about one and one-half times as long as broad, slightly
broader behind than in front, with tubercles arranged as in
echinatinodis but much smaller. In profile the petiole is longer
than high, the node rather low, the peduncle short and indistinct;
the ventral surface concave, with a small, downwardly directed
tooth at the anterior end. Postpetiole broader than long, broader
1921] Wheeler: Tachigalia Ants 161
than the petiole, high and convex above; its sides with two pairs
of small tubercles, the anterior pair scarcely visible. Gaster and
legs as in echinatinodis.
Mandibles shining, very finely striatopunctate. Clypeus
nearly smooth in the middle, shining, longitudinally rugulose on
the sides. Head, thorax and pedicel subopaque, the head densely
and finely reticulate-rugulose, the rugules longitudinal; thorax
with similar sculpture, the rugules on the pronotum somewhat
more pronounced. Postpetiole with a few longitudinal ruge.
Gaster shining, the basal half of the first segment very finely
shagreened and punctate, so that it appears slightly opaque or
sublucid.
Hairs white, abundant, short, clavate and erect on the dorsal
surface of the body. Such hairs are lacking, however, on the
appendages where they are replaced by short, appressed
pubesence.
Brown; upper surface of head, mesonotum, mesopleure,
epinotum, nodes of pedicel and antennal clubs dark brown; man-
dibles, clypeus, scapes and tarsi yellow; pronotum and gaster
brownish yellow.
Female (deadlated). Length about 3 mm.
Resembling the worker. Pronotum more rectangular, its
anterior border very straight and sharp, the corners acute.
Mesonotum and scutellum flat; base of epinotum convex, bluntly
angled on each side anteriorly, declivity longer, abrupt and
concave, the spines of the worker represented by small, blunt
angles. Petiole and postpetiole similar to those of the worker,
but the tubercles on the sides of the latter longer and more acute.
Gaster longer.
Sculpture much as in the worker; mesonotum as finely and
regularly rugulose-punctate as the head; scutellum densely punc-
tate; base of epinotum transversely rugulose-punctate. Gaster
very finely striolate-punctate at base of first segment.
Pilosity like that of the worker.
Yellowish brown; mandibles, clypeus, tarsi and gaster yellow;
ocellar region, scutellum and mesopleure dark brown.
162 Zoologica: N. Y. Zoological Society [III; 4
Described from several workers and a female taken from a
single colony nesting in a Tachigalia petiole at the Penal Settle-
ment, near Bartica, August 10.
This species, though related to L. tristani Emery of Costa
Rica, appears to be distinct both from it and the various forms
of echinatinodis Forel in the structure of the clypeus, and petiole,
in sculpture and coloration.
19.—Allomerus octoarticulatus Mayr.
Of this extraordinary little ant a single small colony with
pupz was found in a young Tachigalia on the Puruni trail at
Kartabo. This I believe to be a very exceptional occurrence,
since the species is the most typical and abundant tenant of
another myrmecophyte, Cordia nodosa, so that I shall de-
scribe its peculiar habits in another publication in connection
with the other ants which inhabit the stem-swellings of that plant
and go to make up what may be called the “‘Cordia biocoenose.”
20.—Atta cephalotes L.
I include this well-known leaf-cutting ant in the Tachigalia
biocoenose, because on one occasion I found that it had completely
defoliated some of the young Tachigalias growing near the Penal
Settlement. The petioles of these plants were either empty or
contained recently fecundated Pseuwdomyrma damnosa queens
that had only begun to lay eggs, so that the plants were quite
unprotected.
SUBFAMILY DOLICHODERIN2.
21.—Dolichoderus attelaboides Fabr.
Workers of this singular ant were occasionally seen attend-
ing Membracids on the shoots of young Tachigalias growing
along the Puruni trail, at Kartabo. The ant is more common in
attendance on Membracids feeding on the terminal shoots of
various Melastomaceze.
1921] Wheeler: Tachigalia Ants 163
22.—Azteca traili Emery.
A few workers and pupe belonging to an incipient colony
of this species were taken during September from a Tachigalia
petiole at Kartabo. The species has been recorded by Forel and
Emery from other myrmecophytes (leaf-sacs of Tococa guianen-
sis and other Melastomacee, root-stocks of Polypodium).
23.—Azteca foveiceps sp. nov.
(Fig. 16, a-c)
Worker. Length 1.7-2.2 mm.
Closely related to A. schumanni Emery; practically mono-
morphic. Head flattened, about one and one-fifth times as long
as broad, distinctly narrower in front than behind, with rather
straight sides and excised posterior border. In the middle of the
head above a small, shallow impression, or fovea. Eyes small, flat,
less than twice their length from the anterior corners of the
head. Mandibles curved, their apical borders with nine teeth,
alternately larger and smaller. Clypeus feebly convex behind,
flattened in front, with nearly straight, feebly bisinuate anterior
porder. Frontal area obsolete; frontal groove absent; frontal
carinz very short. Antenne short and thick; scapes enlarged
and somewhat flattened at the tip, reaching only to the middle
of the distance between the posterior orbits and the posterior
corners of the head; joints 2-10 of funiculi very short and trans-
verse, the more basal fully three times as broad as long. Thorax
short and stout, about twice as long as the transverse diameter
of the pronotum. Sutures, especially the mesoépinotal, pro-
nounced and impressed; mesonotum as long as broad, convex
but rising very little above the pronotum. Epinotum rounded
subcuboidal, the base longer than the declivity, the former hori-
zontal, the latter steep. Petiole small, the node high, squamiform,
inclined forward, elliptical from behind, its border narrowed
above, only moderately sharp in profile. Gaster elongate elliptical,
the first segment short, with a deep impression anteriorly for the
accommodation of the petiole. Legs stout.
Shining; very finely but not densely punctate; the punctures
on the mandibles more distinct.
164 Zoologica: N. Y. Zoological Society [III ; 4
FIG. 16. AZTECA FOVEICEPS SP. NOV.
a, worker, lateral view; 6, head from above; c, head of female.
Body and appendages covered with rather long, grayish,
appressed pubescence, not sufficiently dense to obscure the
shining surface; hairs yellowish, erect, rather short, abundant on
the thorax, sparser on the head and gaster, short on the appen-
dages.
Piceous black; borders of gastric segments paler; mandibles
red; tarsal joints 2-5 and sides of clypeus yellowish brown;
antenne dark brown, their tips blackish, their insertions and the
mouth yellowish.
Female. Length 4.5-4.8 mm.
Head decidedly longer and more nearly oblong than in the
worker, about one and one-half times as long as broad, the sides
straight and parallel, narrowed only near the clypeus. Eyes
large, feebly convex, their distance from the anterior corners of
the head less than half their length. Antennal scapes reaching
1921] Wheeler: Tachigalia Ants 165
only one-third the distance from the eyes to the posterior corners
of the head. Thorax broader than the head; mesonotum convex;
epinotum rounded, convex, without distinct base and declivity.
Petiole thicker, more erect, with much blunter superior border
than in the worker.
Sculpture, pilosity and color as in the worker but the erect
hairs are shorter and less abundant on the thorax. Wings
distinctly and uniformly infuscated, with strong, dark brown
veins and pterostigma.
Male. Length 1.6-1.8 mm.
Head through the very large eyes a little longer than broad,
with extremely short cheeks, growing narrower behind the eyes,
where the sides are nearly straight. Mandibles very small,
atrophied, bluntly acuminate, edentate. Clypeus short, convex
in the middle, with broadly and evenly concave anterior border.
Scapes very small and short, scarcely more than twice as long
as broad, constricted basally, somewhat swollen apically; first
funicular joint globose, broader than the scape, succeeding joints
somewhat longer than broad, growing successively shorter and
narrower, longer at the tip. Thorax shaped much as in the
female, but the epinotum more sloping; petiole broader, erect
with its superior border blunter than in the worker but more
acute than in the female. Legs rather slender.
Sculpture as in the worker and female; pubescence and
especially the pilosity much shorter and sparser; appendages
without erect hairs; antennal funiculi beyond the first joint
clothed with fine, dense, erect, white hairs.
Dull, dark piceous brown; head darker; mouthparts and
genitalia sordid yellowish. Wings as in the female.
Described from numerous specimens taken at Kartabo from
the petioles of large, vigorous Tachigalia trees.
This species is so close to Emery’s schumanni, that it may
prove to be merely a subspecies of that form which was taken
on the Rio Guainia, an affluent of the Cassiquiare, in Venezuela,
166 Zoologica: N. Y. Zoological Society [IIl;4
in the leaf-sacs of a Rosaceous myrmecophyte, Hirtella guainiz
Spruce (ex Hooker fil.) +
Emery saw only workers and a few very imperfectly pre-
served females of schwmanni. His figures show, however, that the
heads of both are decidedly shorter than in foveiceps, that the
head of the worker is not so much narrowed anteriorly and that
the antenne are longer, with the median joints of the funicle
less transverse, and the vetiole of the worker more erect and with
a smaller node.
SUBFAMILY FORMICINAE.
24.—Brachymyrmex heeri Forel.
Colonies of this minute honey-yellow ant with brood were
repeatedly found in the petioles of young Tachigalias along the
trails near Kartabo. More frequently it nests in dead stems ot
a very common Rubiaceous weed (Spermacoce verticillata), or in
the twigs of bushes. It also occurs under the bark of old logs.
25.—Brachymyrmex heeri var. basalis var. nov.
Worker. Honey yellow, like the type, except the first gastric
segment, which is black.
*This plant is cited by Emery in his Azteca monograph (1893) as
“Chrysobalanea hirtella Guainie Hooker fil. but the “Index Kewensis” gives
the name as cited above. It may be of interest to quote in this connection
Spruce’s remarks (1908, p. 395) on another species of Hirtella with ant-
inhabited leaf sacs: ‘‘Examples of sac-like ant-dwellings exist in the leaves
of plants of other orders, so like those already described in Melastomes, that
it is scarcely worth while to do more than indicate some of the species. The
solitary instance known to me in Chrysobalans is that of Hirtella physophora
Mart., a slender arbuscle growing just within reach of inundations in the
forest about the mouth of the Rio Negro. The distichous, oblong, apiculate
leaves are nearly a foot long, and at the cordate base have a pair of com-
presso-globose sacs tenanted by ants. On cutting open the sacs I was rather
surprised to find them lined with cuticular tissue and hairs, just like the
underside of the leaf; which seems to show that they are produced by a
_recurvation of the ale of the leaf, through the ants nestling at first (Aphis-
like) under the leaf and causing it to become bullate, and that the recurved
margins have at length reached and coalesced with the midrib so as to
form a pair of scas.” The fanciful explanation in the concluding sentence
was evidently in part responsible for the unfortunate refusal of the Linnean
Society to publish Spruce’s very valuable paper when it was presented
in 1869.
1921] Wheeler: Tachigalia Ants 167
A single small colony with brood inhabiting the petiole of a
small Tachigalia on the Puruni trail at Kartabo.
26.—Camponotus (Myrmothrix) femoratus Fabr.
On several occasions I found this very pugnacious ant attend-
ing Membracids on the young terminal shoots of small Tachigalias
along the Puruni trail at Kartabo. As a rule the femoratus
workers were accompanied by the workers of Crematogaster
limata subsp. parabiotica. These two ants regularly live in para-
biosis in the ant-gardens on the trees of the jungle, as I have
shown in a previous paper (1921).
27.—Camponotus (Myrmobrachys) zoc Forel
This very active but timid species occasionally nests in the
petioles of young Tachigalias but is more frequently found in
the cavities of the dead twigs of other trees.
28.—Camponotus (Myrmobrachys) pittieri Forel var.
poenalis var. nov.
Worker. Of a darker, more piceous brown color than the
type from Costa Rica, the gaster being concolorous with the
thorax and legs and not paler and more yellowish; the darker
brown area on the top of the head is deeper and more extensive.
The erect pilosity on the upper surface is distinctly sparser and
the hairs on the legs and especially on the scapes are much more
oblique, or reclinate. The base of the epinotum is more sloping
so that the thorax is distinctly lower behind than in the type.
Female. Length about 6.5 mm.
Darker than the worker; front as well as the vertex of the
head and also the scutellum, pleurze and posterior portion of
epinotum, dark chocolate brown; mandibles red, with black apical!
borders. Antennal funiculi not infuscated at the tip. Mesonotum
rather flat above. Head, pro- and mesonotum subopaque, more
strongly punctate than in the worker; scutellum, postscutellum
and epinotum distinctly shining. Petiole with broader, more trans-
168 Zoologica: N. Y. Zoological Society [III;4
verse superior border when seen from behind, more acute in
profile. Wings faintly infuscated, with light brown veins and
conspicuous dark brown pterostigma. Pilosity similar to that
of the worker but somewhat shorter, and the hairs on the funiculi
and legs more appressed.
Described from specimens from a single colony found nesting
with larve and cocoons in a petiole of a young Tachigalia at the
Penal Settlement, near Bartica, August 10. This ant is evidently
rare or very local, as I did not again take it in the vicinity of the
Tropical Laboratory.
—
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III, NUMBER 5
(Tropical Research Station Contribution Number 99)
—_— S| —<_
NOTES ON THE HABITS OF EUROPEAN AND
NORTH AMERICAN CUCUJIDAE sens. auct.)
By WILLIAM MoRTON WHEELER
Pee ukS HED BY THE S Ot Boe
HE ZOOLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
ook “EL
Volume III, Number 6
NOTES ON THE HABITS OF EUROPEAN AND
NORTH AMERICAN CUCUJIDAE (sens. auct.)
By WILLIAM MoRTON WHEELER
Family Silvanidxe Boving.
' Oryzephilus surinamensis L. (Fig. 7.) The “saw-toothed
grain-beetle” the most abundant, most widely distributed and
best-known species of the family.’ It is cosmopolitan and gre-
garious; living in nearly all stored human foods of vegetable
origin; cereals (rice, wheat, maize, barley, etc.), ground or un-
ground or in the form of paste (macaroni), bread etc.; dried
fruits, nuts, copra, more rarely in sugar, starch, drugs, tobacco,
snuff or dried meats. The larva, which is very active, is also
gregarious, living with the beetles and evidently capable of
thriving on most of the substances mentioned (Glover, 1869;
Guillebeau, 1890; Chittenden, 1895, 1897, 1911; J. B. Smith,
1909; Girault, 1912). When ready to pupate it may make a
rude cocoon by agglutinating particles of food detritus with an
oral secretion. The pupa is sometimes free, however, i. e.,
_not inclosed in a cocoon, and is attached by its hind end to
the shrivelled larval skin which has been previously attached
to the substratum (Blisson, 1849; Coquerel, 1849; Chittenden,
1895).. During the summer the whole life-cycle requires about
twenty-four days, in spring from six to ten weeks. There are
six or seven generations a year in the latitude of Maryland, and
in that latitude it winters over as an adult (Chittenden, 1895).
It seems to be present wherever the Indian-meal moth (Plodia
interpunctella) is found (Chittenden, 1897) and has often been
found living with another common grain-pest, the Curculionid
Calandra oryzx (Perris, 1853; Ganglbaur, 1899). In England
and Scotland the beetle has been repeatedly taken out of doors
*I find that this or one of the closely allied species is figured by Redi
(1671) who refers to it as “vermiculus qui condita arrodit” and “vermis
conditorum et pharmacorum.” Redi also figures the grain weevil, Calandra
oryzae.
174 Zoologica: N. Y. Zoological Society [TES
under the bark of trees (Fowler, 1889; Champion, 1896). “As
an instance of unusual trouble caused by this insect may be
mentioned the case cited by Taschenberg of the beetles having
invaded sleeping apartments adjoining a brewery where stores
were kept and annoying the sleepers at night by nipping them
in their beds” (Howard and Marlatt, 1896).
Oryzaephilus mercator Fauvel—The ‘merchant grain-
beetle,” very similar to the preceding, cosmopolitan, and recorded
as living in and feeding on pea-nuts, English walnuts, wheat,
corn-meal, cereipo fruit (WVyrospermum frutescens), candle-nuts
(Aleurites moluccana), and dried currants (Guillebeau, 1890;
Chittenden, 1897, 1911). It has been taken under the bark of
plane-trees in France, in the neighborhood of mills (Guillebeau).
Champion (1896) records it as occurring in shipments of pea-
nuts at Rouen in company with another grain-beetle, the Tene-
brionid Palorus subdepressus Woll. ‘The close relationship of
mercator and surinamensis makes reasonably certain their
identity as regards development, nor is it probable that they
differ in any degree in food-habits” (Chittenden, 1896).
Oryzaephilus bicornis Erich.—Cosmopolitan, but less widely
distributed and more southern than surinamensis. Lives and
breeds in wheat and dried figs in France; also taken in rubbish
at the base of fig-trees (Guillebeau, 1890; Chittenden, 1897).
Oryzaephilus gossypii Chitt—Cosmopolitan; breeding in
cotton-seed (Chittenden, 1897).
Silvanus bidentatus Fabr.—Europe and United States (N.
J., Ind., Conn., So. Cala., Fla.). Under chestnut bark in the
United States (Glover, 1869). Recorded as occurring under
bark of oaks and firs in England (Fowler, 1889), of firs in
Germany (Reitter, 1911), of poplars, elms and figs in France
(Picard, 1919). J. B. Smith (1909) records it as occurring
under bark throughout New Jersey, as not rare, and as taken
most of the season. According to Picard it has the same mode
of life as the species of Laemophloeus (q. v.).
Silvanus gemellatus Duv.—Cuba and Southeastern States to
New York. Lives and breeds in maize in the field as well as in
granaries, also in wheat and over-ripe or dried fruits. It nearly
1921] Wheeler: Habits of Cucujidae 175
always first destroys the germ of the kernel and hence causes
considerable injury to seed maize in the Southern States. “It is
essentially an out of doors species, but when conditions favor its
increase may become a serious pest in granaries, as it is capable
of breeding from egg to adult in the short period of three weeks’”’
(Chittenden, 1897, 1911).
Silvanus unidentatus Fabr.—Europe. Perris (1853, 1876)
states that the larva is common in France under the bark of oak,
poplar, chestnut and willow, rarer under pine bark. It is agile
and photophobic and lives gregariously with the adult beetles
among the detritus left by the larve of Bostrychus, Cerambycids
and Buprestids. In Germany under elm bark (Kaltenbach,
1874) ; in Britain under bark of beech, oak, horn-beam, etc.
(Fowler, 1889), of deciduous trees in general (Reitter, 1911).
Silvanus fagi Guérin.—Europe. Under beech bark; in
England in dead branches of fir (Fowler, 1889); in Germany
under fir bark and fir-cones (Reitter, 1911).
Silvanus planatus Germ.—Eastern United States (N. J.,
Ind., Fla.). Under pine-bark in New Jersey (J. B. Smith, 1909).
Cathartus advena Waltl.—Cosmopolitan. Lives and breeds
in stored wheat, rice, corn in stack, grain, meal, middlings, flour,
dates, figs, lichi nuts, table beans, cacao-beans, edible tubers, etc.,
but apparently only when these substances are not kept dry and
clean. “In breeding experiments recently conducted by the writer
it failed to develop in fresh grain or meal, but bred freely in
corn-meal which was moistened and produced mold. The beetles
particularly fed freely on the molds, of which there were three
or four species, and it would appear that this is the normal habit
of the insect’? (Chittenden, 1897). Pierce (1917) cites C. advena
among the insects infesting teak in India, stating that it “breeds .
in leaves, forming galls, causing leaves to drop” (sic!). Perris
(1876) found the larva in lichi nuts in company with larve of
O. surinamensis. Fowler (1889) states that C. advena is some-
times found out-of-doors in England and that Waterhouse took
it under cut grass. In New Jersey it is “rare under bark; more
common in stored grain, fruits, nuts, etc., particularly such as are
spoiled” (J. B. Smith, 1909)
176 Zoologica: N. Y. Zoological Society [III;5
Cathartus’ cassiz Reiche—Cuba and Southern States.
Glover (1869) describes the larva as feeding on maize kernels,
near the germ, and also on the exposed seeds of cotton bolls.
Doran (1892) found this beetle breeding in bran and middlings
and producing a temperature 42 deg. F. higher than that of the
room in which it was living!
Cathartus longulus Blatchley.—Indiana. “Sifted from bor-
ders of Sphagnum marsh” (Blatchley, 1910).
Cathartus excisus Reitter—Taken in Germany in Havana
tobacco (Ganglbaur, 1899; Reitter, 1911).
Cathartus quadricollis Guérin.—Imported into Germany in
Havana tobacco and also in the fruits of Cassia fistula (Gangl-
baur, 1899) ; imported in Havana cigars (Reitter, 1911).
Nausibius clavicornis Kug.—Cosmopolitan. ‘Under bark
and in sugar” (Glover, 1869). “Often found on ships in plant
wares. It also lives in old nests of South American bees” (Gang]l-
baur, 1899). Introduced into Britain, found in sugar, ete.
(Fowler, 1889). In New Jersey under bark and also occasionally
in store-houses (J. B. Smith, 1909). In Indiana “‘two specimens
were taken with a dozen other species at sap beneath the bark
of a soft maple tree. Leconte states that it occurs usually in
rice, sugar and other articles of commerce throughout the United
States” (Blatchley, 1910).
Telephanus velox Haldem.—United States (N. J., Ia., Ind.,
Conn.). “Very common under rubbish of all sorts and as its
name implies, runs with remarkable swiftness” (Casey, 1884).
“Occurs more often beneath stones, chunks and dead leaves than
beneath bark. When exposed it usually remains quiescent with
antenne folded against sides; but if touched it runs with great
swiftness, whence its specific name” (Blatchley, 1910).
“Throughout the State (N. J.) under stones and old leaves;
rarely under bark; may be sifted out from fall to late spring’
(J. B. Smith, 1909).
Family Cucujide (Boving emend.)
Cucujus clavipes Fabr.—Atlantic States to Illinois and Iowa;
a beautiful scarlet red insect, represented in the Pacific States
1921] Wheeler: Habits of Cucujidae LaF
by the var. puniceus Mann. The larva is said to be predatory
(Le Baron, 1874; J. B. Smith, 1909). Le Baron (1874) figures
the larva; Dimmock (1884) records the adult beetle as occurring
under decaying butter-nut bark. Hamilton (1886) gives the
following account of the insect: ‘‘The very depressed form of
this well known beetle indicates, a priori, its subcortical habit,
and no other has power to adapt its tastes to a greater variety
of timber—locust, maple, oak, hickory, gum, buckeye, ete.—are
all alike to it. The larve do not eat the wood nor the bark,
living apparently on the moisture existing between the two. They
are elongate, much depressed, brownish yellow, and scarcely to
be distinguished from those of Dendroides canadensis. Some-
time in September, the larva having matured, constructs a circu-
lar cell from small particles-of the decaying bark and wood, and
in this completes its transformations before severe frost, but the
beetle does not quit the cell till the following spring. I have
never known any of these insects to be taken elsewhere than
under bark, though they undoubtedly fly, being possessed of a
good pair of wings. On the 10th of October, fifteen newly dis-
closed individuals and several pup# were taken under the bark
of a gum log; the latter are depressed like the beetle, pale at first,
the eyes, antenne and portions of the legs gradually changing
to black, and the elytra becoming red after disclosure. This
insect is annual.”
Cucujus haematodes Erich.—Europe. In Germany under
bark of maples, rarer under bark of fir-stumps (Reitter, 1911).
Cucujus cinnaberinus Scop—Europe. Habits like those of
haematodes (Reitter, 1911).
Dendrophagus crenatus Payk.—Europe. In Germany under
oak bark; gregarious (Kaltenbach, 1874) ; in Britain under bark
of fir (Pinus sylvestris), less frequently under bark of larch
(Larix europxa) (White, 1872; Fowler, 1889). According to
White the larva has been supposed to be carnivorous, but is truly
phytophagous, feeding on the decaying inner bark of dead and
usually prostrate trees of the species mentioned. The beetles are
agile and seem to come out at night and run about on the bark.
The eggs seem to be laid in the spring by hibernating females;
the larve feed twelve to fourteen months, becoming pup during
178 Zoologica: N. Y. Zoological Society [III;5
the second summer, the beetles emerging in August. The larva
is very quick and agile and when disturbed moves the hinder part
of its body quickly from side to side. When about to pupate “it
attaches itself firmly to a piece of bark by the thirteenth segment,
and the pupa remains attached by its anal segment to the larva-
skin.” Perris (1876) believes that White is mistaken in his
account of the food-habits of the insect, and that it is carnivorous
or coprophagous, like Brontes planatus (q.v.).
Emporius signatus Frauenf.—Imported into Germany from —
the tropics in Havana cigars (Ganglbaur, 1899).
Pediacus depressus Herbst.—Recorded from Europe, Van-
couver and the United States (So. Cala., Mich., Lake Superior,
Colo., Pa., S. C.). In Britain under oak-bark, in mold-like fungus;
also in stores on board a yacht (Fowler, 1899). Under bark of
deciduous trees in Germany (Reitter, 1911).
Pediacus fuscus Erich.—Europe and North America, Alaska,
Labrador, British America, Mich., Neb., Colo., N. Mex., Lake
Superior, and probably cosmopolitan. In Germany under pine
_and oak bark (Reitter, 1911), and under the bark of firs (Kalten-
bach, 1874). In Britain under bark and in chunks of freshly
cut oak, beech and horn-beam (Fowler, 1889).
Brontes planatus L.—KEKurope. Perris (1853) describes the
larva as livid reddish, punctate with carmine dots, and as being
very agile and supple. In France it is common throughout most
of the year under the bark of various trees, but most often
under oak or pine bark. Perris believes the larva to be carni-
vorous, because it is always found among the larve of Tomicus
or with Poduri or mites ‘dont elle fait, sans doute, sa nourri-
ture.” ‘When about to pupate it attaches itself to a flat surface
by means of the tubercle of its last segment, then the skin splits
along the back and is pushed back to the posterior end of the
body, where it remains much wrinkled.” In Germany under
bark of deciduous trees, especially of oaks (Kaltenbach, 1874;
Reitter, 1911); in Britain under bark of dead beech trees
(Fowler, 1889). According to Perris (1876) the larva is
predaceous and a scavenger, living under the bark of various
trees, including, chestnut, especially where the larve of other
1921} Wheeler: Habits of Cucujidae 179
beetles have lived. All the stages are passed through in several
months, the young larve being found in the spring, the adults
in late summer. The latter hibernate under the bark. According
to Ganglbaur (1899) the larva is “undoubtedly carnivorous.”
Psammoechus bipunctatus Fabr.—Europe. Recorded in
Britain from marshy places at roots of grass and in refuse
(Fowler, 1889) ; in Germany on the banks of streams or ponds
under dead rushes (Ganglbaur, 1899; Reitter, 1911).
Psammoechus desjardinsi Guérin.—Cosmopolitan (Casey,
1884) ; Florida (Leng, 1920).
Family Laemophloeide Boving.
Laemophloeus alternans Erich.—Cosmopolitan; ‘found
everywhere” (J. B. Smith, 1909). In Germany under bark of
conifers in galleries of beetles of the genus Pityogenes (Reitter,
1911). In France occurring occasionally in the galleries of
Hypoborus ficus, a bark beetle of fig-trees, but also in galleries
of other bark beetles in other plants (Perris, 1853; Picard,
1919).
Laemophloeus ater Oliv—EKurope. In France in dead or
dying stems of broom and gorse, preying on the larve of the
bark-beetle Hylesinus rhododactylus (Perris, 1853). In dead
stems of broom and under elm-bark in Britain (Fowler, 1889).
Rare under bark; common in bran and middlings in Germany
’ (Reitter, 1911). Occurs in France not only in the galleries of
Hypoborus ficus in fig-bark, but also in the galleries of other
Scolytids in other plants (Picard, 1919).
Laemophloeus biguttatus Say.—United States (Ind., Neb.,
Fla., So. Cala., N. J.). Throughout the State of New Jersey
under bark; fall to mid-summer. (J.B. Smith, 1909.)
Laemophloeus bimaculatus Payk—Europe. Under bark of
oak, beech and horn-beam in Britain (Fowler, 1889); in Ger-
many under beech bark (Reitter, 1911).
Laemophloeus castaneus Erich.—Europe. Under birch and
chestnut bark in Germany (Reitter, 1911).
180 Zoologica: N. Y. Zoological Society [TSS
Laemophloeus corticinus Erich.—Europe. Under oak-bark
in Germany (Reitter, 1911).
Laemophloeus clematidis Erich—Europe. In Clematis
vitalba. A constant companion of Xylocleptes bispinosus, feeding
on the refuse and dejecta in its galleries (Perris, 1853; Kalten-
bach, 1874; Reitter, 1911). In dead stems of Clematis vitalba
in Britain (Fowler, 1889). é
Laemophloeus denticulatus Preysl—Europe. ‘The larve
probably merely accompany the larve of really injurious
xylophagous beetles. According to Hartig they are found in
the cones of conifers, according to Ratzeburg also under dead
bark, according to Hellwig and Panzer under the bark of linden
trees’”’ (Kaltenbach, 1874).
Laemophloeus dufouri Laboulb.—Europe. The larve were
found by Perris (1853) under pine-bark hibernating in January
with the pupe and beetles in the galleries of Crypturgus pusillus.
He believed that the larve feed on those of the Crypturgus, but
more probably they merely eat the dejecta in the burrows.
Laemophioeus duplicatus Waltl—Europe. Under bark of
beech, oak, etc., in Britain (Fowler, 1889); in Germany under
beech bark (Reitter, 1911).
Laemophloeus fasciatus Mels.—United States (Ind., Pa.,
Minn.). In Minnesota it “is found under bark, but is also very
common near saw-mills, especially in those in which hardwood
is sawed. In running over persons engaged in mills it will bite,
even without provocation, and may cause bad sores” (Lugger,
1899).
Laemophloeus ferrugineus Steph.—Cosmopolitan (Casey,
1884). In Britain in hay-stack refuse, rarely under bark; also
in granaries (Fowler, 1889). In Germany in rice, bran, mid-
dlings and meal (Ganglbaur, 1899; Reitter, 1911; Chittenden,
1911). “Its larva, which does not differ from that of other species
of the genus, has been described by Carpentier (Bull. Soc. Linn.
Nord. France, April, 1877, 3 pp. 239-241). It lives under the
bark of oaks, cherry trees and, it is said, in fig-trees inhabited
by wood-boring insects, but also in grain attacked by Calandra,
in bran, flour, etc.” (Picard, 1919).
1921] Wheeler: Habits of Cucujidae 181
Laemophloeus hypobori Perris.—Europe. (France). Living
only in the galleries of a bark-beetle Hypoborus ficus, in fig-trees.
Both the adult beetles and the larve live gregariously and feed
on the dejections, exuvie and all kinds of detritus which they
find in the empty galleries. They occur more rarely in the
brood galleries that are still inhabited by the parent Hypobori.
Both larve and adults are found throughout the year and both
stages, but especially the adults, are found hibernating (Picard,
1919). This author gives an excellent account of the beetle and
its habits. .
Laemophloeus junipert Grouv.—Europe. Under bark of
Juniperus in galleries of the bark-beetles Phloeosinus bicolor Br.
and thujze Perris (Reitter, 1911). ‘At Monpellier this is the
rarest species in fig-trees; I know of only one capture made in
August by J. Lichtenstein. L. juniperi is nevertheless common
in Herault, as in the whole South. I often found it under the
bark of elms riddled by Pteleobius vittatus and kraatzi; it is also
found in the galleries of Hypoborus (Liparthrum) mori of the
mulberry, of Phloeosinus thujx and bicolor infesting Juniperus,
Cupressus, Thuja and other conifers, of Phloeotribus scarabaeo-
ides of the olive, etc.” (Picard, 1919). The larva, which is copro-
phagous and detritivorous, has been observed and described from
Thuja infested with Phloeosinus thujx by Decaux (Bull. Soc. Ent.
France, June 25, 1890, p. 125) and is redescribed by Picard.
Laemophloeus modestus Say.—United States (Tex., Fla.,
D. C., N. J.). “Taken in hemp-seed, the interior of which had
been entirely eaten out” (Glover, 1869). In New Jersey
occurring under bark and in siftings (J. B. Smith, 1909).
Laemophloeus monilis Fabr.—Europe. In Germany under
beech and linden bark, gregarious in the burrows of Tachrorhy-
chus bicolor (Reitter, 1911).
Laemophloeus perrisi Grouv.— Europe. Occurring in the fig.
“It is also known from the Lentiscus and pines infested with
Pityophthorus. This is a Corsican insect, very rare in France,
where it has scarcely been seen, except in the Provence” (Picard,
1919).
182 Zoologica: N. Y. Zoological Society — [III;5
Laemophloeus pusillus Schén—Cosmopolitan and common
(Casey, 1884). In Britain in granaries, imported with grain
(Fowler, 1889). Occurs commonly in cereals, but probably in
the main predaceous and scavengering (Chittenden, 1911). In
rice, grocery wares, etc., in Germany (Reitter, 1911).
Laemophloeus testaceus Fabr.—Europe. Similar to L. denti-
culatus but occurring under linden bark (Kaltenbach, 1874), and
under dry beech bark (Reitter, 1911). Perris (1853, 1876) found
the larve under oak bark in galleries of Tomicus and under
chestnut bark with the larve of the bark beetle Dryocoetes capro-
natus on which it was preying, but also living on the dejections
and capable of developing to maturity in the absence of the
Dryocoetes. Also found under bark of dead oaks with the larve
of the same Scolytid and in the elm in the galleries of Hylesinus
vittatus and kraatzt.
Laemophloeus turcicus Grouvy.—Cosmopolitan. Imported
into Germany in dried fruits, especially prunes, from the orient
(Rettter> 1911).
Laemotmetus ferrugineus Gerst——Cosmopolitan. Imported
into Germany in rice (Ganglbaur, 1899; Reitter, 1911).
Laemotmetus rhizophagoides Walker.—Cosmopolitan. Found
in stored rice in Berlin, Germany (Chittenden, 1911).
Lathropus vernalis Erich.—United States (Fla., Miss., N. J.,
Conn.). Throughout New Jersey, secured by beating dead
branches, May to July (J. B. Smith, 1909).
Lathropus sepicola Miller—Europe. According to Perris
(1876) the larva lives under elm-bark in the galleries o£ the bark
beetles Scolytus multistriatus, Hylesinus vittatus and H. kraatzi
and feeds on their dejections. The perfect insect emerges in
May. Reitter (1911) often beat the latter from dry wood
fences.
Phloeostichus denticollis W. Redtb—Europe.- Under the
inner bark of maple-trees, together with the larval stages
(Weisse, 1897; Reitter, 1911).
Prostomis mandibularis Fabr.—Europe and Pacific States
(Or., Nev., Cala.) and Vancouver. In very rotten, damp wood
1921] Wheeler: Habits of Cucujidae 183
of deciduous trees. The larve are often present in great numbers
with the beetles (Ganglbaur, 1899; Reitter, 1911). Perris (1876)
found the larve in rotten chestnut wood and states that Curtis,
Chapuis and Candéze took them in rotten oak wood.
Hemipeplus marginipennis Lec.—Cited only from Georgia
and Florida (Leng, 1920). According to Schwarz a rather
abundant species under palmetto bark (Casey, 1884).
Hemipeplus microphthalmus Schwarz.—Cited only from
Florida (Leng, 1920), where it was taken flying at night by
Schwarz (Casey, 1884).
Inopeplus praeustus Chevr.—Antilles. Adults and larvze
living gregariously in the burrows of a Scolytid in branches of
cacao. The larva is peculiar in having the prolongations of the
ninth abdominal segment in the form of forceps (de Peyerimhoff,
1903).
Family Scalidiide Boving.
Catogenus rufus Fabr.—United States (Ind., Ia., Md., N. C.,
Fla.). Fiske (1905) has shown that the larva is an external
parasite on the larve of Braconid Hymenoptera and Cerambycid
pupe; “its habits differing in no essential feature from those of
many species of external Hymenopterous parasites. The adult is
fairly common throughout the South, and is found beneath the
loose bark of recently dead and dying trees, both conifers and
deciduous. It occurs at nearly all seasons of the year, but is
especially common during the late fall and early spring, and is
found hibernating in situations similar to the above mentioned.”
Fiske seems to have overlooked a note on this beetle by Dimmock
(1884), who says: “In Connecticut it is common beneath the
loose bark of the trunks of hickory trees, and I have reared its
larva which fed upon a pupa of Elaphidion parallelum, a borer
in hickory.”
‘
¢
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 6
(Tropical Research Station Contribution Number 100)
DESCRIPTIONS OF NEW SPECIES OF
COLEOPTERA
By E. A. SCHWARZ and H. 8S. BARBER
United States Department of Agriculture
«
on Bos BE. D eed a SO Ce SO: Co ie 2
THE ZOOLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
Volume III, Number 6
DESCRIPTIONS OF NEW SPECIES OF
COLEOPTERA
By E. A. SCHWARZ and H. S. BARBER
United States Department of Agriculture
Specimens of three species of beetles from Kartabo, Bartica
District, British. Guiana, have been submitted to us by their
discoverer, Dr. W. M. Wheeler, for identification. Two of the
species belong to the family Silvanide, while the third is an incon-
spicuous species of the ubiquitous Coccinellid genus Scymnus.
The two first are, however, very interesting forms, and it has
been necessary to erect a new genus to receive the larger and
more abundant species. These, Dr. Wheeler informs us, were
occupying the petioles of a tree (Tachigalia).
Coccidotrophus gen. nov. (Silvanide).
Eyes small, not emarginate, not prominent, occupying
middle fourth of side of head. Antenne inserted laterally, in
deep cavity between frontal margin and prominent gena, close
to and equidistant from base of mandible and front margin of
eye; compactly eleven-jointed, club not abruptly enlarged, all
joints traverse except basal joint; joints 3-9 rapidly increasing
in size, the three-jointed club densely covered with fine short
hairs. Antennal grooves short, distinct, parallel, bordered by
lower edge of eye and carina from gena. Mouthparts free, maxil-
lary palpi four-jointed, labial palpi three-jointed. Elytra with
lateral costa and nine distinct series of strial punctures.
Pygidium usually covered by elytra when abdomen is con-
tracted (when distended the pygidium is often exposed).
Intercoxal process of prosternum about one-seventh as
wide as prothorax, wider behind, its sides overlapping the
epimera; front coxae separated from hind margin of pro-
thorax by about their own length. Intercoxal process of
mesosternum slightly narrower than that of prosternum, truncate .
apically and nearly reaching hind margin of coxae, metepimeron
reaching the middle coxae. Hind coxae separated by strong,
190 Zoologica: N. Y. Zoological Society iise
acute process of first abdominal sternite, which fits into median
notch in metasternum. First ventral abdominal segment with
coxal lines very fine, short, curved, becoming almost parallel
behind, but not reaching posterior margin; four following
sternites each slightly shorter than the first, the last rounded
apically, concave beneath, with carinate posterior margin. Legs
short, stout, femora incrassate, first three tarsal joints inflated,
hairy beneath, fourth small but distinct, fifth joint as long as the
three basal joints together. Genotype: Coccidotrophus socialis
sp. nov. .
This genus is so unlike other Silvanids known to us that its
true affinity was not recognized until Dr. Boving’s studies of its
larva proved it to agree well with typical Silvanid larve and not
to be related to Hapalips, with which we were trying to associate
the adults. Once suggested this relationship was readily confirmed
by the characters used by Ganglbauer (Die Kafer von Mitteleu-
ropa, 1889, vol. 3, p. 577) to distinguish the adults of this family:
front coxal cavities closed behind, metepimeron reaching middle
coxae, etc.
We cannot identify Coccidotrophus with any of the genera
considered by Grouvelle (Ann. Soc. Ent. Fr., vol. 81, 1912, pp.
313-386), but it seems most nearly to approach Synoemis Pascoe
(1866), the only species of which was described from the Malayan
Peninsula.
Coccidotrophus socialis sp. nov.
(Plate Vi; figs.'1 to 5):
Very elongate, parallel, depressed, shining, castaneous,
glabrous (except for sparse, microscropical, decumbent
pubescence) .
Length, 3.5-4.5; width, 0.6-0.8 mm.
Habitat—British Guiana.
Head slightly wider than prothorax, widest at the very
slightly prominent eyes, one-sixth longer than wide, feebly
constricted into a neck in basal fourth, sides convergent in front,
front margin broadly, shallowly notched; upper surface finely
granulose except a small occipital smooth area, feebly convex,
transversely somewhat tumid between and behind eyes, which
1921] Schwarz and Barber: New Coleoptera 190
are situated about middle of sides; clypeus not separated from
front, produced into a strong marginal carina, surface convex
medially, concave laterally, especially over antennal sockets;
labrum very short, transverse, feebly chitinized, emarginate in
front and with a row of about six stiff hairs; gular region
strongly pilose, feebly concave; mandibles moderately prominent,
strongly bidentate apically; antenne stout, shorter than width
of front at their point of insertion. Pronotum three-fourths as
wide as long, widest at apical angles which are subacute; narrow-
est just before middle and at base; sides feebly sinuate, sub-
parallel, front margin straight, hind angles obtuse, hind margin
arcuate, surface longitudinally flat, transversely feebly convex,
sparsely punctate, the punctures elongate, side margins with fine
marginal line. Scutellum transverse, widest behind, finely punc-
tate. Elytra slightly wider than pronotum, more than three
times as long as wide, base emarginate, humeral angles subacute,
sides subparallel to apical fourth, thence conjointly rounded;
surface longitudinally somewhat convex, transversely rather
strongly convex, strial punctures moderate, the interstices each
with a series of slightly smaller punctures supporting fine,
decumbent, short hairs. Under surface shining but with irregular
microscopic sculpture between the sparsely scattered, nearly
obsolete punctures which bear the short, fine decumbent hairs.
Described from thirty-two examples from a large series
collected by Dr. Wm. M. Wheeler at Kartabo in July and August,
1920. The sexes are almost indistinguishable unless the “palps’”’
‘of the ovipositor are extruded.
Characters of legs, antennze and mouth are shown in accom-
panying figures. Type, allotype and paratypes.—Cat. No. 24070,
U.S.N.M.
Eunausibius Grouvelle, 1912.
Although we have seen neither of the two species (Nausi-
bius tenebrionides and N. elongatus Grouv.) upon which Grou-
velle, (Ann. Soc. Ent. Fr., vol. 81, 1912, p. 314) established this
genus, his generic diagnosis applies so well to the species here
described that his genus is adopted. The postcoxal lines are
very feeble and difficult to see but are bent abruptly forward
to the coxal cavity enclosing a small area under the trochanter:
192 Zoologica: N. Y. Zoological Society [III ;6
the antennal club seems to be more abrupt and larger, and the
shape of the produced clypeus differs also from the figures Grou-
velle, (Ann. Soc. Ent. Fr., vol. 65, 1896, p. 193) has given of
the two previously known species.
Eunausibius wheeleri sp. nov.
(Plate VI, figs. 6 to 10).
Elongate, parallel, moderately convex, smooth, shining, gla-
brous, pale castaneous.
Length, 3.0-3.5 mm.; width, 0.6-0.75 mm.
Habitat.—British Guiana.
Head wider than prothorax; eyes rather large and promi-
nent, coarsely granulated, front widest in front of the rather
strongly impressed fovea opposite which the margin is some-
what thickened; front margin broadly, feebly emarginate, front
angles obliquely truncate; surface finely granulate, feebly con-
vex. Labrum membranous, concealed beneath front. Gular
area feebly concave, opaque, finely pubescent, the lateral carinz
bordering the antennal grooves convergent posteriorly and pass-
ing middle of eye. Antenne as long as width of clypeus, club
abruptly widened, oval. Pronotum about three-fourths as wide as
long, widest at the slightly acute front angles, sides almost
straight and parallel, finely carinate; front margin straight
except small sinuation near angles, hind margin arcuate at mid-
dle, on each side straight, hind angles obtuse. Surface trans-
versely convex, longitudinally flat except for a pair of feeble
impressions at basal fourth. Scutellum transversely oval, twice
as wide as long. Elytra slightly wider than prothorax, three
times as long as wide, sides parallel, apices evenly rounded;
surface nearly smooth, strial punctures feebly impressed but
conspicuous by coloration below surface. Callow specimens dis-
play interstrial rows of microscopic appressed hairs. Under
surface of body sparsely clothed with microscopic decumbent
hairs, each set in a broad obsolescent puncture.
Described from eighteen specimens submitted by Dr.
Wheeler, to whom the species is dedicated. Two of the specimens,
dissected and mounted on slides, are males. One _ speci-
men, the allotype, is somewhat crushed and displays female sex
ed
1921] Schwarz and Barber: New Coleoptera 193
organs. We are unable to distinguish the sex of the other fifteen
paratypes.
Type, allotype and paratypes.—Cat. No. 24071, U. S. N. M.
The antenne and legs are shown in accompanying figures.
Scymnus Kugelann 1794 (Coccinellide)
After some hesitation we have decided to offer the following
description of what appears to be a new but very commonplace
species of this genus, since we have failed to find any description
applicable thereto. It will, perhaps, be long before the multitude
of tropical species of Scymnus will be determinable.
Scymnus (Diomus) xantholeucus sp. nov.
Oval, very convex, shining, pubescent, pale yellow except
basal two-thirds of elytra, meso- and metasternum and median
third of first two abdominal segments, which are infuscate.
Length, 1.7 mm.; width, 1.2 mm.
Habitat.—British Guiana.
Head and pronotum finely, rather densely punctate, elytra
slightly more coarsely punctured; pubescence rather dense, the
short, silky, suberect hairs bent in all directions forming no pat-
tern. Scutellum pale. Elytra piceous in basal two-thirds, apic-
ally flavescent, the pale area not sharply limited. Under surface
moderately, densely and finely punctate, the pubescence decum-
bent and regular. Prosternum with carinae convergent anterior-
ly, reaching front margin and uniting in an arc; the enclosed
area feebly concave and supporting moderately long hairs. Post-
metacoxal line as in other species of Diomus (Group A of Horn
1895). Penultinate abdominal sternite very broadly and feebly
emarginate in the male.
Type (male), allotype and paratypes.—Cat. No. 24084, U.S.N.M.
Described from two males and two females reared by Dr.
Wheeler from larvae found among coccids in company with the
two species of Silvanide.
Although not related to semiruber Casey, the foregoing
species is similar in shape and plan of coloration. A prescutellar
infuscate area is suggested in one example.
Fig. 1. Coccidotrophus socialis sp. nov.
parts, ventral view. X 58.5.
Fig. 2. Fore leg of same. X 36.5.
Fig. 3. Middle leg of same. X 36.5.
Fig. 4. Hind leg of same. X 36.5.
Fig. 5. Hind wing of same. X 36.5.
Fig. 6. Hunausibius wheeleri sp. nov.
Fig. 7. Fore leg of same. X 36.5.
Fig. 8. Middle leg of same. X 36.5.
Fig. 9. Hind leg of same. X 36.5.
Fig. 10. Hind wing of same. X 36.5.
PLATE VI.
Antenna.
Right side of head, with antenna and mouth
X 58.5.
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 7
(Tropical Research Station Contribution Number 101)
THE LARVAE AND PUPAE OF THE SOCIAL BEETLES
COCCIDOTROPHUS SOCIALIS (SCHWARZ AND BARBER) AND
EUNAUSIBIUS WHEELERI (SCHWARZ AND BARBER) WITH
REMARKS ON THE TAXONOMY OF THE FAMILY
CUCUJIDAE
By ADAM GIEDE BO6vING, PH.D.
Bureau of Entomology Department of Agriculture, Washington, D. C.
aoe at ste Ds BY THE S:0: C1. LE TY
THE -ZOOLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
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Volume L111, Number 7
THE LARVAE AND PUPAE OF THE SOCIAL BEETLES
COCCIDOTROPHUS SOCIALIS (SCHWARZ AND BARBER) AND
EUNAUSIBIUS WHEELERI (SCHWARZ AND BARBER) WITH
REMARKS ON THE TAXONOMY OF THE FAMILY
CUCUJIDAE
By ADAM GIEDE BOVING, PH.D.
Bureau of Entomology Department of Agriculture, Washington, D. C.
Professor W. M. Wheeler has kindly given me for study
larvae and pupae of the two species of social beetles discovered by
him in British Guiana. The material was plentiful and in excel-
lent condition. In working out the relationship of these larve
it was necessary to examine larve of several related groups
and often to avail myself of studies as yet unpublished, under-
taken by Dr. F. C. Craighead and myself on the classification of
the Coleopterous families based on larval characters.
In addition to descriptions and illustrations of the two
species, mentioned above, the present contribution therefore con-
tains a brief discussion of the taxonomy of the entire family
Cucujidae.
The different subjects are arranged as follows:
A. The family Cucujidae, authorum; taxonomic remarks
with reference to their larvae.
B. The subfamily Silvaninae as based on larva) characters.
B. 1. Subfamily description of the larvae of the Sil-
vaninae. j
B. 2. The genera of the Silvaninae, systematically de-
fined by larval characters.
C. Coccidotrophus socialis and Eunausibius wheeleri.
C. 1. General morphological description of the larvae.
C. 2. Specific description of the larvae.
C. 3. General and specific description of the pupae.
D. Bibliographical notes.
LOSS 7 = Zoologica: N. Y. Zoological Society BRE ieeys
A.—THE FAMILY CUCUJIDAE (authorum).
The larvae of Coccidotrophus socialis Schw. & Barb. and
Eunausibius wheeleri Schw. & Barb. show remarkable identity
in general morphological structures with the larvae of the genera
Oryzaephilus Gangl., Silvanus Latr., Nausibius Redt. and Cathar-
tus Reiche. All these forms differ from each other only in small
details such as the proportional length of the second antennal
joint, the arrangement of the individual ocelli in the two ocellar
groups, the thickness and color of the chitin of the body or the
number and development of the tergal setae. The larva of
Telephanus (Plate X, figs. 34-38) has a well developed third
antennal joint and differs in this and some other less important
characters from the genera mentioned. However, it approaches
them closely and constitutes together with them a well defined
taxonomic unit distinct from the larvae of the rest of those genera
which have generally been combined into the single family Cucu-
jidae.
This family, however, as hitherto limited, mainly on adult
characters, includes larval types which are rather heterogeneous;
and according to these different larval types the family can be
divided and I propose here to divide it, into several new families
and subfamilies.
Thus it appears appropriate to establish the above mentioned
unit of genera as a distinct family, the Silvanidae.
This family is, through the genus Telephanus, closely associ-
ated with another unit of genera, namely Brontes Fabr. and
Dendrophagus Schon. (by Gilbert F. Arrow 1901 united into a
single genus Hyliota Latr.), Psammoecus Latr., Pediacus Shuck,
Platisus (according to description and figure by A. M. Lea of
P. integricollis Reitter; Proc. Linn. Soc. N. 8. Wales, vol. 29,
1904, p. 88) and Cucujus Fab., which together form a second
family, the Cucujidae (in restricted sense) with Brontes, Dendro-
phagus, with Psammocus in one subfamily, the Hyliotinae, and
the other genera in another subfamily, the Cucujinae. More re-
mote is the relationship between these two families and the genera
Prostomis Latr, and Dryocora (according to my own unpublished
1921] Boving: Larvae and Pupae of Social Beetles 199
notes and figures of four larve of Dryocora howitti, from New
Zealand; in Zool. Mus. Cambridge, England), which two genera
form a third unit and may constitute a subfamily, the Prostominae
(Plate IX, figs. 22, 24-26 and 27-33). Asa fourth unit I consider
the genera Prostominia (according to the excellent description
and figures by M. P. de Peyerimhoff of P. convexiuscula Grouv.,
Trans. Lin. Soc. London—2, Zool.—vol. 17, 1914, p. 156), Narthe-
cius Lec., Lathropus Erichs., Laemophloeus Laporte, Dysmerus
Csy., Hemipeplus Latr. and Inopeplus Smith (according to de-
scriptions and figures by M. P. de Peyerimhoff of P. praeustus
neve (2 0.1. Ann. Soc. Ent. Fr., vol. 71, 1902-3, p. 115):
According to an imperfect description by Weisse (Deutsch. Ent.
Zeitsch., vol. 41, 1897, p. 393) of the larva of Phloeostichus denti-
collis Redtb., this genus also belongs to the present unit. All these
larvae are closely related to the Prostominae, but separated from
this subfamily by good characters; they may constitute another
subfamily, the Laemophloeinae and together with the Prosto-
minae the family Laemophloeidae. The Laemophloeidae are, as
the following key will show, separated from the Silvanidae and
Cucujidae (in restricted sense) by exactly the same main char-
acter by which such families as the Cryptophagidae are separated
from the Mycetophagidae or the Monotomidae (represented by
the larva of Hurops) are separated from the Smicripidae (repre-
sented by the larva of Smicrips palmicola Lec.), namely the dif-
ferent shape of the maxillary mala. The genera Catogenus Westw.
and Scalidia Er., constituting the family Scalidiidae, differ to such
an extent from all the genera mentioned that their larvae
must be placed in another series of families, the Cleroidea,
proposed by myself in a recent paper (Adam G. Béving and
A. B. Champlain, Larvae of North American beetles of the family
Cleridae, Proc. U. S. Nat. Mus., vol. 57, pp. 588-95). They are,
according to Dr. F. C. Craighead (Biology of Colydiidae and
Bothrideridae, Proc. Ent. Soc. Wash., vol. 22, pp. 1-13, pl. I, II),
closely connected with his new family, the Bothrideridae, and
appear also to be near to the family Cleridae, having like this
family protracted ventral mouthparts, large cardo region and
elongate gula region (The figs. 39-44, Plate X, are reproduced
from unpublished drawings by Dr. Craighead).
200 Zoologica: N. Y. Zoological Society [TIL; 7
The four families with subfamilies, into which the old
family Cucujidae may have to be divided according to larval
characters, are characterized briefly in the following conspectus:
I. Mandible with molar structure and complicated hypo-
pharyngeal chitinizations. Ventral mouthparts retracted ;
cardo in some forms indistinct, fused with enlarged mar-
gin of epicranium.
A. Maxillary mala (possibly lacinia) falciform, with
terminal uncus; externally to uncus a more or less
sharply defined, small setose region (possibly
reduced galea). Maxillary articulating area distinct.
1. Without cerci. Eighth abdominal segment nor-
mally developed; ninth small, ventrally rudi-
mentary; tenth long, extending far behind ninth
SOG MENT SIRIOCEL I .0.kccsceskt nee Fam. Silvanidae
a. Antenna with second joint large and clavate,
third joint very small, or wanting; ocelli in
two distinct groups (Plate VIII, figs. 10, 12
EW Che Wf) eee epee Eee Subfam. Silvaninae
b. Antenna with three well developed joints;
ocelli not in two distinct groups (Plate X, fig.
24. Sen 2... SUbTam. Telephiaanams
2. NVA CORDS. A028 occa, aoe ctor sae Fam. Cucujidae
a. Tenth abdominal segment long, conical, ex-
tending far behind ninth abdominal seg-
PIVOT oy ratectoes sec stioxescnssucantess Subfam. Hyliotinae
b. Tenth abdominal segment short, wartshaped,
not extending behind ninth abdominal seg-
p02) NY A. 5, Cea AT Subfam. Cucujinae
B. Maxillary mala (possibly galea) obtuse, with or
without well defined uncus, which when present is
laterally placed on inside of mala (possibly termi-
nating a reduced lacinia) ........ Fam. Laemophloeidae
1921] Boving: Larvae and Pupae of Social Beetles 201
a. Head larger than prothorax; without ocelli;
maxillary articulating area present, cardo
distinct; hypostoma normal (Plate IX, figs.
2) S'S | ee ae Subfam. Prostominae
b. Head as large as or narrower than pro-
thorax; with ocelli; usually having maxillary
articulating area and cardo fused into a
joint region, more or less amalgamated with
broad enlargement of hypostoma,
Subfam. Laemophloeinae
II. Mandible without molar structure or hypopharyngeal
chitinizations; ventral mouthparts protracted; gular
region large (Plate X, figs. 41, 42).......... Fam. Scalidiidae
B.—THE SUBFAMILY SILVANINAE.
B. 1.—Systematic Description of the Subfamily Sivaninae
Three pairs of legs. Fused tarsus and claw. Body orthoso-
matic, dorsally not plicate. Intersegmental membrane present.
Ninth abdominal segment small, ventrally reduced. Cerci not
developed. Tenth abdominal segment extending farther back
than ninth, conical, elongate, a locomotory organ. Holopneustic;
spiracles annular; first spiracle plainly mesothoracic. Heau
porrect and exserted, or slightly invaginated. Clypeus large,
fused with frons. Labrum small, fleshy. Antenna two-
(three-) jointed, with large clavate second joint, very small or
indistinct apical joint. Mandible with flat, broad, molar part;
accessory condyle present; retinaculum hamate; apex tri- (or
quadri-) fid. Hypopharyngeal chitinizations strong, with paired,
ovate, large maxillular area carrying longitudinal series of long
setae. Ventral mouthparts retracted. Maxilla with cardo small,
transverse, in some forms distinctly bidivided ; stipes proximally
and internally attached to large maxillary articulating area; mala
extending from distal end of stipes, large, simple, falciform,
terminated by single uncus; externally to base of uncus with
small setose region, in some forms distinctly defined, being white
and membranous in contrast to otherwise brown and well chiti-
nized rest of mala. Maxillary palpus three-jointed. Labium
202 Zoologica: N. Y. Zoological Society [Tits
with broad ligula; without or with slightly indicated paraglossae;
two jointed labial palp. Mentum—probably fused with submen-
tum—large, oarrelshaped, at base attached to maxillary articu-
lating area, otherwise free. Gula large, hexagonal, transverse,
interposed between postmaxillary margins of epicranium, the
mental-submental area and the anterior prothoracic region.
Apart from the above mentioned, particularly close associa-
tion .with several genera of the old family Cucujidae, the
Silvaninae also exhibit a pronounced relationship to the genera
of the family Cryptophagidae, to the Monotomidae as represented
by the larva of Europs pallipennis Lec. and to the genus Languria,
which together with a few other genera, according to the larval
structures, may constitute a distinct family. These larvae have
all, like those of the Silvanidae and Cucujidae (restricted) a
normally chitinized mandible (compare the Lathridiidae) and a
simple and falciform maxillary mala; but they differ from the
Silvanid and Cucujid larvae in having bifore spiracles.
B. 2—The Genera of the Silvaninae Systematically Defined by
Larval Characters.
The Silvaninae are represented in the collections of the U. 5.
National Museum by the larvae of Carthartus advena Waltl,
Nausibius clavicornis Kugelann, Silvanus quadricollis Guérin,
Oryzaephilus surinamensis Linnaeus, Coccidotrophus socialis
Schwarz and Barber, and Eunausibius wheeleri Schwarz and
Barber.
The genera which these larvae represent can be separated as
follows:
1. Second antennal joint as long as head (Plate VIII,
fie; UD) cls ated late ach Cathartus Reiche
Second antennal joint half as long as head or
STRONGER» ss seer ecaee eoahew chin es bes Soees Oka kan eee Agee 2
2. Second antennal joint half as long as head; upper
ocellar group with four ocelli, lower group with
two ocelli, or individual ocelli of each group
COT AUTRE, az Goes oe atk fous soo Suk cava eoucieeree ee eee eee 3s
1921] Boving: Larvae and Pupae of Social Beetles 203
Second antennal joint one-third as long as head;
upper ocellar group with two ocelli, lower with
BER Ie cores cosivnavciovsnsdeneedonckeas ot ntottie se eer anas 5
3. Mandible apically quadrifid; individual ocelli in each
group well separated; body well chitinized, with
dark brown chitinous shields (Plate VIII, fig.
18), Nausibius Redt.
Mandible apically trifid; individual ocelli in each
group almost confluent; body thinly chitinized,
Waitin pale youowisi Shields....:...../:.c.tccssesssnosssssuaes 4
4. Maxillary palp with length of basal, second and apical
PRUNE CHiet NO ieee eae nee occ sneccesiconavenvons Silvanus Latr.
Maxillary palp with length of basal, second and apical
TOUTS BRN EG) oir ie i Oryzaephilus Gangl.
5. Dorsal shields of abdomen with one long seta on
each side; body very thinly chitinized, creamy
white (Plate VII, fig. 1),
Coccidotrophus Schw. and Barb.
Dorsal shields of abdomen with two long and a few
short setae on each side; body thinly chitinized,
light grey (Plate VII, fig. 3),
Eunausibius Grouvelle
C.—COcCIDOTROPHUS SOCIALIS AND KUNAUSIBIUS WHEELERI.
C 1.—General morphological description of the larvae.
(Plate VII, figs. 1-8, Plate VIII, figs. 9-11, 13, 16-17)
Orthosomatice with dorsal and ventral sides of body equaily
developed; thoracic segments not twice as long as seven anterior
abdominal segments; lateral sides of body nearly parallel, taper-
ing from eighth abdominal segment posteriorly. Legs ambula-
tory, of equal size, of medium length and strength; five-jointed,
with tarsus and claw fused. Thoracic and abdominal terga not
plicate, tergal areas forming shieldlike region. Segments
smooth, without gills, thorns or thick pubescence, setae on both
thoracic and abdominal segments. Ten abdominal segments
204 Zoologica: N. Y. Zoological Society | Ee
present; ninth small, ventrally reduced (9 Plate VII, figs. 1 and
2), without cerci; tenth (10 Plate VII, figs. 1 and 2) elongate
conical, developed as a locomotory organ with extrusible, mem-
branous anal lobes (1 Plate VIII, fig. 2).
Head exserted beyond, or slightly retracted into anterior
portion of prothorax; type of head porrect with frons horizontal
and mouthparts stretched forward; form of head subtriangular
with posterior part broader. Occipital foramen posterior, annu-
lar, dorsally limited by hind margin of epicranium, ventrally
by hind margin of gular plate (gu Plate VII, fig. 7). No collum.
Frons lyriform, one pair of oblong spots where frontal sutures
anteriorly begin to diverge (Plate VIII, fig. 9). Frons and
clypeus fused. Clypeus large, projecting to near tip of mandible,
trapezoidal with sides converging forward, chitinized. Labrum
short, soft and whitish, developed as a fleshy anterior margin of
clypeus (Plate VIII, fig. 9). Epicranical halves dorsally entirely
separated by frons; no median, epicranical suture; hypostoma
(the epicranial margin between ventral articulation of mandible
and attachment of posterior tip of cardo) (hy Plate VII, fig. 7)
longitudinal with transverse curvation below cardo; epicranial
margin longitudinally continued from tip of cardo to foramen
occipitale. Gula median, unpaired, hexagonal, without paragular
regions, chitinous, forming the ventral base of cranium between
mentum (or probably fusion of mentum and submentum), the
maxillary articulating area (7 Plate VII, fig. 7), the epicranium
and the presternal region of prothorax. Hypopharyngeal chitini-
zation (hpy. ch. Plate VIII, fig. 16) (= lingua, Folsom), at en-
trance to pharynx, strong, triangular, considerably wider than
long, with tuft of long sete at each end; between tufts surface —
covered with numerous small, irregularly arranged rugosities.
Coming from hypopharyngeal chitinization, creating firm support
for same, is the following system of five pairs of chitinous rods:
First pair (= lingual stalks, Folsom) (1 Plate VII, fig. 8, Plate
VIII, fig. 16) connecting hypopharyngeal chitinization with an-
terior corner of gular plate, perpendicular, parallel, imbedded in
- membrane between articulating area of maxilla and lateral side
of base of mentum (or probably submental part of fused men-
tum—submentum). Second pair of rods (=hypoharyngeal
1921] Boving: Larvae and Pupae of Social Beetles 205
bracons, Hopkins) (2 Plate VII, figs. 6, 8, Plate VIII, fig. 16)
connecting hypopharyngeal chitinization with that region of
epicranium which carries fossa for mandibular condyle; hori-
zontal, transverse, imbedded in buccal membrane between
mandible and dorsal side of maxillary stipes. Third pair of
rods (3 Plate VII, figs. 6, 8, Plate VIII, fig. 16) situated im-
mediately in front of and parallel with anterior margin of hypo-
pharyngeal chitinization ; horizontal, converging anteriorly, meet-
ing in middle line. Fourth pair (4 Plate VII, figs. 6, 8, Plate VIII,
fig. 16) supporting and externally limiting the maxillular areas,
vanishing distally near glossa (= buccal surface of ligula) (glos.
Fig. 16) ; horizontal, longitudinal and parallel. Fifth pair (5
Plate VII, figs. 6, 8, Plate VIII, fig. 16) forming a posterior
prolongation of fourth pair of rods, imbedded in sides of
oesophagus; horizontal, longitudinal and parallel; proximally,
near hypopharyngeal chitinization, with fossa for accessory
mandibular condyle (fos. a. c. m. Plate VIII, fig. 16), distally
gradually vanishing.* Maxillular areas (sensu H. T. Hansen, =
superlingua Folsom) (mal Plate VIII, fig. 16) fleshy, ovate, an-
teriorly fused together, posteriorly well separated, limited ex-
ternally by fourth and internally by third pair of rods; along
third pair with a series of well developed, closely set sete. Glossa
(= buccal surface of ligula-lingua, Schiddte) (glos. Plate VIII,
fig. 16) unpaired, fleshy, in front of maxillular area, laterally sup-
ported and limited by a pair of longitudinal, horizontal, chitinous
rods (stp. li. Plate VIII, fig. 16); similar to the fourth hypo-
pharyngeal pair. Epipharynx (Plate VIII, fig. 11) consisting of
anterior and posterior part. Anterior part a direct ventral con-
tinuation of labrum; soft, membranous, medially on the hind
margin with a small triangular asperity, which fits in between the
scissorial parts of the closed mandibles; well developed tendons
(t Plate VIII, fig. 11) extending from each back-corner; some
long sete. Posterior part of epipharynx slightly chitinized,
with several transverse, long, curved, parallel wrinkles. Ten-
torium an internal cranial structure, differing in that re-
spect from the two mentioned systems of chitinizations in
*The pair of rods is not present which in many other Coleopterous
larvae extends from proximal ends of fifth pair, along sides of epipharynx,
towards end of large movable labrum.
206 Zoologica: N. Y. Zoological Society iteT
the buccal membrane above labium and above the mental-
submental area; consisting of a median broad and _ short,
slightly chitinized, transverse tentorial bridge (ten. b. Plate
VII, fig. 7), situated a short distance above gular plate
and of two pairs of tentorial arms; the first pair of arms
lateral (t. l. a. Plate VII, fig. 7), between bridge and tentorial
pits (tp. Plate VII, fig. 7) where gular plate and epicranium*
meet; the second pair anterior (t. a. a. Plate VII, fig. 7), reach-
ing from tentorial bridge toward the dorsal side of cranium,
their external ends distally indistinct. Ocelli (Plate VIII, fig. 10)
six, placed in two groups, a lower behind mandible and an upper
straight behind antenna and right above first group. Antenna
with large basal membrane, two- (three-) jointed; basal joint
(1 Plate VII, fig. 5) short, second joint (2 Plate VII, fig. 5) long,
claviform, with tip (3 Plate VII, fig. 5) more or less distinctly
separated as a small additional joint. Mandible (Plate VII, figs.
4 and 5) with molar part, retinaculum and accessory ventral
condyle; half as long as head; apical part (p. sc. Plate VII, fig. 5)
distinctly constricted, incurved, vertically compressed, fornicate,
cleft with some toothlike ends; rest of mandible horizontally flat,
subtriangular with wide base; retinaculum present (7 Plate VII,
fig. 5), hamate, short, pointed, with enlarged socket; molar part
broad (m. Plate VII, fig. 5, Plate VIII, fig. 17), compressed, with
dorsal surface smooth, ventral surface roughened by numerous
granules, these being anteriorly rather large and irregularly dis-
tributed, posteriorly minute and densely placed in longitudinal
rows; basal molar fringed with a series of anomalous, chitinous,
stiff, filamentous processes (fil. pr. Plate VIII, fig. 17), which
gradually diminish in length from inner corner of basis toward
accessory condyle; accessory condyle (acces. Plate VIII, fig. 17),
as mentioned above, fitting into groove (fos. a. c. m. Plate VIII,
fig. 16) in fifth hypopharyngeal rod.* Ventral mouthparts
* This articulation between the mandibles and the whole system: of
amalgamated chitinous hypopharyngeal and epipharyngeal structures indi-
cates that the side to side movements of the mandibles caused by extensor-
flexor— (= abductor-adductor)—muscles, always must be coincident with
the forward-backward movements of the following closely united structures:
the anterior portion of oesophagus, the hypopharynx with other buccal
structures, and the ventral mouthparts, and also coincident, when a free, well-
developed labrum and movable clypeus are present, with their up and down
nodding motions. All forward and upward directed movements are caused
by blood pressure combined with special arrangements of the articulations;
all backward and downward directed movements by retractor muscles.
1921] Boving: Larvae and Pupae of Social Beetles 207
retracted with tip of cardo articulating at considerable distance
behind the ventral condyle of the mandible (vc. Plate VII, fig. 7) ;
hypostomal curvation adjacent to both stipes and cardo. Max-
illary articulating area large, membranous, placed between
stipes, maxillae, cardo, gular plate and posterior part of the
mental-submental region. Cardo maxillaris approximately
transverse, bidivided, much narrower than long, tip articulating
with hypostoma at distinct distance from occipital foramen.
Stipes maxillaris connected with the articulating area along the
proximal two-thirds of its inner margin; distal part free. Mala
(=maxillary lobe) (lac Plate VIII, fig. 16) simple, projecting as.
a direct anterior continuation of stipes, apically attenuate, ter-
minating with a single, well developed uncus (wu Plate VIII, fig.
16) ; external apical part of mala (gal Plate VIII, fig. 16) soft-
skinned with some strong, straight setae behind uncus:* inner
margin of mala set with strong setae. Palpiger maxillaris (plg.
mz. Plate VIII, fig. 16) well developed, subtriangular. Maxil- —
lary palp well developed, three-jointed. Submentum probably
fused with mentum, forming together a barrelshaped, free unit
(m Plate VII, fig. 7). Labium (proper) posteriorly limited by
a chitinous bow, extending between the ends of those rods which
laterally support and limit the glossa; palpiger labii not distinct-
ly chitinized. Ligula (lig Plate VII, fig. 7) broad.** Labial palp
short, two-jointed. Between head and prothorax there is a
well developed cervical membrane, capable of being invaginated
into and protruded from the anterior part of prothorax.
Thoracic segments similar in size and development. Prothor-
acic presterna (prst 1 Plate VII, fig. 7) large broad, oval,
* Possibly corresponding to galea, while chitinized rest of mala is lacinia
(lac Plate VIII, fig. 16).
** The term “ligula” (= glossa, Folsom) is here applied to the median,
terminal labial lobe, which is composed of the fused right and left labial
malae. For descriptive purposes it appears practical to use special terms
for the ventral and buccal surfaces of this lobe; the term “ligula” is here
applied only to the ventral surface (lig Plate VII, fig. 7), while the buccal
surface is mentioned as “glossa” (glos. Plate VIII, fig. 16). The structure
which Schiddte calls “ligula” is not identical with the entire labial lobe in
question, as he designates as the ligula only a special, jointed, terminal part
of the lobe; the rest, or, when no jointed terminal part is developed, the
entire lobe Schiddte calls “lingua,” and he applies this term both to the
ventral and buccal surfaces.
208 Zoologica: N. Y. Zoological Society [Tike
adjacent to a medium, simple, elongate, suboval eusternal sclerite
(eu Plate VII, fig. 7); furcal pits (fur Plate VII, fig. 7) near
inside of legs. A well developed triangular poststernellum (pstl
Plate VII, fig. 7) present. Between pro- and mesothorax and
between meso- and metathorax a well developed intersternal
ring consisting of the poststernellar area (pstl. Plate VII, fig.
7) of the anterior and the presternellar areas (prst Plate VII,
fig. 7) of the posterior or two connected segments. Presternal
areas of meso- and methathorax, subtriangular, dorsally only
slightly separated from the spiracle-bearing preépipleural areas
of the segments. Hypopleural areas, above coxae, separated
by a distinct, short, perpendicular, chitinous line extending from
coxal hinge into an anterior subdivision, prehypopleurum (=epis-
ternum authorum), and a posterior one, posthypopleurum
(=epimeron authorum). Legs inserted widely apart (leg Plate
VII, fig. 7). Abdominal segments, from first to eighth, differing
only slightly in size and shape, about as wide as thoracic segments,
somewhat shorter. Intersegmental membranes, indistinctly
developed dorsally and laterally, more distinctly ventrally. Each
abdominal segment with tergal areas fused, with poorly defined,
low epipleural area without lobe or other subdivisions, with
hypopleurum large, somewhat bulging, dorsally clearly limited
by a ventro-lateral suture, ventrally less sharply defined; sternal
areas more or less fused. Ninth and tenth abdominal segments
mentioned above (p. 204). Spiracles annular (Plate VII, fig.
2, Plate VIII, fig. 18), not very conspicuous; mesothoracic
twice as large as abdominal; metathoracic rudimentary; both
meso- and metathoracic spiracles—as already mentioned—
located in preépipleurum; abdominal spiracles all lateral and
of same size, located in the tergal regions medially or poster-
iorly near the rather indistinct dorsolateral suture. One pair
of small chitinous spots present dorsally on the thoracic seg-
ments; one pair dorsally and one pair ventrally on most of
the abdominal segments. Rounded, soft organs, shaped like
cloverheads attach internally to these spots; function unknown.
1921] Boving: Larvae and, Pupae of Social Beetles 209
C. 2.—Specific Description of Larvae of Coccidotrophus socialis
and Eunausibius wheeleri.
Coccidotrophus socialis Schwarz and Barber
(Described from specimens preserved in U. S. National
Museum.)
(Plate VII, figs. 1 and 2.)
Mature Larva:
Length about 4 mm. Width of prothorax about 1 mm.
Whitish, very thinly chitinized. Head with width to length as
1:1144. Length of head and thorax, together in proportion to
abdomen as 3:5. Ocelli four in a lower, square group, two in
an upper group. Frons with short setae sparsely scattered
over the surface; the paired spots at frontal sutures yellow.
Clypeal region anteriorly with transverse series of four long
setae, laterally with one long seta on each side, several short setae
scattered over entire surface. Labrum, with several very short
setae. Another part of epipharynx with four long setae along
the front margin and one long seta on each side near the attach-
ment of the epipharyngeal tendon; posterior part of epipharynx
without setae. Hypostoma about half as long as lateral outline
of epicranium. Epicranial postmaxillary margin about half as
long as cardo. Epicranium, with a few long and some short
setae scattered over the surface. Antenna about half as long
as head from front margin or labrum to occipital foramen;
basal antennal joint about half as long as second joint (soft-
skinned apical part excluded), subcylindrical, twice as long as
wide; second antennal joint clubshaped, with end twice as wide
as base, almost one-third as long as length of head; tip of
antenna membranous, whitish, low, rounded, not distinctly set
' off from second joint; no supplementary appendix; a ring of
densely set spinules present on distal end of basal joint, numer-
ous thin and medium long or short setae irregularly scattered
over the whole surface of second joint. Mandible with apical
part trifid, several setae of different sizes on exterior lateral
mandibular region. Each maxillular area ovate, membranous;
individual setae belonging to series along inner margin of
medium size, somewhat curved, bending outward. Maxilla, with
end of cardo articulating at a distance from occipital foramen,
210 Zoologica: N. Y. Zoological Society [Tere
somewhat more than half the length of cardo; free distal part of
stipes dorsally with one long seta; mala with inner margin set
with two parallel series of strong, somewhat curved setae; pal-
piger maxillaris with a few minute setae; maxillary palp well
developed, surpassing tip of mala by one-half the length of thc
apical joint, with basal joint cylindrical, somewhat wider than
long; second joint cylindrical, as wide as long; apical joint
slightly conical, almost as long as the two other joints together,
half as wide; front margin of two posterior joints with minute
spines in a ring; apical joint with one long seta and terminally
with several sensory papillae. Submentum with two pairs of
setae. lLigula anteriorly rounded, with several rather short
setae; no paraglossae. Labial palp about as long and wide as the
apical joint of the maxillary palp; basal joint about two-thirds
of the entire length of labial palp; apical joint anteriorly
obtusely rounded and provided with many minute sensorial pro-
jections. Thoracic segments almost equally developed, each
about as wide and long as head; weak line present between
furcal pits, separating an anterior eusternal region from a
posterior sternellar region; poststernellar areas of pro- and
mesothorax well developed, marked with a median chitinous
spot. Prothorax with eusternal sclerite as wide and somewhat
longer than each prothoracic presternum. Thoracic tergal
shields with one seta on each side, the prothoracic rather short;
meso- and metathorax with one yellow spot anteriorly to the
seta. Legs with coxa about as long as clypeal region, slightly
longer than thick; trochanter half as long as coxa; femur and
tibia of equal size, slender, each slightly longer than second
antennal joint, about four times as long as thick; clawshaped
tarsus not quite as long as basal antennal joint. Eight anterior
abdominal segments with tergal shields carrying along the hind
margin a series of seven to eight setae on each side of middle
line; one seta, externally placed, very long, the others very
small; scattered all over the whole surface of shield, numerous,
extremely minute setae. In front of long seta with yellow,
chitinous spot. Spiracles posteriorly placed, below the shields.
Epipleurum narrow, without seta. Hypopleurum with large
median lobe and one long seta; sternal areas on each side with
two long setae and one yellow chitinous spot. Ninth abdominal
segment only represented by small tergal part, about as large
1921] Boving: Larvae and Pupae of Social Beetles 211
as hypopleurum of eighth abdominal segment, with one small
seta and some very minute scattered over whole surface. Tenth
abdominal segment as long as one of the well developed abdomi-
nal segments, conical, one and one-half times as long as wide; a
ring of small setae right above the truncate end; also with very
minute setae scattered over whole surface. Mesothoracic
spiracle about as large as base of claw, situated on top of small
softskinned conical tube; abdominal spiracles half as large and
not on tubes.
Newly hatched larva:
Length 1 mm. Width of prothorax 0.25 mm. Length of
head and thorax together about as long as abdomen. One very
long seta on each side of hind margin of tergum; setae corre-
sponding to the small setae along hindmargin comparativeiy
longer than those of mature larva; no setae corresponding to
minute setae on the whole tergal surface.
Eunausibius wheeleri Schwarz and Barber.
(Described from specimens preserved in U. S. National
Museum.)
Mature Larva:
Length, about 3.5 mm. Width of prothorax, about 0.75
mm. Length of head and thorax combined, in proportion to
length of abdomen, about as 3:5. Shields thinly chitinized, light
grey, shiny. Both thoracic and abdominal segments along hina-
margin with a series of six to seven setae on each side; two
sets very long, the others short; rest of tergal shield smooth
with a few very minute setae. Abdominal hypopleurum with
one long and one short seta. Otherwise like Coccidotrophus
socialis, to which larva it is very closely related.
C 3—General and specific description of the Pupae of
Coccidotrophus socialis and Eunausibius wheeleri.
(Plate IX, figs. 19-21, 23)
General Description.
Body somewhat depressed ; about five times as long as width
of prothorax. Head large and wide; not to be seen from above;
212 Zoologica: N. Y. Zoological Society (Mi<7
with four or five very minute spinules on dorsal surface. Pro-
thorax flat, subrectangular; with length to width at 1.25:1.00;
anterior third slightly broader than rest; anterior corner some-
what rounded; posterior corner rectangular; with one setiferous
protuberance at each anterior corner and a few along the sides,
otherwise entirely smooth. Meso- and metathorax smooth.
Abdomen with third segment the widest and as wide as pro-
thorax; first and second abdominal segments slightly narrower;
posterior segments gradually decreasing to the seventh, which
anteriorly is half as wide as the third and posteriorly only two-
thirds as wide as anteriorly; eighth to ninth abdominal seg-
ments small, forming together a rounded almost semicircular
termination of body. Second to seventh abdominal segments
laterally with small protuberances, without setae; ninth abdomi-
nal segment dorsally terminating with two small, slender,
cylindrical, divergent cerci. Spiracles annuliform, located on
meso- and metathorax and on the first to eighth abdominal seg-
ments, where lateral protuberances develop spiracles placed
immediately above and in front of these. Pouch covering
elytron extending to posterior margin of fourth abdominal
segment, smooth, with four well marked longitudinal ribs.
Tarsal cover of hind legs ventrally extending to middle of
fourth abdominal segment; the entire leg-pouch smooth.
Antennal pouch short, clubshaped, directed backward and out-
ward; ends of the last three or four joints marked by a ring of
small projections. :
Specific Characterization.
Coccidotrophus socialis Schwarz and Barber.
(Pupa described from specimens preserved in the U. S. National
Museum).
(Plate IX, figs. 19-21)
Length, about 4 mm. White. Prothorax with anterior
iateral protuberance small, but distinct, three posterior lateral
protuberances much reduced, not to be seen with naked eye or
ordinary lens magnification.
1921] Boving: Larvae and Pupae of Social Beetles 213
Eunausibius wheeleri Schwarz and Barber
(Pupa described from specimens preserved in the U. S. National
Museum)
(Plate IX, fig. 23)
Length, 3-4 mm. Grey. Prothorax with five lateral pro-
tuberances well developed, to be seen with naked eye or ordinary
lens. (On specimen figured one protuberance was—abnor-
mally—not developed on left side.)
D.— Bibliographical Notes.
A careful list of the descriptions and figures of the larvae
of the family Cucujidae (awth.), including reference to P. de
Peyerimhoft’s key for the determination of the larvae of the
Cucujid genera, is given by F. H. Gravely (in Records of the
Indian Museum, Calcutta, vol. II, 1915, pp. 353-358).
To this list might be added the part dealing with this
family in L. Ganglbauer: Die Kafer von Mitteleuropa, vol. III,
part 2, 1899. Since the list of Gravely was published an impor-
tant work has appeared by U. Saalas: Die Fichtenkafer
Finnlands (in Annales Academiae Scientiarum Fennicae Ser.
A, vol. VIII, 1917 pp. 508-528, figs. 119-130). In Saalas’ book are
described and splendidly figured with habitus and detail draw-
ings the following larvae (and pupae): Pediacus fuscus Er.,
Laemophloeus abietis Wank, Laemophloeus alternans Er.
Finally I may mention that the larva described and figured
by C. V. Gernet as “Cucujiden-Larve” (Dendrophagus crenatus
Payk) is in my opinion a Staphylinid larva of the group
Aleocharini. The figure shows the combination of a mandible
without mola and jointed cerci, movable at base, which is char-
acteristic of the Staphylinids; and also a large movable labrum,
a single ocellus and broad ligula, which characters define the
Aleocharini.
The newest taxonomic arrangement of the Cucujidae is
presented by Charles W. Leng in: “Catalogue of the Coleoptera
of America, North of Mexico,’ Mount Vernon, N. Y., John D.
Sherman, 1920. The arrangement of the larvae given in the
present paper does not agree so well with Mr. Leng’s list as
with the system presented in 1899 in L. Ganglbauer’s “Kafer von
Mitteleuropa.”
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
aor Wh
Plates Vii;
(Figures drawn by A. G. Boving).
. Coccidotrophus socialis. Dorsal view of larva.
. Coccidotrophus socialis. Side view of larva.
. Hunausibius wheelert. Dorsal view of larva.
. Coccidotrophus socialis. Right mandible from above.
. Coccidotrophus socialis. Right mandible from below.
ac, accessory condyle; f, chitinous appendices from
base of mola; fla, tendon of flexor mandibule; m,
mola; psc, pars scissoria; 7, retinaculum; v, ventral
condyle of mandible; 1-2-3, first-second-rudimentary
third antennal joints.
. Coccidotrophus socialis. Hypopharyngeal rods in
buccal membrane. Dorsal surface of ventral mouth-
parts. 1-2-3-4-5, see explanation of Fig. 16, Plate
Vit.
. Coccidotrophus socialis. Head, Prothorax and Meso-
thorax from below. a1-a2, first and second antennal
joints; b, bracon (= second hypopharyngeal rod) ;
bm, basal membrane of antenna; est, eusternum; ew,
eusternal plate of prothorax; fur, pit indicating
attachment of furca; gu, gula; leg, basis of leg; lig,
ligula; m, mentum and submentum fused; prst, 1, 2
and 3, presternum; pstl 1 and 2, poststernellum of
first and second thoracic segments; 7, sp., rudimen-
tary spiracle of metathorax; sp, spiracle of meso-
thorax, stl, sternellum; taa, anterior arm of tento-
rium; tenb, bridge of tentorium; tla, lateral arm of
tentorium; tp, tentorial pit, longitudinal groove,
indicating attachment of lateral arm of tentorium;
stl, sternellum; vc, mandibular fossa of epicranium.
Fig. 8. Coccidotrophus socialis. Ventral surface of ventral
mouthparts. 41-9-3-4-5, five hypopharyngeal rods,
see explanation of Fig. 16, Plate VIII.
f wii
Ry,
Plate VII
Plate VIII
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
10.
bt.
12.
13.
14.
15.
16.
Plate VIII.
(Figures drawn by A. G. Boving).
. Coccidotrophus socialis. Anterior part of head from
above, showing frons with chitinous spots, clypeus
fused with front, membranous short labrum, basis
of antenna. ;
Coccidotrophus socialis. Anterior part of head from
right side, showing basis of antenna basis of man-
dible, upper group of two ocelli, lower group of
four ocelli.
Coccidotrophus socialis. Epipharynx.
Cathartus advena. Head from left side.
Coccidotrophus socialis. Spiracle, highly magnified.
Cathartus advena. Anterior part of left mandible and
maxilla from below.
Cathartus advena. Anterior part of left mandible
from above.
Coccidotrophus socialis. Maxilla, hypopharynx, max-
illular area and glossa (= dorsal surface of ligula)
facing buccal cavity; bracon, second chitinous rod
from hypopharyngeal chitinization; fos. ac. m, fossa
for the accessory mandibular condyle in base of
hypopharyngeal chitinization (acces, fig. 17, Plate
VIII) ; gal, possibly rudimentary galea; glos, glossa;
hyp. ch, hypopharyngeal chitinization; lac, lacinia;
meal, maxillular area; oes, oesophagus; plg. max, pal-
piger maxille; w, uncus; setae, setae along chitinous
rod number three; stipes, dorsal surface of stipes
maxille; stpli, chitinous rod between glossa and
palpiger labii; 1-2-3-4-5, chitinous rods from hypo-
pharyngeal chitinization; J-JJ-IJJI, basal, second,
apical joints of palpus maxillaris.
Fig. 17. Coccidotrophus socialis. Posterior part of left man-
dible from below; acces, accessory condyle fitting
into fossa in base of hypopharyngeal chitinization
(fos. ac. m, fig. 16, Plate VIII) ; fil, pr, stiff chitinous
filaments; fl, tendon of flexor mandibule; mola, gran-
ular ventral surface of the molar structure; 7, tendon
of retractor mandibule; vc, ventral condyle of man-
dible fitting into fossa of epicranium (ve, fig. 7, Plate
VII).
Fig. 18. Nausibius repanda. Head and anterior part of thorax.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Plate IX.
(Figures drawn by A. G. Boving).
. Coccidotrophus socialis. Pupa; left side.
. Coccidotrophus socialis. Pupa; dorsal side.
. Coccidotrophus socialis. Pupa; ventral side.
. Prostomis mandibularis. (Denmark, Europe), Maxilla;
max. art. a., maxillary articulating area.
. Hunausibius wheeleri. Pupa; left side. (Specimen
slightly abnormal, having only four lateral projec-
tions on left side).
. Prostomis mandibularis. Larva from above.
. Prostomis mandibularis. Head from below.
. Prostomis mandibularis. Terminal part of abdomen
from below.
. Dryocora howitti (New Zealand). Maxilla.
. Dryocora howitti. Mandible.
. Dryocora howitti. Ninth abdominal segment from
above.
. Dryocora howitti. Ninth abdominal segment, left side
. Dryocora howitti. Head; dorsal side.
. Dryocora howitti. Larva from above.
. Dryocora howitti.. Head; ventral side.
Plate IX
4|
fi 2
44
42 “3
LJ dorm Ft bratghead
Plate X
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44,
Plate X.
(Figures 34-38 drawn by A. G. Boving).
(Figures 39-44 drawn by F. C. Craighead).
Telephanus pallidulus (Rio Piedras, Porto Rica).
Head; left side.
Telephanus pallidulus. End of abdomen; left side.
Telephonus pallidulus. Left mandible; ventral surface.
Telephanus pallidulus. Mature larva; dorsal side.
Telephanus pallidulus. Maxille, glossa, maxillular
area, hypopharyngeal chitinization, oesophagus;
facing buccal cavity.
Scalidia linearis Lec. Head; dorsal side. (Note: No
distinct clypeus, large movable labrum).
Scalidia linearis. Leg.
Scalidia linearis. Right mandible; dorsal surface.
(Note: No molar structure).
Scalidia linearis. Head; ventral surface. (Note:
Protracted mouthparts; large region formed by
fusion of cardines, mentum and submentum; also
large region formed by fusian of epicranial halves
and gula).
Scalidia linearis. Spiracle; annuliforme.
Scalidia linearis. Left side of the mature, parasitic
larva.
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
Now YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 8
(Tropical Research Station Contribution Number 102)
A NEW DIADIPLOSIS
By ‘E. P. FELT
erat SHED. BY THE 50 C.D EE
foo 6~ ZOOLOGICAL PARK, NEW: YORK
DECEMBER 24, 1921
Volume III, Number 8
A NEW DIADIPLOSIS
By E. P. FELT
The midge described below was received from Prot.
Wheeler accompanied by the statement that the larvae were
devouring mealy-bugs (Pseudococcus bromeliae Bouché) in a
cavity of a peculiar myrmecophilous tree, Tachigalia, in British
Guiana. The coccids and the fly larve live in a cavity of the
leaf petiole.
This species approaches closely that West Indian
Diadiplosis cocci Felt, which was reared from larve preying
upon the eggs of Saissetia nigra Nietn., a scale insect fre-
quently abundant upon the stems of Sea Island cotton. The
female of the West Indian species has a distinct knob upon the
terminal antennal segment and the lobes of the oviposter are
somewhat narrower, two characters which serve to distinguish
the species, though it is frequently very difficult to find charac-
teristic structures in female gall midges.
Diadiplosis pseudococci sp. nov.
Female. Length 1.25 mm. Antennae extending to the base
of the abdomen, sparsely haired, yellowish brown, of fourteen
segments, the fifth with a stem about one-fourth the length of
the cylindrical basal enlargement, which latter has a length
about two and one-half times its diameter and is slightly con-
stricted near the basal third; low, broad circumfila occur on the
enlargement at the basal third and apically; basally there is a
thick whorl of rather long, stout setae and on the ventral face
near the distal third a rather thick. group of long, rathei
strongly curved, slender setae; terminal segment somewhat
produced, with a length nearly three times its diameter and
tapering gradually to a sub-acute apex (no knob as in D. cocci)
Palpi: the first segment short, the second with a length about
twice its diameter, and the third a little longer than the second.
Mesonotum dark reddish brown; scutellum and postscutellum:
yellowish orange; abdomen “orange red”; the sclerites yellowish
226 Zoologica: N. Y. Zoological Society [III;8
brown; wings hyaline; halteres pale yellowish, fuscous sub-
apically; coxae and femora basally pale yellowish, the remainder
of the legs dark straw; claws moderately stout, strongly curved,
unidentate; the pulvilli about one-half the length of the claws;
ovipositor short, the lobes broadly oval and clothed sparsely
with rather coarse hairs.
Pupa. Length 1.45 mm. Rather stout, yellowish orange,
the antennal cases extending to the middle of the thorax; the
wing cases to the third abdominal segment and the leg cases
to the fourth and fifth abdominal segments; posterior extremely
broadly rounded, with a median furrow, the abdominal seg-
ments dorsally each with a rather broad, transverse band of
scattering weak spines.
Larva. Length 1.5 mm. Moderately stout, yellowish
orange (probably reddish orange in life); head short, mostly
retracted; the breastbone weakly developed, bilobed anteriorly,
the lobes rather broad, broadly rounded and roundly excavated
at the internal basal angles; the shaft weakly and irregularly
developed; the posterior extremity obsolescent; near the middle
of each segment there is a transverse row of moderately long,
tapering, setose processes; skin rather finely shagreened, pos-
terior extremely broadly rounded.
Type Cecid. A3176, New York State Museum.
The larve, as noted by Prof. Wheeler, live under a web in
small groups. The pupae occur intermixed and can be distin-
guished from larvae only with difficulty. The midges doubtless
emerge directly from these shelters.
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 9
(Tropical Research Station Contribution Number 103)
A NEW BLEPYRUS
By C. T. BRUES
_—ee Ltt SHhED--BY THE 5:0 C15; ae
mae ZOOLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
Volume III, Number 9
A NEW BLEPYRUS
By C. T. BRUES
Blepyrus tachigalice sp. nov.
Female. Length 1.5 mm. Head metallic green, thorax
black with an eneous cast, abdomen bronzed black; antennz
vellow and black, the scape and funicle yellow, darker on the
scape basally, the pedicel and club black; coxe black, femora
black with brown tips, tibize brownish yellow with a black
streak at base, tarsi pale, with the last joint dark. Wings
deeply infuscated basally, hyaline on apical half. Front one-
fourth the width of the head at middle, broader above and
below, with large punctures forming about four vertical rows;
malar furrow shallow and not very distinct; cheeks finely, ver-
tically aciculated, eyes with short pubescence. Antenne inserted
close to the mouth, the scape reaching barely halfway to the
vertex, excavated and distinctly, although not broadly, produced
below; pedicel triangular, longer than thick and half as long
as the funicle; funicle joints about equal in length, the apical
joint more than twice as broad as the basal one; club enlarged,
flattened and obliquely truncate at apex, as long as the pedicel
and funicle together. Thorax, including scutellum, highly
convex when seen in profile, the mesonotum nearly as long as
the scutellum, the surface of both shining, minutely punctulate
and finely hairy; axillary suture apparently obsolete; scutellum
very long, the side margins straight till near the apex, which
is therefore rather sharply rounded. Abdomen very short,
concave above, with long, scattered hairs at the sides. Legs
stout, the spur of middle tibia strong, as long as the first tarsai
‘joint, the tarsal spinules strong on all four basal joints. Mar-
ginal vein short, not over twice as long as thick; stigmal vein
long, postmarginal slightly shorter, their angle of separation
about 100 deg.; wing cilia short, those on the costa longer.
230 Zoologica: N. Y. Zoological Society [III;9
Type and paratype bred from Pseudococcus bromelix
Bouché, occurring in cavities in the petioles of Tachigalia sp.
at Kartabo, British Guiana.
This species agrees well with Howard’s original generic
diagnosis’ except that the antennal scape is slightly enlarged
below as in certain species of Bothriothorax (e. g., B. minor
Silvestri). The color of the body, appendages and wings will
distinguish the present form.
‘Proc. U. S. Nat. Mus., Vol. 21, p. 233 (1898).
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE '
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 10
(Tropical Research Station Contribution Number 104)
TWO TACHIGALIA MEMBRACIDS
By HERBERT OSBORN
esr lisHED BY THE SOC Fits
THE ZOOLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
ia is
ti Oe AY
Bo
Volume IIIT, Number 10
TWO TACHIGALIA MEMBRACIDS
By HERBERT OSBORN
Endoastus (?) productus sp. nov.
(Fig. 17.)
Head produced before the eyes, about as long as width
between the eyes, distinctly furrowed above and below and bifid
at tip, ocelli close to front margin of eye, antennze below the
eye in distinct sockets with raised margin; pronotum strongly
convex, sloping to head, posterior angles subacute; scutellum
triangular slightly longer than basal width; elytra narrow,
scarcely reaching end of abdomen; legs short cylindric, all of
nearly equal length.
The color is uniformly dark brown except tibiz, which are
lighter, the surface of body and elytra densely and minutely
punctured.
Female genital segment three times as long as preceding;
the ovipositor extruded; male subgenital plate narrow, upcurved.
Length of female 5.5 mm., of male 4 mm.
This species has a more produced head than E. caviceps
Fowl., but there seems to be no good structural character to
warrant the formation of a distinct genus for it, and at least
until the study of related species shows such distinction 1t
seems proper to place it here.
Superficially it has a strong resemblance to the Cicadellids
but in structural characters is obviously Membracid.
Eight specimens, seven females, one male. Kartabo, Bartica
District, British Guiana, 1920, feeding on the terminal shoots of
young Tachigalia trees.
234 Zoologica: N. Y. Zoological Society [Tits 7e
FIG. 17. ENDOASTUS (?) PRODUCTUS SP. NOV.
FEMALE.
a, dorsal; b, lateral aspect.
Microcentrus (?) sp. Nymph.
A single nymph agreeing closely in structural characters
with nymphs of Microcentrus sps. of Eastern U. S.
Body flattened, slightly convex above, head short, the
anterior border with broad, fringed plates each side; abdominal
segments 2-6 with flattened, fringed, plate-like expansions and
the terminal segment broadly fringed. Length 6.5 mm., widtn
of thorax 3 mm.
Feeding on the terminal shoots of young Tachigalia trees
Kartabo, B. G., July, 1920.
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 11
(Tropical Research Station Contribution Number 105)
A NEW ENTOMOBRYA
By J. W. FOLSOM
Mir ilisHED-BY THE SOCIETY
mene ZOOLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
Volume III, Number 11
A NEW ENTOMOBRYA
By J. W. FoLsom
Entomobrya wheeleri sp. nov.
(Fig. 18)
Head and body mottled with blue pigment, with no definite
color pattern (Fig. 18a). Sternum white. Antenne blue; first
three segments darker apically. Coxe slightly pigmented, also
femora distally and tibiotarsi proximally. Furcula unpig-
mented. Eyes 8-8, on black patches, the two inner proxima!
eyes of each side being smaller than the others. Antenne one
and three-fifths times as long as the head, with segments in
relative lengths about as 9:22:19:30; basal ring large, resem-
bling a segment. Mesonotum not projecting anteriorly. Fourth
abdominal segment almost five times as long as the third.
Unguis (Fig. 180) with a pair of lateral teeth and with two
inner teeth, the proximal tooth being doubled. Unguiculus
three-fifths as long as the unguis. Furcula attaining the ventral
tube. Manubrium and dentes subequal in length. Dentes
crenulate dorsally. Mucrones (Fig. 18c) subequally bidentate,
the basal spine being absent; two fringed setz projecting from
each dens extend almost to the end of each mucro. Corpus of
tenaculum with a single curving ventral seta. General clothing
of dense short setz. Clavate, ringed setz occur on the anterior
part of the head, on the last abdominal segment, and on the dor-
sal region of the manubrium. Pointed, fringed sete are present
on the last two abdominal segments and on the furcula dorsally
and ventrally. A few long, stout, fringed sete occur on the
antenne, two or three on each coxa, a few on femur and
tibiotarsus, and several on the anterior surface of the ventral
tube. Length 1 mm.
The preceding description applies only to the largest speci-
men as regards proportions and pigmentation, for these
characters vary according to the age of the individual, as usuai.
Thus in a specimen 0.39 mm. in length, there is scarcely any
blue pigment, the fourth antennal segment being, however,
-
238 Zoologica: N. Y. Zoological Society (Teas
FIG. 18. ENTOMOBRYA WHEELERI SP. NOV.
a, lateral view, X 41.5; b, left hind foot, X 386; c, left mucro, X 386.
faintly tinged apically with blue; the eyes are pigmented sepa-
rately instead of collectively; the antennze are subequal to the
head in length, with segments short and stout, in relative lengths -
as 4:5:5:11; while the fourth abdominal segment is only two
and two-thirds times as long as the third. In an individual 0.6
mm. in length, the ratio between the third and fourth abdominal
segments is as 1:4.
Described from four cotypes, which have been deposited
in the Museum of Comparative Zoology, Cambridge, Mass.
I take pleasure in naming this new collembolan after
Professor W. M. Wheeler, who found it living with colonies
of a peculiar social beetle, Coccidotrophus socialis Schwarz and
Barber, at Kartabo, in British Guiana.
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| 'THE PRESERVATION OF 01
“FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
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Js VOLUME III. NUMBER 12
en : (Tropical Research Station Contribution Number 106)
MONKEY
By ApotpH H. ScHULTZ
rch Associate, Department of Embryology, Carnegie. Institution
Bey of asain
LISHED. BY ‘THE SOCIETY
PAO OLOGICAL PARK, NEW YORK
DECEMBER 24, 1921
New York Zonlonircal Socivty
General Office: 111 Broadway, New York City
Officers
President, HENRY FAIRFIELD OSBORN;
Vice-Presidents, MADISON GRANT and FRANK K. STurRGIs;
Secretary, Chairman, Exec. Committee, MADISON GRANT;
Treasurer, PERCY R. PYNE.
Board of Managers
Glass of 1922 >
Prercy R. PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DODGE,
AG: LEDYARD BLAIR, EMERSON McMILLIn, ANTHONY R.
KUSER, WATSON B. DICKERMAN, Mortimer L.
SCHIFF, FREDERIC C. "WALCOTT, BEEKMAN
WINTHROP, GEORGE C. CLARK,
W.-REDMOND CROSS.
@lass of 1923
HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F.
DIETERICH, GEORGE F. BAKER, WM. PIERSON HAMILTON,
ROBERT S. BREWSTER, EDWARD S. HARKNESS,
WILLIAM B. OsGooD FIELD, A. BARTON
HEPBURN, WILLIAM WOODWARD,
EDWIN THORNE, PERCY A.
ROCKEFELLER.
Glass nf 1924 :
MADISON GRANT, WILLIAM WHITE NILES, FRANK K. STURGIS, ,
GEORGE J. GouLp, OGDEN MILLS, LEWIS RUTHERFURD
Morris, ARCHER M. HUNTINGTON, GEORGE DUD.
Pratt, T. CoLEMAN DUPONT, HENRY D. WHITON,
EDWARD HATCH, JR. , CORNELIUS R. AGNEW
Srivutific Stait
WILLIAM T. HORNADAY, Director of the Zoological Park;
CHARLES H. TOWNSEND, Director of the Aquarium;
RAYMOND L. DITMARS, Curator of Reptiles;
WILLIAM BEEBE, Honorary Curator of Birds and Director of the
Tropical Research Station;
« LEES. CRANDALL, Curator of Birds;
‘GEORGE S. HUNTINGTON, Prosector;
GEORGE A. MACCALLUM, Pathologist;
W. REID BLAIR, Veterinarian;
ELWIN R. SANBORN, Photographer and Editor. °
Editorial Committer
HENRY FAIRFIELD OSBORN, Chairman; .
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND.
Corrected to December, 1921
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
MEW VORK: ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBER 12
(Tropical Research Station Contribution Number 106)
FETUSES OF THE GUIANA HOWLING
MONKEY
By ADOLPH H. SCHULTZ
Research Associate, Department of Embryology, Carnegie Institution
of Washington.
Pave bask Sh DB Ys.) To 5 ONC Ee NG
Mite 7OOLOCTGAL PARK, NEW YORK
DECEMBER, 1921
sti BRITISH
<4 BRITISH GUIANA A
SCALE 5a142'W.
———
5S MILES
_- =
LOCATION OF THE TROPICAL RESEARCH STATION OF THE
NEW YORK ZOOLOGICAL SOCIETY
The circle represents a radius of six miles.
Volume III, Nuwber 12
FETUSES OF THE GUIANA HOWLING
MONKEY
By ADOLPH H. SCHULTZ
Research Associate, Department of Embryology, Carnegie Institution
of Washington.
Our knowledge of the development of the monkey is still very
limited. This is especially true in regard to platyrrhines, most
of the scanty literature dealing with Old World monkeys. Just
as in the case of human development, more work has been done
on the early than on the later stages of fetal development. It is
only through a complete knowledge of the entire intrauterine
development of the different monkey genera, however, that we
may hope to understand fully their position in the system of
primates, their relation to each other, and their various speciali-
zations and differences, problems of equal importance to the
zoologist, as well as to the comparative anatomist and scholar of
evolution. Furthermore, such knowledge will prove of great
interest to the embryologist and physical anthropologist, who
may derive therefrom a clearer insight into the laws governing
growth and the conditions of development of the various parts of
the body.
The following notes form a small contribution in this direction ;
they comprise a description of two older fetuses of a howling
monkey from South America. These specimens were given by
Mr. William Beebe, Director of the Tropical Research Station
of the New York Zoological Society, to the Carnegie Laboratory
of Embryology. In addition to the fetuses, use has been made
of observations on the skeleton of an adult male Alouatta, lent
by Mr. Beebe, and another skeleton of an adult and two
preserved bodies of juvenile male Alouattas from the anatomical
collection of the University of Ziirich. The author wishes to take
this opportunity to express his sincerest thanks to Mr. Beebe for
this valuable material, and to Prof. W. Felix for his kind permis-
sion to study the specimens of the Anatomy in Ziirich.
The sub-species of monkey to which the fetuses and one of the
skeletons (No. 4) belong is Alowatta seniculus macconnelli Elliot,
the Guiana howling monkey (Beebe, 719). The other skeleton
244 Zoologica: N. Y. Zoological Society [ Libs
(No. 3) and the two preserved bodies (Nos. 1 and 2) are of the
Species Alouatta seniculus L.t. The rarity of the fetuses is
increased by the fact that they are twins, twinning in monkeys
being to all appearances not any more frequent than in man.’
From an examination of the fetal membranes it is evident that
the fetuses are monozygotic or single-ovum twins. Both are
males and one of them is slightly larger than the other, their
respective sitting-heights (crown-rump lengths) being 111 mm.
(twin A) and 105 mm. (twin B). To what actual age this size
corresponds it is impossible to tell. The duration of pregnancy
in Alouatta is probably not more than five months,* and the
fetuses in question had reached certainiy the second half of their
intrauterine development. Through a careful comparison with
the human it is found that the development of these monkey
fetuses corresponds most closely with that of a human fetus of
twenty weeks, but no doubt they are actually younger. In respect
to the state of development of the lanugo, the ears, genitals, hands
and feet, and of ossification, the Alouatta fetuses are analogous
to human fetuses of the twentieth week, but the latter are consid-
erably larger, their sitting-height being on an average 154 to
164 mm.
1 The sub-species of these cannot be determined.
* Selenka (92) mentions one case of twins in a Cercocebus cynomolgus
and Fitzsimons (’19) records two instances of twins in Papio porcarius and
one in Cercopithecus pygerythrus.
* Bluntschli (13) gives the duration of pregnancy for Cebus and Chry-
sothrix as 4% to 5 months.
TABLE 1.
ABSOLUTE MEASUREMENTS OF THE ALOUATTA FETUSES
No. Measurement (in millimeters) : twin A. twin B.
Se ee EE ———ee——e
1. Sitting height: Top of head to lowest point on buttocks 111 105
2. Thoraco-abdominal height: Symphysion (upper border
of symphysis pubis) to suprasternal notch.......... 525 47
3. Symphysion to nipple (the latter projected on midsa-
gittal plane)’ Soi see wee cis bes ats pia sree k aul nae 2 ea AT 40
4. Symphysion to omphalion (center of attachment of um-
Pobre: eo): PSE eatere soars tahoe tens vas eee fails te ences Ore erecta 18 14.8
5. Biacromial diameter: Distance between the acromial
PTOCESSES,. os ore eterna ct aaar oe ehetariet 3) 5) 5) 5 lagen Ue eae Ea en 3 28.5
6. Bimammillary diameter: Distance between nipples..... 21 18.8
7. Bitrochanteric diameter: Distance between the great
tROCHANLErS: Genoese eect etig © ole ihereeeiade aac ean emerea 21, rag
8. Transverse diameter of chest (at nipple height)....... ZT 26.4
1921 ]
9.
10.
19
12.
13.
14.
15.
16.
iy B
18.
a 19.
20.
21.
22.
23.
24,
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
Sagittal diameter of chest (at nipple height)
Circumference of chest (at nipple height) .............
Length of upper arm: Top of caput humeri to humero-
Pn PPL IOPUIU MU TRUIGIC) oie oda ccc vod uv ccwadectny oe
Length of forearm: Radiale to tip of styloid process
SAME ase Oidionass's sc con bo a’e2c’s bay dk Soe ae
Length of hand: Middle of line combining styloid pro-
cesses of radius and ulna to tip of middle finger
Length of thumb: Stylion to tip of thumb.............
Breadth of hand (across metacarpo-phalangeal joints
ee I RAs ave y end oe OWA ae lew
Length of thigh: Top of great trochanter to lateral
Parr EMEC TORING «ie on i's oa Sek aca v's Die x wate «ck a LB
Length of leg: Medial point of knee joint (tibiale) to
Bier aevoMnal WiAllPOIis ... 55. u 4 oss + eed sane aac
PS) Spe) OA a 0 a rr rr
Length of foot: Heel to tip of longest toe..............
Breadth of foot (across metatarso-phalangeal joints II
to V + breadth of this joint on great toe)...........
Greatest length of head: Glabella to most distant point
SN Se Pe cae eee hase ais Gram x © noon site \owkg 2, chee
Greatest breadth of head: (over temporal or parietal
pie eee rick Sts oT Ss ceo \x-s os a nies loka <i wie ate
Auricular height of head: Tragion (upper border of
tragus) projected on midsagittal plane to vertex (per-
pendicular LO eGar-eye HOrizon)..26 6.6... s ce deen eae
Nasion-inion diameter: Point over middle of naso-frontal
suture (nasion) to occipital protuberance (inion)....
Biauricular breadth: Width between the tragion points
Horizontal circumference of head (greatest circumfer-
eneepassine’ through. clabella)) ... 6.60. d ce ae
UPL rehome NASON DOMINION: ct. e.< esa * «l0-sle/acd sie aiv = mi slele
Transverse arc: Tragion to tragion (perpendicular to
enV MMA (OTN) mctere eter Re. cSe aon 'cb0l) 0.0) ow, 6/Qle ele als iey «is
Total head height: Lowest point of chin (gnathion) to
vertex (perpendicular to ear-eye horizon)...........
Total face height: Nasion to gnathion................
Upper face height: Nasion to middle of mouth.........
Bizygomatic breadth: Greatest breadth between zygo-
TIPU CM AC Camere Aree rela tsis, givtsualie’s\ ele: o oysve's diene wnheie myers
Nasal height: Nasion to subnasal point (where nasal
septum and upper lip meet)....... ae Sta te eee wt hep A act
Nasal breadth: Greatest breadth between nasal wings..
Breadth of nasal septum: Smallest distance between
SRE Me odie visisle fa eaikis ovis caan déeerens vas
aS. awl e
Re MNES Eee lp A) Fee ccrele! aie oye os, 0 0 0's 6 a's miareoisie.e
ATE ICU O GR IRLOUID Mo riecnet Pate ia eieicrclsas cis sso sia's see ow eialecie
Length of ear: Highest point on helix to lowest point on
DUS Ly aay Seca iC fy Beas gtd BIO POON ene eee ac
Breadth of ear: Greatest breadth between anterior and
MBSteMOM DOrGeh OL NGlIX ce. 5 cece ese et agentes as
Schultz: Fetuses Guiana Howling Monkey
17.2
12.3
12.7
245
23.8
84
dl
26.9
23.3
14.8
11.8
29.6
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29.9
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248 Zoologica: N. Y. Zoological Society ea ee
For the purpose of comparing the outer form of the Alouatta
fetuses with that of the human fetus and of older Alouattas, and
in order to describe the proportions of the former fetuses in an
exact, numerical way, a series of.measurements has been taken
and indices of these have been formed. The technique of meas-
Sunrasternale__ :
Sunrasternale
Sithng Height
Sumphysion
ae Symphysion
FIG. 19. SCHEMATIC DRAWING OF ALOUATTA
FETUS (DOTTED HALF) AND NEGRO
FETUS OF TWENTY WEEKS, REDUCED TO
SAME SITTING - HEIGHT.
uring and the formulae for the indices, as well as the measure-
ments and indices themselves, are compiled in tables 1 and 2.
In drawing conclusions from these tables one has to be careful on
account of the scarcity of the material and the considerable
variability thereof. When more material is available, especially
1921 | Schultz: Fetuses Guiana Howling Monkey 249
different genera and different stages of development, such conclu-
sions will become safer and more extensive. In this paper I
intend chiefly to place on record a detailed and accurate
description of the two monkey fetuses and also to sketch in a
preliminary way the changes during growth and some results
of the comparison with human fetuses. The averages of the
above tabulated measurements and of those of the human fetuses
were used for the construction of a schematic drawing (Fig. 19)
which may serve to illustrate the following remarks concerning
the body proportions of the Alouatta fetuses.
The length of the trunk relative to the sitting-height is the
same in the Alouatta fetus as in the human fetus; however, the
different transverse diameters of the trunk, and the circumfer-
ence of the chest are all relatively considerably smaller in Alou-
atta, which accounts for the slender appearance of the trunk in
the latter. The adult condition is well expressed in the fetuses by
the fact that, in relation to the length of the trunk, the width
between the shoulders, as well as between the hips, is very much
smaller in the Alouatta than in the human fetus. We know that,
with the exception of the gorilla, man has the widest shoulders
and hips of any of the adult primates. The shoulders of the
Alouatta fetus are not only relatively closer together but they
are also very much higher than in the human fetus, which fact
constitutes a very marked difference between the two types. The
thoracic index shows only a slight difference; in both types the
transverse diameter of the chest surpasses the sagittal diameter.
The somewhat larger average in human fetuses, however, points
to a much more marked difference in later stages of growth, when
the thorax is much broader in man than in any platyrrhine
monkey, the latter having a deep and narrow chest. The nipples
of the Alouatta fetus are considerably higher and more laterally
situated than in the negro;' this close proximity of the mammae
to the axillae is characteristic for most New World monkeys.
The extreme in the very high and lateral position of the nipple in
Alouatta is not reached until postnatal life, when the nipple may
lie above the level of the suprasternal notch. The umbilicus of
Alouatta lies relatively much higher than in the human fetus.
From our findings on the trunk it is apparent that there exist
‘ The nipple of the Alouatta fetus lies over the third rib, in the human
fetus between the fourth and fifth rib.
250 Zoologica: N. Y. Zoological Society [ 13a
differences between the Alouatta and the human fetus in every
point except in the relative length of the anterior wall of the
trunk. The greatest differences consist in the shorter transverse
diameters of the trunk, and the higher position of the shoulders,
the nipples, and the umbilicus in the Alouatta.
The upper extremity is relatively considerably longer in the
Alouatta than in the human fetuses; it reaches practically no
farther down in Alouatta but is, as shown above, inserted higher
than in human fetuses. In this difference in length the various
parts composing the arm participate to a different degree. The
relatively greatest difference exists in the length of the hand;
somewhat less is the difference in the forearm, and least of all
in the upper arm, but all three are relatively longer in the
Alouatta. The relations of these parts to each other are expressed
in the humero-radial and forearm-hand indices; according to the
former, the radius, relative to the humerus, is longer in Alouatta,
and according to the latter the hand, in relation to the forearm,
is also very much longer in the monkey fetus. The humero-radiai
index in the adult Alouatta amounts to 91 according to Mollison
(710) and to 88 to 91.4 according to table 2. These figures are
higher than the corresponding ones for the fetuses; therefore,
the forearm in Alouatta has a greater rate of growth than the
upper arm. According to the forearm-hand index, which in
Alouatta decreases during growth, the hand has a slower rate
of growth than the forearm. The human hand, in contrast to
the hand of other primates, is characterized by its relatively -
greater breadth. This difference is already present in our fetuses,
the hand index being larger in the human. During postnatal
development the hand of Alouatta becomes still more slender.
The length of the thumb in relation to the length of the hand
is considerably less in the Alouatta than in the human fetuses,
and this relation does not seem to change markedly during
growth. The reduction of the thumb, typical for most monkeys,
is, therefore, recognizable in fetal life. In the Alouatta fetuses
fingers II to V are, in relation to the metacarpus, very much
longer than the human fingers. Finger III is the longest but
IV is almost as long, and finger II reaches about as far as
finger V.
1921 ] Schultz: Fetuses Guiana Howling Monkey “B61
The lower extremity is relatively little shorter in the Alouatta,
both femur and tibia being slightly shorter than in the negro
fetuses. This minute difference is at first rather surprising, in
consideration of the fact that man’s lower extremity is relatively
by far the longest of all primates. However, this distinction
does not fully appear until some time during postnatal growth.
The relation of the tibia to the femur is also only slightly differ-
ent, the relative length of the tibia of the Alouatta being some-
what greater than of the negro fetus. During postnatal
development the femoro-tibial index in Alouatta increases
steadily. Martin (’14) states that this index increases during
growth in all human races; it may, therefore, be concluded that
the lower leg has a more intense rate of growth than the thigh,
not only in man but also in Alouatta. The foot of the Alouatta is
very much longer than that of the human fetus, and this, indeed,
is one of the most marked differences between the body propor-
tions in the two types. This difference is very pronounced in the
relation of the foot length to the length of the tibia. The human
foot is relatively the shortest of all the primates and this most
probably holds true in fetal stages also. In both Alouatta and
man the leg-foot index decreases markedly during growth. The
fetal Alouatta foot is narrower and more slender than the foot of
the human fetus. In the former the great toe is very much
shortened, the second and third toes are of equal length, and the
heel is not prominent.
In summarizing the results of this comparison of the extremi-
ties in Alouatta and human fetuses the greater length of the
upper extremity in the Alouatta fetus and the approximately
equal length of the lower one are points especially noteworthy.
This different behavior in the relation of the upper to the lower
extremity is precisely expressed in the intermembral index,
which amounts to 136.7 in the Alouatta and to only 109.7 in
negro fetuses. This index decreases during postnatal develop-
ment in Alouatta as well as in man. The most distal portions
of the extremities, the hand and the foot, in the Alouatta surpass
in length the corresponding members of the human fetus to a
greater extent than the more proximal parts. The thumb and
the great toe especially is less developed in the Alouatta than
in the human fetus.
252 Zoologica: N. Y. Zoological Society | | Tita
It remains to consider briefly the proportions of the head. Here
the most striking feature consists in the smaller size of the brain
part of the head of the Alouatta as compared with the human
fetus, a difference which is especially manifest in the height.
There is a greater difference in the breadth than in the length of
the head, as shown by the smaller cephalic index of the Alouatta,
and a very much greater difference in height than in either of
the other diameters, as shown in the length-height index, which
is very much smaller in the Alouatta; the length, therefore,
shows the least difference of any of the head diameters. The
height of the face, from nasion to chin, is equal in the two types;
the upper-face height is somewhat greater in the monkey, and
the anterior part of the mandible is therefore less developed in
height than it is in the human fetus. The breadth of the face
shows but little difference, but the mouth is very much broader
in Alouatta. The external nose is higher as well as broader in
the monkey fetuses, the greater difference existing in height, so
that the nasal index becomes considerably larger in the human
fetuses. The low nasal index of our monkey fetuses is not
restricted to platyrrhines, but is also found in fetuses of catarr-
hines and apes (Schultz, ’20), in which also the nose is high
relative to its width. In regard to the fetal nasal index, there-
fore, man seems to occupy an exceptional position among the
primates. The relative interocular breadth is greater in adult
man than in any adult monkeys or apes; it is therefore not
surprising that the human fetus surpasses in this respect the
Alouatta fetus, although not to such a degree as it would in the
adult stage. The relative interocular breadth decreases in both
Alouatta and man during growth. The nasal septum is very
much broader in the Alouatta fetuses, which already show the
éypical features of the platyrrhine nose with laterally pointing
nostrils. The ear of the Alouatta is considerably larger than
that of the human fetus, a difference which becomes very evident
when the size of the ear is expressed in percentage of the size
of the head. The ear of the juvenile Alouatta is relatively almost
twice as large as that of the fetus. In relation to its length. it
is somewhat broader in the latter than in the average human
fetus of that stage. The external meatus is situated farther
back on the head in the monkey fetus. A further point of interest
1921 | Schultz: Fetuses Guiana Howling Monkey — 253
FIG. 20. FRONT AND RIGHT SIDE VIEW OF HEAD OF ALOUATTA
FETUS (TWIN A). APPROX. NATURAL SIZE
in the latter is the finding, on the lateral surface of the auricular
fold, immediately behind the anthelix, of two low and not very
distinct longitudinal folds, which without doubt, correspond to
the (five) folds found by Schwalbe (’97) in human fetuses of
four months. Figure 20 illustrates the typical features of the
head of the Alouatta fetuses.
Following is a condensed description of points of interest on
the integument of the Alouatta fetuses. In the latter, in contrast
to human fetuses, there is no philtrum nor labial tubercle and
the visible part of the mucous lips is extremely narrow. The
lanugo of the Alouatta fetuses at this stage of development is
restricted to the head. Very fine and short sparse hair is found
on the forehead and over each zygoma in front of the ears.
Longer, and somewhat more strongly developed hair occurs in
the region of the chin and on the upper lip. These, with the
exception of a few black ones on the upper lip, are very light.
The eyebrows are formed by long, almost bristle-like sinus hairs,
which are entirely black on the medial portions of the brows;
the lateral parts consist of hairs black in their lower part and
light at the end. A few of the outermost hairs in the brows are
entirely light. There are no anlagen for sinus hairs on the
cheeks of these twin fetuses, but such were found by Frédéric
(706) in three out of five Alouatta fetuses. No papillary ridges
can be made out on the palms, soles, or ventral side of the tail;
these apparently do not occur until later in fetal development.
254 Zoologica: N. Y. Zoological Society { (itis
The arrangement of pads (touch balls) and epidermal folds on
the palm and sole is shown in figure 21. The finger nails, as well
as the toe nails, are well developed and curved, in both longitu-
dinal and transverse directions, especially in the latter, and
resemble claws.
FIG. 21. RIGHT HAND AND FOOT OF ALOUATTA FETUS
(TWIN A), TWICE NATURAL SIZE
A few remarks concerning the degree of resemblance between
these twin fetuses of Alouatta may be of interest. One frequently
finds the assumption that monozygotic twins are “identical”, and
this is especially expected in fetuses in which environmental
conditions have not exerted an influence, as they do in postnatal
life. Newman (’17) has collected sufficient proofs to show that
absolute identity is never found, even in single-ovum twins. In
comparing the columns. for twin A with those for twin B in
tables 1 and 2, it is at once apparent that there is no identity in
the proportions of their bodies. All of the absolute measure-
ments of A are larger than the corresponding ones of B with two
exceptions—the upper-face height and the nasal height, which
are slightly greater in twin B. The degree of resemblance
between twins is most accurately obtained by figuring out the
1921 ] Schultz: Fetuses Guiana Howling Monkey 255
average percentage difference for all the absolute measurements
taken. This is done according to the following formula:
f m A —m B |
> a S100 b Fs i
% (m A+m B) J
A stands for twin A, B for twin B, m for measurement, and n
for the number of measurements used—in our case 39. The
result thus obtained is 4.81, i. e.. a measurement of twin B
differs on an average from the corresponding measurement of
twin A 4.81 percent. This rather high percentage is naturally
affected by the difference in absolute size in general between the
twins, but even if this were equalized, considerable difference
would remain, as shown by the fact that the indices, likewise
differ. The author’s experience with human monozygotic twins,
especially those of fetal stages, is analogous to that gained on
these monkey twins, inasmuch as human twins also show upen
closer examination a great number of more or less marked
deviations. Finally, I may state that so far, I have never found
human single-ovum twins, of any state of development, with ex-
actly the same general size. This is also the case in the Alouatta
twins, one being larger than the other.
For a study of the ossification and of the cartilaginous parts
of the skeleton, X-ray photographs were taken of one of the
Alouatta fetuses and the other one (twin A) was stained with
toluidin blue and cleared in a three percent solution of potassium
hydroxide and afterward placed in glycerine, a process which,
in addition to the ossified parts, shows the cartilage in a dark
blue color. For some points it became necessary also to partly
dissect one of the fetuses in order to observe in detail certain
conditions of the skeleton. Figure 22 is an exact drawing of the
cleared specimen and may serve to illustrate the following
description.
The spinal column consists of 57 vertebrae; 7 cervical, 14
thoracic, 5 lumbar, 3 sacral, and 28 caudal. These numbers
occurred also in all the other Alouattas examined, with the excep-
tion of skeleton 3, which has only 27 caudal vertebrae. In table
3 the lengths of the different spinal regions are expressed in
percentages of the praecaudal length of the spine. From the
(IIT; 12
Zoologica: N. Y. Zoological Society
Cr orm n onnregg
r i =
. SIDE VIEW OF CLEARED ALOUATTA FETUS, SHOWING THE
SKELETON. NATURAL SIZE
Se
FIG. 22.
figures in this table it can be concluded that the cervical and the
sacral regions of the Alouatta fetus are shorter than in human
fetuses, whereas the thoracic region is considerably longer in
the former than in the latter. It is furthermore of interest to
note that in both Alouatta and man during growth the relative
1921 ] Schultz: Fetuses Guiana Howling Monkey 257
length of the thoracic region decreases, while the lumbar and
sacral regions increase. In Alouatta the length of the caudal
region is relatively greater in adults than in fetuses. This
relatively greater rate of growth of the tail is also well expressed
in the following percentage relations of the length of the latter
to the sitting-height: Fetus A 109.0, fetus B 113.3, juvenile
Alouatta (No. 1) 165.0, and juvenile Alouatta (No. 2) 165.7.
Toldt (03) found that the relative length of the tail changed
very little during growth in case of Macacus cynomolgus L. This
suggests the possibility that, whereas the tail of Alouatta is in a
state of progressive evolution, that of Macacus is stationary, if
not regressive, a suggestion which may be supported by the fact
TABLE 3. LENGTHS OF THE DIFFERENT SPINAL REGIONS IN
PERCENTAGES OF THE PRAECAUDAL LENGTH OF
THE SPINE IN ALOUATTA AND MAN.
Region of spine:
cervical thoracic lumbar sacral caudal
Alouatta seniculus macconnelli.... 17.0 49.3 22 11.0 Iles as.
fetus (twin B)
Alouatta seniculus juv. (No. 1)... 13.4 airs, 26.0 ilsyal 186.7
Alouatta seniculus ad. (No. 3).... 14.8 45.1 Piles 12.8 160.0
Alouatta seniculus macconnelli.... 17.4 43.2 26.6 12.8 161.3
adult (No. 4)
_ Human fetuses of 20 weeks....... 21.0 41.5 22.0 525
Human adults (Martin, ’14)...... 16.0 39.0 25.0 19.0
that in the genus Macacus there are several species with an
almost rudimentary tail. In the spine of the Alouatta fetus each
vertebra contains one ossification center for the body and, down
to the fifth caudal vertebra one for each hemiarch. There are
14 ribs on each side, which are well ossified, their osseous shafts
having about the same proportional length in regard to the costal
cartilages as they have in the adult stage. The first eight pairs
of ribs reach the sternum; the next two pairs are asternal ribs,
and the remaining four pairs are floating ribs; even the last pair
is still fairly long. This arrangement of ribs is the rule in the
juvenile and the adult Alouatta also, with the exception of skele-
ton 8, in which only seven pairs of ribs reach the sternum and
five pairs are floating ribs, the last pair being very short. The
258 Zoologica: N. Y. Zoological Society [ TER sei
fd i eee
epee
_ *
if
XS
~
a
ZS s
FIG. 23. SKETCH OF STERNUM OF ALOUATTA FETUS
sternum as yet shows no ossification centers. It is a slender car-
tilaginous structure in which only three ring-shaped zones and
one uppermost, V-shaped zone are stained. The sternum of the
adult Alouatta is distinguished by a unique condition—a splitting
of the manubrium, due to the enormous development of the hyoid
and larynx. It is extremely interesting to find this condition
already present in the fetus. The cranial end of the fetal sternum
forks into two diverging processes to which the two uppermost
pairs of ribs and the clavicles are attached. Between these halves
of the manubrium emerges the trachea from the thorax, and
immediately above and almost in front of them lies the large
hyoid (see figure 23). In the adult the second rib inserts some-
what lower on the sternum, not, as in the fetus, on the lateral
process itself, but on the base of the latter, or even slightly below
the forking of the two processes. The clavicles are well ossified
1921 |] Schultz: Fetuses Guiana Howling Monkey 259
and are curved S-shaped, very similar to that in the human fetus.
The scapula has a greatest length equal to its greatest breadth,
whereas in the adult state the breadth surpasses the length (from
the glenoid cavity to the vertebral margin). The vertebral
margin above the dorsal end of the spina scapulae is approxi-
mately one-half the length of the portion of this margin below
the spine of the scapula. The latter portion of the vertebral
margin has a concave contour, thus forming a scaphoid scapula.
The acromial and coracoid processes are cartilaginous without
ossification centers, as are also the dorsal edge and glenoid cavity
of the scapula. The pelvis contains two pairs of ossification
centers, one in the ilia and the other in the ischia; the pubic
portion of the pelvis as yet shows no sign of ossification. The
blades of the ilia are long and slender.
All the shafts of the long bones of the extremities are well
ossified ; there are no ossification centers in any of their epiphy-
seal ends. The humerus has no foramen entepicondyloideum,
which is found in Cebus and other platyrrhines. On the humeri
of the skeleton No. 3 there is a foramen supratrochleare on each
side, but this is missing in the other specimens of Alowatta ex-
amined. The torsion of the fetal humerus, i. e., the angle between
the axes of the caput and of the trochlea, amounts to 90 degrees,
which is less than the torsion in human fetuses of corresponding
development, in which I found this angle to vary from 98 to 150
degrees. The proximal end of the olecranon projects consid-
erably beyond the incisura semilunaris, whereas the olecranon
in the human fetus ends abruptly at the proximal end of the
latter incisura. An analogous difference in the olecranon is
found between modern adult man and adult monkeys, and it is
interesting to see this distinction already clearly defined in fetal
stages. The tibia in its upper portion shows a rather marked
backward bend (proximal retroflexion). The carpus consists
entirely of cartilage and contains a well-developed centrale,
which at this stage of human development has disappeared from
the wrist. Among the tarsial cartilages the calcaneus possesses
a rather extensive ossified zone. In the human fetus this ossifi-
cation center normally does not occur until the sixth month; this
seems to be the only point in which the state of ossification of
the Alouatta fetuses does not coincide with that of the human
260 Zoologica: N. Y. Zoological Society [a
FIG. 24. SKULL OF AN OLD ALOUATTA SENICULUS WITH
FORAMEN TEMPORALE AT X
fetuses of twenty weeks. At the distal end of each metacarpus
and of the first metatarsus sesamoid cartilages are to be found.
Most of the elements of the skull of the Alouatta fetuses are
already ossified to a considerable extent; apparently only the
petrosum forms an exception in this respect. The great fonta-
nelle sends a graduaily narrowing arm almost as far as the
naso-frontal suture and posteriorly communicates by a fairly
broad arm with the occipital fontanelle; both are rather large.
The two pairs of lateral fontanelles are small. From the frontal
bone a process reaches toward the alisphenoid separating the
parietal from the malar bone, a condition usually found in
Alouatta, but contrary to the rule in other platyrrhines. The
lacrimal fossa is situated almost outside of the orbit and is in
full view when looking at the skull from in front. The foramen
zygomatico-faciale is very wide. Where the orbital plate of the
zygomaticum and the alisphenoid meet there is a foramen zygo-
matico-temporale of considerable size. This foramen has been
described by Joseph (’76), who found it in all adult New World
monkeys. It represents a vestige of the complete communication
between temporal and orbital fossae found in Lemuroidea. The
foramen is closed by a true membrana obturatoria orbitae. In
192f | Schultz: Fetuses Guiana Howling Monkey 261
our fetuses this foramen communicates by a narrow arm with
the fissura orbitalis inferior, thus actually forming a continua-
tion of the latter, which constitutes a more conspicuous remnant
of the former full communication between the two fosse. In
the squama temporalis of the Alouatta fetus, over the root of the
zygomatic arch, a fine foramen is to be found (x in figure 22).
This foramen may correspond to that noted by v. d. Broek (’08)
in the squama temporalis of Ateleus. I observed that the pres-
ence of this foramen is not restricted to the genus Ateleus but
occurs also in many other platyrrhines.» Among the skulls
of New World monkeys of my collection I found the foramen
at a corresponding place and of relatively large size in eight
Alouatta seniculus L. (see figure 24), including both juvenile and
very old animals, in two species of Atelews, and in one Aotus
boliviensis Elliot. It was missing in all skulls of Cebus and of
Hapale which I examined: This foramen is formed by an emis-
sary vein; it may be called foramen temporale. In the skull of
the very old Alowatta seniculus macconnelli (No. 4) no trace of
it could be seen. This, and the fact that in the fetus of the same
sub-species there is only an extremely fine foramen in the squama
temporalis, which may be merely a foramen nutritium, makes
it possible that this sub-species of Alouatta seniculus has no true
foramen temporale. The ramus mandibule is broad and rather
high; in later stages of growth this portion of the mandible of
Alouatta increases enormously in size, enclosing the greatly en-
larged hyoid. The hyoid capsule in the fetus is still cartilaginous,
but already of a quite extraordinary size. The nasal cartilages are
well developed and differentiated. The lateral nasal cartilage is
of triangular shape and the greater alar cartilage encircles the
nostril almost entirely, whereby its greatest surface is directed
forward instead of sidewise, as in catarrhines.
The most interesting conclusion of this study is the fact that
most of the typical differences existing between adult man and
adult Alouatta—be it those of outer form or those of the skele-
ton—are already well defined in fetal stages, although not yet as
pronounced as in the adult.
‘> A rather large foramen on the corresponding place was found by the
author in two human adult skulls of his collection (Nos. 266, white, and
216, negro).
BIBLIOGRAPHY
BEEBE, W.
1919 The higher vertebrates of British Guiana. Zoologica. - Scientific
contributions of New York Zool. Soc., Vol. II, No. 7
BLUNTSCHLI, H.
1913 Entwicklungsgeschichte platyrrhiner Affen, von Didelphis mar-
supialis, Tamandua . . . Anat. Anzeiger, Vol. XLIV,
Ergzh., p. 196.
V..D: BRONK. Aeon.
1908 Ueber einige anatomische Merkmale von Ateles, in Zusammen-
hang mit der Anatomie der Platyrrhinen. Anat. Anzeiger,
Vol XXXITIE, pp, 111:
FITZSIMONS, F. W.
1919 The natural history of South Africa. Vol. I. London.
FREDERIC, J.
1906 Nachtrag zu den “Untersuchungen tiber die Sinushaare der
Affen.” Zeitschr. f. Morph. u. Anthrop., Vol. IX, p. 327.
JOSEPH, G.
1876 Ueber die dussere Seitenwand der AugenhOhle bei den ameri-
kanischen Affen. Morphol. Jahrb., Vol. I, p. 454.
MARTIN, R.
1914 Lehrbuch der Anthropologie. Jena.
MOLLISON, TH.
1910 Die K6rperproportionen der Primaten. Morphol. Jahrb., Vol.
XLII, p. 97:
NEWMAN, H. H.
1917 The biology of twins (mammals). University of Chicago Science
Series.
ScHULTZ, A. H.
1920 The development of the external nose in whites and negroes.
Contrib. to Embryology, No. 34. Pub. 272, Carnegie In-
stitution.
SCHWALBE, G.
1897 Das dussere Ohr. Handb. d. Anatomie d. Menschen, Bardeleben.
Vol. V, p. 180.
SELENKA, E.
1892 Affen Ostindiens. Studien iiber Entwicklungsgesch., H. 5, 2nd
half. Wiesbaden.
TOLDT, C.
1903 Ueber die Aussere KOrperform zweier verschieden grosser Em-
bryonen von Macacus cynomolgus L. Arch. f. Anthrop., Vol.
XXVIII, p. 277.
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LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III], NUMBER 13
(Tropical Research Station Contribution Number 108)
MAMMALS COLLECTED BY WILLIAM BEEBE
AT THE BRITISH GUIANA TROPICAL
RESEARCH STATION
By H. E. ANTHONY.
Associate Curator of Mammals of the Western Hemisphere
American Museum of Natural History
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Volume IIT, Number 13
MAMMALS COLLECTED BY WILLIAM BEEBE
AT THE BRITISH GUIANA TROPICAL
RESEARCH STATION
By H. E. ANTHONY.
Associate Curator of Mammals of the Western Hemisphere
American Musewm of Natural History
When the New York Zoological Society established a Trop-
ical Research Station in British Guiana, with Mr. William
Beebe as Director, an arrangement was made whereby the De-
partment of Mammals of the American Museum was to receive
such specimens of mammals as might be collected from time to
time.
The first work of the Station was done in 1916, and has
been carried on at intervals ever since, resulting in the acces-
sion of some five hundred and twenty-one specimens of mam-
mals. The collecting has been done, for the most part, at three
points, Kartabo, Kalacoon and the Penal Settlement, all, as
shown on the accompanying map, lying within a small area
forty-five miles inland from the coast. This region is included
in the humid, tropical, rain forest zone’ and is a most important
locality, not alone for the richness of the mammalian fauna
but because of the great historical value which attaches to
specimens secured from northeastern South America. Many
‘of the classical species, many of the forms described by Lin-
_neus, have their habitat somewhere within this general region,
so that a series from the Guianas may be considered as typical.
The work of the Station staff was so occupied by their own
particular problems that the collecting of mammals was an in-
cidental feature and, in consequence, the list of species secured
there is far from complete. On the other hand, the aggregate
amount of time spent at the Station has resulted in the accu-
mulation of large series of some species, and a very gratifying
— —
*For photographs and details of this region see, “Tropical Wild Life in
7 British Guiana”, William Beebe, N. Y. Zool. Soc., 1917, and Zoologica, III,
1921, No. 1, by Henry Fairfield Osborn.
BRITISH
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LOCATION OF THE TROPICAL RESEARCH STATION OF THE
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4921] Anthony: British Guiana Mammals 267
number of rarities. Four forms new to science were taken and
in addition there are ten species new to the museum collections.
In all fifty-six species and subspecies are represented. No
small part of the value to be attached to this collection lies in
the number of skeletons preserved, since there are skeletons
for nearly all of the species represented by a series of any size.
In the identification of this collection, I have received valu-
able assistance from Mr. Gerrit S. Miller of the United States
National Museum, through the loan of comparative material,
while I am indebted to Dr. Wilfred H. Osgood of the Field
Museum, for opinions on some nomenclatural points.
1. Didelphis marsupialis marsupialis Linneus.
1758. Didelphis marsupialis Linneus, Syst. Nat., I, p. 54, (part).
1902. Didelphis marsupialis Allen. Bull. Amer. Mus. Nat. Hist., XVI,
p. 207.
Six specimens: Kartabo, 5 skins, 4 skulls, 1 skeleton.
2. Marmosa chloe Thomas.
1907. Marmosa chloe Thomas. Ann. and Mag. Nat. Hist., (7) XX,
p. 167.
Seven specimens: Kartabo, 7 skins, 6 skulls, 4 skeletons.
These animals are practically topotypes of chloe, since this
species was described from the Demerara river, 29 miles above
Georgetown.
3. Marmosa cinerea demerarae Thomas.
1905. Marmosa cinerea demerarae Thomas. Ann. and Mag. Nat. Hist.,
(7), XVI, p. 313.
Eight specimens: Kartabo, 7 skins, 4 skulls, 5 skeletons.
The series, which includes both adults and half grown
young, agrees quite closely with the type description of de-
merarae, type locality, Comackka, eighty miles up the De-
merara River.
268 Zoologica: N. Y. Zoological Society [TITs%3
4. Metachirus nudicaudatus nudicaudatus (Geoffroy).
1803. Didelphys nudicaudata E. Geoffroy. Cat. Mus., p. 142.
Immature specimen: Kartabo, skin with skeleton.
5. Monodelphis brevicaudata brevicaudata (Schreber).
1778. Didelphys brachyuros Schreber. Sédug. III, 549 pl. cli (plate
published in 1777).
Peramys brevicaudata auctorum.
One specimen: Kartabo, skin with skeleton.
The material for comparison with this specimen of Peramys
is too inadequate to enable me to do more than assign it pro-
visionally to brevicaudata, on the assumption that the animal
of the Guiana lowlands is Schreber’s species. Judging from the
limited series of red Peramys in the collection from Venezuela
and British Guiana, there exists either a very great degree of
individual variation or else a need for additional new species.
6. Bradypus cuculliger Wagler.
1831. Bradypus cuculliger Wagler, Isis, p. 605.
1871. Arctopithecus cuculliger Gray, Proc. Zool. Soc. London, p. 440.
Eight specimens: Kartabo, 5 skins, 5 skulls,-2 skeletons;
Kalacoon, 1 skin; Kyk-over-al, 1 skin, 1 skull.
This series is referred to cuculliger upon the basis of the
descriptions given in the two references cited above, and with
regard to the fact that Gray had a specimen from Demerara
which he called cuculliger. The agreement with the descrip-
tions is fairly close and the series averages darker in color
than a series of tridactylus flaccidus from Venezuela; but the
color pattern is rather similar to that of flaccidus, in fact so
similar that, should my identification of cuculliger be correct, I
believe that flaccidus should stand as a subspecies of cuculliger
and not of tridactylus. Compared with specimens of tridac-
tylus from Santarem, Brazil, the Guiana specimens are radi- |
eally different in the coloring of the head, throat and neck.
1921] Anthony: British Guiana Mammals 269
7. Choloepus didactylus Linneus.
1766. Bradypus didactylus Linneus. Syst. Nat., I, p. 51.
Two specimens: Kalacoon, 1 skin, 1 odd skull.
8. Myrmecophaga tridactyla Linneus.
1758. Myrmecophaga tridactyla Linneus, Syst. Nat., p. 35.
One of these anteaters was brought back alive and placed
in the New York Zoological Park. It has since died and is
now in the Museum collection. It was taken at Kartabo and
was only about half grown.
9. Tamandua tetradactyla tetradactyla (Linneus).
1766. Myrmecophaga tetradactyla Linneus, Syst. Nat. I, p. 52.
Twelve specimens: Kartabo, 9 skins, 10 skulls, 2 skeletons;
Kalacoon, 1 skin, 2 skulls.
The color of the pelage of these anteaters varies so consid-
erably that, were the two extreme examples to be considered
alone, they might well be thought to be distinct from one an-
other. The dark dorsal area is almost completely absent from
one specimen which in consequence greatly resembles the yel-
low longicaudata. However, the apparent gap between this
yellow specimen and the darkest of the series, is well bridged
over by the specimens of intermediate coloration. The shape
of the nasals, used as a character of separation between longi-
caudata and tetradactyla, varies almost as much as does the
color of the pelage, the narrowest examples being no wider
than the nasals of longicaudata from Maripa, Venezuela.
10. Cyclopes didactylus didactylus (Linneus).
1766. Myrmecophaga didactyla Linneus, Syst. Nat. I, p. 51.
One specimen: Penal Settlement, skin and skeleton.
11. Dasypus novemcinctus novemcinctus Linneus.
1766. Dasypus novemcinctus Linneus, Syst. Nat. I, p. 54.
Four specimens: Kartabo, 4 skins, 4 skulls, 2 skeletons.
These are all half grown young.
270 Zoologica: N. Y. Zoological Society [TIT 343
12. Tatu kappleri (Krauss).
1862. Dasypus kappleri Krauss, Archiv. Naturg., Vol. I, p. 24.
Three specimens: Kartabo, 2 skins, 1 skull; Kalacoon, 1
skin with skeleton.
These are the first specimens of kapplevi to be received in
the Museum’s collection. This species is widely different from
Dasypus novemcinctus, which it resembles somewhat super-
ficially. Aside from a slight difference in size, kappleri being
the larger, it has only eight movable bands instead of nine, it
has two rows of spur-like scales on the hind legs which are
entirely wanting on novemcinctus, a rudimentary fifth toe on
the fore foot, longer ears and noticeably different skull charac-
ters, the most important of which is the peculiar flange-like
margins of the posterior palate.
13. Layassu pecari beebei Anthony.
1921. Tayassu pecari beebei Anthony, Amer. Mus. Novitates, No. 19,
pl.
Eight specimens: Kartabo, 7 skins, 7 skulls, 2 skeletons.
General Characters."—Closely related to pecari pecari, but differing in
the extent of white on the snout and lower jaw.
Description—Coloration about as in p. pecari but white of face and
throat markings more yellowish; long hairs of upper parts brownish black;
snout, above, only slighter lighter in color than rest of upper parts and not
with strongly contrasting whitish of p. pecari; chin and throat patch
restricted and not in such marked contrast to the surrounding areas; feet
dark to hoofs. Skull as in p. pecari.
Measurements.—Taken from animal in flesh: total length, 1090 mm.;
tail vertebrae, 60; hind foot, 224; weight 80 pounds.
This subspecies was described upon the basis of the re-
stricted white areas upon the nose and throat. It is closely re
lated to true pecari of Brazil, and was named in honor of Mr.
William Beebe, the Director of the Tropical Research Station.
14. Pecari tajacu macrocephalus Anthony.
1921. Pecari tajacu macrocephalus Anthony, Amer. Mus. Novitates,
INO? 19, nao.
Eight specimens: Kartabo, 5 skins, 6 skulls.
*This description, together with those of the other three new species,
appeared first in American Museum Novitates, No. 19, by H. E. Anthony.
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Zoologica
Vol. ITI, No. 13.
Fuce page
FIGURE 25
(b), Pecari t
(c) Pecari torvus
halus
All figures three eighths natural size.
QAjJACUuU MACTOCE p
(a), Pecari tagacu tajicu
1921] Anthony: British Guiana Mammals Sil
General Characters.—Similar to tajacu but with skull larger and mark-
edly different in structure.
Description.—Pelage about as in tajacu, grizzled yellowish and black,
with black dorsal area; collar fairly well outlined.
Skull larger than that of tajacu, with more massive build, the forward
extension of the zygomatic flange continued to canine alveolus and forming
a heavy rostrum; outline of entire skull noticeably subtriangular viewed
either from above or below, due to extended zygomatic flange; palate
throughout anterior portion wider than distance across the molar series of
that portion.
Measurements.—Taken in the flesh: total length 948 mm.; length
of hind foot, 195.
Macrocephalus has been set off from typical tajacu because
of important cranial differences. The Kartabo skulls have wide
zygomatic flanges, which extend well out on the rostrum and
give to the skull a subtriangular outline, when viewed from
above or below. The skulls of true tajacw from Brazil have
much slenderer rostra and the outline is flask-like.
15. Mazama americana tumatumari Allen.
1915. Mazama americana tumatumari Allen, Bull. Amer. Mus. Nat.
Hust., XXXIV, p. 5386.
Two specimens: Kartabo, 1 skin with skull; Kalacoon, 1
skin, spotted.
The specimen in the adult pelage appears to agree with the
type, from Tumatumari, which is at no great distance from
Kartabo and Kalacoon.
The very young specimen, which is only a flat skin with-
out skull, is very brightly colored with numerous and conspicu-
ous buffy spots.
16. Mazama nemorivaga (F. Cuvier).
1817. Cervus nemorivagus F. Cuvier, Diction. Sci. Nat. VII, p. 485
(part, the Cayenne specimens only). j
1915. Mazama nemorivagus Allen, Bull. Amer. Mus. Nat. Hist.
XXXIV, p. 548.
-Nineteen specimens: Bartica, 1 skin; Kartabo, 11 skins,
9 skulls, 1 skeleton; Kalacoon, 4 skins, 2 skulls, 2 skeletons.
There is considerable variation in color shown by this large
series, although all are some shade of light brown. The varia-
272 Zoologica: N. Y. Zoological Society [III;13
tion consists chiefly in a more or less extensive darkening of
the dorsal area and, to a lesser degree, in the intensity of the
dark coloring on the legs. Four young, in the spotted coat, are
included in the series, the youngest of which is very conspicu-
ously spotted but the oldest is only very faintly marked and
about to assume a pelage like that of the adult.
17. Hydrocherus hydrocherus (Linneus).
1776. Sus hydrocherus Linneus, Syst. Nat., I p. 1038.
Two immature specimens: Kartabo, skins with skeletons. —
These specimens are too young, being about the size of
Sylvilagus, to give any characters.
18. Dasyprocta aguti flavescens (Thomas)
1898. Dasyprocta rubrata flavescens Thomas, Ann. and Mag. Nat.
Hist., (7), Il, p. 274.
Thirty-four specimens: Kartabo, 20 skins, 33 skulls, 1
skeleton; Kalacoon, 1 skull.
This large series exemplifies the degree of variation found
in the genus. The intensity of coloration on the rump varies
from ochraceous—orange to Sanford’s brown (Ridgway, Color
S‘andards and Nomenclature) ; the extent of the bright area is
often considerably reduced by the encroachment of the darker
colored hairs of the upper dorsal region; the nape and shoul-
ders, while normally quite dark, are sometimes much lighter
and the degree of punctulation is far from constant.
Mr. Oldfield Thomas has referred* the northern Guiana
agoutis to the above species, mentioning among his specimens
a large series from Demerara. Specimens in our collection
from Tumatumari, British Guiana, are indistinguishable from
the Kartabo series, although they had previously been identi-
fied as lucifer cayennae. I agree with Mr. Thomas that’the
Guiana specimens have nothing to do with Wagler’s prymnolo-
pha, since none of the large series of flavescens before me
°1917, Ann. and Mag. Nat. Hist., (8), XX, p. 259.
FIGURE 26
(a), Dorsal aspect of skull of Echimys armatus, Trinidad.
(b), Dorsal aspect of skull of Echimys longirostris, Kartabo.
(ce), Dorsal aspect of skull of Echimys armatus, Demerara.
(d), Ventral aspect of skull of Echimys longirostris, same skull as shown in b.
e), Ventral aspect of skull of Echimys armatus, same skull as shown in ¢.
All figures natural size.
Zoologica Vol. III, No. 13. Face page 275
1921] Anthony: British Guiana Mammals 273
shows any tendency toward the development of a black rump
patch, which is a very conspicuous feature of prymnolopha.
19. Agoutt paca paca (Linneus).
1766. Mus paca Linneus, Syst. Nat., I, p. 81.
Two specimens: Kartabo, skins with skulls.
These specimens are provisionally identified as paca but it
is possible that they represent fulvus of Cuvier. Owing to a
scarcity of suitable material from northeastern South America,
and to the rather confusing status of the group as set forth
in literature, the wide range of individual variation making
identification from written descriptions most difficult, I have
thought it best to assign the Guiana material to paca.
20. Proechimys cayennensis (Desmarest).
1817. Echimys cayennensis Desmarest, Nouv. Dict., X, p. 59.
Eleven specimens: Kartabo, 4 skins, 3 skulls, 4 skeletons;
Kalacoon, 1 skin; Samiri Island, Mazaruni River, 6 skins.
The relationships of cayennensis, as given by Thomas’,
are with trinitatis, and this series of spiny rats from Guiana
bear considerable resemblance to the rats from Trinidad, both
superficially and in cranial characters.
21. Echimys longirostris Anthony.
1921. Echimys longirostris Anthony, American Museum Novitates,
No. 19, p. 5.
One specimen: Kartabo, skin and skeleton.
General Characters.—Most like armatus, but differing in characters of
pelage and in significant details of cranial structure, having much longer
nasals and shallow postpalatal notch.
Description—Pelage spiny, but with many unmodified hairs which
partially mask the spines; hairs on crown only slightly spinous; color above,
a mixture of black, ochraceous and buff, the ochraceous strongest on nose
and face and posterior to shoulders along dorsal area; black strongest on
neck and shoulders; flanks lighter than dorsal area and merging insensibly
into the grayish under parts; hairs of underparts subspinous, gray at base
and tipped with buff; pectoral area more brightly colored than posterior
“1903, Ann. and Mag. Nat. Hist., (7), XI, p. 491.
274 Zoologica: N. Y. Zoological Society (iiss
under parts; hands and feet grizzled gray, buff and ochraceous, dirty white
distally; tail haired at base for about 50 mm., colored same as rump, scaly
for rest of its length, sparsely haired, practically unicolor, ashy in color.
Skull elongate with convex superior outline; nasals long, slender, sub-
cylindrical; lateral margins of temporals forming straight lines, not concave;
postpalatal notch U-shaped, reaching scarcely beyond the posterior margin
of last molar; molar pattern typical of the genus.
Measurements.—Taken from dried skin; total length 466 mm., tail
vertebrz 225; hind foot 38.
Although no less than three different names have been
employed for the Echimys of British Guiana, the types have
been determined to be specifically identical so that guianae and
castaneus stand as synonyms of armatus. The Kartabo Echi-
mys could be identified under none of these names and it was
necessary to make it a new species. A fair amount of com-
parative material representing all three of these names has
been available but no specimen was found which had such long
nasals nor such a great interorbital breadth.
22. Mus musculus musculus Linneus.
1758. Mus musculus Linneus, Syst. Nat., I, p. 62.
Twelve specimens: Kartabo, 1 skin; Georgetown, 11 skins,
5 skulls.
23. Rattus rattus alexandrinus (Geoffroy).
1818. Mus alexandrinus Geoffroy, Descr. Egypt, II, 733.
Twenty-three specimens: Georgetown, 17 skins, 9 skulls,
3 skeletons; Penal Settlement, 6 skins, 3 skulls, 1 skeleton.
24. (Ccomys guiane Thomas.
1910. Ccomys guiane Thos. Ann. and Mag. Nat. Hist., (8) VI, p.
187.
Two specimens: Kartabo, 2 skins, 1 skeleton.
These specimens agree with the description of guiane closely
enough to be so identified and they were taken sufficiently near
to the type locality of guiane, River Supinaam, to be consid-
ered topotypical.
1921] Anthony: British Guiana Mammals 275
25. (Mcomys nitedulus Thomas.
ae Gicomys nitedulus Thos. Ann. and Mag. Nat. Hist., (8) VI, p.
505°
Three specimens: Kartabo, 3 skins, 1 skull, 1 skeleton.
The type locality of nitedulus is the lower Essequibo River,
thirteen miles from mouth, which is not very distant from
Kartabo. The specimens from Kartabo are none of them old
adults and consequently appear to be a trifle smaller than the
measurements given by Thomas.
26. Mcomys rutilus Anthony.
1921. C£comys rutilus Anthony, Amer. Mus. Novitates, No. 19, p. 4.
One specimen: Kartabo, skin with skeleton.
General Characters.—A small, brightly colored species, with very short
tail and clear white under parts.
Description.—Color above, between amber brown and hazel (Ridgway),
darkest along dorsal area and on crown, the hairs slaty black for basal
two-thirds; below, clear white, the hairs white to the base; hands and feet
dirty white, almost dusky; dark orbital ring with small dark area at pos-
terior corner of the eye; tail brownish, unicolor. Skull small and broad,
rostrum very short, zygomata flaring, a low supra-orbital beading.
Measurements.—Taken in the flesh: total length, 171 mm.; tail ver-
tebrae, 94; hind foot, 20. Greatest length of skull, 24.2; zygomatic breadth,
13.5; length of nasals, 7.7; interorbital breadth, 4.4; breadth of brain case,
11; palate, to incisors, 10; palatal foramina, 3.7x2.2; length of upper
molar series, 3.4.
This is a small, brightly colored mouse, of the genus
comys, quite distinct from the other @comys collected there,
nitedulus, and possibly a relative of rosilla Thomas, from
which it differs however in clear white underparts.
27. Neacomys guiane Thomas.
1905. Neacomys guiane Thomas, Ann. and Mag. Nat. Hist., (7) XVI,
p. 310.
Two specimens: Kartabo, 2 skins, 1 skull, 2 skeletons.
These specimens are essentially topotypes since Thomas
gives the type locality as the Demerara River, altitude 120
feet.
28. Nectomys squamipes melanius Thomas.
1910. Nectomys squamipes melanius Thomas, Ann. and Mag. Nat.
Hist., (8) VI, p. 185.
276 Zoologica: N. Y. Zoological Society [III;13
Twenty-three specimens: Kartabo, 7 skins, 6 skulls, 6
skeletons; Kalacoon, 1 skin; Kyk-over-al, 11 skins, 8 skulls, 4
skeletons; Samiri Island, Mazaruni River, 4 skins. .
The series agrees quite closely with the description ‘of the
type, and as the type locality is given as the lower Essequibo
River, twelve miles from mouth, the Beebe specimens are
practically topotypes.
29. Oryzomys velutinus?
1893. Oryzomys velutinus Allen, Bull. Amer. Mus. Nat. Hist., V, p. 214.
Ten specimens: Kartabo, 10 skins, 3 skulls, 3 skeletons.
The short-haired Oryzomys of the Beebe collection are
provisionally referred to velutinus, although it may be ques-
tioned whether velutinus is not truly insular and the name not
to be used for mainland forms. Without specimens of Lund’s
laticeps for comparison, and because of the confusing status
of the Oryzomys of northeastern South America, these speci-
mens are so named now, merely for the sake of convenience,
but it is quite possible that more material will show them to
be a subspecies of laticeps. These specimens agree quite closely
with velutinus from Trinidad.
30. Oryzomys sp.?
One specimen: Bartica, skin, without skull.
This is a large species, strongly ochraceous above and
buffy white below. It is not unlike trinitatis in general ap-
pearance, and on the other hand it agrees fairly well with the
type description of macconnelli®? but appears to be rather too
small in size.
31. Oryzomys sp.?
Two specimens: Kartabo, 2 skins, 1 skull.
°1910. Thomas, Ann. and Mag. Nat. Hist., (8) VI, p. 186.
a
1921 | Anthony: British Guiana Mammals 277
These two specimens are of the meridensis group and pos-
sibly are closely related to caracolus Thomas® described from
near Caracas, Venezuela.
32. Guerlinguetus wstuans wstuans (Linneus).
1766. Sciurus xstuans Linneus, Syst. Nat., I, p. 88, (Surinam).
1915. Guerlinguetus xstuans xstuans Allen, Bull. Amer. Mus. Nat
Hist., XXXIV, p. 256.
Five specimens: Kartabo, 5 skins, 3 skulls, 2 skeletons.
This series is especially acceptable since this species of
the Guiana lowlands has hitherto been very poorly represented
in the Museum collection.
33. Procyon cancrivorus cancrivorus (Cuvier).
1798. Ursus cancrivorus Cuvier, Tab]. Elem. Hist. Nat., p. 113.
Two specimens: Penal Settlement, 2 skins, 2 skulls, 1
skeleton.
34. Potos flavus flavus (Schreber).
1775. Lemur flavus Schreber, Saug. I., p. 145, pl. 42.
Two specimens: Kalacoon, 1 skin, 1 odd skull.
35. Lutra mitis Thomas.
1908. Lutra mitis Thomas, Ann. and Mag. Nat. Hist., (8), I, p. 393.
Three specimens: Kartabo, 1 skin, 1 skull, 1 skeleton;
Kalacoon, 1 skull.
The skin and the skulls seem to agree fairly well with
Thomas’s description of the type.
36. Tayra barbara barbara (Linneus).
1766. Mustela barbara Linneus, Syst. Nat., I, p. 67.
Four specimens: Kartabo, 2 skins, 2 skulls, 2 skeletons;
Kalacoon, 1 skull.
*1914. Ann. and Mag. Nat. Hist., (8) XIV, p. 242.
278 Zoologica: N. Y. Zoological Society (TItsee
One specimen has the head and neck above grizzled gray
and the chest area dirty whitish; the other has the corre-
sponding areas yellowish above and pale ochraceous below.
The odd skull is unusually large, with a very high sagittal
crest, and measures, greatest length, 130 mm.; zygomatic
breadth, 78.5 against 107 and 65, the dimensions of an adult
female from Kartabo.
37. Nasua pheocephala Allen.
1904. Nasua phxocephala Allen, Bull. Amer. Mus. Nat. Hist., XX, p.
4,
Four specimens: Kartabo, 3 skins, 3 skulls, 1 skeleton;
Kalacoon, 1 skeleton.
These specimens agree well in coloration with the type of
pheocephala, from Suapure, Venezuela.
38. Panthera onca (Linnzus).
1766. Felis onca Linneus, Syst. Nat., I, p. 61.
One specimen: Kartabo, skin and skull, adult male.
Skull measurements: Greatest length, 238 mm.; length of
nasals, 57; zygomatic breadth, 160; mastoid breadth, 99;
breadth of rostrum, 67; length of upper tooth row to in-
cisors, 95.
39. Margay tigrina vigens (Thomas).
1904. Felis weidii vigens Thos., Ann. and Mag. Nat. Hist., (7) XIV,
DLs
1919. Margay tigrinw vigens Allen, Bull. Amer. Mus. Nat. Hist., XLI,
1h Se
One specimen: Kartabo, skin with skeleton.
This specimen agrees, in most essential characters, with
the type description of vigens (loc. cit.). The skull measure-
ments are a trifle larger for the Kartabo animal and the color
pattern varies slightly from that given by Thomas, in the
lesser number of dark rings on the tail and the whiter under-
parts.
1921] Anthony: British Guiana Mammals 279
40. Herpailurus yaguarondi unicolor (Traill).
1819. Felis unicolor Traill, Mem. Wernerian Soc., III, p. 170.
1919. Herpailurus yaguarondi unicolor Allen, Bull. Amer. Mus. Nat.
Hist., XLI, p. 383.
One specimen: Kartabo, skin and skeleton, adult male.
This rare cat is in the black phase and is a glistening
black all over, except about the head and neck which is grizzled
with gray.
Measurements, taken in the flesh; total length, 1150 mm.;
tail vertebre, 470; hind foot, 155; weight, 19 pounds.
Skull, greatest length, 112 mm.; basal length, 99; zygo-
matic breadth, 71; breadth of braincase, 43.5; length entire
upper tooth row, 42.7.
41. Saccopteryx bilineata (Temminck).
1839. Urocryptus bilineatus Temminck, Van der Hoeven, Tij\'sch.
Natur., p. 33.
Three specimens in alcohol: Kalacoon.
42. Rhynchiscus naso (Wied).
1821. Vespertilio naso Wied, Schinz’s Thierreich, Vol. I, p. 179.
Thirteen specimens in alcohol: Kaow Island, Essequibo
River, 12; Kartabo, 1.
The collector’s notes state that these bats were found “on
bark of tree.”
43. Glossophaga soricina soricina (Pallas).
1766. Vespertilio soricinus Pallas, Miscell. Zool., p. 48.
Sixty specimens: Creeklands, Berbice, 28 skins, 20 skulls,
20 skeletons; Georgetown, 6 skins, 16 alcoholics; Kartabo, 10
alcoholics.
This large series of Glossophaga presents but little varia-
tion in color and appears to be typical soricina in every char-
acter.
280 Zoologica: N. Y. Zoological Society [Tlis¥e
44. Hemiderma perspicillatum perspicillatum (Linnezus).
1758. [Vespertilio] perspicillatus Linneus, Syst. Nat., I, p. 31.
Fifty-seven specimens: Georgetown, 10 skins; 29 alco-
holics; Kalacoon, 4 alcoholics: Kartabo, 14 alcoholics.
This series is quite uniform in character and presents no
points worthy of comment.
45. Mesophylla macconnelli Thomas.
1901. Mesophylla macconnelli Thomas, ‘Ann. and Mag. Nat. Hist., (7),
VIX, p: 145:
Seven specimens in alcohol: Kartabo, July 22, 1920.
This genus has hitherto been unrepresented in the Museum
collection, but there is little difficulty in identifying it from
the description given by Miller in “The Families and Genera
of Bats,” p. 158. The most conspicuous features of the skull
are the swollen maxillaries and depressed nasal region, while
the pelage of macconnelli is very light colored.
These specimens were taken at no very great distance
from the type locality of the species which is Kanuku Moun-
tains, British Guiana.
46. Phyllostomus hastatus hastatus (Pallas).
1767. Vespertilio hastatum Pallas, Spici. Zool., III, p. 7.
Three specimens in alcohol: Kartabo, 2; Kalacoon, 1.
47. Vampyrus spectrum spectrum (Linnzus).
1766. Vespertilio spectrum Linneus, Syst. Nat., I, p. 46.
One specimen: Kartabo, skin and skeleton.
This very large species is represented by only one speci-
men which appears to be typical in all respects. The forearm
is 104 mm. long.
48. Furipterus horrens (F. Cuvier).
1828. Furia horrens Cuvier, Mem. Mus., XVI, p. 150.
Six specimens: Kartabo, April 26 to August 25.
1921] Anthony: British Guiana Mammals 281
This series is a valuable addition to the Museum collec-
tion since Furipterus is exceedingly rare. The series is uni-
form in coloration and the average measurement of the fore-
arm is 34.9 mm.
49. Humops milleri (Allen).
1900. Promops milleri Allen, Bull. Amer. Mus. Nat. Hist., XIII, p. 91
Two specimens (1 imm.) in alcohol: Kartabo.
The adult specimen agrees fairly well in all characters but
size with the type of milleri. In cranial characters the two
are identical with the following exceptions, the Kartabo speci-
men has slightly smaller upper incisors and less extensive
basicranial pits. The following measurements are of the
Kartabo bat, contrasted with the type of milleri in paren-
theses; forearm, 55 mm. (58.7); greatest length of skull, 24.7
(25.2) ; zygomatic breadth, 14.2 (14.2): length of upper tooth
row, C-M°, 9.3, (9.8).
This specimen is not unlike the type of Eumops barbatus
(Allen) which differs from milleri mainly in size only. Addi-
tional material may show that barbatus should stand either
as a subspecies of milleri or as its synonym, and the older
name of milleri is followed, because of the inadequate material
representing barbatus, the type being unique.
50. Molossus obscurus Geoffroy.
1805. Molossus obscurus Geoffroy, Ann. du Mus., VI, p. 154.
Twenty-two specimens: Georgetown, 9 alcoholics; Kala-
coon, 1 skin, 9 alcoholics; Kartabo, 3 alcoholics.
Only one specimen of the entire series is in the red phase.
51. Molossus rufus Geoffroy.
1805. Molossus rufus Geoffroy, Ann. du Mus., VI, p. 154.
Six specimens: Georgetown, 3 skins; Kartabo, 1 alcoholic;
Penal Settlement, 2 alcoholics.
282 Zoologica: N. Y. Zoological Society [I11;13
These specimens are in the dark phase, and the Kartabo
example has a forearm 48 mm. in length:
52. Saimiri sciureus (Linneus).
1758. Simia sciwrea Linneus, Syst. Nat. I, p. 19.
1913. Saimiri sciureus Elliot, Review of the Primates, I, p. 310.
Five specimens: Kartabo, 3 skins, 2 skulls, 1 skeleton;
Kalacoon, 1 skin, 1 skull.
Of the Saimiri taken in British Guiana, all but one ap-
pear to be typical sciwreus, the series being fairly uniform in
coloration. The exception, a skin without skull, differs in
having black lateral stripes on the head and behind the eye,
while the dorsal region is a much brighter color than in the
other specimens, being quite yellow. This specimen, from
Kartabo, I have provisionally identified as cassaquiarensis, a
considerable extension of range, if the identification proves to
be correct, since the range of this species is to the west of
British Guiana.
538. Saimiri cassaquiarensis (Humboldt).
1811. Chrysothrizx sciureus cassaquiarensis Humboldt, Rec. Obs. Zool.
I, p. 334, (1815)
1913. Saimiri cassaquiarensis Elliot, Review of the Primates, I, p. 311.
One specimen: Kartabo, skin without skull.
54. Alouatta seniculus macconnelli (Elliot).
1910. Alouatta macconnelli Elliot, Ann. and Mag. Nat. Hist., (8), V.
p. 80.
1916. Alouatta seniculus macconnelli Allen, Bull. Amer. Mus. Nat.
Hist., XXXV, p. 233.
Thirty-three specimens: Kartabo, 22 skins, 23 skulls, 8
skeletons; Kalacoon, 3 skins, 7 skulls.
This large series displays a very considerable range of
individual variation in color. The general tone of the upper
parts varies from yellow to bright orange red, with a corre-
sponding lack of uniformity in the coloration of the limbs and
tail, which in some specimens are much darker than the upper
1921] Anthony: British Guiana Mammals 283
parts, although this condition is not as conspicuous as in true
seniculus.
A group of eight individuals, six adults and two young,
has been mounted and placed on exhibition in the American
Museum of Natural History.
55. Pithecia pithecia (Linneus).
1766. Simia pithecia Linneus, Syst. Nat., I, p. 40.
1918. Pithecia pithecia Elliot, Review of the Priniates: I, p. 2938.
Twenty-seven specimens: Kartabo, 17 skins, 19 skulls, 5
skeletons; Kalacoon, 4 skins.
The status of the species of the genus Pithecia is far from
satisfactory, to judge by the literature and the specimens in
the Museum collection, and the present series focuses atten-
tion upon ‘this fact. The specimens all were taken in a small
area and exhibit considerable variation in coloration. In the
males, the color of the head varies in the amount of ochraceous;
in one specimen this color extends from the under side to the
median line of the upperparts, but the normal color above is
a dirty white.
56. Cebus apella apella (Linneus).
1758. Simia apella Linneus, Syst. Nat. I, p. 28.
1913. Cebus apella Elliot, Review of the Piihiates: TT, p18.
Twenty-five specimens: Kartabo, 15 skins, 23 skulls, 2
skeletons.
Although the range of color variation shown by this series
is considerable, the average is darker than that of a series of
wpiculatus from Venezuela. The series shows but little of the
fulvous which characterises apiculatus and is much darker
than apella brunneus.
APPENDIX A
By H. E. ANTHONY
The following four species of mammals, a rat, a mon-
goose and two bats, were collected at Georgetown, sixty miles
away on the coast, and have not as yet been taken at the
Station.
Rattus norvegicus (Erxleben).
1777. [Mus] norvegicus Erxlebon, Syst. Regni Anim., Vol. I, p. 381.
Four specimens: Georgetown, 4 skins, 4 skulls, 2 skeletons.
Mungos birmanicus (Thomas).
.1886. Herpestes auropunctatus birmanicus Thomas, Proc. Zool. Soc.,
London, p. 58.
1911. Mungos birmanicus G. M. Allen, Bull. Mus. Comp. Zool., LIV,
Deselile
One specimen: Georgetown, skin and skeleton.
It is to be hoped that the mongoose will find enough ene-
mies upon the continental mainland to prevent the widespread
distribution and destruction which have followed its introduc-
tion into the West Indies.
Artibeus planirostris planirostris (Spix).
1823. Phyllostoma planirostre Spix, Sim. et Vesp. Bros., p. 66.
1908. Artibeus planirostris planirostris Andersen, Proc. Zool. Soc.,
London, p. 2387.
One specimen: Georgetown, skin and skeleton.
This specimen is best considered as p. planirostris on the
basis of measurements, although on geographical grounds it
should be p. fallax. The following measurements, total length
of skull, 28.5 mm.; mastoid width, 15.5; width of braincase,
12; maxillary width across m', 17.5; upper tooth row, c-m?, 10;
forearm, 61; all come closer to those of the former subspecies
than they do to those of the latter, as set forth by Andersen
in his monograph of the genus op. cit. page 246.
——
1921] Anthony: British Guiana Mammals 285
The skull lacks the m* on both sides and it is interesting
in this connection to note that three out of four of Andersen’s
only specimens of the planirostris group which lacked m* came
from British Guiana, page 234, op. cit. as did also an “unusu-
ally small specimen” of p. fallax, page 243.
Macrophyllum macrophyllum (Wied).
1825. Phyllostoma macrophyllum Wied, Beitr. zur Naturgesch. Brasil-
ien, Vol. II, p. 188.
One specimen (male) : Georgetown, April.
This specimen is in alcohol and is in excellent condition.
The skull has been removed and cleaned for examination. This
rare species is well characterized externally by the very high
and broad noseleaf, long, slender tragus and very extensive
interfemoral membrane, which has a series of longitudinal
rows of minute dermal papille along the proximal half of the
under surface. The most striking cranial characters are the
shelf-like surface just anterior to the external nares, the en-
larged median incisors above and the greatly reduced second
lower premolar, which is scarcely visible to the naked eye.
Length of forearm, 34.5 mm.
APPENDIX B
By WILLIAM BEEBE
The following unmistakable mammals have been observed
by me at the Station, or in some cases collected alive or dead,
and not preserved. I add them to complete the tentative list
of mammals of the Station.
57. Charonectes, Spr. ee ae eee (Water Opossum)
58: Desmoedus sp. #2 Is 10noot me ieee (Vampire Bat)
59/0 Jeticoyow BPs tL. DSK E8I9. AES: VS (Cae (Crab-dog)
60io Felis sp: YL Alaa DORN e ae eee (Puma)
612) Leopardus splot snes 20el Team (Ocelot)
62(; Myourotia sp. a... 280) Boia (Short-tailed Agouti)
68° 1 Coenduspn 320) 300s SiG ad i (Tree Porcupine)
64; Sevaritids spl cs) Shou Te thom (Dwarf Squirrel)
65 Prodovtessp: 201..9) TOs es (Giant Armadillo)
66 > Tapirus( spe i487 Oks Sy0as a Ipeie (Tapir)
OT Trichechus spt efor Boe: Bt Ore (Manatee)
68° 1g - AD) 2.2 ee (Fresh-water Dolphin)
69 -Leontocebus? spn ee (Marmoset)
10: Atéles Spi. ee eee eee (Spider Monkey)
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FROM THE TROPICAL RESEARCH
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es
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VOLUMB III, NUMBERS 14 AND 15
Tropical Research Station Contributions Numbers 134 and 135 aif
14. NEW BATRACHIANS
15. NEW LIZARDS
FROM THE TROPICAL RESEARCH STATION
BRITISH GUIANA
By G. KINGSLEY NOBLE
Associate Curator of Herpetology, in Charge,
‘American Museum of Natural History
owt one D BY j%+THE SOCIETY
SEPTEMBER 10, 1923
At
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Corrected to August, 1923
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
mew YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III, NUMBERS 14 AND 15
Tropical Research Station Contributions Numbers 1384 and 135
14. NEW BATRACHIANS
q 15. NEW LIZARDS
FROM THE TROPICAL RESEARCH STATION
BRITISH GUIANA
By G. KINGSLEY NOBLE
Associate Curator of Herpetology, in Charge,
American Museum of Natwral History
MEEBLISHED BY THE SO: O bh Bele
HE ZOOLOGICAL PARK, NEW YORK
SEPTEMBER 10, 1923
BRITISH
SCALE 58°942'W.
Oo!
5S MILES
LOCATION OF THE TROPICAL RESEARCH STATION OF THE
NEW YORK ZOOLOGICAL SOCIETY
The circle represents a radius of six miles. °
First form on press September 10, 1923
Volume III, Number 14
NEW BATRACHIANS
FROM THE TROPICAL RESEARCH STATION
BRITISH GUIANA
By G. KINGSLEY NOBLE
Associate Curator of Herpetology, in Charge,
American Museum of Natural History
‘Among the Amphibia secured by Mr. William Beebe or by
members of his staff while at the Tropical Research Station and
substations maintained by the New York Zoological Society in
British Guiana, there are included a number of new forms. The
reptiles and amphibians collected by Mr. Beebe have already been
reported upon in part (Beebe, 1919, Zoologica II, No. 7). Mr.
Beebe has been kind enough to place in my hands his new species
for description. The present paper deals with only the Amphibia
collected. A second paper will consider the new reptiles obtained
by Mr. Beebe. It is Mr. Beebe’s intention to publish later a full
account of the reptiles and amphibians found near the Tropical
Research Station, together with a series of colored plates illus-
trating most of the forms described below. I have added to the
following descriptions a diagnosis of one species found at the
Zoological Society’s Research Substation near the Kaieteur
Falls, but represented in the collections at my disposal only by
specimens secured by a former expedition.
Hyloxalus beebei sp. nov.
DIAGNOSIS
Readily distinguished from all other species of the genus by
its small size, rudimentary webs and brilliant coloration in life.
Preserved specimens are straw-color above with a dark stripe on
either side of the head and body, and with an irregular stippling
of dark brown on the back. From the species of Phyllobates,
which it closely resembles, this species may be distinguished by
the short but well-defined webs between the toes, and by its dis-
tinctive coloration.
* Tropical Research Station, Contribution Number 134.
290 Zoologica: N. Y. Zoological Society (IIT; 14
TYPE
A.M.N.H. No. A-18683; adult 9; near Kaieteur Falls,
British Guiana; February 18, 1921; William Beebe.
DESCRIPTION OF TYPE
Size very small; snout rounded, a trifle longer than the
greatest diameter of the eye; distance between nostril and tip of
snout a trifle less than the distance between nostril and eye;
interorbital space one and one-half times as broad as the upper
eye-lid; tympanum nearly two-thirds the diameter of the eye,
rather indistinct, less than its own diameter from the eye. Tibio-
tarsal articulations of either side in contact when the legs are
placed at right angles to the body; tibiotarsal articulation reach-
ing the posterior border of the eye. Digital dilations very small,
the discs less than one-half the diameter of the tympanum;
fingers free, the first finger not as long as the second; toes with a
short but very distinct rudiment of a web; no fringe extending
the length of the toes as in most species of Hyloxalus. Two
metatarsal tubercles, the outer pointed and one-third the
diameter of the inner which is not very distinct; a well-defined -
tarsal fold. Tongue ovate, slightly nicked behind. Skin smooth
above, slightly granular on the sides; no well-defined folds on the
back.
Ground tone in alcohol a pale straw-color; streak of dark
brown from tip of the snout through the eye to the groin, this
stripe very narrow on the head, broadening out behind the
tympanum until it equals the greatest diameter of the latter,
narrowing again on the sides of the body to disappear in the
lumbar region; dorsal surface slightly stippled with the same
brown tone; two pale streaks on either side of the back, extending
from the eye to the pelvic region, free of this dark stippling;
jaws and ventral surface immaculate; hinder and upper portion
of the thigh indistinctly stippled with dark brown; upper sur-
faces of the lower limbs stippled with the same dark tone, the
stippling tending to form three or more dark cross-bands. In
life, the ground tone was a bright yellow and the pattern was
much more distinct.
1923] Noble: New Batrachians, British Guiana 291
MEASUREMENTS
Sumer mneur to Vent 16.5 mm.
Tip of Snout to Posterior Border of Tym-
Li ee A TS oer 7
Greatest Breadth of the Head............... O25.
Distance from Axilla to Tip of Longest
SS Aes SS cc ee Leos
Distance from Vent to Tip of Longest Toe... 26,005
ae te.
REMARKS
This distinctive species of Hyloxalus is of special interest
because it seems to bridge the gap supposed to exist between
Hyloxalus and Phyllobates. The species agrees entirely with
Phyllobates in regard to its skeletal characters, and in most
external features is similar to that genus, but the presence of the
short web, much shorter than in any hitherto described species of
Hyloxalus, prevents our referring it to Phyllobates. Un-
doubtedly, Hyloxalus is the more primitive genus, and it would
be interesting to find the stock from which it arose. As I have
pointed out elsewhere (Noble, 1922, Bull. A.M.N.H., Vol. XLVI,
pp. 1-87), the brachycephalids have been derived from bufonid
ancestors. It is highly probable that the stock from which
Hyloxalus has been derived will be found in the Bufonidae.
The type specimen described above, although only 16.5 mm.
in total length, possessed seven eggs within the ovaries,
averaging 2 mm. in diameter. These eggs were heavily pig-
mented. All Salientia which lay eggs of such enormous size
deposit them on land. It is therefore very probable that
Hyloxalus, in spite of the web between its toes, is purely
terrestrial and lays its eggs in moist situations similar to those
utilized by Hleutherodactylus.
Hyla ornatissima sp. nov.
DIAGNOSIS
A medium sized Hyla possessing arched vomerine teeth
which form a shallow N; fingers two-thirds webbed, toes nearly
292 Zoologica: N. Y. Zoological Society [III; 14 |
entire webbed. Tibiotarsal articulation extending beyond the
snout. Gaudy coloration of pinks and browns; two dark, pink-
edged spots on the snout; a dark interorbital bar and a diamond
shaped spot just anterior to the pelvis, similarly edged with pink.
TYPE
A.M.N.H. No. A-13491; adult ¢ ; Meamu, Mazaruni R.,
British Guiana; June 10, 1920; William Beebe.
DESCRIPTION OF TYPE
Tongue as long as broad, unemarginate behind; vomerine
teeth in two slightly arched series nearly in contact with each
other and directed slightly forward; the anterior margin of the
arch on a level with the posterior border of the choanae. Head
exactly as long as broad; nostrils near the end of the snout;
interorbital space slightly more than half as great as the distance
between nostril and eye; head greatly flattened with concave
loreal region and prominent snout; interorbital space twice the
diameter of the upper eye-lid, slightly greater than the greatest
diameter of the eye; no ossification in the derm of the head;
tympanum distinct, its greatest diameter less than half the great-
est diameter of the eye. Tibiotarsal articulation extending
beyond the tip of the snout. Digits with well-defined discs, these
a trifle less than the greatest diameter of the tympanum; fingers
two-thirds webbed; toes fully webbed, except that the web hardly
extends beyond the base of the penultimate phalanx of the fourth
toe; no external evidence of a rudiment of the prepollex. Skin
smooth above, slightly granular on the belly ; no folds on the back
or sides of the body.
Ground color, in preserved specimens, pale straw-color or
yellowish; dorsal surface finely sprinkled with small pink spots;
a dark brown spot on either side of the snout edged with pink;
a dark spot on each upper eye-lid and an interorbital bar of the
same color, these outlined by the same pinkish tone; a dark
diamond-shaped figure just anterior to the pelvis, continued
posteriorly in a narrow coccygeal stripe; two small spots anterior
to the diamond-shaped figure of the same tone, this figure broadly
edged with pink, a few irregular pink blotches along the coccy-
1923] Noble: New Batrachians, British Guiana 293
geal stripe and near the anterior spots; tips of the toes brownish;
two dark spots on the forearm; a few small pinkish spots on the
‘upper surface of the limbs; a few small pinkish spots and three
or four dark ones on the upper surface of the hind limbs; ventral
surface immaculate except for the tips of the fingers and toes
which are brownish.
MEASUREMENTS
emer ent 10 Vent. 40.0 mm.
Tip of Snout to Posterior Border of Tym-
TE aa A a A ES a 12.5555"
Greatest Breadth of the Head..........0.--0. ow... Tar
Distance from Axilla to Tip of Longest
re ee ee eae eo
Distance from Vent to Tip of Longest toe______. 70.0 “
al i a a Zo
REMARKS
The species is represented in the collection by only a single
specimen which was captured by an Indian and brought to the
Zoological Station. Its gaudy coloration in life readily dis-
tinguishes it from the other Guianan hylas.
Leptodactylus stictigularis sp. nov.
DIAGNOSIS
A medium sized Leptodactylus, having a broad head; chest-
nut or reddish brown dorsal coloration; conspicuous whitish or
pink upper lip and a dark throat studded with white spots.
Tibiotarsal articulation extending to the middle of the eye; toes
bearing narrow dermal fringes; skin smooth or granular; two
well-defined dorso-lateral folds extending from eye to groin.
TYPE
A.M.N.H. No. A-10398; adult ¢ ; Kartabo, British Guiana;
1919; William Beebe.
DESCRIPTION OF TYPE
Size moderate; head flattened, a little broader than long;
snout distinctly longer than the greatest diameter of the eye; dis-
294 Zoologica: N. Y. Zoological Society [III; 14
tance between tip of snout and nostril contained twice in the ©
distance between nostril and eye; interorbital space one and —
one-third times as broad as the upper eye-lid, about equal to the
greatest diameter of the eye; canthus rostralis rounded but |
rather distinct; the loreal region concave; tympanum two-thirds
the greatest diameter of the eye, separated from the eye by a
space equal to half its own diameter. Tibiotarsal joints of either
side overlap when the legs are placed at right angles to the body;
tibiotarsal articulation, when extended forward, marks the mid-
dle of the eye. Digits slender, terminating in very small discs;
first finger very long; second finger only two-thirds as long as the
first; toes slender, bordered with a narrow fringe; sub-articular
tubercles pronounced; two metatarsal tubercles, the outer two-
thirds the length of the inner; a well-defined tarsal fold. Tongue
large, slightly nicked behind; vomerine teeth in two arched
groups nearly in contact with each other and touching the pos-
terior border of the choanae; the most anterior part of each arch
anterior to the posterior edge of the choanae. Skin smooth or
slightly granular; a well-defined dorso-lateral fold extending
from the eye to groin; ventral surfaces smooth, no abdominal
disc.
Ground tone chestnut brown; upper lips broadly edged with
gray; a grayish interorbital bar; the canthus and the dorso-
lateral fold edged with dark brown; the supra-tympanic fold of
the same color; posterior surfaces of thighs very dark brown;
three or four rows of irregular whitish spots studding this dark
area; three ill-defined dark bars on the dorsal surface of the
thighs, three or four on the lower limbs; soles of feet very dark
brown, parts of the lower leg suffused with the same color; sides —
of the body milky, stippled with the chestnut ground tone;
_ ground tone of ventral surface white; throat a dark brown
studded with numerous white spots; periphery of abdomen ~
densely stippled with brown; middle of abdomen and distal por-
tions of the thighs lightly stippled with the same color.
MEASUREMENTS
Tip of Snout:to, Vent. = eee 56.0 mm.
Tip of Snout to Posterior Border of Tym-
panuM |... ee 20:5
1923] Noble: New Batrachians, British Guiana 295
Srentest Isreadtin of Head...........................-- 25:0: «¥
Distance from Axilla to Tip of Longest
ee 34.
Distance from Vent to Tip of Longest Toe....84.0 “
Se ee 200. *
REMARKS
The species is represented in our collections by a single
paratype. It differs but little from the type in color. The dark
throat color is indistinctly continued on the anterior part of the
abdomen, but here the white spots have given place to white
blotches. The pale stripe on the upper lip is pink in this speci-
men, and the interorbital bar is of the same color.
L. stictigularis seems to be most closely aliied to L. rhodomy-
stax Boulenger, from which it differs in its longer leg, somewhat
differently arranged vomerine teeth and different coloration. L.
rhodomystax has been recorded from British Guiana by Ruthven
(1919 Occ. Papers Mus. Zool., Univ. of Mich., No. 69, p. 4.) I
have recently had the opportunity of examining Ruthven’s speci-
mens and find them to be unquestionably the young of L. penta-
dactylus. This leaves the question open as to whether or not
L. rhodomystax might not have been based upon a juvenile speci-
men of that species. This question can only be answered by
examination of the types.
Leptodactylus minutus sp. nov.
DIAGNOSIS
A minute Leptodactylus lacking fringes to the toes and with-
out dorsal folds; skin glandular, but never warty; an irregular
series of dark spots above, these sometimes forming an interor-
bital bar and a symmetrical pattern on the shoulders. Appar-
ently closely allied to L. pulcher Boulenger, from which it is
distinguished by its different color pattern and shorter leg;
somewhat similar to the immature L. caliginosus and L.
typhonius, distinguished from these by the characters already
mentioned.
296 Zoologica: N. Y. Zoological Society [IIIl; 14
TYPE
A.M.N.H. No. A-13495; adult ¢?; Bartica District, British
Guiana; January 8, 1916; William Beebe.
DESCRIPTION OF TYPE
Size very small; head distinctly longer than broad; distance
between tip of snout and nostril contained twice in distance be-
tween nostril and eye; interorbital width one and one-half times
as broad as the upper eye-lid, equal to the greatest diameter of
the eye; canthus rostralis rounded, the loreal region sloping
gradually; tympanum very distinct, one-half the greatest
diameter of the eye, separated from the eye by a trifle less than
half its own diameter. Tibiotarsal joints of each side strongly
overlap when the legs are placed at right angles to the body;
tibiotarsal articulations mark the anterior corner of the eye when
the legs are extended forward. Tips of digits not swollen into
dises, only slightly larger than the diameter of the penultimate
phalanges; first finger a trifle shorter than the second; toes
slender, not fringed; subarticular tubercles pronounced; two
well-defined metatarsal tubercles; a tarsal fold. Tongue ovoid,
slightly nicked behind; vomerine teeth in two slightly arched
series well behind the choanae, separated from each other by a
space equal to their distance from the choanae. Skin slightly
glandular but not warty above, a few feeble warts on the
sides of the body tending to form an ill-defined dorso-lateral
fold; ventral surface smooth, a very pronounced abdominal disc.
Ground tone of dorsal surface dull olive-gray; three pale
stripes running the length of the back, the two outer pale stripes
irregularly suffused with pink; the whole dorsal surface, except-
ing the pale bands, irregularly spotted with dark brown; about
twelve spots on the back and four on each side of the body ventral
to the pale streak; two or three dark spots on each upper jaw;
a series of dark spots forming cross bars on the limbs; hinder
surface of thighs suffused with brownish; ventral surfaces im-
maculate excepting for a delicate suffusion of brown on the
throat and hind limbs.
Bah) Aly 4
1923] Noble: New Batrachians, British Guiana 297
MEASUREMENTS
SS | 22.0 mm.
Tip of Snout to Posterior Border of Tym-
MEER ee Si
Sepmremirsreadiuh Of Plead: :2 2.02. s8 oes. ee ‘fe: Werk
Distance from Axilla to Tip of Longest
OL ee ae 2 ee eee 12.6.. “
Distance from Vent to Tip of Longest Toe........ a1.) &
SI a ee i | eet
REMARKS
The six paratypes show a considerable range of variation
in color; the ground tone may be very pale or somewhat darker
than in the type; a well-defined interorbital bar may be present
forming a regular cross with a scapular marking; the other spots
on the back may have a very symmetrical or irregular arrange-
ment; the dark spots on the upper jaw usually form a very
well-defined series; the three dorsal stripes are not as well-
defined in the paratypes as in the type; in three of the speci-
mens there is no indication of these light streaks.
Leptodactylus rugosus sp. nov.
DIAGNOSIS
A small species, very similar to L. caliginosus, but with a
broader head, shorter leg and very rugose dorsum. It is further
distinguished from that species by its large tympanum, different
coloration, and absence of nuptial spines in the breeding male.
TYPE
A.M.N.H. No. A-1169; adult 4; near Kaieteur Falls, British
Guiana; August 13, 1911; F. E. Lutz.
DESCRIPTION OF TYPE
Size small; head flattened, as long as broad; snout longer
than the greatest diameter of the orbit; interorbital space nar-
_ rower than the breadth of the upper eye-lid, much less than the
greatest diameter of the eye; canthus rostralis rounded; loreal
298 Zoologica: N. Y. Zoological Society (III; 14
region slightly concave; tympanum two-thirds the greatest
diameter of the eye, separated from the latter by a space less
than half its own diameter. Tibiotarsal joints of either side
barely overlap when the legs are placed at right angles to the
body; tibiotarsal articulation when extended forward reaches the
anterior border of the tympanum. Digits slender, without ter-
minal dilations; first finger longer than the second; toes slender,
without fringes; subarticular tubercles pronounced ; two metatar-
sal tubercles, the outer two-thirds the length of the inner; a well-
defined tarsal fold. Tongue large, slightly nicked behind;
vomerine teeth in two arched series on a level with the posterior
border of the choanae, and nearly in contact with each other in
the mid-line; anterior edge of each arch not extending forward
beyond the choanae. Skin very rugose above, some of the
tubercles forming short folds which run cephalo-caudad on the
back; upper eye-lids covered with numerous warts; a few warts
on the snout and dorsal surface of the limbs. ;
Ground tone above reddish brown, a few indistinct paler
marks on the upper lip, and numerous pale mottlings on the
posterior surfaces of the thighs; some indication of dark cross-
bands:on the upper and lower legs; ventral surfaces straw-
colored, heavily blotched with brown, the spotting darkest on
the throat and palest on the ventral surfaces of the thighs.
MEASUREMENTS
Tip of Snout-to Vent... —. 2 eee 38.0 mm.
Tip of Snout to Postrior Border of Tym-
panum 7 eee ee 15.5>
Greatest Breadth-of Head =.= aa 160733
Distance from Axilla to Tip of Longest
Finger 1.2.2 455 Ses eee 21525
Distance from Vent to Tip of Longest Toe........ 55.07
Tibia: .20..4.4,5 22 Oe eee 17.5
REMARKS
The four paratypes in our collection range from 16.5 to
41.0 mm. head and body length. They show very little variation
fede Oty,
1923] Noble: New Batrachians, British Guiana 299
in either coloration or structural characters. The pale bars on
the upper lip are very distinct in the largest specimen and some
indication of an interorbital bar is present. The venter of all
four paratypes is either immaculate or lightly spotted with
brown.
Volume III, Number 15
NEW LIZARDS
FROM THE TROPICAL RESEARCH STATION
BRITISH GUIANA
By G. KINGSLEY NOBLE
Associate Curator of Herpetology, in Charge,
American Museum of Natural History
Among the reptiles secured by Mr. William Beebe or by
his staff at the New York Zoological Society’s Tropical Research
Station in British Guiana, there are included a number of rare
and interesting forms. Two lizards in the collection are found
to be undescribed, while two others are reported from British
Guiana for the first time. The latter are Neusticwrus bicarinatus
(Linné), from Kartabo, and Cercosaura ocellata Wagler from
both Kartabo and the Bartica District. Many of the species
secured by Mr. Beebe, such as Sphaerodactylus molei Boettger,
were very imperfectly known. These Mr. Beebe is planning to
discuss in a later paper. The present paper is limited to
merely a description of the new forms in the collection. It
may be added that the species described below will be figured
in Mr. Beebe’s general account of the reptiles of the Research
Station.
Gonatodes beebei sp. nov.
DIAGNOSIS
A large Gonatodes of uniform reddish brown color above
and without spots on the throat or venter; a species having
not one but a series of spines over the eye, and having the
nostril not indenting the rostral.
TYPE
A.M.N.H. No. R-21251; adult ¢; Kartabo, British Guiana;
1921; William Beebe.
DESCRIPTION OF TYPE
Size large; distance from tip of snout to ear contained
exactly four times in the distance from snout to vent; greatest
diameter of the eye contained one and one-half times in the
* Tropical Research Station, Contribution Number 135.
302 Zoologica: N. Y. Zoological Society [Tiss
distance from tip of snout to eye; distance from tip of snout to
nostril contained twice in the distance between nostril and eye;
distance from snout to middle of eye decidedly greater than the
distance from ear to latter point; ear opening oval in outline,
about one-third the greatest diameter of the eye. Digits
cylindrical, proximal scales of their ventral surfaces very much
larger than distal ones, grading rather abruptly into the latter;
dorsal surfaces of head and body covered with coarse granules,
those of the head not larger than those of the body; granules
of the occiput slightly smaller than those of the snout; posterior
superciliaries slightly more pointed but not distinctly larger
than the supraorbitals; anterior superciliaries very much larger
than the supraorbital granules, three or four of these enlarged
superciliaries sharply pointed and forming a series of low spines
which project from the upper eye-lid just anterior to the mid
point of the eye; a large cleft rostral bordered posteriorly by
three small median granules, two large lateral scales and the
nostrils; nostrils in contact with the rostral but not indenting
it; five upper and four lower labials; mental large, pointed hbe-
hind, followed by two small scales larger than the smallest
labial, gular region with coarse granules of the same size as
those of the snout; ventral surfaces of the body and posterior
appendages covered with large cycloid, overlapping scales, these
of about three times the diameter of the dorsal granules; under
surface of the tail covered proximally with scales similar to
those of the abdomen, covered distally by one or two series of
very broad scales,—these scales three to five times as broad as
the abdominal scales.
Uniform reddish brown above, whitish immaculate below,
except for a slight suffusion of brown on the abdomen and
appendages, this suffusion tending to form dark edges to the
scales of the ventral surface of the thigh.
MEASUREMENTS
Tip of Snout to*Vent.{ eS eee 47.0 mm.
Tip-of ‘Snout:towiar i. 2 et eee LES es
Tip of Snout tetOrbit... as ae ae ee 5b
Greatest Width ofsHead 2.24) .33 29 10°35
Vent to Tip of Tall Us. 2 eee ATO
1923] Noble: New Lizards, British Guiana 303
REMARKS
The species is represented in our collections by only a
single specimen. The species may be readily distinguished
from other forms of Gonatodes by its large size, narrow head
and uniform coloration. It is perhaps allied to G. ferrugineus
_ described by Cope from Trinidad.
‘a> i Pe, Se eee ©
Leposoma taeniata sp. nov.
DIAGNOSIS
Very closely related to L. scincoides and L. dispar; inter-
mediate between these two forms in scutation, very different
_ from either in coloration; fronto-nasal obtusely angular poster-
- jorly; one anterior and three pairs of chin shields, the posterior
pair separated from each other by a single scale; scales of the
body strongly keeled, mucronate; dorsal scales forming trans-
- verse and oblique rows; ventral scales forming transverse and
longitudinal rows on the abdomen. Reddish brown above,
_ whitish below, a broad band of dark brown extending along
_ either side of the head and body.
TYPE
P A.M.N.H. No. R-21266; adult 2; Kartabo, British Guiana;
June 19, 1919; William Beebe.
DESCRIPTION OF TYPE
|
: Head narrow; fronto-nasal obliquely angular behind; a pair
of small prefrontals, not half as long as the frontal, slightly
larger than the fronto-parietals; two lateral parietals and an
4 enormous inter-parietal, the former about one-third as wide as
the latter; four supraoculars; all dorsal head shields roughened,
as in the other species of Leposoma; no loreal but two freno-
orbitals, the dorsal having twice the diameter of the ventral;
six upper and five lower labials; chin shields large, one anterior
and three pairs, the two anterior pairs in contact, the posterior
Pair separated by a single scale; a few enlarged scales posterior
_ to the chin shields; separated from the gulars by a single row
of small scales which extends across the throat from ear to
304 Zoologica: N. Y. Zoological Society [III ; 15.
ear; chin shields similar to the ventrals, but narrower and more
pointed. Body covered with uniform scales which form trans-
verse and oblique rows on the back; transverse and longitudinal
rows on the ventral surface; the scales about as broad as they
are long; strongly keeled, mucronate; 27 scales around the
middle of the body, 38 scales from the occiput to the base of
the tail and 40 from the third pair of chin shields to the vent;
4 pre-anals, three of these slightly larger than the ventral
scales; caudal scales like those of the body but the scales tend-
ing to form regular longitudinal and transverse rows as on the
venter; the keels of the caudal scales forming a series of ridges;
14 of these ridges around the tail, fifteen scales from the base.
Ground tone above, reddish brown, two broad stripes of
dark brown extending from the tip of the snout along the
entire length of head and body and about one-third the length
of the tail; no spotting on dorsal surface; a few dark spots on
the labials and sides of the head; sides of the body below dark
stripe, brownish, somewhat spotted; ventral ‘surface white,
immaculate except for four small spots on the chin shields and
a suffusion of brown on the ventral surface of the tail.
MEASUREMENTS
Tip o Snout' to Vent...2 = eee 34.0 mm.
Tip of Snout te Har: 2) eee tia
Tip. of Snout,.to Orbit. > ae eee 30'S
Greatest Width of Head... ... = eee 45-%
REMARKS.
The eight paratypes of this species in the collection differ
only slightly in color. They all exhibit the dark band on either
side of the head and body. In a few specimens there is some
indication of a pale, narrow band dorsal to this dark stripe. In
a few of the specimens there are a few flecks of dark brown
on the dorsal surface, but these never form the dark spots found
in the other species of the genus. This flecking is perhaps —
most distinct at the base of the tail.
Se ee
4
1923] Noble: New Lizards, British Guiana 305
There is very little variation in scutation. The scale counts
of six of the paratypes (two others are badly damaged) is as
follows:
Seales around the middle of the body average... 27.0
(Max. 29. Min. 26.)
Seales from occiput to base of tail average _._...__. 38.2
(Max. 39. Min. 37.)
Seales from 3d pair of chin shield to vent average 39.6
(Max. 40. Min. 39.)
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OSBORN, JR., ARTHUR A. FOWLER.
Glass uf 1926 re
HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES rie
DIETERICH, GEORGE F. BAKER, Wo. PIERSON HAMILTON,
ROBERT S, BREWSTER, EDWARD 8. HARKNESS, WILLIAM
B. Oscoop FIELD, EDWIN THORNE, PERCY A.
ROCKEFELLER, JOHN E. BERWIND,
IRVING K. TAYLOR.
Srientific Staff /
Witeriaki T. HorNabDAyY, Director of the Zoological Park;
W. REID BLAIR, D iN: S., Assistant to Director;
CHARLES H. TOWNSEND, Dircenoe of the Aquarium;
RAYMOND L. DITMARS, Curator of Reptiles; By
WILLIAM BEEBE, Honorary Curator of Birds and Director aS th p
Tropical Research Station;
LEE S. CRANDALL, Curator of Birds;
GEORGE S. HUNTINGTON, Prosector;
GEORGE A. MACCALLUM, Pathologist;
ELWIN R. SANBORN, Photographer and Editor.
Enditurial Committee
HENRY FAIRFIELD OSBORN, Chairman; set
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND.
Corrected to October, 1923
LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
16.
eT.
18.
29.
20.
21.
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III, NUMBERS 16-21
(Tropical Research Station Contributions 136-141)
DESCRIPTIONES TERMITUM IN ANGLORUM
GUIANA By F. SILVESTRI
NEW GENERA AND SPECIES OF TERMITOPHILOUS
COLEOPTERA By WILLIAM M. MANN
GLANDULAR STRUCTURE OF THE ABDOMINAL
APPENDAGES OF A TERMITE GUEST
By N. E. McINbDoo
TERMITOPHILOUS APTERYGOTA
By Justus W. FOLSOM
THREE APPARENTLY NEW SPECIES OF
TERMITAPHIS By HAROLD MORRISON
FOUR TERMITOPHILOUS MILLIPEDS
By RALPH W. CHAMBERLIN
mo BLIS HE D ay THE 5 O° C ig ts
foo ZOOLOGICAL PARK, NEW YORK
OCTOBER 1, 1928
BRITISH
BRAZIL
SCALE 58942'W.
=
5 MILES
Fig. 27. LOCATION OF THE TROPICAL RESEARCH STATION OF THE
NEW YORK ZOOLOGICAL SOCIETY
The circle represents a radius of six miles.
First form on press October 16, 1923
Volume III, Number 16
DESCRIPTIONES TERMITUM
IN ANGLORUM GUIANA REPERTORUM*
By F. SILVESTRI.
(Plates XI-XV).
Eutermes parvellus
(Plate XI).
Femina alata.—Corpus castaneum clypeo _isabellino,
pronoto macula isabellina T-formi signato, mesonoti et metanoti
parte antica isabellina, sternorum 1-6 parte mediana isabellina,
pedibus rufescentibus, alis rufo-brunis.
Caput fere 1/4 longius quam inter oculos latius, supra setis
sat numerosis brevibus et brevioribus instructum, fenestra per-
parva angustiore cum linea antica, clypeo bene inflato sat setoso,
‘ ejusdem dimidia parte c. 1/5 latiore quam longiore; oculis sat
magnis, bene convexis, prominulis, ocellis parvis, ab oculis
aliquantum minus quam ocelli diametros longitudinalis remotis,
antennis 14-articulatis, articulo tertio quam secundus fere
dimidio breviore et quam quartus parum longiore.
Pronotum subsemiellipticum, quam caput cum oculis 1/3
_minus latum, medium antice haud sinuatum, spatio brevissimo
gursum vergente, lateribus gradatim convergentibus, margine
postico rotundato; meso-et metanotum lateribus gradatim ali-
quantum convergentibus, angulis posticis rotundatis, margine
postico medio parum late et parum profunde inciso. Alae in
superficie tuberculis percrebris 5-6 radiatis et setis brevibus
sparsis instructa; venas vide Plate XI, 4, 5.
Pedes bene setosi; tibiae secundi paris setas vide Plate
mat, 7.
Abdominis tergita et sternita setis numerosis brevibus et
brevioribus, pleuris setis brevioribus sat numerosis instructis.
Cerci breviores setas vide Plate XI, 8.
* Tropical Research Station, Contribution No. 136.
308 Zoologica: N. Y. Zoological Society [Liss
Plate XI. EUTERMES PARVELLUS.
1, feminae caput pronum; 2, idem lateraliter inspectum; 3, scuta thoracalia;
4, 5, alae; 6, alae particula multo ampliata; 7, tibia pedum paris secundi;
8, cercus; 9, 10, militis caput pronum et lateraliter inspectum; 11, militis pes
paris secundi a tibia.
Long. corp. cum alis mm 8, sine alis 5, long. capitis 0,65,
ejusdem lat. inter oculos 0,52, diametros longitud. oculi 0,28,
long. antennarum 1,20, alae anticae 6,7, ejusdem lat. 1,7, long.
tibiae III, 0, 78.
Mas. Feminae similis.
Miles. Corpus stramineum abdomine cibi contenti causa maxima
pro parte cinereo.
Caput (Plate XI, 9-10) parum minus quam duplo longius
quam latius, subovale, naso sat longo et sat attenuato, super-
ficie setis brevioribus sat numerosis et setis nonnullis brevibus
1923 ] Silvestri: Descriptiones Termitum 309
instructa, antennis 12-articulatis, articulo tertio quam secundus
multo angustiore et c. 1/3 breviore, a quarto haud bene
separato et quam idem aliquantum angustiore et eidem longitu-
dine subaequali.
Pronoti lobus anticus parvus, margine medio paullum pro-
funde sinuato, antice setis nonnullis spiniformibus instructus,
postice setis 3+3 sat longis et nonnullis brevioribus; meso- et
metanotum parce setosa lateribus rotundatis.
Pedes sat setosi; secundi paris vide Plate XI, 11, tarsorum
articulis 1-3 apice infero elongato, acuto.
Abdominis tergita et sternita setis sat numeroris breviori-
bus et brevibus instructa. Cerci apice acuto instructi.
Long. corporis mm. 2,5, long. capitis 0,98-1,04, ejusdem
lat. 0,56, long. nasi ab antennarum foveae margine antico mensi
0,39, antennarum 0,90, tibiae III, 0,52.
Operarius militi similis capite aeque longo atque lato, antennis
13-articulatis, articulo tertio a quarto vix distincto et quam
secundus angustiore et c. dimidio breviore.
Long. corp. mm. 2,8-3, lat. capitis 0,58, long. antennarum
0,78, tibiae III, 0,38.
Observatio. Species haec ad Eutermes microsoma Silv. per-
proximum sed imaginis oculis parum majoribus et magis
prominentibus, militum capite, praesertim naso, breviore facile
distinguenda est; ab Eutermes holmgreni Banks militis capitis
latitudine etiam distincta.
Habitat. Anglorum Guiana: Exempla vidi alata, nec non
milites et operarios apud Kartabo cl. A. Emerson collecta, et
milites et operarios cl. G. E. Bodkin lecta.
Hamitermes excellens sp. nov.
(Plate XII).
_ Femina alata. Corpus supra brunum clypeo latericio, ore et
.
abdominis tergito ultimo rufescentibus, urosternitis lateraliter
brunis, cetera superficie pallide isabellina, pedibus rufescenti-
bus, alis brunis.
310 Zoologica: N. Y. Zoological Society [III; 16
Plate XII. HAMITERMES EXCELLENS SP. NOV.
1, feminae caput pronum; 2, idem lateraliter inspectum; 3, scuta thoracalia;
4, 5, alae; 6, alae particula; 7, tibia paris secundi; 8, militis caput pronum;
9, ejusdem labrum; 10, militis tibia secundi paris postice inspecta; 11, militis
pes primi paris a tibia.
Caput paullum longius atque inter oculos latius, supra setis
brevibus et brevioribus numerosis instructum, fenestra magna
subovali, parum longiore quam latiore, clypeo bene inflato et
bene setoso, ejusdem dimidia parte c. 1/4 longiore quam latiore;
oculis sat magnis, convexis, prominulis, ocellis parvis, ab oculis
quam ocelli diametros longitudinalis parum minus remotis,
antennis 15-articulatis, articulo tertio quam secundus c. dimidio
et quam quartus c. 1/3 breviore, articulo quarto quam quintus
parum longiore.
Pronotum quam caput cum oculis parum minus latum,
medium antice haud sinuatum, brevi spatio sursum vergens
lateribus parum longe ab angulo antico multo convergentibus
margine postico subrecto, meso-et metanotum lateribus ali-
are
1923] Silvestri: Descriptiones Termitum 311
quantum convergentibus, margine postico medio late et parum
profunde sinuato, angulis posticis rotundatis.
Alae superficie setis brevioribus sparsis et aliis per venas
dispositis et tuberculis percrebis 5-6 radiatis instructa; venas
vide Plate XII, 4, 5.
Pedes bene setosi, secundi paris tibiis, praeter calearia duo
interna, seta spiniformi parum robusta praeapicali externe
aucta.
Abdominis tergita et sternita setis pernumerosis breviori-
bus et setis brevibus instructa. Cerci breviores, bene setosi.
Long. corp. cum alis mm 12, sine alis 6,8, long. capitis 1,23,
ejusdem lat. inter oculos 1,17, diametros long. oculi 0,39, long.
antennarum 1,68, alae anticae 10,2, ejusdem lat. 2,7, long. tibiae
III, 1,38.
Mas feminae similis.
Miles. Corpus cremeum vel ochroleucum capite ferrugineo,
mandibulis parte distali nigrescente, abdomine cibi contenti
causa cinereo. .
Caput parum longius quam latius, late convexum, lateribus
partem anticam versus parum convergentibus, angulis posticis
rotundatis, superficie supera setis paucis instructa, fontanella
labro subsemielliptico, paullum longiore quam ante basim latiore,
basi ipsa parum angustiore, setas vide Plate XII, 8, mandi-
bulis quam capitis latitudo aliquantum brevioribus, dente sat
magno ad basim partis distalis instructis, parte apicali bene
arcuata, attenuata, antennis 15-articulatis, articulo tertio quam
secundus et quam quartus 1/3 breviore, articulo quarto quam
quintus parum breviore, in antenna nonnulla a tertio haud
bene separato.
Pronotum quam capitis latitudo c. dimidio minus latum,
lobo antico sursum vergente, medio parum sinuato, margine
postico subrecto, lateribus rotundatis, superficie supera setis
nonnullis sat longis et brevibus instructa, lobi antici super-
ficie antica setis nonnullis brevissimis sat robusti instructa;
meso- et metanotum lateribus rotundatis, supra setis nonnullis
sat longis et brevibus praesertim posticis instructa.
als Zoologica: N. Y. Zoological Society [IT ais
Pedes sparse setosi, primi paris tibiis setis nonnullis inter-
nis robustis, secundi paris tibiis etiam seta spiniformi attenuata
externa instructa.
Abdominis tergita et sternita setis sat longis sat numerosis
et setis nonnullis brevibus et brevioribus instructa. Cerci
articulo secundo quam primus c. 1/3 longiore, conico, acuto,
setis consuetis.
Long. corp. (cum mandibulis) mm 6, long. capitis (sine
mandibulis) 2,10, ejusdem lat. 1,76, long. antennarum 2, mandi-
bularum 1,28, tibiae Th, 1,°32:
Operarius. Corpus cremeum capite ochroleuco, abdomine cibi
contenti causa cinereo.
Caput fere 1/9 latius quam longius, supra setis sat longis et
brevibus instructum, clypeo sat inflato ejusdem dimidia parte
paullum longiore quam latiore, antennis 15-articulatis, articulo
tertio quam secundus ec. dimidio et quam quartus parum
breviore.
Thorax et abdomen eisdem militis similia.
Long. corp. mm 5, long. capitis 1,10, ejusdem lat. 1,8, long.
antennarum 1,30, tibiae III, 1,10.
Habitat. Anglorum Guiana: exempla alata et militem et
operarium ad Kartabo (ab A. Emerson) lecta et exempla alia
ad Potaro a Bodkin vidi.
Observatio. Species haec magnitudine et militis forma ab
Ham. hamifer Silv. distinctissima est.
Capritermes bodkini sp. nov.
(Plate XIII.)
Nympha feminilis. Corpus stramineum.
Caput aliquantum longius quam inter oculos latius, supra
setis paucis brevibus instructum, oculis parvis, fuscis, parum
prominentibus, antennis 16-articulatis, articulo tertio divi-
sionem proximalem vix distinctam monstrante, secundum longi- —
tudine aequante et quam quartus vix longiore.
Silvestri: Descriptiones Termitum
Plate XIII. CAPRITERMES BODKINI SP. NOV.
1, militis caput pronum; 2, idem lateraliter inspectum; 3, ejusdem
labrum; 4, pronoti lobus anticus a facie postica inspectus; 5, pes
paris secundi a tibia; 6, nymphae tibia primi paris; 7, nymphae
pedis primi paris a tibia apex.
313
314 Zoologica: N. Y. Zoological Society [III;16
Pronotum quam caput cum oculis c. 1/5 minus latum,
postice aliquantum late et parum profunde sinuatum. Ap-
pendices alares long. mm 2,4.
Pedes primi paris (Plate XIII, 7) ut ceteri calcaribus
duobus instructi, parce setosi.
Long. corp. mm 8, long. capitis 1,62, ejusdem lat. inter
oculos 1,35, long. antennarum 3, tibiae III, 2,2.
Miles. Corpus ochroleucum capite ochroleuco vel ochraceo
mandibulis nigris. Caput fere 1/4 longius quam postice latius
supra parce convexum lateribus partem posticam versus
gradatim parum divergentibus, labro (Plate XIII, 3) sub-
rectangulari apice subtriangulari, parum minus quam duplo
longiore quam latiore, setis paucis brevibus distalibus et setis
sat numerosis brevissimis instructo, mandibula laeva quam
caput parum breviore, forma vide (Plate XIII, 1-2), mandibula
dextera quam laeva aliquantum breviore, apice triangulari
acuto, antennis 16-articulatis, articulo tertio secundum longi-
tudine aequante et quam quartus paullum breviore.
Pronotum quam caput c. dimidio minus latum, lobo antico
magno, margine supero medio parum inciso, facie antica setis
brevissimis conicis sat numerosis, margine supero setis nonnullis
brevibus et brevioribus, facie postica setis nonnullis brevioribus
instructa; mesonotum postice subrotundatum, metanotum mar-
gine postico late et sat profunde sinuato.
Pedes longi, parce setosi, primi paris ut ceteri calcaribus
duobus armati.
Abdominis tergita setis sat numerosis brevibus et sternita
etiam setis sat numerosis brevioribus instructa. Cerci bene
setosi, parte apicali longiuscula attenuata.
Long. corp. mm 11, long. capitis 3,8, ejusdem lat. 3, long.
mandibulae laevae 3, antennarum 4,2, tibiae III, 3.
Operarius. Corpus ochroleucum, abdomine cibi contenti causa ~
cinereo.
Caput subaeque longum atque latum, fenestra magna, setis
nonnullis brevioribus instructum, clypeo bene inflato, antennis
i
.
; 1923] Silvestri: Descriptiones Termitum 315
16-articulatis articulo tertio nudo a quarto haud bene separato,
quam secundus 1/3 breviore et quartum longitudine subaequante.
Thorax et abdomen eisdem militis similia.
Long. corp. mm 5,5, long. capitis 1,35, long. antennarum
2,20, tibiae III, 2.
Habitat. Anglorum Guiana: Potaro River (Bodkin legit. Dec.,
1915).
_ Observatio. Species haec pedibus primi paris calcaribus duobus
tantum instructis, militis capitis et labri forma, pedum longi-
tudine distinctissima est.
Capritermes bodkini Silv. modestior var. nov.
(Plate XIV.)
Femina alata. Caput et thoracis tergita luride testacea, clypeo
aliquantum pallidiore, abdominis tergita rufescentia, sternita
ochraceo-ferruginea, pedibus testaceo rufescentibus, alis? (in
exemplo typico abruptis).
Caput parum longius (c. 1/11) quam inter oculos latius,
supra setis sat numerosis minimis et nonnullis_ brevibus,
fenestra magna, obcordiformi, bruna, prominula,_ clypeo
-parum inflato transverse subrectangulari ejusdem dimidia parte
aliquantum latiore quam longiore; oculis sat magnis, bene con-
vexis, ocellis sat parvis ab oculis ocelli dimidia longitudine
distantibus, antennis?-articulatis (in exemplo typico haud
integris, articulis 12-sistentibus), articulo tertio secundo sub-
aequali et quam quartus paullum longiore.
Pronotum quam caput cum oculis aliquantum minus latum,
parum magis quam duplo latius quam longius, medium antice
_ vix sinuatum, brevi spatio sursum vergente, angulis anticis late
4
.
rotundatis, lateribus aliquantum convergentibus, angulis posticis
late rotundatis, margine postico late et parum profunde sinuato ;
meso-et metanotum lateribus postice aliquantum conver-
gentibus, margine postico parum late et aliquantum profunde
- sinuato, angulis posticis obtusis.-
Alae? (abruptae).
4
316 Zoologica: N. Y. Zoological Society [II]; 16
a
> =
Po ees
f
} /
fifi
Plate XIV. CAPRITERMES BODKINI SILV. MODESTIOR VAR. NOV.
1, feminae caput pronum; 2, idem lateraliter inspectum; 3, scuta thoracalia; 4, pes
paris secundi a tibia; 5, militis caput pronum; 6, idem lateraliter inspectum;
7, ejusdem labrum; 8, militis pronoti lobus anticus postice inspectus; 9, militis
pes secundi paris a tibia; 10, operarii tibia primi paris.
Pedes bene setosi, tibiae setis internis sat robustis.
Abdominis tergita et sternita setis pernumerosis breviori-
bus et setis brevibus nonnullis instructa. Cerci breviores, bene
setosi.
Long. corp. cum alis ?, sine alis 9, long. capitis 1,70,
ejusdem lat. inter oculos 1,56, diametros long. oculi 0,59, long.
antennarum ? , alae anticae ? , ejusdem lat. ? , long.
tibiae lil, 2.
Miles. Corpus stramineum, capite cremeo, mandibulis nigris.
1923] Silvestri: Descriptiones Termitum 317
Caput 1/6 longius quam postice latius, supra parce con-
vexum, lateribus a parte antica ad posticam aliquantum diver-
gentibus, labro subrectangulari, apice tantum subtriangulari
parum minus quam duplo longiore quam postice latiore, parte
distali supra setis nonnullis brevioribus et aliis brevissimis
instructa, mandibula laeva capitis longitudinem aequante, forma
vide (Plate XIV, 5-6), mandibula dextera quam laeva ali-
quantum breviore apice triangulari, antennis 16-articulatis,
articulo tertio quam secundus c. 1/4 breviore et quam quartus
parum longiore.
Pronotum quam capite c. dimidio minus latum, lobo antico
magno, margine supero late rotundato medio vix inciso, facie
antica setis brevissimis conicis sat numerosis, margine supero
setis nonnullis brevibus et brevioribus, facie postica setis
nonnullis brevioribus instructa, mesonotum postice vix sinu-
atum, metanotum late et sat profunde sinuatum.
Pedes longi, parce setosi, secundi paris vide (Plate XIV,
9).
Abdominis tergita et sternita setis nonnullis brevibus et
aliis parce numerosis brevioribus vel (in dorso) brevissimis
instructa. Cerci apice subconico longiusculo.
Long. corp. mm 10, long. capitis (sine mandibulis) 3,
ejusdem lat. 2.5, long. antennarum 3, 6, mandibulae laevae 3,
tibiae III, 2,5.
Operarius. Corpus stramineum, abdomine cibi contenti causa
cinereo.
Caput c. 1/6 longius quam latius, ad antennas latius, supra
seta nonulla breviore instructo, clypeo bene inflato, ejusdem
dimidia parte parum latiore quam longiore, antennis 15-arti-
culatis, articulo tertio quam secundus aliquantum angustiore et
parum breviore, quam articulus quartus haud longiore et parum
angustiore.
Thorax et abdomen ejusdem militis similia.
Long. corp. mm 4, long. capitis 1, ejusdem lat. 1,24, long.
‘ antennarum 1,85, tibiae III, 2,4.
318 Zoologica: N. Y. Zoological Society [III ; 16
Habitat. Anglorum Guiana: exempla typica ex Kartabo ab A.
Emerson lecta.
Syntermes parallelus sp. nov.
(Plate XV).
Femina alata. Corpus castaneum, capitis parte antica testacea,
mandibulis praeter apicem nigrum testaceis, thoracis pleuris et
sternitis testaceo-latericiis areis avellaneis interruptis, antennis
pedibusque testaceis, alis fuligineis, abdominis ventre ochraceo.
Caput subaeque longum atque inter oculos latum, supra —
setis paucioribus sparsis instructum, fenestra magna, circulari, ;
clypeo convexiusculo, ejusdem dimidia parte fere 1/5 latiore
quam longiore, oculis sat parvis valde convexis, ocellis (capite
prono) ab oculis quam oculi diametros longitudinalis parum ~
minus distantibus, antennis 19-articulatis,.articulo secundo :
quam tertius haud vel paullum breviore et quam quartus ali- i
quantum longiore. 4
Pronotum quam caput cum oculis vix minus latum, medium
antice paullum, postice aliquantum sinuatum, angulis. anticis
acutis haud productis, angulis posticis late rotundatis, meso- et
metanotum postice lateribus parum convergentibus, mesonoti
margine postico latissime, metanoti vix sinuato.
‘Alae superficie setis brevioribus numerosis vestita; venas —
vide Plate XV, 4-5.
Pedes setuloli, tibiae latere interno setis brevibus sat
robustis numerosis instructo, calearibus apicalibus elongatis.
Abdominis tergita setis nonnullis praesertim posticis, —
sternita setis numerosis brevioribus instructa. .
Long. corp. cum alis mm 29, sine alis 17, long. capitis cum —
mandibulis 3,7, sine mandibulis (labro excluso) 2,8, ejusdem —
lat. inter oculos 2,6, diametros longitudinalis oculi 0,59,. long.
antennarum 5, alae anticae 24,5, ejusdem lat. 5,8, long. tibiae
TT, As 4
1923] Silvestri: Descriptiones Termitum 319
Plate XV. SYNTERMES PARALLELUS SP. NOV.
1, feminae caput pronum; 2, idem Jateraliter inspectum; 3, scuta thoracalia;
4, 5, alae; 6, alae particula multo ampliata; 7, militis caput pronum; 8, ejusdem
labrum; 9, militis scuta thoracalia; 10, operarii scuta thoracalia.
Mas ignotus.
Miles. Corpus fulvo-testaceum capite ochraceo-ferrugineo,
mandibulis parte distali nigra.
Caput ce. 1/3 longius quam latius lateribus parallelis,
angulis posticis late rotundatis, setis brevioribus paucis sparsis
instructum, supra pone fontanellam bene convexum, antice a
fontanella ad labrum discendens et utrimque ad antennas ali-
quantum inflatum, fontanella supra tuberculum minimum tubi-
formem sita. Labrum (deplanatum) parum latius quam
longius, antice angustatum, subtriangulare, utrimque ad partis
triangularis basim excisum, lateribus ceteris postice parum con-
vergentibus, setis vide (Plate XV). Mandibulae sat robustae,
quam capitis latitudo c. 2/9 breviores, dextera dente minimo ad
partis distalis basim, laeva a dente sat magno et dentibus
———
320 Zoologica: N. Y. Zoological Society BON gags
duobus minimis, praeter basalem, armata, parte distali
attenuata, apice breviter arcuato, acuto. Antennae 19-
articulatae, articulo secundo quam tertius vix et quam quartus
aliquantum longiore, articulo decimo duplo latiore quam
longiore.
Scuta thoracalia in processum lateralem brevem (metanoti
mm 0,48) acutum, spiniformem, extrorsum et aliquantum sur-
sum vergentem producta; pronoti lobus anticus margine medio
vix sinuato.
Pedes longi, breviter setosi.
Abdomen tergitis setis paucis praesertim posticis, sternitis
setis sat numerosis intructis.
Long. corp. mm 12, long. capitis cum mandibulis 6,8, sine
mandibulis 4,5, antennarum 4,6, tibiae tertii paris 3, lat.
metanoti cum spinis 2,5.
Operarius major. Corpus fulvo-testaceum antennis latericiis,
abdomine cibi contenti causa fusco-cinereo. Caput subaeque .
longum (mandibulis exclusis) atque latum lateribus postice
parum convergentibus, fenestra circulari manifesta, supra setis
paucioribus sparsis instructum, clypeo subplano setis 3-4
instructo; antennis 19-articulatis, articulo secundo quam
tertium haud vel vix breviore et quam quartus aliquantum
longiore, articulo decimo magis quam duplo longiore quam
latiore.
Scuta thoracalia in processum lateralem brevem (metanoti
mm 0,20) acutum, spiniformem producta; lobus anticus margine
medio vix sinuato.
Abdomen tergitis setis paucis praesertim posticis, sternitis
setis sat numerosis intructis.
Long. corp. mm 8, lat. capitis 2,6, long. antennarum 6,
tibiae paris tertii 2,6, metanoti lat. 2.
Operarius minor. Corpus ochroleucum, abdomine cibi contenti
causa cinereo.
Caput parum latius quam (mandibulis exclusis) longius
fenestra magna, antennis 18-articulatis, articulo secundo quam
1923] Silvestri: Descriptiones Termitum 321
tertius haud vel parum breviore et quam quartus paullum
longiore.
Scuta thoracalia in processum lateralem perbrevem (meta-
noti mm 0,12) acutum producta.
Long. corp. mm 7, lat. capitis 1,9, long. antennarum 3,5,
tibiae paris tertii 2, metanoti lat 1,56.
Habitat. Anglorum Guiana: Cattle Trail Survey, Canister Falls
VI. 1920. A. A. Abraham legit.
Observatio. Species haec ad S. molestus Burm. proxima est,
sed magnitudine majore, corporis et alarum colore, militis
capitis forma et spinis thoracalibus bene distincta est.
we
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Volume III, Number 17
NEW GENERA AND SPECIES
OF TERMITOPHILOUS COLEOPTERA
FROM NORTHERN SOUTH AMERICA*
By WILLIAM M. MANN.
Bureau of Entomology, Department of Agriculture, Washington, D. C.
(Figs. 28-40)
Mr. Alfred Emerson has had the kindness to loan me for
study a most remarkable series of beetles that he collected
in termite nests, chiefly at the Tropical Research Station at
the New York Zoological Society at Kartabo, British Guiana,
but containing also several species from Trinidad.
The present paper includes merely descriptions of some of
the forms It is intended at a later date, to publish a more
extended account of these, together with descriptions and notes
of others.
Holotypes of the new species have been deposited in the
U. S. National Museum collection.
The accompanying habitus drawings have been made by
Mr. Robert E. Snodgrass, Mr. Harry B. Bradford and Miss
Julia Ellen Edmonson.
Family—STAPHYLINIDAE
Subfamily—ALEOCHARINAE
Tribe—CoOROTOCINI
Spirachtha mirabilis sp. nov.
(Fig. 28)
Female.—Length 2-2.50 mm. Membranous portions white,
in some of the larger specimens irregularly infuscated; chitin
on abdomen and tarsi pale brown, on head and thorax darker
brown, antennae and legs black. Feebly shining.
* Tropical Research Station, Contribution Number 137.
FIG. 28. SPIRACHTHA MIRABILIS-SP. NOV.
1923] Mann: Termitophilous Coleoptera 325
Basal abdominal segment, the central surface of thorax and
the appendages with erect hairs, which are pale yellow in color
and longer on the abdomen and more sparse and black else-
where.
Head about as long as broad, barely broader in front than
behind, with a narrow median impressed submembranous area
for nearly the length of front; sides posterior to eyes moder-
ately convex; anterior border of front truncate; clypeus trans-
verse, broadly concave at anterior border. Eyes one-third as
long as sides of head, situated at front of sides. First antennal
joint nearly as long as head, slender at base then gradually
thickened and feebly sinuate to apex; second joint longer than
broad and distinctly longer than the third; third joint twice as
long as broad, joints 4-10 elongate, each a little shorter than
the one preceding; terminal joint a little thicker than the others,
and one and two-thirds times as long as the penultimate. Pro-’
notum transverse, surface evenly and feebly convex, sides
rounded, anterior and posterior borders broadly arcuate;
separated from head by a membranous collar and with a
smaller membranous pad visible at each side of the posterior
border. Scutellum large, broadly triangular. Elytra together
broader than long, separated for their entire length, posterior
angles narrowly rounded, posterior borders nearly straight and
strongly oblique. Abdomen nearly covering the remainder of
body, second and third segments strongly inflated, rounded at
apex; segments 3-7 with narrow transverse bands of chitin
above; segments 4, 5, 6 with massive, constricted exudatorial
developments, those on segments 5-6 terminating as large
irregular masses, tuberculate and each tubercle bearing a stiff
hair; on the fourth segment long, irregularly cylindrical,
sharply bent at middle and with numerous constrictions, not as
distinctly tuberculate as the other two pair Legs short and
rather stout
Host.—Nasutitermes (Constrictotermes) cavifrons (Holm-
gren).
Type-locality—Kartabo, British Guiana.
Differs from S. ewrymedusa Schiédte in its longer head, the
first antennal joint is less thickened and less arcuate, the pro-
326 Zoologica: N. Y. Zoological Society [Ts 4%
notum lacks the broad, posterior membranous band, the elytra
are separated for their entire length and their posterior borders
are much more strongly oblique. The large masses at the
ends of the two posterior pairs of exudatoria are characteristic.
In eurymedusa they are slender and similar in form to the other
pair and the posterior pair has one strong constriction instead
of many feeble ones. The amount of membranous structure
on the pronotum is variable in mirabilis and probably also in
eurymedusa.
Spirachtha schiodtei sp. nov.
Female.—Length 1.50 mm.—Membranous portions white;
chitin on abdomen pale brown, on head, thorax and appendages
darker. Moderately shining, finely coriaceous.
Erect, stiff yellowish hairs scatered on abdomen (except
on chitinous plates). Fine, black, semi-recumbent hairs on head,
thorax and appendages.
Head a little broader than long; median membranous area
broader than in mirabilis; front broadly impressed. Antennae
stout, first joint nearly as long as the head, suddenly thickened
near base and only slightly and very gradually enlarged toward
tip, two and one-half times as long as the second joint; second
joint as long as the third, less than twice as long as broad,
narrow at base; joints 4-10 subequal, a little longer than broad;
terminal joint one and two-thirds times as long as the penul-
timate. Pronotum broader than long, widest in front of middle.
Elytra together broader than long, posterior corners project-
ing and narrowly rounded. Abdomen recurved as in ewry-
medusa and mirabilis but (in the majority of the specimens)
extending only to anterior third of pronotum; exudatoria well
developed, the anterior two pair constricted at middle into two
portions; subequal in length, the apical part somewhat pear-
shaped, not tuberculate but beset with bristles similar to those
on second and third abdominal segments; posterior pair of
exudatoria similar but with three ‘‘joints,” the middle of which
is small and the terminal a little larger than on the others.
Legs a little more slender than in mirabilis.
1923]. Mann: Termitophilous Coleoptera 327
Host.—Nasutitermes (Constrictotermes) cavifrons (Holm-
gren).
Type-locality.—Kartabo, British Guiana.
Other locality —Kalacoon, British Guiana.
This is very distinct from the preceding species in its much
smaller size, more robust antennae, broader head with more
strongly impressed front and in the radically different struc-
ture of the abdominal exudatoria.
One specimen in a vial with a series of the females repre-
sents a phase of the male of this species. The structure of the
head and thorax is similar to that of the female, except the
median membranous band on the head is much narrower. The
elytra are separated for their entire length (in the female of
schiodtei they are connate as in euwrymedusa) and less narrowed
at the tips. The abdomen is small, entirely chitinized, com-
pressed dorso-ventrally and margined at sides, quite a typical
Staphylinid abdomen. The antennae of the specimen are
broken but the basal joint is proportionately shorter than in
the other specimens.
Among the series with inflated abdomens there is some
variation in the length of the antennal joints and in individuals
cleared in xylol I have noted differences in the internal chitinous
structures in the terminal abdominal segments, which indicates
that both sexes are present. So the one that I take for the
male may be a phase in the imaginal development of the female.
In fully developed specimens it is probable that both sexes
are very similar in habitus.
It appears certain that we have two species of Spirachtha
living in the same locality with the same host termite, another
character similar to that of Corotoca, which Schiddte records
in the same manner.
Corotoca guyanae sp. nov.
Female.—Length (in alcohol, with abdomen recurved)
2.60 mm.
328 Zoologica: N. Y. Zoological Society [Tis a7
Chitinous portion dark brown, except on median abdominal
segments, where it is higher; lateral part of gaster yellow.
Shining and every finely coriaceous.
Erect hairs fine on head, thorax and elytra, course on
abdomen and shorter on appendages; one pair on vertex, numer-
ous on ventral surface of head; scattered and sparse on thorax
and elytra; abundant on gaster and arranged in three rows on
each dorsal band, more scattered on the sides.
Head, excluding eyes, a little longer than broad, front
shallowly impressed and, between eyes, with a v-shaped suture
and a median vertical impression extending about one-third the
length of head. Clypeus transverse, broadly concave at anterior
border. Eyes large and convex, occupying two-thirds the sides
of head. First antennal joint elongate, clavate, concave at
outer border, about two-thirds as long as head; second joint
half as long as the third, which is three times as long as broad,
remaining joints gradually shorter, the penultimate about twice
as long as broad; apical joint slender, shorter than the two
preceding joints together. Pronotum a little broader than long,
separated from head by a membranous collar, anterior border
broadly concave; sides very broadly and obtusely angulate at
anterior third, straight and moderately convergent posterior
to this; surface trifoveolate, the anterior pair of foveae more
elongate and less impressed than the posterior, median one;
surface between the impressions low and rounded. Elytra
together broader than long, sides nearly straight, posterior
angles narrowly rounded, posterior border convex. Abdomen
greatly inflated and projected forward; apical segment com-
pletely chitinized, penultimate segment chitinized on apical half,
remaining segments (visible from above) largely membranous,
each with a narrow chitinous band. Legs slender, posterior
tibiae slender.
Host.—Nasutitermes (Constrictotermes) cavifrons (Holm-
gren). :
Type-localityKartabo, British Guiana.
Described from seven specimens taken from several nests
of the host species.
1923] Mann: Termitophilous Coleoptera 329
C. guyanae resembles C. melantho Schiddte in having the
front foveolate, and C. phyllo Schiddte in having the dise of
pronotum trituberculate and the posterior tibiae linear instead
of fusiform. It differs from phyllo in its broader pronotum,
the much less pronounced tubercles and from both of the
described species in the shape of the elytra, which have the
outer, posterior angles broader and not acuminately produced.
The maxillary palpi are markedly different in guyanae, the
third joint instead of being thickly oval and about as long as
the second, is greatly enlarged, nearly three times as long and
twice as broad as the second.
Thyreoxenus gen. nov.
Female.—Small, robust species, largely membranous,
abdomen strongly inflated and permanently projected over
meso- and metanotum. Head very small, broader than long,
not constricted behind. Labrum membranous, broad, rather
narrowly concave at middle of anterior border. Mandibles
small, slender, arcuate, acute at tips. Mentum broad, trape-
zoidal, nearly straight in front. Maxillary palpi 4-jointed, basal
joint small, second joint slender, third joint thicker and oval,
terminal joint small, awl-shaped. Ligula short and broad, in-
distinctly bilobed. Labial palpi 3-jointed. Antennae 11-jointed,
first joint scapiform, remaining joints (except the apical)
short; apical joint elongate, its sides parallel to near apex, then
converging into a short, connate tip. Eyes small, oval, situated
at sides of head. Prothorax large and convex, three times as
broad as head and partly enveloping it in front; membranous
except for a transverse chitinous plate on anterior half of
dorsum and very narrow lateral bands that connect it with a
transverse band on the sternum; dorsal plate divided at middle
by a longitudinal incision, which does not extend to the anterior
or posterior borders; prosternum with a median, elongate,
lobe-like membranous production. Meso- and metasternum
strongly chitinized, not separated by a suture, elongate, sub-
quadrate, obtusely carinate at middle, the carina terminating
anteriorly i in a flat, triangular area; posterior border projecting
as a broad, obtuse triangle between the posterior coxi. Elytra
small, strongly divergent posteriorly, each about twice as long
330 Zoologica: N. Y. Zoological Society (Ita
as broad, with straight, subparallel sides, posterior border
oblique, with corners rounded. Wings reduced to elongate, oval,
pad-like vestiges. Abdomen strongly inflated, elevated and
projected forward, five ventral segments visible from above;
dorsal sclerites (not visible except when gaster is straightened)
chitinized; ventral sclerites largely membranous but with thinly
chitinous plates in front at middle; apical and penultimate
sclerites chitinous, between them with a large, rounded collar-
like membranous inflation.
Legs short and rather stout, femora irregularly constricted
at middle. Tarsi 4-4-4 jointed; metatarsi short. Anterior and
middle coxi widely separated, elongate, conical; posterior coxi
less widely separated and shorter.
Male.—Similar in form to female, but smaller and more
slender, with the chitin on the ventral (upper) abdominal
sclerites limited to well defined, transverse bands at the anterior
border of each segment except the terminal, which is chitinous
throughout.
Genotype.—Thyreoxenus parviceps, sp. Nov.
Though evidently a member of the tribe Corotocini Thyre-
oxenus in an aberrant genus, even in this aberrant tribe, and
sharply distinct from the others in the structure of the abdomen,
the unusual development of the membranous areas, in the collar-
like development near the tip of the abdomen and especially in
its very small head.
It is represented in the collection by three species, two
small and largely white in color and one large form with dark
brown pigmentation over nearly the entire body. Each appears
to be a guest of a different species of Nasutitermes.
Thyreoxenus parviceps sp. nov.
Length.—(In alcohol) 2.50-2.75 mm.
Membranous portion white; chitin of head, pronotum and
the femora and tibia dark brown, elsewhere pale brown tarsi
1923] Mann: Termitophilous Coleoptera 331
white. Appendages and abdomen with rather short, erect black
hairs, their points of insertion on the abdomen marked by
small, fuscous dots.
Head a little broader than long, sides behind eyes nearly
straight. Eyes occupying anterior half of sides of head. Basal
antennal joint as long as the four following joints together;
SS
i hi
i
|
nie
~~
oo Na
FIG. 29. THYREOXENUS PARVICEPS SP. NOV.
joints 2-10 subequal, only slightly larger apically, a little
broader than long; terminal joint a little shorter than the two
preceding joints together. Median membranous projection of
mesosternum longer than broad (seen from behind) and
rounded at apex; from beneath, about twice as broad as long.
Posternum transversely impressed posterior to middle. Chitin-
ous bands on upper (ventral) abdominal sclerites narrow,
arranged in three sided figures.
Host.—Nasutitermes (nasutitermes) costalis (Holmgren).
Type-locality.—St. Joseph, Trinidad.
Other localities.—Georgetown and Kartabo, British Guiana.
332 Zoologica: N. Y. Zoological Society [TUL $17
Thyreoxenus pulchellus sp. nov.
(Fig. 30)
Female.—Length 2.50 mm.
Differing from Th. parviceps in the arrangement of the
chitinous areas on the upper surface of the abdomen; the penu-
litimate segment is evenly chitinous, the third (from the apex)
nea)
\
TT,
Nee mn
\
a) 2 NS gee eee
ret
FIG. 30. THYREOXENUS PULCHELLUS SP. NOV.
is chitinous at the sides and in front and membranous at middle,
the latter portion not bordered by a strongly chitinized margin;
the fourth segment is thinly chitinous, except for a narrow,
median surface rounded in front; the fifth segment is still more
thinly chitinized except in front and the median, entirely mem-
branous portion is oval in shape; the last segment (visible
from above) has a transverse bar in front, similar to parviceps.
The collar-like, membranous projection between the apical and
penultimate segments is proportionately larger than in the
other species.
Host.—Nasutitermes (Nasutitermes) ephratae (Holm-
gren).
Type-localityx—Kartabo, British Guiana.
Thyreoxenus major sp. nov.
Length.—(In alcohol) 3 mm.
Larger and more robust than parviceps and much darker
in color, with the upper surface of the abdomen brown and the
1923] Mann: Termitophilous Coleoptera 333
second, third and fourth (from the apex) segments each with a
large, quadrate median blotch considerably darker than the rest.
Sides of prothorax thinly chitinized and infuscated.
Head subopaque; chitinized portions of thorax and abdomen
shining. Median plate on pronotum with sparse, fine, erect
hairs, much shorter than those on abdomen. Hairs on abdomen
long, rather stiff and erect. Sides of prothorax uneven, im-
pressed at middle, roundly tuberculate in front. Membraneous
collar-like process on abdomen nearly absolete. Each segment
of the abdomen (above) broadly impressed posteriorly to the
front border; front borders elevated at middle into a rather
coarse and blunt carina, which is distinctly lighter in color than
the surrounding surface.
Host.—Nasutitermes (Nasutitermes) guayanae (Holm-
gren).
Type-locality—Kartabo, British Guiana.
Described from a unique specimen.
Eburniola gen. nov.
Head small, elongate, thin in profile; only slightly narrowed
behind; vertex with a strong, narrow, longitudinal impression
that terminates on the front in a broad impression, which is
bordered arcuately in front; anterior portion of front not pro-
duced, truncate. Clypeus membranous, strongly transverse,
concave at front border. Labrum corneous, broad, anterior
border scarcely concave. Submentum nearly oval, disc with a
pair of conspicuous setae. Mentum small, slightly trans-
verse; excised at middle of anterior border. Ligula very small
and slender, apparently simple. Labial palpi 3-pointed, minute.
Maxillary palpi 4-jointed; basal joint very small, second joint
oval, a little longer than broad; third joint twice as long as
broad, moderately compressed, about as broad basally as at
apex, the sides little convex, rather strongly setose; terminal
joint less than half as long as the third, broadly awl-shaped.
Eyes situated in front of sides of head, little convex, their outer
border on a plane with outline of sides of head, in outline
4-sided rather than oval, with the nearly straight sides
334 Zoologica: N. Y. Zoological Society (Tits ae
separated by rounded angles. Antennae situated in very small
sockets at sides of anterior border of front, 11-jointed, very .
thick; first joint nearly as long as the three following joints
together, thickly clavate and about half as long as head; second
joint transverse; joints 3-10 thick, about as long as broad,
subequal, quadrate in outline and not separated by constric-
tions; terminal joint elongate oval, one and two-thirds times
as long as the penultimate.
Pronotum strongly impressed over the greater portion of
the disc, anterior border elevated and narrowly rounded,
posterior border sloping; sides inflexed, concealing side pieces.
Elytra well developed, separated from front to rear by a narrow,
triangular fissure. Chitinous portion of posternum transverse,
truncate in front and behind. Mesometasternum longer than
broad, median portion flat; posterior border truncate and not
at all projecting between coxae. Abdomen permanently re-
curved, large and deeply and broadly inflated, largely mem-
branous, nearly flat above, the marginal line weakly impressed ;
ventral segments strongly convex, with strong impressions
between; six segments visible from above. Legs short, femora
and tibae not flattened, the middle and posterior pair somewhat
arcuate. Anterior coxae elongate, conical, posterior and middle
shorter and conical, all moderately separated, the posterior more
widely than the others. Tarsi 4-4-4 jointed, middle and
posterior metatarsi elongate, but shorter than the remaining
joints together.
Genotype—Eburniola leucogaster sp. nov.
Eburniola leucogaster sp. nov.
(Fig. 3i)¢
Length.— (With abdomen straightened) 1.50-1.75 mm.
Pale brown, abdomen mostly ivory white, with the chitinized
portions a little darker brown than the head and thorax.
Gaster, especially at sides, strongly shining, remainder feebly
shining and coriaceous. Head, thorax and elytra without erect
hairs; pubescence sparse and exceedingly minute. Dorsal
abdominal segments with an inconspicuous, erect hair on each
eS
1923 ] Mann: Termitophilous Coleoptera 335
side; apical and penultimate segments thinly covered with
recumbent, very fine yellow hairs. Basal ventral segments
with abundant long and fine erect hairs, which become sparser
and shorter on the apical segments and are lacking on the
penultimate and terminal segments. Appendages with moder-
ately abundant erect hairs.
Oe,
eae:
== Wa os
FIG. 31. EBURNIOLA LEUCOGASTER SP. NOV.
Head distinctly longer than broad, slightly narrowed in
front. Eyes less than half as long as head. Antennae reaching
to posterior end of elytra, very stout, nearly one-third as
broad as head. Pronotum a little broader than long and a
little broader than the head; anterior and posterior corners
rounded. Elytra at base slightly broader than the pronotum,
broader than long, sides divergent, arcuate at basal half, nearly
straight at apical half, posterior border straight, inner angles
broadly rounded, outer angles subacute. Abdomen longer than
the remainder of body, more than twice as broad as the elytra;
dorsal chitinous plates 1-4 very short, extending the width of
the segments, each with the anterior margin broadly concave
at middle; penultimate and apical segments entirely chitinized,
subquadrate in shape, strongly transverse, less than half as
long as the segments, extending at sides as thin lines which
become obsolete before attaining the dorsum. Anterior femora
of subequal width throughout; middle and posterior tibial very
gradually enlarged from base to apex.
336 Zoologica: N. Y. Zoological Society [Tlie
Host.—Nasutitermes (Nasutitermes) guayanae (Holm-
gren)..
Type-locality—Kartabo, British Guiana.
Other locality.—St. Joseph, Trinidad.
Described from a series of ten specimens.
Perinthus tarsatus sp. nov.
Length.—2.50 mm.
Dark brown, abdomen and appendages paler than head and
thorax, moderately shining, with abundant very fine, short
yellow pubescence and in addition with coarse*’and long black
hairs arranged as follows: five on pronotal margins; two on
elytral margins; one on each elytron; a series of four (inter-
spersed with shorter and finer hairs) at the posterior border of
each abdominal segment above and from four to six beneath.
Antennae with a series of long hairs at the apex of each joint.
Head a little broader than long, front not margined, rather
flat, vertex convex. Antennae distinctly longer than the head
and pronotum together, basal joint slightly shorter than the
second and third together, joints 4-11 strongly compressed, a
little longer than broad; terminal joint suboval, one and one-
third times as long as the penultimate. Pronotum about one-
third broader than long, narrowest in front, sides convex,
anterior and posterior corners broadly rounded, anterior border
concave, posterior border concave at sides, slightly convex at
middle. Elytra basally as broad as the pronotum, sides feebly
convex, posterior angles obtuse, posterior border nearly straight.
Abdomen longer than remainder of body, connate, first five
segments margined. Tarsal claws long and very slender.
Host.—Nasutitermes (Nasutitermes) surinamensis (Holm-
gren).
Type-locality—Kartabo, British Guiana.
P. tarsatus is nearest to silvestrii, but that species has
antennal joints 7-10 slightly transverse, the terminal joints is
twice as long as the penultimate and the body above is not
setose.
1923] Mann: Termitophilous Coleoptera 337
Perinthus wasmanni sp. nov.
Length 1.60-2 mm.
Dark reddish brown, elytra darker than the rest and the
abdomen and appendages paler. Moderately shining and
microscopically punctate and covered with rather abundant,
fine and short yellow pubescence and in addition fine, long and
erect hairs arranged as follows: six on lateral margins and
twelve on the disc of pronotum; two at the lateral margins
and two on the disc of each elytron; a row of four at the
posterior border of each abdominal segment above and a row
of six to eight beneath. Antennae with shorter and rather stiff
hairs. Legs with dense short hairs.
Head broader than long, rounded at sides, front flat, vertex
convex. Antennae slightly shorter than the head and prothorax
together ; the first joint a little shorter than joints 2-3 together,
joints 2-3 suboval and equal in length; joints 4-10 strongly
compressed, subequal, slightly broader than long, terminal joint
nearly as long as the two preceding joints together. Pronotum
about one and two-fifths as broad as long, narrowest in front,
sides moderately concave, anterior and posterior corners
rounded, posterior border concave at sides, convex at middle.
Elytra as broad basally as the pronotum, together more than
twice as broad as long, sides nearly straight, anterior corners
broadly rounded, posterior corners obtusely angulate.
Host.—Nasutitermes (Nasutitermes) ephratae (Holmgren).
Described from a small series.
This species in habitus closely resembles dudleyanus Casey
from Panama but may be distinguished by the relatively short
elytra. In dudleyanus the length of the suture is nearly two-
thirds that of the pronotum; in wasmanni it is less than half.
Perinthus vestitus sp. nov.
Length 1.90-2 mm.
Reddish brown, elytra darkest, abdomen and legs lightest.
Head, thorax and abdomen very distinctly, though shallowly
338 Zoologica: N. Y. Zoological Society (TEs
punctate and thickly covered with semi-recumbent, yellow pile
much longer than that of wasmanni or tarsatus and in addition
with a series of four strong black hairs on the lateral borders
of pronotum, two on the outer margins of the elytra and a row
on the ventral abdominal segments; abdomen also with long,
fine and yellow hairs on the posterior margins of the segments.
Head barely broader than long; front flat, vertex convex.
Antennae much shorter than head and pronotum together, first
joint nearly as long as the second and third together, second
and third joints elongate, sub-cylindrical, remaining joints
compressed, all a little longer than broad, terminal joint nearly
as long as the two preceding joints together. Pronotum about
one-fourth broader than long, only slightly narrower in front
than behind, sides feebly convex, anterior border nearly
straight, posterior border feebly convex at middie. Elytra a
little narrower than pronotum, together much broader than
long, sides nearly straight, posterior corners angulate, posterior
borders straight. Abdomen rather broadly margined.
Host.—Nasutitermes (Nasutitermes) octopilis (Banks).
Type-locality—Kartabo, British Guiana.
The comparatively long and dense pubescence and the
arrangement of the setae, with the more elongate and different
shaped pronotum distinguish vestitus from the other species of
Perinthus. The long hairs on the dorsal segments of the ab-
domen are unusually fine and are yellow in color, similar to
the shorter hairs.
Tribe Oxypodini
Termitogaster simulans sp. nov.
(Fig. 32).
Length (with abdomen straight) 3 mm.
Head, thorax and elytra dark brown; front of head,
abdomen, ventral surface and antennae reddish brown, legs and
palpi yellowish brown. Rather strongly shining and finely
alutaceous; outer and inner inflexed portion of elytra densely
1923] Mann: Termitophilous Coleoptera 339
punctate. Erect hairs on head, thorax and elytra, short and
fine, several at sides of occiput, four rows of 4-5 each on
pronotum and a row of four on each elytron; each dorsal
sclerite of abdomen with two pairs of longer, black, erect hairs,
one pair at middle of basal border and one at middle of poster-
ior third, the apical borders with a row of long recumbent
FIG. 32. TERMITOGASTER SIMULANS SP. NOV.
yellow hairs; sides of dorsum and the sides and ventral surface
with moderately dense yellow pubescence; appendages with
short and rather stiff, flattened, oblique yellow hairs.
Head about as long as broad, distinctly widest at occipital
border which is broadly arcuate; front concave on either side,
longitudinally convex at middle to opposite antennal surface,
where there is a broad, transverse elevation with an oblique
anterior face and truncate borders. Labrum distinct, convex
at sides, strongly excised at middle of anterior border. Eyes
prominent, oval, a little less than half as long as head. First
340 Zoologica: N. Y. Zoological Society [IITs 27
antennal joint a little shorter than head, in cross section sub-
quadrate; second joint nearly four times as long as broad and
one and two-thirds times as long as the second; third joint
more than half as long as the second, distinctly longer than
road; joints 4-10 gradually shorter, the tenth scarcely longer
than broad; terminal joint elongate-oval, narrowed and rounded
at tip. Pronotum a little longer than broad, with nearly
straight subparallel sides, with strongly arcuate anterior border
and rounded angles, posterior border, narrowly and shallowly
concave at middle; surface with a longitudinal very shallow
median impression in front of middle and a flattened triangular
area at middle near posterior border. Elytra at base as broad
as pronotum, similar to those of simopelta but with the posterior
margin concave and the outer angles less projecting. Abdomen
large, evidently not held erect, in all the specimens before
me it is strongly compressed and about twice as long as broad.
it may not be as strongly compressed in life, as a distinct
marginal line is present, but it is so in the alcoholic as well as
in the dried specimens before me); segments comparatively
long, each with a transverse carina posterior to middle; mem-
branous portions narrow, not visible in all specimens. Legs
slender, middle and posterior femora strongly flattened and
bent; tibiz moderately flattened; metatarsi shorter than the
remaining tarsal joints.
Host.—Nasutitermes (Nasutitermes) costalis (Holmgren).
Type-locality—Kartabo, British Guiana.
Described from nine specimens taken with several colonies
of the host termite.
In three of my specimens the sixth abdominal segment has
the border evenly emarginate, as described by Casey in the
male of Termitogaster fissipennis, the others, presumably fe-
males, have the border produced, but rounded instead of
angulate.
The resemblance to a worker termite is strong even in
dried and pinned specimens and must be strikingly so in the
J
2
1923] Mann: Termitophilous Coleoptera 341
living beetle. From the structure of the abdomen, it is evidently
carried straight, or but slightly elevated.
Termitogaster simopelta sp. nov.
(Fig..33).
Length (with abdomen straightened) 2.50 mm.
FIG. 33. TERMITOGASTER SIMOPELTA SP, NOV.
Color pale yellowish brown, elytra a little darker than the
remainder; exposed membraneous portions white. Integument
alutaceous, shining, the gaster more than the rest.
Erect hairs short and rather stiff, black in color, lacking
on head, ten on disc of pronotum, two on each elytron; a row
of 4-6 at the middle of each dorsal abdominal sclerite (the
posterior border of each sclerite with a thin row of recumbent
hairs) ; more abundant on ventral surface of abdomen; short
on appendages where there are also abundant, fine and short,
erect yellowish hairs. Ventral surface of thorax with dense
and very fine yellow pubescence.
Head about as long as broad, vertex transversely convex,
front concave, except near anterior margin where there is a
transverse ridge, thick basally and acute above (appearing
dentiform from the side), anterior border obliquely concave at
sides, acutely excised at middle, with a thin, sharp triangular
erect tooth at either side of incision. Clypeus very short and
broad, strongly concave at middle of anterior border. Labrum
342 Zoologica: N. Y. Zoological Society [IITs ee
transverse, longitudinally impressed at middle and convex at
sides. Eyes oval, rather strongly convex, nearly half as long
as head, situated in front of middle of sides. First antennal
joint a little shorter than head, in cross section subquadrate,
the sides somewhat flattened, and seperated from each other
by an angle; second and third joints a little thicker than the
others and about twice as long as broad, following joints
searcely longer than broad and decreasing in length to the
penultimate which is as broad as long; apical joint oval, less
than twice as long as broad and narrowed at tip. Pronotum
about as long as broad, sides and anterior border nearly
straight, posterior border rounded at middle, dise broadly con-
cave in front of middle. Elytra together longer than broad at
base; narrowly connected at base, the remainder broadly’
separated; each elytron two and one-half times as long as
broad, with feebly concave outer border, straight innner border
and concavely oblique posterior border with the outer angles
slightly produced. Abdomen widely inflated, two and one-half
times as broad as the pronotum, all segments, except the apical,
margined; membranous portions narrow, in some specimens
not visible between the dorsal sclerites. Legs moderately long;
femora and tibiae strongly flattened; posterior femora on upper
border concave at basal five-eighths, then oblique, the margin
forming, in outline, a broad triangle posterior to middle;
anterior tibiae with a brush of coarse, rather flat yellow hairs,
arranged in two rows; each metatarsus a little shorter than the
other tarsal joints together.
Host.—Nasutitermes (Nasutitermes) costalis (Holmgren).
Type-locality—Kartabo, British Guiana.
Termitogaster emersoni sp. nov.
(Fig. 34).
Female.—Length (in alcohol and with abdomen straight-
ened) 3 mm. Color brown, pronotum dark brown to black,
membranous portions white. Shining. Sparse erect hairs on
head, three rows of four each on pronotum, one on each elytral
4 eee a. ee
1923] Mann: Termitophilous Coleoptera 343
disc, one at the side of each dorsal abdominal sclerite (a row
of recumbent hairs on the apical margins) and scattered and
more abundant on the vertical surface.
Head a little broader than long, front broadly concave,
sides posterior to eyes feebly convex, occipital border broadly
rounded; anterior border very slightly concave. Labrum trans-
FIG. 34. TERMITOGASTER EMERSONI SP. NOV.
verse, broadly concave in front. Eyes oval, less than half as
long as head and situated near anterior border. First antennal
joint as long as head, second and third joints nearly three
times as long as broad, joints 3-10 proportionately shorter, but
all distinctly longer than broad, decreasing in length apically;
terminal joint less than twice as long as the penultimate.
Maxillary palpi with the basal joint very small, second joint
broader than long, its greatest width at anterior border, which
is nearly straight and rather pointed at the angles, outer sur-
face convex, inner surface concave; third joint as long as the
second, longer than broad, gradually narrowed apically; apical
joint minute and spiniform, less than half as long as the third.
Joints of labial palpi very small and slender. Mandibles stout,
acuminate apically, thickened at middle. Pronotum wider
than the head, about as broad as long and slightly broader in
front than behind, sides and posterior border convex, the angles
344 Zoologica: N. va Zoological Society [LI 5 aa
broadly rounded, anterior border moderately projecting and
rounded at the middle; surface in front of middle with a pair
of broad and rather shallow impressions and behind middle
with a pair of smaller and less distinct pits. Scutellum distinct,
broadly triangular. Elytra shorter than the pronotum, sep-
arated at apex by a V-shaped area about half their length,
together much broader than long, humeri subgibbous, sides
feebly arcuate, posterior angles projecting and narrowly
rounded, posterior border concave. Abdomen widely inflated,
longer than the remainder of body, more than twice as broad
as elytra, sides arcuate, broadly margined, the segments sep-
arated by broad bands of membrane. Legs moderately long
and slender, middle and posterior metatarsi long, anterior
metatarsus one and one-half times as long as the second joint.
Host.—Nasutitermes (Nasutitermes) ephratae (Holmgren).
Type-locality—Kartabo, British Guiana.
Other locality.—St. Joseph, Trinidad.
Described from a series of specimens taken from several
nests.
This species is near T. brevis but distinct in its larger
size, more elongate structure, the more shallowly impressed
front of head and in the strongly impressed pronotum and the
separated elytra, as well as in having the membranous portions
more developed than in the other species of Termitogaster.
Termitogaster brevis sp. nov.
Female.—(in alcohol and with abdomen straightened)
2.75 mm.
Chitinous parts brown, pronotum dark brown to black,
membranous portions white. Shining, microscopically punctate.
Erect hairs black, fine, scattered on anterior portion of head;
arranged in four rows of five on the pronotum; one on each
elytron at base; four on each abdominal sclerite above, shorter
on ventral surface; posterior margin of each abdominal seg-
ment with a row of very fine recumbent hairs.
tt @ alte
1923 ] Mann: Termitophilous Coleoptera 345
Head broader than long, vertex rather strongly longitud-
inally impressed at middle and with more shallow impressions
lateral to this; anterior portion of front broadly and shal-
lowly impressed tranversely and separated from the remainder
by a carinae that extends transversely in an irregular arc
across the front; anterior border broadly and strongly excised.
Eyes oval, convex, occupying anterior half of sides of head.
First antennal joint as long as head, somewhat flattened
basally; second and third joints subequal, less than twice as
long as broad, remaining joints gradually shorter to the penul-
timate, which is scarcely longer than broad, terminal joint
shorter than the two preceding joints together. Pronotum
slightly transverse, a little broader in front than behind, sides
and posterior border feebly convex, anterior border nearly
straight, posterior angles broadly rounded; surface with a
strong pit at middle and a more shallow pair in front of this,
posterior portion with a shallow transverse groove near the
border. Elytra connate, together strongly transverse, broadest
behind, sides nearly straight, posterior border very broadly
concave, posterior angles rounded, not produced. Abdomen
short and thick, more than twice as broad as elytra, with
rather weak lateral margins. Legs short and rather slender.
Femora and tibiae moderately flattened.
Host.—Nasutitermes (Nasutitermes) costalis (Holmgren).
Type-locality—Kartabo, British Guiana.
Other locality.—St. Joseph, Trinidad.
Described from a series taken in nests of the host.
Superficially this species appears identical with the geno-
type, T. insolens Casey from Panama, but comparison with the
type shows a number of differences in the structure of the
head, which, in insolens, is comparatively longer; the median
vertical impression is feeble, the anterior portion of front is
not so distinct from the remainder and the anterior border is
broadly arcuate, instead of excised as in brevis.
The maxillary palpi of brevis are shorter and broader than
in emersoni, the third joint is oval, much less than twice as
long as broad and distinctly shorter than the second.
336 Zoologica: N. Y. Zoological Society (III; 17
Host.—Nasutitermes (Nasutitermes) guayanae (Holm-
gren)..
Type-locality—Kartabo, British Guiana.
Other locality.—St. Joseph, Trinidad.
Described from a series of ten specimens.
Perinthus tarsatus sp. nov.
Length.—2.50 mm.
Dark brown, abdomen and appendages paler than head and
thorax, moderately shining, with abundant very fine, short
yellow pubescence and in addition with coarse*’and long black
hairs arranged as follows: five on pronotal margins; two on
elytral margins; one on each elytron; a series of four (inter-
spersed with shorter and finer hairs) at the posterior border of
each abdominal segment above and from four to six beneath.
Antennae with a series of long hairs at the apex of each joint.
Head a little broader than long, front not margined, rather
flat, vertex convex. Antennae distinctly longer than the head
and pronotum together, basal joint slightly shorter than the
second and third together, joints 4-11 strongly compressed, a
little longer than broad; terminal joint suboval, one and one-
third times as long as the penultimate. Pronotum about one-
third broader than long, narrowest in front, sides convex,
anterior and posterior corners broadly rounded, anterior border
concave, posterior border concave at sides, slightly convex at
middle. Elytra basally as broad as the pronotum, sides feebly
convex, posterior angles obtuse, posterior border nearly straight.
Abdomen longer than remainder of body, connate, first five
segments margined. Tarsal claws long and very slender.
Host.—Nasutitermes (Nasutitermes) surinamensis (Holm-
gren).
T ype-locality—Kartabo, British Guiana.
P. tarsatus is nearest to silvestrii, but that species has
antennal joints 7-10 slightly transverse, the terminal joints is
twice as long as the penultimate and the body above is not
setose.
1923] Mann: Termitophilous Coleoptera 337
Perinthus wasmanni sp. nov.
Length 1.60-2 mm.
Dark reddish brown, elytra darker than the rest and the
abdomen and appendages paler. Moderately shining and
microscopically punctate and covered with rather abundant,
fine and short yellow pubescence and in addition fine, long and
erect hairs arranged as follows: six on lateral margins and
twelve on the disc of pronotum; two at the lateral margins
and two on the disc of each elytron; a row of four at the
posterior border of each abdominal segment above and a row
of six to eight beneath. Antennae with shorter and rather stiff
hairs. Legs with dense short hairs.
Head broader than long, rounded at sides, front flat, vertex
convex. Antennae slightly shorter than the head and prothorax
together ; the first joint a little shorter than joints 2-3 together,
joints 2-3 suboval and equal in length; joints 4-10 strongly
compressed, subequal, slightly broader than long, terminal joint
nearly as long as the two preceding joints together. Pronotum
about one and two-fifths as broad as long, narrowest in front,
sides moderately concave, anterior and posterior corners
rounded, posterior border concave at sides, convex at middle.
Elytra as broad basally as the pronotum, together more than
twice as broad as long, sides nearly straight, anterior corners
broadly rounded, posterior corners obtusely angulate.
Host.—Nasutitermes (Nasutitermes) ephratae (Holmgren).
Described from a small series.
This species in habitus closely resembles dudleyanus Casey
from Panama but may be distinguished by the relatively short
elytra. In dudleyanus the length of the suture is nearly two-
thirds that of the pronotum; in wasmanni it is less than half.
Perinthus vestitus sp. nov.
Length 1.90-2 mm.
Reddish brown, elytra darkest, abdomen and legs lightest.
Head, thorax and abdomen very distinctly, though shallowly
348 Zoologica: N. Y. Zoological Society [TIT ; ee
as broad; joint longer than the two preceding joints together.
Pronotum longer than broad, widest at anterior corners which
are rounded, sides feebly concave, anterior, posterior borders
arcuate, surface moderately and evenly convex. Scutellum
small, transverse. Elytra together much broader than long
and broadest behind, each less than twice as long as broad;
FIG. 35. CORYMBOGASTER MIRANDA SP. NOV.
humeri elevated and convex, sides at middle exceedingly shal-
lowly concave, behind straight and divergent, and forming,
with the oblique, nearly straight posterior border a distinct
angle; inner corners broadly angulate. Abdomen nearly four
times as broad as elytra and in profile three times as deep as
metathorax; dorsal surface with an elevated median portion,
feebly convex and separated from the flat lateral areas by a
poorly defined longitudinal impression; sides separated by a
strong marginal impression.
Host.—Cornitermes (Cornitermes) pugnax, Emerson.
Type-locality—Kartabo, British Guiana.
Described from six specimens.
—- °°
1923] Mann: Termitophilous Coleoptera 349
The description and figure based on the specimens which
have the abdomen the least distended. In these the mem-
branous portions are of limited to small portions along the
margins of the dorsal surface. Other specimens have the
membrane developed to such an extreme that the chitonized
portions appear as small plates, the large gibbosites at the sides
are greatly reduced, with the chiton showing only as a pair of
-very narrow, parallel plates. In one the abdomen is at least
three times as long as broad and even on the dorsum the plates
extend scarcely half its width.
Termitophya punctata sp. nov.
(Fig. 36).
Length.—2.75-3 mm.
Head, thorax, elytra and appendages dark brown; abdomen
pale brown, each segment with a darker median blotch. Shin-
ing. Head with coarse regular separated punctures; smooth
between. Pilosity stiff and erect, moderately long, very sparse
except at the tip and on the ventral surface of the abdomen
and on the legs, two of them near the inner border of eyes,
three near the lateral borders of pronotum and two pairs on
each elytral disc; one at middle of margin, stout semi-recumbent
black hairs in a row of six at the posterior margins of the
dorsal abdominal segments; head, thorax and elytra with very
minute recumbent whitish hairs.
Head a little broader than long, front and vertex broadly
convex, sides immediately in back of eyes subparallel, anterior
border nearly straight. Labrum broad, strongly though very
narrowly excised at middle, membraneous at middle, corneous
at sides, the sides elevated into low tubercles in front. Eyes
moderately convex, occupying front half of sides of head.
Antennae a little longer than head and pronotum together;
first joint swollen, much broader than the others and nearly
as long as the second and third together; second joint shorter
than third, remaining joints subequal, terminal joint shorter
than the two preceding joints together. Pronotum somewhat
broader than head, a little longer than broad, broadest in front
350 Zoologica: N. Y. Zoological Society (IIT; ie
of middle and as broad in front as behind; anterior border
straight, posterior border evenly convex, anterior corners
rather narrowly rounded, posterior corners very broadly
rounded, sides only feebly convex. Elytra as broad as prono-
tum, together broader than long, sides nearly straight, posterior
corners rounded, border concave at middle. Abdomen con-
FIG. 36. TERMITOPHYA PUNCTATA SP. NOV.
siderably longer than the head and thorax together, broadly
margined at sides for entire length; feebly convex above and
rather strongly below; penultimate tergite trapzoidal, twice
as broad as. long and straight at the posterior border; last
tergite subtriangular, with the apex narrowly rounded.
Femora strongly compressed, tibiae less compressed;
posterior metatarsus one and three-fourths times as long as
the remaining joints together; claws moderately long and
slender.
Host.—Nasutitermes (Nasutitermes) guayanae (Holmgren).
Type-locality—kKartabo, British Guiana.
Described from four specimens.
1923] Mann: Termitophilous Coleoptera 351
This, and the following two species belong to the genus
Termitophya, though none of the specimens before me bear strong
cerci on the last dorsal segment, as figured by Wasmann.
(Tijdschr. Ent., vol. 45, 1902, pl. 9, fig. 1) and possibly repre-
sent the opposite sex.
T. punctata is near T. heyeri Wasm. but has the pronotum
proportionately longer and rounded instead of angulate at the
anterior corners. The punctation of the head is similar in
both species.
Termitophya amica sp. nov.
Length.—(with abdomen straightened) 2.75-3 mm.
Head, thorax and elytra dark brown to black; appendages
and abdomen pale brown, the latter with the median portion
of dorsum dark. Shining; head with fine, though distinct,
separated punctures and the intervals between with short and
irregular, fine and dense striolae. Pronotum and elytra smooth;
abdomen and appendages coriaceous. Erect hairs fine, black,
sparse on appendages, a pair on head at inner border of eyes,
three on the pronotal margins, two on each elytron. Dorsal
abdominal segments 1-4 with a row of six coarser, recumbant
hairs at apical margin; segment 5 with a pair of erect hairs;
ventral segments with sparse, erect black hairs and, apically
with elongate recumbant yellow hairs. Microscopic yellowish
pubescence sparse on head and body, more distinct on legs.
Head broader than long, sides posterior to eyes slightly
convex, occipital border broadly arcuate, anterior border
straight, front flattened, at middle very shallowly impressed.
Labrum large, entirely corneous, impressed at middle, narrowly
excised at tip. Eyes moderately convex, a little longer than
their distance to the posterior margin of head. First antennal
joint stout, nearly as long as second and third together, third
joint distinctly larger than the second, scarcely longer than
broad, remaining joints a little longer than broad aad becoming
slightly narrower toward apex; terminal joint a little shorter
than the two preceding joints together. Pronotum a little
longer than broad, as broad in front as behind, with feebly
352 Zoologica: N. Y. Zoological Society [111 3%
arcuate sides and posterior border, straight anterior border
and broadly rounded angles; surface evenly convex. Elytra at
base a little narrower than the pronotum at its widest part,
together broader than long, sides straight, little divergent,
posterior angles evenly rounded, the borders feebly convex
and meeting at broad angle. Abdomen elongate, flat above,
sides elevated as broad margins. Legs short, the femora
broader than in TJ. punctate and distinctly though narrowly
concave at middle of flexor border.
Host.—Nasutitermes (Nasutitermes) guayanae (Holmgren).
Type-locality—kKartabo, British Guiana.
Described from five specimens.
This species is very close to T. punctata, differing, in addi-
tion to the broader legs, in the punctation of the head, in the
arrangement of the erect hairs on the elytra and in having
the pubescence more conspicuous.
Termitophya flavivertris sp. nov.
Length.—(with abdomen straightened) 3 mm.
Head, thorax and elytra rather pale brown; ventral surfaces
and appendages lighter; dorsal surface of the abdomen yellow-
ish, except for a median brown stripe. Shining, head finely
and shallowly punctate; abdomen coriaceous. Erect hairs fine
and shorter than in the other species; lacking on dorsal portion
of head; four on sides of pronotum, two on each elytron.
Coarser, black, semi-recumbent hairs arranged in rows of six
unequal in size at the apical margin of abdominal segments
1-4 and one on either side of the same segments, one pair on
penultinate segments and a row, mixed with long yellow hairs
at the tip of apical segment; arranged in four irregular series
on each ventral segment. Long and very fine yellow recumbent
hairs scattered on dorsal surface of abdomen and arranged in
rows at apices of ventral segments. Pubescence fine and re-
cumbant, yellow in color, more conspicuous than in punctata
and amicus and sparsely and regularly distributed on head,
thoraze and appendages.
1923] Mann: Termitophilous Coleoptera 353
‘Head transverse, similar to that of amicus; clypeus
strongly and broadly convex. Labrum deeply and acutely
incised at apical borders. First antennal joint much thicker
than the others, a little shorter than joints 2-3 together; second
joint thicker and somewhat shorter than the third, which is
about twice as long as broad; remaining joints gradually decreas-
ing in length; terminal joint one and three-quarter as long as the
penultimate. Pronotum a little broader than long, straight in
front, feebly convex at sides and posterior border, with the
corners broadly rounded; surface moderately convex except
in front of middle when it is very shallowly impressed. Elytra
at base slightly narrower than pronotum; sides straight.
Posterior angles broadly rounded; posterior borders obliquely
convex and meeting at an obtuse angle. Abdomen elongate,
the sides elevated as strong margins. Legs short; femora and
tibiae broad and strongly compressed.
Host.—Nasutitermes (Nasutitermes) costalis (Holmgren).
Type-locality.—Kartabo, British Guiana(Aug. 1), St. Joseph,
Trinidad.
Distinct from T. amica in its paler coloration, more distinct
pubescence, the arrangement of the erect hairs and in the
more slender antennae, all joints of which are more elongate in
flaviventris. The two species are otherwise very similar.
Trachopeplus gen. nov.
Head suboval, broader than long, front with two strong
lateral impressions confluent with antennal fossae, and a small
median impression. Eyes rather large, oval and convex. Clypeus
membranous, very short and broad, anterior border straight at
middle. Labrum coriaceous, short, divided into two lobes by a
strong, triangular median incision. Gula very short, with sides
straight and divergent behind and not separated from sub-
mentum by a suture. Submentum large, a little broader than
long, flat behind, oblique and a little narrowed in front, sides
of posterior portion and the anterior border straight. Labium
transverse, subquadrate. Ligula large broadly incised apically.
Labial palpi 3-jointed; basal joint longer than bro.” and a
354 Zoologica: N. Y. Zoological Society [Tits ae
little thicker than the second; second joint distinctly longer
than the basal; apical joint minute. Maxillary palpi 4-jointed,
basal joint small, second joint twice as long as broad, very
narrow at base and gradually thickened toward apex, outer
surface convex, inner surface concave, the third joint a little
longer than the second, elongate oval in outline and two and
one-half times as long as broad, with convex outer surface and
shallowly concave inner surface; apical joint exceedingly short
and blunt at tip. Antennae 11-jointed, not thickened apically,
first joint strongly scapiform, remaining joints elongate.
Pronotum broad, strongly excised at middle, sides inflexed.
Pronotum transverse, anterior border membranous; anterior
half composed of a transverse chitinous plate that extends
laterally as narrow bands joining to the side pieces; posterior
half membranous. Meso- and metathorax together a little
longer than broad, anterior border strongly concave, posterior
border projecting between coxae and truncate at tip, surface
flat in front, convex behind. Scutellum distinct, transverse.
Elytra elongate and widely separated. Abdomen inflated, car-
ried forward over the elytra and posterior portion of prono-
tum; six segments visible from above; all segments margined
at sides.
Legs short and stout, femora and tibiae strongly flattened.
Tarsi 5-5-5 jointed; metatarsus broad, a little shorter than the
other joints together; fourth and fifth joints anchylosed; tarsal
claws very slender.
Genotype.—Trachopeplus setosus, sp. nov.
Trachopeplus setosus sp. nov.
(Fig. 37).
Length.— (with abdomen straightened) 3 mm.
Head, antennae, pronotum and elytra dark brown to black,
abdomen and ventral surface red-brown, legs yellow-brown.
Shining, very finely punctate, pronotum and elytra with small
tubercles, each bearing a hair. Front with sparse, rather
strong punctures, elytra densely, rather coarsely but shallowly ~
in ie
yet
1923] Mann: Termitophilous Coleoptera 355
punctate. Hairs black, very stiff, erect and moderately long
and abundant on head, thorax, elytra, and appendages; on the
abdomen they are semirecumbant and irregular in length and
thickness, arranged in two rows of 10-12 on segments 2-5, one
row on the basal segment; ten rows on the apical segment,
and abundant on ventral surface (where arranged in three ir-
regular rows) and legs.
i Wik *
FIG. 37. TRACHOPEPLUS SETOSUS SP. NOV.
Head broader than long, sides in back of eyes convex,
posterior borders rounded; border feebly arcuate; antennal
foveae extending to the vertex as a pair of broad, shallow
depressions; anterior portion of front flat, the border a little
produced and truncate apically. Clypeus broad and narrow,
entirely membranous. Labrum distinctly concave at middle
of anterior border. First antennal joint about as long as the
head, third joint twice as long as broad, a little longer than
the second and one and one-half times as long as the third;
joints 3-10 distinctly longer than broad, gradually decreasing in
length; terminal ‘joint elongate, shorter than the two preceding
together, with parallel sides and rounded tip. Eyes nearly
half as long as head, convex, situated near front of sides.
Pronotum transverse, as broad in front as behind. Sides very
feebly concave, anterior border convex at middle, narrowly
concave at sides; posterior border broadly arcuate at middle,
nearly straight at sides; disc in front of middle broad with a
deep transverse impression two-thirds its width, the surface
356 Zoologica: N. Y. Zoological Society [IITs oF
anterior to this nearly flat, posterior to it, convex. Elytra at
base as broad as pronotum and about as long, widely separated
from front to rear; each elytron nearly twice as broad as base
as at apex, with straight outer border, feebly concave inner
border and oblique, slightly arcuate posterior border; exterior
corners subangulate. Abdomen a little less than twice as broad
as pronotum, dorsal surface nearly flat, sides elevated as
prominent margins Legs rather short, femora and tibiae
broadly flattened.
Host.—Nasutitermes (Nasutitermes) acajutlae (Holmgren).
Type-locality——Kartabo, British Guiana.
Described from two specimens. (Mouth parts on slide.)
The shape of the head, the profoundly impressed pronotum,
the structure of the maxilles and tarsi and the tuberculate
nature of the pronotum, as well as the abundant and vary
coarse and stiff hairs are distinctive and separate Trachopeplus
from Termitogaster, with which is resembles somewhat in
habitus and in the form of the antennae.
While there are five tarsal joints, the fourth and fifth are
closely jointed and appear superficially as one, but in a balsam
mount the suture is distinct, and furthermore marked by the
arrangement of the hairs, which, with the other characters,
place the genus in the Coloderae group.
Xenopelta gen. nov.
Robust; gaster moderately inflated and held elevated. Head
elongate, front strongly and broadly impressed between eyes,
abruptly elevated and longitudinally carinate at middle in front
of antennal sockets, anterior border broadly angulate at middle
and slightly projecting over base of clypeus. Clypeus membran-
ous, strongly transverse, concave at anterior border. Labrum
entirely membranous, strongly and broadly excised at middle
of anterior border, with the surface of the lateral portions
convex. Mandibles stout basally, slender and acute at apical
half, unidentate on inner border. Mentum longer than broad,
its surface flat, front border moderately elevated. Ligula very
small, apparently simple. Labial palpi 3-jointed, the basal
1923 ] Mann: Termitophilous Coleoptera 357
and second joints much thicker than the terminal. Maxillary
palpi 3-jointed; basal joint very small, second joint with the
upper surface greatly produced into a chitinous, asymmetrical,
arcuate lobe, that extends nearly to the apex of the third joint,
basal portion flat; third joint thick, strongly convex above and
nearly straight beneath. Antennae 11-jointed, not thickened
apically; first joint scapiform. Eyes large, situated at sides
of head. Prothorax elongate; disc of pronotum strongly im-
impressed, sides elevated into thick rounded margins, anterior
border elevated, projecting, narrowly rounded; posterior mar-
gin with flat surface and broadly rounded border. Elytra well
developed, a little broader than long. Abdomen moderately in-
flated, more convex beneath than above; seven segments visible
from above, all segments, except the terminal, broadly mar-
gined, lateral borders and the areas between the segments, both
dorsally and ventrally, with narrow bands of white membrane.
Prosternum in front of coxae elevated into a ridge which
is projecting and triangular at middle. Metasternum trans-
verse, surface at middle convex; posterior border truncate
between coxae. Anterior coxae large and conical, approximate.
Middle and posterior coxae short,, rather broad, separated.
Legs flattened, tibiae especially so, the hind tibiae with the
border very thin and with an indication of a submarginal line
on inner surface. Tarsi 5-5-5 jointed, the joints sharply dis-
tinct; hind metatarsus a little shorter than the remaining joints
together.
Genotype.—Xenopelta cornuta, sp. nov.
Xenopelta cornuta sp. nov.
(Fig. 38)
Female.—(length with abdomen straightened) 2.75 mm.
Dark reddish brown to black, gaster lighter, projecting tip
of front and the apical half of tarsi yellow brown; feebly
shining, the appendages more than the rest. Head and body
rather coarsely, densely punctuate; appendages similarly, but
much more finely sculptured; tibae with coarse setigerous
punctures.
358 Zoologica: N. Y. Zoological Society [LIT saa
Hairs black in color, lacking on head, a row of three on
lateral borders of pronotum, two on each elytron, an oblique
row of 10-12 on the posterior margins of abdominal segments
2-5 and a few near the basal border; one pair on penultimate
abdominal segment; finer and rather abundant on ventral sur-
face of abdomen; shorter on appendages.
FIG. 38. XENOPELTA CORNUTA SP. NOV.
Head longer than broad and a little broader behind than in :
front, sides nearly straight, occipital corners and border
broadly rounded. First antennal joint nearly as long as the
head; joints 2-4 somewhat thicker than the following, the
second twice as long as broad, remainder gradually decreasing
in length to the penultimate which is about one and one fifth
times as long as broad; terminal joint elongate oval, shorter
than the two preceding joints together. Eyes less than half as
long as head, situated a little in front of middle. Pronotum a
little broader than head, broadest at anterior third; anterior
angles rounded; depression occupying the larger part of the
surface, deepest a little in front of middle with a large, shallow
fovea at bottom on either side. Elytra broadest behind, with
straight sides, rounded posterior corners and concave border.
Abdomen one and three fourths times as broad as elytra.
Host.—Nasutitermes (Nasutitermes) guayanae (Holm-
gren).
Type-locality—Kartabo, British Guiana.
Described from two females.
The basal point of the maxillary palpi is very small and
visible only when the mouth-parts are dissected out.
< mig, p-—eaaenelieins amel,
he eS
ee a ©
1923] Mann: Termitophilous Coleoptera 359
Xenogaster fossulata sp. nov.
Female.— (length) —2.80 mm.
Head, pronotum and gaster coriaceous shining, elytra
densely punctate and less shining. Hairs yellowish, fine and
silky, moderately abundant on head, thorax and elytra and
appendages. Dorsal segments of abdomen with a row of longer,
nearly recumbant hairs at posterior border; ventral surface
with fine long hairs.
Ferruginous, abdomen and legs lighter.
Head longer than broad and broader behind than in front,
flat above, except for shallow elongate impressions that extend
from antennal insertions a little more than half the distance
to occipital border. Labrum broad, nearly straight in front.
Maxillary palpi large, with the second joint distinctly longer,
though more narrow than the basal; apical joint small,
subulate and less than half as long as the penultimate. Eyes
large, nearly flat, occupying anterior three-eights of sides of
head. Antennal scapes moderately thick, extending a little less
than the distance to posterior border of head, first funicular
joint subcylindrical and twice as long as broad, second joint
similarly shaped but shorter, joints 3-9 subquadrate in profile,
slightly transverse, apical joints but little broader than the
others; terminal joint oval, about as long as the two preceding
joints together. Pronotum longer than broad, sides in tront of
middle slightly convex, behind middle straight and very feebly
convergent, anterior corners subangulate, posterior border
truncate; dorsum strongly impressed at middle with the sides
forming rounded borders to the pit, which is deepest in front
and sloping behind with the surface of the posterior, sloping
portion very feebly convex toward the basal half. Scutellum
large. Elytra at base a little broader than pronotum, divergent
behind, widely separated, the inner border slightly concave,
outer borders nearly straight, posterior border rounded at tip.
Abdomen not strongly inflated, sides only slightly convex,
dorsal surface flat, strongly margined and elevated at sides.
Host.—Nasutitermes (Nasutitermes) intermedius (Banks).
Type-locality—Kartabo, British Guiana.
360 Zoologica: N. Y. Zoological Society [Ui siy
Ceratoxenus gen. nov.
Physogastric species with the abdomen carried upward and
forward over the thorax. Head broader than long, not con-
stricted behind, margined in front, with projecting, thick
spinose developments. Mandibles rather thick, elongate-
triangular, curved at tips. Labrum entirely membranous, very
strongly transverse, concave at anterior border. Mentum sub-
quadrate, a little broader than long, broadly concave at anterior
border. Ligula membranous, simple, transverse. Maxillary
palpi 4-jointed, the joints coarse, second joint thicker and
about as long as the third, second and third less than twice
as long as broad, their outer surfaces convex, fourth joint
short, subulate. Labial palpi 2-jointed, the basal joint dis-
tinctly thicker than the second. Antennae 11-jointed, first
joint scapiform, apical portion slightly enlarged. Eyes a little
less than half as long as sides of head. Pronotum with sides
inflated vertically; anterior, inferior angles with an elongate
acute spine. Prosternum elevated and margined at anterior
border; prothoracic hypomera concealed: Elytra well de-
veloped, tapering and widely separated behind. Mesosternum.
as long as prosternum and longer than metasternum, surface
behind coxae evenly convex, triangular, projecting, coxae nar-
rowly separated, acetabuli closed. Abdomen about as long as
the thorax, strongly convex beneath, plane above, six segments
visible from above, margined at sides for entire length. Legs
long, rather slender. Tarsi 5-5-5-jointed. Posterior metatarsus
much shorter than the remainder of tarsus.
Genotype—Ceratoxenus tricornis sp. nov.
Ceratoxenus tricornis sp. nov.
(Fig. 39)
Female.— (length) —2.60 mm.
Head a little broader than long, sides and posterior border
moderately rounded, vertex and front broadly impressed; an-
terior border of front strongly margined, the margin project-
ing at middle as a broad triangular tooth, at sides as elongate,
1923] Mann: Termitophilous Coleoptera 361
bluntly pointed, laterally compressed spines. Labrum broad,
concave at anterior border, from above entirely concealed by
the frontal margin. Mandibles rather thick and not very acute
at tips. Antennae extending to posterior end of elytra; scape
thick, as long as head from above; funicular joints 1-3 elongate,
subcylindrical and equal in length, remaining joints more
rounded and, toward apex becoming transverse; terminal joint
FIG. 39. CERATOXENUS TRICORNIS SP. NOV.
oval, less than twice as long as broad. Eye large, moderately
convex, slightly emarginate at the border approximate to
antennal insertions, occupying most of the front half of sides
of head. Pronotum a little broader than long, side pieces
nearly perpendicular, inferior anterior angles produced as
spines; dorsum with lateral and front borders roundly mar-
gined and with a transverse ridge a little in front of middle,
the anterior surface evenly concave and the posterior surface
impressed and plane. Elytra a little narrowed at base and
about as long as pronotum, very broadly separated, (the distance
between their tips equal to their length) tapering and narrowly
rounded at tips. Abdomen strongly inflated and carried above
thorax; dorsal surface flat, sides and venter evenly convex,
dorsal segments 1-4 subequal and less than half as long as the
corresponding ventral segments. Legs moderately long and
slender.
362. Zoologica: N. Y. Zoological Society [TIL saa
Finely coriaceous and opaque throughout. Rather stiff
erect brown hairs sparsely distributed on head and body and
appendages. Maxillary palpi with dense yellowish white hairs.
Color ferrugineous, with the abdomen, legs and palpi
lighter than the rest; membranous portions pure white.
Host.—Nasutitermes (Nasutitermes) guayanae (Holmgren).
Tribe—Gyrophaenae
Blapticoxenus gen. nov.
Small, rather flattened species, tapering posteriorly. Head
concealed beneath pronotal margin, small, not constricted, front
subtriangular, front flat. Antennal fossae with acute margins
which terminate at inner border of eye and approximate at
middle of anterior border of front. Genae acutely margined.
Labrum elongate, nearly straight at anterior border, tri-
angularly depressed at sides, leaving a median portion shaped
somewhat similar to an hour-glass, with broad and thin lateral
margins. Mandibles small, arcuate, acute at tips, the right one
dentate at middle of inner margin. Maxillary palpi 4-jointed,
the first three joints elongate and subequal in length, terminal
joint short, subulate. Labial palpi small, 2-jointed, the joints
subequal. Ligula simple, elongate, rounded in front. Eyes
large and convex, with rather large facets, occupying nearly
all of the exposed sides of head. Antennae 11-jointed, not
geniculate, the first joint much shorter than the second and
third together. Pronotum broadly convex, inflexed at sides and
in front Prothoracic hypomera not visible. Anterior coxae
very large and conical. Mesosternum much larger than pro-or
metasternum, not carinate, slightly elevated posterior to coxae,
separated from metasternum, coxal cavities closed, coxae ap-
proximated. Metasternum very short and conical. Elytra well
developed, as broad as pronotum. Abdomen longer than head
and pronotum together, tapering behind, first four segments
strongly margined, seven segments visible from above, the sixth
not impressed and the seventh not elongate. Legs short and
1923] Mann: Termitophilous Coleoptera 363
rather stout, tibiae not spinose; 4-4-5 jointed. First joint of
posterior tarsi elongate.
Genotype—Blapticoxenus brunneus sp. nov.
Blapticoxenus brunneus sp. nov.
Length.—1.75-1.90 mm.
Pale brown, shining, minutely punctate. Head, thorax
and elytra devoid of hairs; abdomen with exceedingly coarse,
black setae at sides and on ventral surface (on ventral surface
arranged in rows near posterior margins of segments); legs
and antennae with finer, shorter and yellow hairs; apex of the
second and third antennal joints with strong, black, hairs.
Head a little broader than long, front flat, triangular,
bordered by the distinct carinae that margin the antennal
fossae; sides posterior to eyes oblique, slightly concave; pos-
terior border straight. Antennae rather slender, extending a
little beyond posterior borders at elytra; basal joint thicker
than the others and longer than the second. Second joint very
slightly clavate and shorter than the third; joints 3-4-subequal,
longer and more slender than the remaining joints which de-
crease in length toward the tip; terminal joint elongate-oval,
rounded at apex, much shorter than the two preceding joints
together. Pronotum broader than long, broadest behind middle
of sides, with slightly convex sides, broadly rounded anterior
corners and border and rounded posterior corners; posterior
borders very feebly bisinuate. Elytra more flattened than
pronotum, together more than twice as broad as long, slightly
broader behind, with nearly straight sides, subangulate pos-
terior corners, posterior border nearly straight at sides and
concave at middle. Dorsal abdominal segments 1-5 broader
than long, segment 6 twice as long as broad and bluntly tri-
angular in shape.
Host.—Nasutitermes (Velocitermes) beebei (Emerson).
Type-locality—Kartabo, British Guiana.
Described from three specimens (one on slide).
364 Zoologica: N. Y. Zoological Society (LI se
This genus belongs in the tribe Gyrophaenae, but the joints
of the maxillary palpi are unusually elongate. The depressed
sides of the front of labrum, the elongate mesosternum are very
characteristic.
Family—HISTERIDAE
Thaumataerius gen. nov.
Near Teratosoma (Lewis).
Body subquadrate, very long and slender, flattened above.
Head elongate, mandibles strong and moderately projecting.
Antennae inserted a little in front of middle of inner border of
eyes; first joint subclavate, broad and thick, joints 2-8 grad-
ually increasing in size; club large, one jointed, as long as the
three preceding joints together, broadly rounded at tips.
Antennal fossae visible from front and sides, thinly mar-
gined by prosternum and pronotum, capable of containing the
entire antennae. Prosternum flat basally, sides between coxae
straight, narrowly margined, posterior border truncate; rather
strongly, transversely impressed at anterior fourth. Mesos-
ternum flat at middle, about as broad in front as behind.
Pronotum with projecting anterior and posterior corners, sides
diagonally impressed, with deep, elongate fossae on either side
of base. Elytra elongate, flat above, anterior and posterior
corners broadly rounded; sutural striae broadly impressed, ex-
tending three-fourths the length of elytra; humeri with deep,
very short striae, and disc with a similar short and deep stria
midway between the humeral and the sutural; anterior border
bigibbous, one tubercle at base of each of the short striae.
Propygidium broad and feebly convex. Pygidium with three
very strong elongate costae, about half as long as the pygidium,
the surface between these unevenly concave. Legs slender,
moderately long; femora not arcuate; anterior tibiae with short
tarsal grooves. Anterior tarsi noticeably shorter and _stouter
than the others.
From Teratosoma, known from a Brazilian species of both
termitophilous and myrmecophilous habit, Thawmataerius
differs in its very slender, elongate form, in the comparatively
1923] Mann: Termitophilous Coleoptera 365
| simple structure of the prothorax, in the shorter and straight
_ legs and in having the pygidium very strongly tricostate. The
two genera resemble each other-in the non-inflated legs and
somewhat in the striation of the elytra, and though distinctly
different genera, together constitute a radically aberrant group
of the tribe Hetaerimorphini.
Genotype.—Thaumataerius emersoni, sp. nov.
FIG. 40. THAUMATAERIUS EMERSONI SP. NOV.
Thaumataerius emersoni sp. nov.
(Figure 40)
Length.—2.10 mm.
Brownish red, shining; minutely punctate; body and ap-
pendages with moderately abundant, fine silky yellow hairs,
which on the outer edge of the posterior and middle tibiae and
shorter and denser and form a well defined brush.
366 Zoologica: N. Y. Zoological Society (ll; a@
Head longer than broad, not margined; vertex broad,
rather strongly impressed. Eyes large, elongate, with very dis- —
tinct facets. Prothorax as.broad as long, slightly broadest
behind, sides in front of middle feebly concave; anterior angles
obliquely truncate, flattened and submargined at tip; posterior
angles prolonged, with convex surface, convex outer border and ©
narrowly rounded tip, separated from disc by strong’ oblique ©
impressions which are deep basally and become shallow
anteriorly; disc convex in front, shallowly concave at posterior
half and separated from sides by a pair of oblique striae, one
of which terminates in the lateral fossa and one of which
extends to form an inner margin to the posterior angulate
processes. Elytra together more than twice as long as broad,
sides straight and parallel for three-fourths their length,
feebly narrowed apically.
Host.—Mirotermes (Mirotermes) nigritus Silvestri
Locality—Kartabo, British Guiana, July 31.
—
~~
Mi gem Si SOME 0
Volume III, Number 18
GLANDULAR STRUCTURE
OF THE ABDOMINAL APPENDAGES
OF A TERMITE GUEST (Spirachtha)*
By N. E. McINpDoo, Pu. D.
Bureau of Entomology, Department of Agriculture, Washington, D. C.
(Figure 41, Plates XVI-XVII)
INTRODUCTION.
Some of the specimens of Spirachtha, collected by Mr.
Alfred Emerson of the University of Pittsburgh in nests of
Nasutitermes (Constrictotermes) cavifrons (Holmgren) in
British Guiana, were given to me to determine whether or not
the abdominal appendages are glandular.
Emerson informs me by correspondence that his specimens
of Spirachtha were identified by Dr. W. M. Mann of the Bureau
of Entomology as two new species—S. schiddtei and mirabilis
Mann. The material sectioned by me belongs to the latter
species, but Emerson made his observations on both species;
however, he says: “As far as I observed, the habits of both of
the Spirachtha were exactly the same, and both species came
from the same nests.”
The live insects were fixed by Emerson in hot corrosive
sublimate plus a little acetic acid and then allowed to cool,
and finally were preserved in 85 per cent alcohol. Parts of
the abdominal appendages and the abdomen with the appen-
dages intact were embedded in 60° parraffin. Sections were cut
five microns in thickness and were stained in Ehrlich’s hema-
toxylin and eosin. The drawings, except fig. 41, are original
and were made at the base of the microscope with the aid of a
camera lucida. Figure 41 was made by Emerson from a speci-
men, but I have added a few hairs and some shading to it.
The following references are all that I can find which
relate directly to the abdominal appendages of Spirachtha.
* Tropical Research Station, Contribution Number 138.
368 Zoologica: N. Y. Zoological Society [III;18
Schiddte (1853) says: “The abdomen is furnished with three
pairs of appendages, which are elongate, cylindrical, 2-jointed,
membraneous, and moveable by muscles at the base. The
appendages are perhaps intended for the same purpose as the
tufts of hairs on the abdomen of the genus Claviger, which are
known to be sucked by the ants.’”’ The same author (1856, p.
181), after having prepared sections from specimens preserved
FIG. 41. SPIRACHTHA MIRABILIS MANN
Dorsal view showing abdominal appendages (1, 2 and 3), which consist
of portions a, b and ¢; only the last two being glandular. Those portions
marked A are the abdomen which conceals all the thorax and head,
except parts of the legs, the antennae and anterior part of the head (all
in solid black). Greatly enlarged. Photographed and originally drawn
by Emerson, but later slightly modified by the writer.
in alcohol, was not able to decide definitely whether or not
these appendages are glandular in structure. He says that
there is a pair of muscles at the base of these organs, and
consequently they are moveable. They are formed from pro-
longations of the abdominal integument, and have thick walls.
Their structure appears to be homogeneous, and their con-
tents is composed of clear granules of an irregular and
globular form.
EXTERNAL STRUCTURE OF ABDOMINAL APPENDAGES.
This staphylinid beetle, about the size of a common pin
head, is most remarkable in that the abdomen bends forward
Rites ine oy dear As,
1923] McIndoo: Glandular Structure, Termite Guest 369
so that the latter lies directly over the thorax and head. Fur-
ther than this, the abdomen (Fig. 41A) bears three pairs of
appendages (1, 2 and 3), the size and shape of which vary
considerably in different specimens. Very few of the speci-
mens, however, have perfectly developed appendages. Emer-
son says that the appendages were of various sizes or were in
various stages of development when observed in the field; some
possessed only small knobs on the abdomen, while in a few the
appendages were well developed as shown in Fig. 41.
The first pair of appendages (Fig. 41, 1) is unusually long
and filamentous, while the second and third pairs are elongate
or club shaped, but when perfectly developed their distal ends
(2 and 3) are more or less globular in shape, as represented in
Fig. 41. The appendages, when fixed in the corrosive sub-
limate solution, are white, soft and fleshy-like structures, and
are easily sectioned. Emerson says that when the insects are
alive, the appendages are held up over the abdomen; sometimes
those of one side touching those of the other side. The first
pair, arising from the fourth abdominal segment, is held up
in the air, each resembling the letter S. The second and third
pairs, arising from the fifth and sixth abdominal segments,
were observed to be slightly moveable through a vertical plane
at right angles to the body. Emerson observed that when the
appendages were moved, all of them moved at the same instant.
He thinks that both sexes develop these extraordinary appen-
dages and that they are postimaginal structures.
INTERNAL STRUCTURE OF ABDOMINAL APPENDAGES.
Each appendage arises from a fleshy prolongation (Fig.
41, a) of the abdomen, and appears to be 2-segmented, but sec-
tions show that the articulations between portions marked a
and b and between b and e¢ are nothing more than constric-
tions with thinner cuticula than elsewhere. The constriction
between a and b is shallow, while the one between b and ¢ is
deep. Several muscle fibers run diagonally across the portion
marked a, but only a few other fibers unite with the integument
in the constriction, and no muscles were seen in the apparent
EEO
370 Zoologica: N. Y. Zoological Society [I11;18
segments b ande. Judging from this arrangement, the appen-
dages certainly cannot be moved very much.
A microscopical examination of the integuments (not
treated with KOH) of these appendages did not show any pores
and only two types of hairs. The cuticula is literally covered
with tiny prickles or pseudohairs and (Plate XVI, 5, Hr’)
many comparatively large hairs (Plates XVI-XVII, 5, 14, Hr)
were observed arising singly from miniature mounds, widely
scattered.
Sections through the portions marked b and e (Fig. 41)
reveal the most peculiar arrangement of tissues that I have
ever observed. These portions (Plates XVI-XVII, 12-14) appear
hollow, but are really filled with a coagulated liquid, apparently
blood (Plate XVII, 14, Bl.). The walls are thick and consist of
four layers; the two outermost ones being the cuticula (Plate
XVII, E, D), the middle one the hypodermis (Hyp), and the in-
nermost one the basement membrane (M) of the hypodermis.
Passing through the hypodermis from the basement membrane
to the outer layer of the cuticula there are many dark strands;
some of these (S) are attached to the bases of the hairs (Hr),
and the others (St) either to the cuticula directly or at the
bases of tiny pores (P). These various structures are colored
very beautifully in sections passed through alcohol containing
iodine, and stained by Ehrlich’s hematoxylin and eosin. The
blood and basement membrane are colored pinkish by the
eosin, but the later takes the deeper color; the hypodermis is
stained purple by the hematoxylin; the inner layer of the
cuticula (D) and the strands are stained brownish by the
iodine; and the outer layer of the cuticula (E) remains un-
stained, being whitish or semi-transparent.
A more careful study of these various structures shows
the following details. The outer and inner layers of the
cuticula are practically equal in thickness. The former con-
tains numerous tiny pores (Plate XVI, 11, P) whose outer
ends are funnel-shaped, while the inner ends are straight or
curved. The pores are practically all single openings, but one
double pore (17) was found. These pores are peculiar in
that the outer cuticula surrounding them is considerably
—~
1923] McIndoo: Glandular Structure, Termite Guest 371
thicker than elsewhere, thus making semispherical projections
which extend into the inner cuticula.
While there is nothing uncommon about the outer cuticula,
except its pores, the inner cuticula is very peculiarly modified.
The latter usually appears porous or spongy (Plate XVI, 4, 6,
8, D), but may be stratified (5, 11), or may occasionally be
S
Plate XVI. S. MIRABILIS MANN
Internal structure of abdominal appendages.
2, end view of columnar hypodermal cells, showing nucleus (N) and stands (S and St)
in cross section; 3, longitudinal view of glandular hypodermal cells (Hyp) and_blood
cells (BC); 4, 5 and 11, wall of appendage, showing original hypodermis (Hyp'),
hair (Hr), tiny prickles (Hr), outer cuticula (E), inner cuticula (D), pore (P),
glandular hypodermis (Hyp), basement membrane “(M), and strands (S and St) in
longitudinal view; 6 to 8, various types of the dermis; 9 and 10, two types of the
basement membrane; 12, cross section (semidiagrammatic) of the smaller bulb-shaped
end of an appendage, showing all of the internal structures; 2 to 11, X 505, and
12528 Ob:
wavy (7). It lies against the outer cuticula and is firmly
anchored there by many tiny prickle-like projections (6, 8), ex-
tending into the outer layer. In order to distinguish the inner
from the outer cuticula in the various drawings, a white space
has been left between these two layers.
Section passing crosswise through the hypodermal cells
show that these are usually five or six sided columnar cells
(Plate XVI, 2), whose cut ends appear net-like and resemble
372 Zoologica: N. Y. Zoological Society [III;18
plant cells more than animal cells. Their nuclei (N) are
scarcely discernible and only appear as faint elliptical areas
which contain a few dark particles with a few radiating lines.
It is strange that these nuclei should be so inconspicuous, be-
cause ordinarily nuclei are the most conspicuous parts of cells.
Perhaps the fixation was not good for these nuclei.
Sections passing lengthwise through the hypodermal cells
usually indicate that the hypodermis consists of more than one
layer of cells, but upon closer examination it becomes evident
that the hypodermis really consists of only one layer (Plate
XVI, 3). In longitudinal sections the contents of the cells
appear to consist of coagulated streaks, somewhat resembling
strings of beads, which generally extend lengthwise through
the cell, but sometimes diagonally across it. The longitudinal
walls of the cells are conspicuous as dark lines, vsually running
in zigzag style, while in cross sections the walls appear as
curved or straight lines. Under a high magnification each dark
line appears double as shown in (2, 3).
Like the inner cuticula and hypodermis, the basement
membrane is also unusually developed. So far as I am aware,
the basement membrane in sections of insects almost always
appears as a single line without nuclei, but in spiders
(McIndoo, 1911) it appears as a double line with nuclei. This
may be due to the fact that the hypodermis in spiders is always
thick, never becomes atrophied, but continues to function as
long as the spider lives. This explanation may possibly serve
to explain why the basement membrane in these abdominal
appendages is so highly developed; here both walls are always
discernible (Plate XVI, 5, 9-11), and nuclei are usually seen
between them. The inner wall is generally more or less
smooth, but occasionally it is very rough and may bear finger-
shaped or papilla-like projections (9), which extend into the
blood.
For some time the dark strands, already mentioned, were
a puzzle to me, but now I believe that I can satisfactorily ex-
plain them. They are very conspicuous in all the sections
made, and appear in four different conditions; fragments may
be seen adhering to the cuticula or bases of the hairs; small
1923] McIndoo: Glandular Structure, Termite Guest 373
isolated portions (Plate XVI, 2, 12, S), may be observed in
either cross or longitudinal sections of the hypodermal cells;
prolongations (12) of various length may be seen extending
from the basement membrane into the hypodermis; and oc-
casionally a complete strand (5, S and 11, St) may be observed.
Their outer end (2, S and St) appear to be spongy and in
structure are similar to the inner cuticula, but their inner ends
seem to be as soft as the basement membrane. Practically
every hair has a strand attached to its base, while only about
one half of the pores have strands attached at their inner
ends. Those strands running to the hairs look darker in
sections and are much narrower than are the other ones
described. The probable function of these strands is stated
under “Interpretation of Results.”
In life these abdominal appendages are probably com-
pletely filled with blood, because in sections the coagulated
remains of the blood almost fill the entire cavity. Two types
of blood cells (Plate XVI. 3, BC) were found in the blood. The
smaller type, although probably not blood cells at all, is the
commoner.
INTERPRETATION OF RESULTS.
After a preliminary examination of the sections prepared,
my first interpretation of these appendages was that the liquid
found in them is a secretion which finds its way to the exterior
through the strands and pores. This interpretation was found
to be incorrect for the three following reasons: 1 The quantity
of liquid contained in all six appendages is more than that of
the blood contained in the remainder of the abdomen; thus,
the source of any secretion must always be greater than the
secretion itself. 2 This liquid contains blood cells, appears the
same in structure, and has the same color as the blood found
elsewhere. And 3, if the strands were efferent tubes they
should be hollow in order to permit the secretion to pass freely
to the exterior.
My interpretation now is that the blood passes freely
through the basement membrane and inner ends of the strands
into the hypodermal cells which act as secreting or gland cells.
374 Zoologica: N. Y. Zoological Society [III;18
If this is true the secretion then passes from the hypodermis
into the outer ends of the strands and into the inner cuticula
which serves as a reservoir to store the secretion. From this
reservoir the secretion passes through the numerous tiny pores
to the exterior where it probably spreads over the entire sur-
face of the appendages and abdomen. This view is supported
by the following facts. Emerson says that the termites carry
these beetles about from place to place and that he often saw
them lick not only the abdominal appendages but also the entire
bodies of these insects. The inner ends of the hypodermal cells
must be extremely active, judging from their deeper staining
capacity. The basement membrane and inner ends of the
strands contain coagulated particles like those in the blood, thus
indicating that the blood passes freely into the hypodermis.
The secretion must be different from the blood, because no
remains of it can be found in the sections prepared. It may
be of an oily or fatty nature and evidently totally soluble in
the reagents used.
It would be interesting to know the exact sequence of
formation of the various structures in the walls of these appen-
dages. The formation is perhaps about as follows: The original
hypodermis (Plate XVI, 4, 5, Hyp’), little of which remains,
first secretes the outer cuticula and hairs, then the inner
cuticula; afterwards instead of its becoming atrophied as usual,
it becomes greatly hypertrophied and secretes an entirely diff-
erent substance which probably serves a nutritive purpose.
Since the hypodermis is a thick, soft and flabby membrane it
needs supports and a means of firmly anchoring it to the
dermis. All of this is accomplished by the semirigid strands.
Those strands attached to the hairs might originally have
been trichogen cells, but now they are entirely different and
certainly have a different function. In fact it seems that all
the strands have originated as outgrowths from the basement
membrane, because they are still attached to it and a large
nucleus is usually present in this membrane where a strand
departs The strands are attached to the bases of hairs and
pores, because these projections serve as good places for attach-
ing them.
——weee
—
a
1923] McIndoo: Glandular Structure, Termite Guest 375
The various structures, as described, are present in all
parts of the appendages marked b and e¢ (Fig. 41), but the
blood chamber is not always centrally located. Sometimes, as
in the smaller bulb-shaped end of an appendage, the blood
chamber (Plate XVI, 12) has shifted to one side, totally
eliminating the hypodermis from that side. In the portion
marked a (Fig. 41) and in the remainder of the abdomen none
Plate XVII. S. MIRABILIS MANN.
Internal structure of abdominal appendages.
13, cross section through portion b (Fig. 41) of first appendage; 14, oblique
section through portion e of second or third appendage, showing following
internal structures: pore (P), outer cuticula (E), inner cuticula (D), glandular
hypodermis (Hyp.), strands (S and St), basement membrane (M), blood cells
(BC), and blood (Bl); both diagrammatic, and both X 105; 15 and 17, various
sizes and types of pores, X 505; 15 to 17, sections similiar to 14; 16, from
abdomen; 15, largest pore; 16, smallest pore found.
of these structures are unusually developed and only traces of
some of them can be found. Starting with the proximal end of
portion b and following the sections into the abdomen, it is
seen that the highly developed hypodermis with its basement
membrane suddenly disappears and that the inner and outer
cuticula gradually become thinner so that in the portion a all
that remains is: 1 a thinner outer cuticula with an occasional
pore; 2 a very thin inner cuticula, slightly porous; and 3 a
trace of the original hypodermis. In the abdomen the outer
376 Zoologica: N. Y. Zoological Society [Tilers
cuticula is still thinner and only a trace of the inner cuticula
and hypodermis can be found. Here an occasional pore (Plate
XVII, 16), the smallest of all, is still present but the inner
cuticula is no longer porous. In these sections there is nothing
unusual about the anatomy of the abdomen. The greater part
of the space is filled with the intestine, cut two or three
times thus showing that it is much convoluted. At certain
places the intestine seems very large and contains many
particles, resembling bits of vegetable matter. Other struc-
tures, apparently eggs in their follicles are present; besides
muscles, blood and the fat cells. The fat body is comparatively
small and seldom lies against the cuticula.
GENERAL DISCUSSION.
In referring to the literature on this subject, I find only
one insect in which the apparent arrangement of tissues is
similar to that already described for Spirachtha; yet in this one
there is no similarity if these tissues have been correctly
named. This staphylinid beetle (Termitomimus) was found in
considerable numbers by Tragardh (1907a) in the Hutermes
colonies of Zululand. Trigardh says that this genus matches
Spirachtha with regard to the peculiar development of the
abdomen and the mouth-parts. The large abdomen bears no
appendages, but curves upward and forward so as to cover
completely the thorax and posterior half of the head. For
description he has divided the abdomen into a “pseudocaput,” a
“pseudothorax” and “‘pseudoabdomen.” Sections through the
“pnseudothoracic” projections show four layers in the thick
body wall. He calls the outer and inner layers of the cuticula
epiostracum and endostracum respectively. In the position of
my glandular hypodermis, he finds a cyanophilous tissue of a
spongy appearance which sometimes exhibits a very distinct
radial structure, sometimes is concentrically stratified and
contains numerous granules which are also to be found in the
trichogen cells. He thinks that this tissue is a fluid, which
has either passed through the hypodermis and is a derivate
from the fat body, or it is a secretion produced by the hypoder- |
mis and is coagulated by the method of fixation. It seems to
1923] MclIndoo: Glandular Structure, Termite Guest 377
me that this tissue, which is poorly fixed, might correspond to
my glandular hypodermis. In the positions of my basement
membrane and the strands, running to the hairs, he finds two
structures which he calls hypodermis and trichogen cells
respectively. He imagines that the secretion passes through
the cuticula, although he saw no pores at this place, but at
other places in the cuticula near which lie fat cells he saw
many extremely fine pores.
Tragardh found two pairs of glands opening into the cuti-
cular folds at the dorsal side of the neck of Termitomimus.
He calls them cephalic and prothoracic glands, because the
unicellular hypodermal glands of the former lie in the
head and those of the latter in the prothorax. Each gland
cell opens to the exterior through a tiny pore. He does not
think that these glands are in any way connected with the
termitophilous life of Termitomimus.
While it is exceptionally rare for any adult insect to bear
appendages or projections similar to those already described,
many larval insects bearing unusual thoracic and abdominal
appendages have already been found.
Silvestri (1920) found unusual thoracic and abdominal
appendages on certain dipterous and lepidopterous larvae, and
also apparently eight pairs of lateral appendages on the
coleopterous larva of Troctontus, all of which were taken from
termite nests in Africa. He seems to have found glands in only
the tiny club-shaped appendages of the last named insect.
These consist of many very large unicellular, hypodermal
glands. They lie in a thick hypodermis and open to the exterior
through pores. Silvestri thinks that they secrete a special sup-
stance for the termites.
Wheeler (1918) found unusual thoracic and abdominal
protuberances or appendages on three species of ant larvae
belonging to the genera, Tetraponera and Pachysima. Speak-
ing of the tubercles of 7. tessmanni, Wheeler (p. 306) says:
“Sections and stained, cleared preparations of the whole
Jarva show that the various tubercles contain portions of the
fat body, at least in the bases of their cavities, and next to
378 Zoologica: N. Y. Zoological Society [1II;18
the hypodermis a dense, granular substance, evidently a coag-
ulated liquid produced by the underlying adiphocytes, or tro-
phocytes. . . . Around the bases of tubercles are muscles
so arranged that their contraction must increase the pressure
on the fat and granular liquid and in all probability cause the
later to exude through the hypodermis and delicate chitinous
cuticle onto the surface. The whole arrangement of the
tubercles, in fact, constitutes a system of exudate organs, or
exudatoria, as I shall call them, adapted to secrete substances
that can be licked up by the.ants when they are feeding and
caring for the larvae.”
Wheeler also studied sections through the appendages of
P. latifrons and found a similar arrangement of tissues, but
in the fat cells in these sections he imagined that he saw urate
crystals, which caused him to believe that these cells function
as a storage kidney till the malpighian vessels are sufficiently
developed to excrete. To me Wheeler has not shown any evi-
dence that these appendages are really exudatory, but it is
very probable that they are. He saw no pores in the cuticula
and does not say how his coagulated liquid differs from the
blood, but in support of his view he claims. that we must
interpret the exudatoria as very primitive glands, which in all
probability have arisen as new formations and not as homo-
logues of the embryonic legs. He (p. 318) says:
“They are, as we have seen, small diverticula like the
embryonic legs, consisting of hypodermis and its overlying
cuticula and containing a portion of the fat-body separated —
from the hypodermis by a granular liquid. Now the fat-body
of the insects may be regarded as a diffuse ductless gland, the
cells (trophocytes) of which take certain substances from the
blood in which they lie, store them in the cytoplasm as fat-
globules or proteid granules and later return them to the blood
in a more finely divided, if not chemically modified form. The
exudate which accumulates in the distal ends of the exudatoria
is therefore merely blood charged with nutrient substances
from the fat-cells, and either filters gradually through the hypo-
dermis and overlying cuticle or is forced through them by
1923] McIndoo: Glandular Structure, Termite Guest 379
muscular pressure. At first sight it would seem that the cuticle
must be impervious to such a liquid, but a consideration of
the more recent work on the minute structure of chitin
* * * shows that there is nothing to prevent the passage
of a thin fatty liquid, even if it were not under pressure and
even if the cuticle were much thicker than it is in the ant
larva. The cuticle is a colloid, either of a reticular structure,
as Kapzow believes, or formed of horizontal layers of very fine
fibrille crossing one another at an angle of 60° as most investi-
gators, including Biedermann and Casper, maintain. Between
the fibrille are regularly distributed and extremely fine openings
or ‘pore canals,’ through which a liquid might readily pass as
if the cuticle were a filter.”
In further support of his view, Wheeler cites the work
done on certain meloid, cantharid, lampyrid, coccinellid and
chrysomelid beetles in which a liquid, usually regarded as blood
plasma charged with cantharidin, is discharged from the
articulations of their legs. Wheeler has overlooked my work
(1916) in which I found gland pores in the femoro-tibial
articulations of meloid and coccinellid beetles, and in E’pilachna
borealis I described two types of gland cells which discharge
the liquid through these pores. It is also possible that the
other named insects have glands which discharge secretions
from their legs.
Wasmann (1903), Trigardh (1907 a and b) and Holm-
gren (1909) have published much concerning the exudate
organs of myrmecophiles and termitophiles, but considerable of
the work done on the finer anatomy is not clear to me, although
Wheeler uses their results to support his view.
Wheeler (1910, p. 399) states that students of myrmeco-
phily observed that true guests of ants generally bear tufts of
hairs or trichomes which are assiduously licked by the ants,
and Wasmann (1903), who has written much about these struc-
tures, shows that the trichomes are borne by the integument
at points or depressions where clusters of unicellular glands
open, and that they function by rapidly diffusing some
380 Zoologica: N. Y. Zoological Society [1II;18
aromatic secretion. Wasmann thinks that the secretion is not
liquid, but perhaps a fatty ether, thus being volatile or
etherial. The ants are so fond of it that he thinks it must
affect them very much as a good cigar affects a smoker.
Wheeler adds: “Perhaps it would be nearer the truth to say
that its fascination is more like that of catnip or oil of
bergamot on the various members of the cat family.”
BIBLIOGRAPHY
HOoLMGREN, N.
1909. Termitenstudien 1. Anatomische Untersuchungen. K. Svenska Vetensk.
Handl., Bd. 44, p. 1-215, 3 pl., 76 text figs.
McInpoo, N. E.
1911. The lyriform organs and tactile hairs of araneads. Proc. Phila.
Acad. Nat. Scei., vol. 63, pp. 375-418, 4 pls.
1916. Reflex ‘‘Bleeding’’ of the coccinellid beetle, Ephilachna borealis.
Ann. Ent. Soc. Amer., vol. 9, No. 2, pp. 201-221.
ScHIODTE, M.
1853. On some Staphylinidae, found in the nests of Termites. Proc. Zool.
Soe. London, Part 21, pp. 101-103.
1856. Observations sur des staphylins vivipares qui in habitent chez les
termites. Ann. Sci. Nat., Zool., quart sér., t. 5, pp. 169-183.
SILVESTRI, F.
1920. Contribuzione alla conoscenza dei Termitidi e Termitofili dell’ Africa
occidentale; II—Termitofili, parte seconda. Portici Bollettino
lab. di Zool. gen. e Agraria, vol. 14, pp. 265-319, 32 figs.
TRAGARDH, Ivar.
1907a. Description of Termitomimus, a new genus of termitophilous physo-
gastric Aleocharini, with notes on its anatomy. Zool. Studier
tillag. T. Tullberg, Uppsala, pp. 172-190, 1 pl., 10 text figs.
1907b. Notes on a termitophilous teneid larva. Ark. f. Zool., Bd. 3, pp. 1-7,
1 pl.
WASMANN, E.
1903. Zur naheren Kenntnis des echten Gastverhiltnisses (Symphilie) bei
den Ameisen und Termitengisten. Biol. Centralbl., Bd. 23,
pp. 63, 195, 232, 261, and 298.
WHEELER, W. M.
1910. Ants, their structure, development and behavior. New York.
1918. A study of some ant larve, with a consideration of the origin and
meaning of the social habit among insects. Proc. Amer. Phil.
Soc., vol. 57, pp. 293-343, 12 figs.
Volume III, Number 19
TERMITOPHILOUS APTERYGOTA
FROM BRITISH GUIANA*
By JusTuS W. FOLSOM.
University of Illinois.
(Plates XVITI-XXITI).
The material upon which this report is based was col-
lected at the Tropical Research Station of the New York
Zoological Society, Kartabo, British Guiana, in 1919 and 1920
by Mr. Alfred Emerson.
The following six species are described, all of which are
new, and one of which represents a new genus of Collembola.
THYSANURA: Atelura guiana, A. lepismoidea, A. crispula,
A. cucullata, Nicoletia emersoni. COLLEMBOLA: Borecus pin-
natus.
Types of these have been deposited with the New York
Zoological Society.
All these forms except the Nicoletia are termitophilous.
The genus Atelura, in particular, is strictly myrmecophilous
or termitophilous, and it is appropriate here to bring together
the little that has been published on the habits of this genus.
Additional observations made by Mr. Emerson, appear beyond.
HABITS OF ATELURA
The statement of Escherich (’04) that all the species of
Atelura are either myrmecophilous or termitophilous is prob-
ably still true. Of twenty-four species that I have found re-
corded up to the present eighteen are myrmecophilous or ter-
mitophilous, as are doubtless also the remaining six (from
’ Africa and Australia), descriptions of which I have not seen.
According to the observations of Janet (96a, ’96b) on
Atelura formicaria Heyd. in a nest of Lasius umbratus Nyl.,
* Tropical Research Station, Contribution Number 139.
384 Zoologica: N. Y. Zoological Society [TIE 3
the Atelura can get along without the ants provided it has
available an appropriate nourishment; but is attracted to the
nests of ants by the nutritive fluid that the ants disgorge and
feed to each other. As a pair of ants stand mouth to mouth,
absorbed in the process of feeding, the Atelwra rushes in, grabs
the drop of food and hurries away. This performance is re-
peated with other pairs of ants until the hunger of the
Atelura is satisfied. The guests are tolerated by their hosts
for the good reason that by their agility they are usually able
to elude the ants, but these interlopers are constantly being
chased by their victims, and are sometimes caught and killed.
In regard to the Brazilian species Atelura termitobia Silv.,
Silvestri (01a) gives the following observations.
“In the royal chamber of Anoplotermes tenebrosus I found
with the king and queen ... three examples of Grassiella
[Atelura], which were running about unmolested among the
workers. I placed the royal pair, some workers and the three
individuals of Grassiella together in a glass tube and observed
that the last-named lived in perfect harmony with the termites,
without at any time being pursued by them. Once I saw a
Grassiella on the back of a queen, while the head of a worker,
which had just cleaned the latter, was quite near the head of
the Grassiella. Perhaps these thysanurans also steal food from
the termites, as others do from ants.”
Silvestri (’0la) observed the minute Brazilian species
Atelura synoeketa Silv. in the nests of Hutermes microsoma
Silv., and says concerning them: “In the galleries of Eutermes
I saw three individuals of Grassiella [Atelwra] rambling around
in the midst of the workers, soldiers and larvae, which I cap-
tured and placed alive in a glass tube, together with their
hosts. They walked about everywhere among the termites,
sometimes encountering them head to head without, however,
either species making the slightest impression upon the other.
I saw also a Grassiella enter a gallery in which there were
Eutermes and emerge from the other side unharmed. Never
did I see a Eutermes pursue a Grassiella, or vice versa.” ~-
ee Fy
1923] Folsom: Termitophilous Apterygota 385
Thus the species of Atelura get food and shelter in the
nests of ants and termites, but the benefit is one-sided, for the
hosts gain nothing from the association with their guests.
The species of Atelwra are not usually limited to a single
species of host, but most of them occur with a number of dif-
ferent species of ants and termites.
In distribution the genus is widespread, occurring in all
the faunal realms. The species are small, as a rule, attaining
at most a length of six or seven millimeters.
Mr. Emerson says, in regard to the species of Atelwra in
the nests of termites: “In all cases they seemed to be ignored
by the termites. They moved swiftly about among the termites,
resting very little. In no case did I observe any hostile actions
toward them on the part of the termites. I observed Atelwra
lepismoidea n. sp. several times running about, closely following
worker termites (Nasutitermes (N.) acajutlae Holmgren) but
did not happen to see them obtain regurgitated food or abdominal
secretions from the termites, although it is very possible that at
times they obtain food in this way. They seem to be pantermito-
philous, as shown by the list of the host species in whose nests
they were found.”
Atelura guiana sp. nov.
(Plate XVIII, figs. 1 to 11).
General color golden yellow, due to the scales; body color
yellowish white. Appendages white except in large specimens,
in which the antennae, bases of the legs, and of the pseudo-
cercus (“median cercus”) may be brownish yellow. Small in-
dividuals are yellowish white throughout. Form subelliptical
(fig. 1), broadest across the mesothorax, with width to length
as 1:2.5. Thorax five-eighths as long as the abdomen. Abdomen
smoothly confluent with the thorax in outline. Body strongly
arched. Thoracic terga with rounded lateral borders, project-
ing far down over the bases of the legs. Antennae (fig. 2)
short, about nine-tenths as long as the thorax, or less than
one-third the total length, with usually 12 segments, and occa-
Plate XVIII. ATELURA GUIANA SP. NOV.
1, dorsal aspect, X 25; 2, antenna, X 66; 3, terminal sense organ of antenna,
female, X 790; 4, maxillary palpus, X 86; 5, labial palpus, male, X 96; 6, sense
organ from labial palpus, X 790; 7, tenth urotergite of male, dorsal aspect; the
pegs are ventral in position, X, 86; 8, extremity of abdomen of male, to show
cerci and styli; the lower cercus in the figure is displaced, X 68; 9, ovipositor
and styli, ventral aspect, X 85; 10, parameres and penis, ventral aspect, X 85;
11, typical scale from dorsum, X 505.
ES OOo eee, rt‘ tT
————————eeee Oo
1923] Folsom: Termitophilous Apterygota 387
sionally 11 or 13; distally moniliform; last five segments each
with a distal subsegment; apical segment oblong-elliptical,
with length to width as 9:5 or 5:3; the three or four segments
immediately preceding the apical segment subovate to sub-
globose; apical segment with a terminal branched sensory
organ (fig. 3); second segment in male without a secondary
sexual process, but with a ventral circular radiate sense organ.
Maxillary palpi (fig. 4) with segments in relative lengths as
8:15:16:18:25; last segment elliptico-cylindrical. Labial palpi
(fig. 5) with segments as 6:12:10:40; last segment elliptical,
‘twice as long as broad, with sensory pegs and an apical cluster
of six branching sensory papillae of the type shown in fig. 6.
Tenth abdominal tergite (fig. 7) with a shallow emargination in
both sexes; distal lobes each with about 35 ventral pegs in the
male only. Cerci(fig.8) short; lateral cerci with about 10 segments
(9-12) ; pseudocercus (“median cercus’’?) one and one-half
times as long as the lateral cerci, with about ten segments
(10-12), and with short dorsal pegs on the proximal two seg-
ments in the male. Styli (figs. 8, 9) three pairs, on the 7th
to 9th abdominal segments, respectively. Dorsal valves of
ovipositor (fig. 9) longer than the ventral; ventral valves
semi-ovate in ventral aspect, unsegmented. Parameres of male
(fig. 10) in ventral aspect subtriangular, blunt, divergent,
armed apically with short spines, and mesally with many short
hooks. Body segments with dorsal rows of long outstanding
hairs, mostly along the posterior border of each segment.
Dorsum of head with several short hairs, apically bifurcate.
Dorsal scales dense, varying in form and size, but typically as
in fig. 11. Maximum length, 3.7 mm.
This species is near the Brazilian Atelura (Grassiella)
synoeketa Silvestri (’0la, ’01b) in which, however, the body
is anteriorly oval, the antennae ten-segmented and one-third
as long as the body, and the latter only 1.6 mm. in length.
Twenty-eight specimens, Kartabo, Bartica District, British
Guiana, June 30, July 20, 27, 28, August 11, 20, 21, October
11, 15, November 4; Barakara, Bartica District, July 15. (Field
Nos. 91, 108, 151, 208, 236A, 242, 248, 274, 403, 426, 476.)
388 Zoologica: N. Y. Zoological Society [III; 19
Atelura guiana occurred in the nests of the following species
of termites: Nasutitermes (N.) guyanae (Holmgren), ephratae
(Holmgren), octopilis Banks, swrinamensis (Holmgren) ; Nasu-
titermes (Angularitermes) nasutissimus n. subg. n. sp.; Anoplo-
termes (A.) sp. (No. 151) ; Mirotermes (Cavitermes) tuberosus
n. subg. n. sp. All these species build conspicuous, well formed
nests above the ground with the exception of N. octopilis and N.
nasutissimus, which have rather loosely constructed nests.
Atelura lepismoidea sp. nov.
(Plate XIX, figs. 12 to 20).
Pale, yellow; appendages white. Body (fig. 12) two-fifths
as broad as long, somewhat lepismiform, broadest across the
metathorax, not strongly arched. Abdomen confluent with
thorax, about 1.4 times as long as thorax, tapering in dorsal
aspect, with lateral outlines straight or slightly concave.
Thoracic terga rounded laterally and covering the bases of the
legs. Head rounded in front. Antennae (fig. 13) as long as the
head plus the thorax, with 15, occasionally 16, segments, of
which all but the first 6 are subsegmented, segments 7 and 8
each having two subsegments, and segments 9-15 having three;
first 6 segments in relative lengths about as 19:17:8:4:5:10;
distal segments elongate, elliptical; last segment more than
twice as long as broad, with a terminal sensory organ of the
type shown in fig. 3; second segment in male without a
secondary sexual process. Maxillary palpus (fig. 14) with
segments as 13:17:19:22:32; last segment narrowly subellipti-
cal, three times as long as broad, with a single apical branching
sensory papilla. Labial palpus (fig. 15) with segments as
9:10:21:46; last segment elliptical, twice as long as broad, with
an apical cluster of six sensory papillae of the type shown in
fig. 6. Tenth abdominal tergite with a median rounded
emargination, deep in the male (fig. 16) and shallow in the
female (fig. 17); posterior lobes each with about 20 (17-21),
ventral cones or pegs (fig. 16) in the male only. Cerci short;
pseudocercus (‘median cercus’’) with ten segments, the proxi-
mal two segments bearing a few dorsal pegs in the male
(fig. 16), usually four pairs, becoming successively longer;
Plate XIX.
ATELURA LEPISMOIDEA SP. NOV.
12, dorsal aspect, X 25; 13, antenna, X 86; 14, maxillary palpus, X 100; 15,
labial palpus, X 100; 16, dorsal aspect of extremity of tenth urotergite and
base of pseudocercus of male; the pegs of the former are ventral in position,
X 175; 17, tenth urotergite of female, X 86; 18, left aspect of ovipositor, X 53;
19, parameres and penis, ventral aspect, X 86; 20, typical scale from dorsum,
X 505.
390 Zoologica: N. Y. Zoological Society [TI ; 79
lateral cerci five-sixths as long as the pseudocercus, with 8-10
segments. Styli four pairs, on the 6th to 9th abdominal seg-
ments respectively. Ovipositor extending as far as, or slightly
beyond the lateral cerci, with both pairs of valves segmented
(fig. 18), the ventral valves smaller and slightly shorter than
the dorsal. Parameres of male stout, subcylindrical (fig. 19),
apically armed with many stiff spines. The long stiff dorsal
hairs of the body are limited to the posterior borders of the
segments; there are possibly more of these than are shown in
fig. 12. Dorsal scales typically as in fig. 20, though vary-
ing in form and size. Length, 3.2 mm.
The usual number of antennal segments is 15 in both
sexes, but one female had 16, and small specimens (1.7 mm.
in length) had only 12, 18 or 14.
Forty-one specimens, Kartabo, Bartica District, British
Guiana, July 13, 20, August 7, 18, 21, September 21. (Field
Nos. 100, 197, 225, 242, 248, 339.)
Atelura lepismoidea had as hosts these five species of
termites: Nasutitermes (N.) surinamensis (Holmgren), costalis
(Holmgren), acajutlae (Holmgren), ephratae (Holmgren) ;
Armitermes (N.) teevani n. sp. These build conspicuous, well
formed nests above the ground, with the exception of A. teevani,
the nest of which is usually on the ground.
Atelura crispula sp. nov.
(Plate XX, figs. 21 to.30).
Pale yellow; appendages white. Broadly elliptical (fig. 21) ;
length to breadth as 1.8:1; body strongly arched, reminding
one of a “sow-bug’”; abdomen smoothly confluent with the
thorax in outline, twice as long as the latter. Lateral borders
of thoracic terga rounded, extending down over the legs.
Antennae short, about as long as the thorax, or about one-third
the entire length, with 14 or 15 segments, of which the last 6
or 8 have each two subsegments (fig. 22) ; last segment ellipti-
cal, twice as long as broad, with a terminal branched sensory
organ (fig. 23). Maxillary palpi (fig. 24) with segments in
relative lengths as 8:12:16:17:20; last segment elongate-conical.
—s-
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Plate XX.
ATELURA CRISPULA SP. NOV.
21, dorsal aspect, X 25; 22, last three segments of antenna, X 110; 23, terminal
sensory organ of antenna, X 790; 24, maxillary palpus, X 86; 25, labial palpus,
X 86; 26, tenth urotergite, cerci and pseudocercus of female, X 53; 27, ovipositor
and styli of right side, ventral aspect, X 85; 28, right aspect of ovipositor,
X 86; 29, dorsal aspect of scale from dorsum, X 505; 30, lateral aspect of
scale from dorsum, X 959.
392 Zoologica: N. Y. Zoological Society (III; 19
Labial palpi (fig. 25) with segments as 5:7:14:26; last seg-
ment elliptical, twice as long as broad, with six branching
sensory papillae near the apex. Tenth abdominal tergite sub-
trapezoidal (fig. 26), with posterior margin entire (feebly
emarginate in one specimen) and with postero-lateral borders
rounded. Cerci short and stout (fig. 26) ; pseudocercus (“median
cercus”) one-fifth longer than the lateral cerci, 10- or 11-
segmented; lateral cerci 7- or 8-segmented. Posterior abdominal
segments short, bringing the styli close together. Styli three
pairs, short and stout (fig. 27), on the 7th to 9th abdominal
segments, respectively. Ovipositor short and stout (fig. 28),
the ventral valves exceeding the dorsal; dorsal valves short
and stout, rounded apically; ventral valves in ventral aspect
suboblong, apically truncate, with a subapical transverse suture
(fig. 27). Most of the hairs of the body are dense, short and
stiff; long suberect hairs occurring on the head. Most of the
scales, which clothe the dorsum densely, are of the peculiar
and characteristic form shown in figs. 29 and 30, each scale
having a pair of long curling branches. Length 2.8 mm.
Six specimens, all females, Kartabo, Bartica District,
British Guiana, August 4, 1920 (No. 182).
Atelura crispula had as host Armitermes (A.) percutiens
n. sp., which builds a fairly well constructed dirt nest on the
ground or very close to the ground.
Atelura cucullata sp. nov.
(Plate XXI, figs. 31 to 39).
Golden yellow; appendages white, with pseudocercus
brownish basally. Form elliptical (fig. 31), twice as long as
broad, thorax to abdomen in length as 5:9. Abdomen smoothly
confluent with thorax in dorsal aspect. Body strongly arched,
as in a “sow-bug”. Thoracic terga projecting down over the
bases of the legs.
Antennae (fig. 32) short and stout, one-tenth the length
of the body, 13-segmented, without subsegments; last segment
ovate, the three preceding segments cup-shaped, about as broad
as long. Maxillary palpi (fig. 33) with segments as 1:3:3::4258
——— rc el
o
33
ae ae a
oS ea
37
38
35
Plate XXI. ATELURA CUCULLATA SP. NOV.
31, dorsal aspect, X 17; 32, antenna, X 68; 33, maxillary palpus, X 85; 34,
left aspect of
labial palpus, X 85; 35, tenth urotergite of female, X 85; 36,
from dorsum, X 505; 38, modified
ovipositor, X 86; 37, typical unmodified scale
scale from dorsum, dorsal aspect, X 505; 39, Modified scale from dorsum,
lateral aspect, X 505.
394 Zoologica: N. Y. Zoological Society (ILT; 38
last segment subconical, three times as long as broad. Labial
palpi (fig. 34) with fourth segment three times as long as the
third and lanceolate-elliptical, with length to breadth as 5:2.
Tenth abdominal tergite (fig. 35) trapezoidal, with posterior
border feebly emarginate and postero-lateral angles broadly
rounded. Cerci short and stout; lateral cerci 8-segmented;
pseudocercus (‘‘median cercus”) twice as long as the lateral
cerci, 1l-segmented. Posterior abdominal segments short,
bringing the styli close together. Styli three pairs, on 7th to
9th abdominal segments, respectively. Ovipositor (fig. 36)
with the ventral valves longer than the dorsal; both pairs
segmented; dorsal valves with a terminal cluster of hooks.
Dorsum of body without hairs, excepting a few stiff lateral
hairs on the posterior abdominal segments (fig. 31). Dorsal
scales dense, there being in each transverse series two types
of scales, alternating with each other in position: the simple
type shown in fig. 87 and a peculiar modified form (figs. 38,
39) in which the distal end is bent downward, forming a kind
of hood. Length, 3.6 mm.
Two specimens, both females, Kartabo, Bartica District,
British Guiana, September 9, 1920 (No. 321).
Atelura cucullata occurred with Cornitermes pugnazx Nn. sp.,
- which builds a nest close to the ground or under the ground,
the nest being rather loosely constructed.
Nicoletia emersoni sp. nov.
(Plate XXII, figs. 40 to 48). (Plate XXIII, figs. 49, 50).
Pale yellow; appendages white. Campodeiform (fig. 40),
five times as long as broad; body parallel-sided, the last four
abdominal segments becoming successivly narrower. Thorax
not broader than abdomen, from two-fifths to two-thirds as
long as the latter. Head much longer than prothorax, with a
median dorsal pentagonal sclerite (fig. 40). Eyes absent.
Antennae long (incomplete in the specimens examined) ; first
six segments (fig. 41) in relative lengths as 28:20:9:5:8:10;
first segment large, subovate; second cylindrical. In one speci-
men the first 12 segments are simple (fig. 41) and each of
EEE
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Plate XXII.
NICOLETTA EMERSONIT SP. NOV.
40, dorsal aspect, X 25; 41, base of antenna, X 100; 42, distal segments of
antenna, each primary segment being divided into two subsegments, X 100;
43, galea of right maxilla, ventral aspect, X 250; 44, lacinia of right maxilla,
ventral aspect, X 250; 45, maxillary palpus, X 86; 46, apical sense organs of
left maxillary palpus, X 677; 47, end of. labial palpus, X 86; 48, subapical
sense organs of left labial palpus, X 505.
396 Zoologica: N. Y. Zoological Society [TI a8
the remaining 14 segments is divided into two equal sub-
segments (fig. 42), making apparently 40 segments in all; total
number of segments unknown; second segment in male without
a secondary sexual process. Galea maxilla (fig. 43) with a
pair of terminal papillae. Lacinia (fig. 44) with large apical
tooth, small inner subapical tooth, and an inner subapical
comb, with series of inner teeth and setae as in fig. 44. Maxil-
lary palpus with segments as in fig. 45; last segment sub-
elliptical, three times as long as broad, with six apical
branched sensory papillae (fig. 46), each consisting of a stalk
bearing curving branches that surround a central sensory lobe.
Last segment of labial palpus (fig. 47) clavate, two-thirds as
broad as long, with a group of six distal sensory papillae
(fig. 48) like those of the maxillary palpus except in being
sessile. Tenth abdominal tergite in both sexes with a shallow
median rounded emargination and rounded posterior lobes
(fig. 49). Cerci and pseudocercus of unknown length (being
broken off). Styli eight pairs, on the second to ninth abdominal
segments, respectively; with accompanying eversible vesicles
(fig. 50) except on the ninth segment. Ovipositor in form and
length as in fig. 40. Parameres of male finger-like in form,
each with a terminal seta (fig. 50). Clothing of dense short
setae of irregular lengths, with long setae on the head, lateral
borders of the thoracic segments, and postero-lateral angles
of the abdominal segments. Length, 4 mm.
Nicoletia emersoni approaches N. neotropicalis Silvestri
(01b), from Argentine, Paraguay and Brazil, but differs from
that species in having wider abdominal segments, and the
pseudocercus of the male not narrower than the cerci, as well
as in other respects. The first three segments of the maxillary
palpus lack the spines shown in the figure by Escherich (’04)
and the terminal sensory papillae of the maxillary palpus differ
in form in the two species.
Two specimens, one of each sex, Penal Settlement, Bartica,
March 24, 1919. Not termitophilous.
I take pleasure in naming this species after Mr. Alfred
Emerson.
1923 ] Folsom: Termitophilous Apterygota 397
Borecus gen. nov.
Eyes and postantennal organs absent. Antennae short,
but longer than the head, four-segmented, without subsegments
or rings. Mouth parts biting. Prothorax reduced but not
rudimentary. Mesonotum large but not produced over the
base of the head. Fourth abdominal segment much longer
than the third. Tibiotarsi two-segmented. Unguis with a pair
of large inner basal lobes and a pair of smaller outer basal
lobes, or pseudonychia. Unguiculus well developed, with a
strong outer basal lobe. One tenent hair. Furcula strongly
developed, appended apparently to the fifth abdominal segment.
Manubrium longer than dentes. Dentes unsegmented, smooth
dorsally, without crenulations or rings, and without chitinous
hooks, but with two dorsal longitudinal series of large feather-
like setae, and clothed ventrally with scales. Mucrones strongly
elongate, non-lamellate, tomocerine in type but without the basal
tooth, with a large apical tooth, a large subapical dorsal tooth,
and a varying number of smaller dorsal teeth between the
subapical tooth and the base of the mucro. Dens with an
apical pair of long hyaline scales extending under the mucro.
Clothing of both scales and setae, the latter mostly fringed and
frequently clavate.
This new genus falls near the peculiar genus Oncopodura,
described from the species hamata, from the Crimea, by Carl
and Lebedinsky (’05), and represented also by a second species,
crassicornis Shoebotham (’11) from England and Poland
(Stach ’21). Oncopodura is unlike Borecus in having the fol-
lowing differentiating characters: Postantennal organs present
or absent. Prothorax rudimentary. Fourth abdominal segment
slightly longer than the third. Unguis simple, without inner
teeth, without pseudonychia, and without large inner basal
lamellae, though narrow pointed lateral lamellae may or may
not be present. Unguiculus simple, without teeth or lobes.
Tenent hairs absent immediately above the unguis, but a strong
clavate hair projects from the tibiotarsus about midway from
the base to the apex on the second pair of legs only. Dentes
with chitinous hooks, with slender fringed hairs dorsally, but
Plate XXIII.
NICOLETIA EMERSONI SP. NOV. BORECUS PINNATUS GEN. ET SP. NOV.
NICOLETIA EMERSONI.—49, dorsal aspect of tenth urotergite of female,
X 70; 50, ventral aspect of male to show Parameres and last three pairs
of styli, X 88.
BORECUS PINNATUS.—51,
53, concave aspect of left hind unguis, X 632; 54,
and end of dens, X 380;
56, left aspect of left mucro and end of dens, X 404; 57, dorsal clavate fringed
1923] Folsom: Termitophilous Apterygota 399
without broad pinnate setae. Mucro with a well developed
hyaline membrane, or lamella.
The genus Borecus may be placed with Oncopodura in the
subfamily Oncopodurinae of the family Entomobryidae. The
affinities of Oncopodura lie with Cyphoderus, as Stach (’21)
has pointed out.
Borecus pinnatus gen. ¢t sp. nov.
(Plate XXIII, figs. 51 to 58).
White throughout (fig. 51). Eyes and postantennal organs
absent. Antennae longer than the head, varying from slightly
longer (in small specimens) up to 1.5 times as long as the
head (large specimens) ; four-segmented, without subsegments;
antennal segments varying greatly in relative lengths, but with
third segment always much shorter than the second or the
fourth; first segment cylindrical; second clavate, becoming
subcylindrical with age; third clavate; fourth subconical,
elongating with age; third segment with a subapical pair of
elliptico-cylindrical sensory pegs lying exposed, not covered
by an integumentary fold; fourth segment distally with short
curving sensory setae; antennal clothing of abundant short
minutely fringed setae, with occasional outstanding simple
setae. Mouth parts biting. Prothorax reduced. Mesonotum
rounded anteriorly, covering the prothorax, the latter ab-
normally exposed in fig. 51, but not projecting over the base
of the head. Metanotum two-thirds as long as mesonotum.
Tibiotarsi with a transverse suture one-third from the apex;
femora with a transverse suture near the apex; these sutures
not always evident, however. Legs clothed with abundant
fringed setae, with one (occasionally two) long stout outstanding
fringed setae on each segment of each leg. Unguis (figs. 52,
53) straight basally, curving apically, with a pair of large
inner basal sublanceolate lobes extending half the length of
the claw; with a pair of smaller basal lateral lanceolate lobes
(pseudonychia) ; and with an obscure tooth (doubled?) or angle
near the middle of the inner margin. Unguiculus (fig. 52)
large, extending three-fourths as far as the unguis, with a
large basal subovate acuminate outer lobe. Hind claws the
400 Zoologica: N. Y. Zoological Society [TI1; 19
largest. Tenent hair single, feebly knobbed. Ventral tube
with a pair of eversible rounded vesicles. Fourth abdominal
segment much longer than the third, varying in relative length
but usually from three to five times as long as the latter. Rami
of tenaculum quadridentate; corpus with a single ventral seta.
Furcula apparently appended to the fifth abdominal segment,
attaining the ventral tube in the larger specimens, but not in
the smaller. Manubrium elongate, scarcely tapering; dorsally
with many setae, either simple, or clavate and fringed; ven-
trally with scales. Dentes two-thirds as long as manubrium,
slightly tapering, one-segmented, smooth dorsally, without
crenulations or rings, with two dorsal rows of large pinnae,
or feather-like setae (figs. 54-56) which, proceeding posteriorly,
become successively larger. The setae of the outer series are
5-10 in number (according to age), the proximal one, two, or
more setae being relatively simple and the remainder modified.
The setae of the inner series are 4-7, the proximal one or two
being simple, and the last of the pinnate setae (subapical)
being exceptionally long (figs. 55, 56), sometimes a little longer
than the mucro. At the base of the dens dorsally is a stout
clavate erect fringed seta (fig. 57). Under the mucro are two
long, narrowly elliptical, hyaline, minutely striated scales (figs.
55, 56), one of which may extend beyond the mucro; these
scales arising near the apex of the dens. Each dens bears a
lateral row of 4-15 simple or feebly fringed setae; ventrally
the dentes are clothed with scales. Mucrones (figs. 54-56)
one-half to three-fifths as long as the dentes, slender and
tapering in dorsal aspect, in form much like those of Tomocerus,
with a large apical tooth, usually hooked, and a dorsal subapical
tooth subequal to the first. Between the base of the mucro
and the subapical tooth is a dorsal row of small intermediate
teeth (fig. 54), one of which is larger than the others. These
intermediate teeth may, however, be absent, as in fig. 56, and
when present vary in number from 1 to 11. Mucronal lamellae
are absent, excepting in some specimens a minute lamella
extending forward from the anteapical tooth as in fig. 56. The
seales that clothe the head and body dorsally differ in size and
vary in form from elliptical to oval, ovate or roundish, but
are commonly elliptical, as in fig. 58. Fringed setae occur on
1923 ] Folsom: Termitophilous Apterygota 401
the head anteriorly, on the fourth abdominal segment poster-
iorly, and on the fifth and sixth abdominal segments, many of
the setae on the genital and anal segments being strongly
clavate. Length of specimens, 0.54 mm. to 1 mm.
This species varies considerably, some of the variation
being correlated with the age of the individual (as indicated
by its size), and some being independent of age. The dorsal
setae of the dentes increase in number with the age of the
individual, and more of them become pinnately modified. In
several small specimens, 0.54 mm. to 0.87 mm. in length, the
dorsal intermediate teeth of the mucrones were absent; in one
individual, 0.9 mm. in length, there was one of these teeth
on each mucro; in five specimens, all 1 mm. long, the number
of intermediate teeth varied from one to eleven.
Fifteen specimens, Kartabo, Bartica District, British
Guiana, June 21, 30, July 13, 30. (Field Nos. 57, 87, 92, 101.)
Hosts.—Rhinotermes (R.) marginalis (L.), Nasutitermes
(N.) costalis (Holmgren), Nasutitermes (N.) octopolis Banks,
Nasutitermes (N.) acajutlae (Holmgren).
Borecus pinnatus was with four species of termites: Nasu-
titermes (N.) octopilis Banks, costalis (Holmgren), acajutlae
(Holmgren) ; Rhinotermes (R.) marginalis (Linné). All but
the last of these species build conspicuous, well formed nests
above ground.
Mr. Emerson reports that this collembolan was found very
often in large numbers in nests, and says, “I think it very
likely that the same species is found outside of termite nests,
but found one striking illustration of their true association with
the termites. One morning I found an entire termite colony
of the species Nasutitermes (N.) costalis (Holmgren) migrat-
ing over a sandy stretch of ground. The king and queen ter-
mites in addition to the workers and soldiers were all going
along in a long file from a clump of bamboos to our dwelling.
In addition to numerous other guests found among the termites,
springtails were also running along the trail and were undoubt-
edly migrating with the termite colony. To my mind, this is a
402 Zoologica: N. Y. Zoological Society [Iil; 3
conclusive proof that the springtails have a distinct liking for
life among the termites. They run rapidly about among the
termites in the nests and I have never observed any hostile
action toward them on the part of the termites. They are also
pantermitophilous, as illustrated by the different host species,
and in the case where they were found associated with Rhino-
termes (R.) marginalis (Linné) there was no particualr nest,
the termite colony being very small. It therefore seems to me
that the association of the Collembola with the termites is
rather more loose than is the case with the species of Atelwra
and most other synoeketes, and it is highly probable that they
are not entirely dependent upon the termites for their existence.”
BIBLIOGRAPHY
CARL, J., AND LEBEDINSKY, J.
1905. Materialen zur Hohlenfauna der Krim. II. Aufsatz. Ein neuer
Typus von HGéhlenapterygoten. Zool. Anz., vol. 28, pp. 562-565,
figs. 1-6.
ESCHERICH, K.
1904. Das System der Lepismatiden. Zoologica, Heft 43, 164 pp., 4 pls.,
67 figs.
JANET, C.
1896a. Sur les rapports des Lépismides myrmécophiles avee les Fourmis.
Comptes rend. Séances Acad. Se., vol. 122, pp. 799-802, 1 fig.
1896b. Sur les Lepismina polypoda, Grassi, et sur ses rapports avec les
Fourmis. Bull. Soc. Ent. France, pp. 131-138.
SHOEBOTHAM, J. W.
1911. Some Records of Collembola New to England, with Description of a
New Species of Oncopodura. Ann. Mag. Nat. Hist., ser. 8, vol. 8,
pp. 32-39, pl. 3.
SILVESTRI, F.
1901a. Descrizione di nuovi Termitofili e relazioni di essi con gli ospiti. Boll.
Mus. Zool. Anat. comp. Univ. Torino, vol. 16, pp. 1-24.
1901b. Materiali per lo Studio dei Tisanuri. Bull. Soe. Ent. Ital. vol. 33,
pp. 204-249, figs. 1-48.
StracuH. J.
1921. Vorarbeiten zur Apterygoten-Fauna Polens. Teil II. Apterygoten
aus den Pieniny. Bull. Acad. Polonaise Se. Lett. (1919), pp.
133-233, pls. 5-7.
Volume III, Number 20
ON THREE APPARENTLY NEW SPECIES OF
TERMITAPHIS (Hem. Het.)* *
By HAROLD MorRISON ?
Bureau of Entomology, Department of Agriculture, Washington, D. C.
(Plate XXIV)
The following new species of Termitaphis were received
for study from Dr. W. M. Mann, to whom they had previously
been sent for examination by Dr. Alfred Emerson of the
University of Pittsburgh.
The genus Termitaphis was first established by Wasmann °
in 1902, but was not accurately characterized until 1911, when
Silvestri ¢ described two new members of the genus, presented a
critical generic diagnosis and erected a new family in the
suborder Heteroptera for its reception. Later, Mjéberg® gave
a short description of a species obtained in Australia, with the
promise to describe it in greater detail at some future date.
This extended description has apparently never been published,
and the original characterization is too brief to permit a com-
parison of Mjéberg’s species with those described below.
Wasmann’s species was obtained in Columbia, and the first
species described below may possibly be identical with it,
although this cannot be definitely established on account of the
incompleteness of the description of the genotype and the
probable inaccuracy of the figures given by Wasmann. It has
accordingly been described as new.
*
* Tropical Research Station, Contribution Number 140.
1 Tropical Research Station, Contribution Number 140.
?The figures illustrating the structural characteristics of these species have
been worked out and prepared by Emily Morrison.
Wasmann, E. Species novae Insectorum Termitophilorum ex America
Meridionali. In Tidj. voor Ent. vol. 45, 1902, pp. 95-107, 9 pl.
*Silvestri, F. Sulla Posizione Sistematica del Genera Termitaphis Wasm.
(Hemiptera), ete. In. Boll. del Lab. di Zool. Gen. e. Agraria della. Rse. Sup.
d’Agr. in Portici. vol. 5, 1911, pp. 231-236, fig. I-VI.
’Mjoberg, E. Preliminary description of a new representative of the family
Termiticorddae Silv. In Entomologisk Tidskrift, ete. vol. 35, 1914, p. 98, 9 fig.
Plate XXIV.
TERMITAPHIS GUIANAE SP. NOV., T. INSULARIS SP. NOV.,
T. TRINIDADENSIS SP. NOV.
T. GUIONAE.—1, outline of body showing shape, lobes and sultures, dorsal X30; 2, antenna,
X57.5: 3, dorsal pore, X1280; 4, dorsal pore, second type side view, X1280; 5, same as 4,
view from above, X1280; 6, marginal lobe of abdominal segment, dorsal, X220; 7, portion
of 6 ventral view, X440; 8, apical abdominal lobes with anal opening ventral, X220; 13,
dorsal seta, X1280; A
T. INSULARIS.—9, marginal flabellum, X640, 11, outline showing marginal lobes and
sutures, dorsal, X17.5.
T. TRINIDADENSIS.—10, marginal flabellum, X1280; 12, apex of anterior tibia, two sides,
X220; 14, outline of body showing marginal lobes and sultures, dorsal, about X17.5.
se eS
1923] Morrison: Three Termitaphis 405
Termitaphis guianae sp. nov.
(Plate XXIV, Figures 1 to 14)
Very similar to T. mexicanus as described. by Silvestri,
differing only in certain details: length 2 mm. width 1.44 mm.,
arrangement of the dorsal sutures, particularly the anterior
pseudo-suture of the mesothorax, differing somewhat from that
shown in Silvestri’s drawing (cf. fig. 1 with fig. III of Silvestri’s
paper) ; the fourteen marginal lobes of each half of the body
bearing tubes and flabella as follows, counting from the cephalic
apex: 7, 3 (head); 9-10 (prothorax) ; 4, 4 (mesothorax) ; 4-5
(metathorax) ; 5-6, 6, 6-7, 6-7, 6, 6, 4, 3 (abdomen) the usual
number for each prothorax lobe being nine, for each meso-
thoracic lobe, four, and for each metathoracic and abdominal
lobe, anterior to the last two, six, differing in these from
mexicanus where the usual number for the mesothoracic lobes
is five, for the anterior abdominal lobes, seven, and for apical
abdominal lobes, two instead of three flabella; these flabella
with minute setae or denticulae over the entire surface as well
as along margin and with similar minute setae very numerous
on the lobes; lateral margins of each lobe conspicuously serrate,
not smooth; derm pores of two sorts, some simple cylindrical
tubes surrounded by a circular clear area, and with a funnel-
shaped opening, others with an intricate radiate arrangement;
with occasional stout setae dorsally, each set at the end of a
tube through the derm; with two larger thoracic and six well
developed but smaller circular abdominal spiracles followed
by an incompletely developed seventh abdominal spiracle;
anterior legs in addition to curved hairs and some stiff spines
with a straight comb of long stiff hairs at apex of tibia; other
characteristics, so far as can be observed, agreeing very
closely with those of mexicanus as described and figured by
Silvestri.
This species has been described from three specimens with
the following information:
Kartabo, British Guiana, No. 48. Emerson.
Host.—Leucotermes crinitus (Emerson).
HoLoTyPE.—(A slide mount) and paratypes. Cat. No.
25034, U. S. National Museum.
406 Zoologica: N. Y. Zoological Society [III ; 20
Termitaphis trinidadensis sp. nov.
Very similar to the preceding, size almost identical, length
2 mm., width 1.5 mm., the chief differences being found in the
somewhat more elongate and more slender marginal flabella
and in the different numbers of marginal flabella for corres-
ponding lobes of the body, the arrangement of these being as
follows: 7:3 (head); 8 (prothorax); 4, 4 (mesothorax); 3-4
(metathorax) ; 4, 4, 4, 4, 4, 4, 4, 3 (abdomen).
This species has been characterized from a single specimen
with the following information:
‘“‘Port-of-Spain, Trinidad. 26-XI-20. Emerson. No. 495a.”
Host.—Leucotermes tenuis (Hagen).
HoLOTYPE.—(A slide mount). Cat. No. 25035. U. 8S. Nat.
Museum.
Termitaphis insularis sp. nov.
Body somewhat more elongate than in the two preceding
species, length, 2.75 mm.; width, 1.6 mm.; legs more elongate
and more slender than with the last, more nearly resembling
those of T. guianae; differing conspicuously from all the other
accurately described species in that the body has only 12 lobes
on each half and from the two preceding species in the much
more elongate and slender marginal flabella, these resembling
those of T. subafra Silvestri; the arrangement of these flabella
as follows: 7, 3 (head); 10-11 (prothorax) ; 11 (meso and
metathorax) ; 6, 7-8, 6-7, 7, 7, 6, 4, 3 (abdomen), the fusion
occurring in the meso and metathoracic lobes, usually three on
each side; in other respects quite similar to the two species
already described.
Characterized from a single specimen with the following
information:
“Port-of-Spain, Trinidad. 26, XI, 20. Emerson. No. 495a.”
Host.—Leucotermes tenuis (Hagen). }
HoLotTyPE.—(A slide mount). Cat. No. 25036. U.S. Nat.
Museum. :
1923] Morrison: Three Termitaphis 407
There may be seen in the posterior portion of the abdomen
of the specimen on which this species is based, some curled
and twisted tubes and some other less definite, apparently
chitinized structures which suggest themselves as_ possibly
retracted male sexual organs. No definitely developed copula-
tory or other external sexual modifications of the apex of the
abdomen, such as are to be found in many of the Heteroptera,
have been noted in these specimens, and nothing appears to be
known regarding the differences in sexes in the genus. Since
the last two species described were received in the same con-
tainer, and were collected from the same nest of termites, the
further possibility presents itself that they are the two sexes
of one species, although the obvious differences in certain of
the comparable structural characters fully justifies their separa-
tion at this stage of our knowledge of these insects.
By way of comment, it may be noted that the writer’s
interpretation of the body segmentation differs from that given
by Silvestri in his generic diagnosis and in his figures of both
mexicanus and subafra. In both species Silvestri shows a total
of fourteen lobes for each half of the body of which two are
allotted to the head, two to the prothorax, two to the meso-
thorax, one to the metathroax, and one each to five complete
and two incomplete abdominal segments. It would appear that
Silvestri failed to observe the anterior thoracic spiracles,
located close to each prothoracic coxa, and highly probable that
his specimens actually possess a partially suppressed seventh
abdominal spiracle, as do all of those examined in connection
with the preparation of the preceding descriptions.
These two additions give a total of nine more or less
developed spiracles for the genus, instead of seven as given by
Silvestri, of which the two anterior are slightly but distinctly
larger and appear to be placed at the anterior margins of the
under surface of the meso and metathorax, leaving seven pairs
for the abdomen and resulting in a reassignment of the
marginal lobes of those species having fourteen lobes to each
half of the body as follows: two to the head; one to the pro-
thorax, two to the mesothorax, one to the metathorax, and
eight to the abdomen.
408 Zoologica: N. Y. Zoological Society [III ; 20
Silvestri’s key to the species of this genus may be modified
as follows to include the newly described species:
a. Flabella of marginal lobes simple entire setae®
circumvallata Wasm.
aa. Flabella of marginal lobes broader, spatulate, margins
minutely serrate.
b. Margin of body with fourteen lobes to each half.
c. Marginal flabella short and broad, at most hardly
more than twice as long as wide.
d. Apical abdominal segments with two flabella on
each lobe; anterior abdominal segments normally
with seven flabella on each margin
’ mexicanus Silv.
dd. Apical abdominal segments with three flabella
on each lobe; anterior abdominal segments nor-
mally with six or fewer flabella on each margin.
e. Anterior abdominal segments normally with
six flabella on each margin; legs relatively
slender... 3:32.09: eee guianae, sp. nov.
ee. Anterior abdominal segments normally with
four flabella on each margin; legs relatively
short and steutt.2c.0ese! trinidadensis, sp. nov.
cc. Marginal flabella elongate, slender, the length much
more than twice the width__..__.........___. subafra Silv.
bb. Margin of body with twelve lobes to each half; fla-
bella elongate. 2 ee eee insularis, sp. nov.
* Based only on figures and description of Wasmann.
Volume III, Number 21
ON FOUR TERMITOPHILOUS MILLIPEDS
FROM BRITISH GUIANA*
By RALPH V. CHAMBERLIN
(Plates XXV, XXVI, XXVII)
The types of the new millipeds here described were taken
by Mr. Alfred Emerson from nests of termites at Kartabo,
Bartica District, British Guiana, in December, 1919, and August,
1920. All three pertain to the Cryptodesmidae in the broad
sense, a group of mostly very small polydesmoid forms which
in tropical regions appear to be rather numerous and wide-
_ spread although commonly overlooked because of their small
size and obscure habits. A number of species have previously
been reported as occurring in termite nests and others in
ant nests.
Leuritus gen. nov.
Body consisting of the head and twenty segments.
Head concealed from above by the collum which projects
widely beyond it. Antennae moderately long, with the fifth
joint much longer than the sixth.
Collum with a broad horizontal border which is divided
by radial sulci into twelve lobes. Central region of collum
moderately convex and tubercular.
The keels of the succeeding tergites wide and horizontal,
the lateral border divided into three areas by radial sulci ex-
cepting the fifth, which shows two, and the most caudal ones,
which show a fourth lobe in some degree developed. The
poriferous keels in respect to lobation not differing from the
non-poriferous. Repugnatorial pores minute, situated near
caudalateral corner of keels of segments V, VII, XI, X, XII,
_ XIII, and XV to XIX; pores not opening on special cones or
lobes and detected with difficulty. Tergites between keels bear-
* Tropical Research Station, Number 141.
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Plate XXV.
GASATOMUS EMERSONI SP. NOV., TIDOTERUS SEQUENS SP. NOV.
1, the four anterior tergites from above; 2, the four posterior tergites from above;
3, caudal end of body from below; 4, keel of thirteenth segment cleared and
view by transmitted light to show the course of the repugnatorial duct; 5, antenna;
6, portion of collum more highly magnified to show hairs; 7, caudal portion of
anal tergite from below under higher magnification to show hairs and setae;
8, a leg; 9, a gonopod of male in subcaudal view.
Dee eee ee EOE Stl
1923] Chamberlin: Termitophilous Millipeds 413
ing three transverse rows of tubercles, none of them specially
enlarged to form more prominent longitudinal series.
Last tergite broad, flat, rounded behind, the margin tri-
lobed; widely surpassing the anal valves.
Second article of legs longer than the third and than the
sixth.
Gonopods of male with the basal joint enlarged as usual,
but the telopodite extending beyond it and clearly exposed. The
telopodite deeply furcate, the two branches slender, smooth
and curved.
Genotype—Leuritus termitophilus sp. nov.
In the general form and proportions of the collum and
keels much resembling the West Indian genus Tridesmus; but
the posterior lobes of the poriferous keels are not at all en-
larged as in that genus and the caudal tergite is broad and
rounded instead of triangular. It is also quite distinct in the
structure of the gonopods.
Leuritus termitophilus sp. nov.
(Plate XXV, Figures 1 to 7)
General color flavous, often appearing in part brownish
because of adherent foreign material. Surface of tergites
densely clothed with very short hairs which are evident par-
ticularly on borders of keels and collum. (Cf. pl. XXV, figs. 6
and 7.)
The head is compressed dorsoventrally and presents a sharp
transverse edge across vertex and down each side. Sulcus
sharply impressed across vertex, fureate below, sending a
branch to base of each antenna. A deep groove on each side
of the head above into which the basal articles of the antenna
fit. Labral margin with three teeth.
Fifth joint of antennal article about two-thirds as thick
as long and twice as long as the sixth article. (Cf. pl. XXV,
fe.5.)
414 Zoologica: N. Y. Zoological Society (TIT; 24
Collum depressed, broad; rim broad, with deep radial
sulcis, but with the corresponding marginal notches slight;
surface within the border tubercular. (Pl. XXV, fig. 1.)
On the keels of the following tergites in general the
anterior border is elevated; the lateral lobes indicated by radial
furrows but the marginal notches slight or obsolete. Dorsum
of tergites between keels densely tubercular, the tubercles not
unequally developed in a way to leave longitudinal series of
more prominent ones; tubercles in general in three transverse
rows on each tergites; the tubercles form also longitudinal
rows which on each tergite tend to run ectad of directly for-
ward from caudal end. (Pl. XXV, figs. 1 and 2.) The course
of the duct from the repugnatorial gland to the pore is indicated
in pl. XX\V, fig. 4.
The anal tergite with caudal margin broad, at middle
nearly straight, the lateral corners rounded; sides subparallel;
surface clothed with very fine short hairs; on ventral surface
a transverse series of seven short setae, with two on median
lobe caudad of these and one cephalad of them. (Cf. pl. XXV,
figs. 3 and 7.)
The gonopods of the male as represented in pl. XXV, fig. 9.
Length, 4.5 mm.; width, 1.5 mm.
Locality—British Guiana, Kartabo, Bartica District, Aug.
16, 1920. Fifteen specimens, some of which are only partly
grown, taken from the nest of Nasutitermes (Nasutitermes)
brevipilus (Emerson).
Stenitus gen. nov.
Consisting of the head and twenty segments.
Fifth joint of antennae much exceeding the sixth.
Collum high and convex, with sides steep; the rim de-
pressed and descending almost in line with the sides above it,
narrow, with ten areas separated by radial sulci.
Keels of tergites in general short and bent downward, the
lateral border of the keels of the second tergite showing three
1923] Chamberlin: Termitophilous Millipeds 415
areas or lobes separated by sulci, the succeeding ones showing
only two. Tergites between keels tubercular, the tubercles
forming distinct longitudinal rows of which one a little each
side of middle and one on each side are higher or more prom-
inent, especially on the more posterior tergites; three tubercles
in each row on each segment. Repugnatorial pores minute as
in Leurodesmus, situated toward posterior lateral corner on
segments V, VII, IX, X, XII, XIII, and XV to XIX inclusive.
Anal tergite large and freely exposed, caudally well
rounded.
Second joint of legs longer than second and than sixth.
Genotype.—_S. guiananus sp. nov.
Stenitus guiananus sp. nov.
(Plate XXVI, Figures 1 to 8)
General color flavous.
Median sulcus across vertex of head deep, furcate below,
a branch running to each antennal socket. Vertex protruding
each side of sulcus and densely granular. Head with a furrow
on each side for the reception of the basal joints of the anten-
nae. Fifth joint of antennae less than two times longer than
wide and between four and five times longer than the sixth
gt. (Pi. XXVI, fig. 7.)
Collum strongly tubercular excepting over the rim. Rim
strongly depressed, the radial lines distinct, the corresponding
marginal notches not deep.
Keels of the second tergite with marginal notches weakly
developed; the notches scarcely evident on following keels when
viewed at right angles to their surface, but when viewed
obliquely the radial furrow may give the appearance of a
marginal notch. Cf. pl. XXVI, figs. 1, 2 and 3.) Four rows
of tubercles along the dorsum more prominent, particularly the
two submedian ones; two rows of smaller tubercles between
each two more prominent rows.
COFCO
Witiee
8
Plate XXVI.
STENITUS GUIANANUS SP. NOV.
1, anterior tergites in lateral view; 2, posterior tergites, dorsal view; 3, posterior
end of another specimen in outline, with tergites depressed to bring keels into a
more nearly horizontal position; 4, caudal end, ventral view; 5, sixth segment,
anterior view, in outline; 6, tenth and eleventh keels viewed at right angles to
surface; with course of repognatorial duct indicated on the tenth by dotted line;
7, antenna in outline; 8, a leg in outline.
1923] Chamberlin: Termitophilous Millipeds 417
The anal tergite much exceeding the nineteenth, caudally
rounded, the margin notched at median line and on each side
as shown in pl. XXVI, fig. 4.
Proportions of joints of legs as shown in pl. XXVI, fig. 8.
Length, 4.4 mm.; width, 7 mm.
Locality.—British Guiana, Kartabo, Bartica District. One
adult female, all but the four anterior segments of another,
and three immature specimens taken Aug. 16, 1920, from a nest
of Nasutitermes (Nasutitermes) brevipilus (Emerson).
Gasatomus gen. nov.
Consisting of the head and twenty segments.
Head completely covered by the collum. Fifth joint of
antennae much exceeding the sixth in length.
Border of collum nearly horizontal; divided by radial
furrows into twelve areas or lobes, with corresponding marginal
notches or crenations. Collum within the border convex and
strongly tubercular.
Keels of succeeding tergites of moderate length; presenting
laterally three lobes or areas separated by sulci and correspond-
ing marginal incisions excepting the fifth, which have but two
lobes, and the seventeenth, eighteenth, and nineteenth, which
have four. Each repugnatorial pore opening at the apex of a
special and prominent process projecting laterad from the
posterior lobe. Pores present on segments V, VII, IX, X, XII,
XIII, XV and XVI. Posterior margin of keels incised.
Last tergite freely exposed; much exceeding the processes
of the nineteenth keels; with six marginal lobes or crenations.
Second joint of legs longer than the third and than the
sixth.
Genotype.—G. emersoni sp. nov.
Much suggesting Cynedesmus, a genus known from Central
America, the West Indies, and the Canary Islands. From that
genus it differs clearly in the lobation of the keels.
ii a0. 75)
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Plate XXVII.
GASATOMUS EMERSONI SP. NOV.,TIDOPTERUS SEQUENS SP. NOV.
GASATOMUS EMERSONI.—1, anterior end, dorsal view; 2, caudal end, dorsal
view; 3, eleventh and twelfth right keels, dorsal view; 4, anal scale; 5, antenna;
6, a leg.
TIDOPTERUS SEQUENS.—7, caudal end of body, dorsal view; 8, leg of eleventh
degment; 9, gonopods of male, subventral view.
1923] Chamberlin: Termitophilous Millipeds 419
Gasatomus emersoni sp. nov.
(Plate XXVII, Figures 1 to 6)
Fulvous, appearing brown from adherent foreign material.
Vertex and frons of head densely granular; head smooth
below level of antennae. Fifth joint of antennae about one
and two-thirds times as long as thick, and not fully twice as
long as the sixth article. (Pl. XXVII, fig. 5.)
Border of collum horizontal, its upper surface a little
concave; the radial furrows sharply impressed and the cor-
responding marginal notches pronounced. Central portion of
collum strongly convex and densely tubercular, with two trans-
verse rows of larger tubercles as shown in pl. XXVII, fig. 1.)
The keels of the following tergites are trilobed laterally
excepting those of the fifth, which are bilobed, and these of the
seventeenth, eighteenth and nineteenth, which are four lobed.
Caudal margin of keels with two principal lobes separated by
narrow incisions, and a minor one proximad of these; the
anterior margin with one distinct incision at base of the
anterior lateral lobe. On the dorsum between the keels four
longitudinal series of enlarged tubercles which greatly exceed
the intervening ones in size, the three tubercles of each of these
series on each tergite more or less confluent at base. Tubercles
of the smaller size are also present on the keels excepting on
the lateral lobes. The posterior tubercles of the two submedian
principal series on the more caudal tergites are enlarged and
project caudad from the plate. See further pl. XXVII, figs. 1
and 2.
Anal tergite with six small lobes or crenuli as shown in
pl. XXVII, fig. 2. The anal scale broadly triangular, the caudal
angle obtuse. (Cf. pl. XXVII, fig. 4.)
Form and proportions of joints of legs as shown in pl.
my Il, fig. 6.
Length, 7 mm.; width, 1.2 mm.
Locality.—British Guiana, Kartabo, Bartica District. Two
females taken Aug. 30, 1920, from nest of Nasutitermes (Sub-
ulitermes) baileyi (Emerson).
420 Zoologica: N. Y. Zoological Society [TIT sam
Tidopterus gen. nov.
Composed of head and twenty segments.
Head nearly wholly covered by the collum from above.
Fifth article of antennae but little longer than. the sixth.
Border of collum divided by radial sulci and marginal
notches into ten lobes.
Keels of succeeding tergites narrower than in Gasatomus,
but the lateral margins lobed as in that genus, excepting that
the sixteenth as well as the seventeenth, eighteenth, and
nineteenth keels, has four marginal lobes. Pores present on
segments V, VII, XI, X, XII, XIII, and XV. The pore processes
as in Gasatomus as to form and position.
Nineteenth tergite with median caudal border bowed caudad
beyond the caudal processes of keels, not forming the bottom
of a quadrate excision between the latter. Last tergite exposed
from above; bearing two large tubercles which conceal or nearly
conceal it from above; margin not caudally incised at middle
behind but with crenations, which are typically setiferous, on
each side.
Second joint of legs not much differing in length from
third, but both the second and third joints shorter than the
sixth.
Telopodite of gonopods conspicuously exposed.
Genotype.—Tidopterus sequens sp. nov.
Related to Cynedesmus and Gasatomus but differing from
both in lacking pores on the sixteenth segment and in the form
of the nineteenth and twentieth tergites.
Tidopterus sequens sp. nov.
(Plate XXVII, Figures 7. to 9)
General color flavous.
Head with vertigial and frontal regions densely granular
or finely tubercular on each side of the sulcus. The vertex of
the head protrudes a little beyond the collum at the middle in
1923] Chamberlin: Termitophilous Millipeds 421
dorsal view. Fifth article of antennae nearly equal in length
and thickness; not much longer than the narrower sixth article,
the lengths of these articles being about as 5:4.
Border of collum nearly horizontal, narrow, the incisions
between the lobes deep. Convex surface densely tubercular,
with two transverse rows of larger, well separated tubercles.
Dorsum of the following tergites in general strongly
convex and densely tubercular, with four principal longitudinal
series of larger tubercles. These higher tubercles are propor-
tionately larger on the more caudal tergites where those at
caudal border project conspicuously caudad. The keels are
obviously narrower than in the type of Gasatomus, showing but
a single lobe on caudal margin; but the lateral lobation and
the form and relations of the pore bodies are essentially the
same as in that genus.
Anal tergite exposed in dorsal view but its margin con-
cealed, or nearly concealed, in dorsal view by the two large,
contiguous, caudally projecting and distally rounded tubercles
which it bears. (Cf. pl. XXVII, fig. 7.) Anal scale triangular.
The telopodites of the gonopods of the male are compara-
tively large and conspicuous. They cross each other at the
middle line, each passing in front of and curving back on the
outside of the enlarged basal joint of the opposite gonopod.
(GE. pl. XXVII, fig. 9.)
Length, about 6 mm.
Locality.—British Guiana, Kartabo, Bartica District. One
male taken, Dec. 18, 1919.
>
eae eat
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[EW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
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VOLUME III. NUMBERS’ 92 AND 23
(Tropical Research Station Coniibaion ued 4nd 143)
A TERMITOPHILOUS BRACONID
TWO MYRMECOPHILOUS PHORIDAS
FROM BRITISH GUIANA
By CHARLES T. BRUES
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LOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
FROM THE TROPICAL RESEARCH
STATION IN BRITISH GUIANA
VOLUME III. NUMBERS 22 AND 23
(Tropical Research Station Contribution Numbers 142 and 143)
22. A TERMITOPHILOUS BRACONID
23. TWO MYRMECOPHILOUS PHORIDAt
FROM BRITISH GUIANA
By CHARLES T. BRUES
meet SHE .D —-B.Y... THE S.0:¢C Le Ts
THe ZOOLOGICAL .PARK, NEW YORK
OcTOBER 1, 1923
BRITISH
SCALE 58742'W.
5 MILES
LOCATION OF THE TROPICAL RESEARCH STATION OF THE
NEW YORK ZOOLOGICAL SOCIETY
The circle represents a radius of six miles.
First form on press October 11, 1923
Volume III, Number 22
TERMITOBRACON,
A TERMITOPHILOUS BRACONID
FROM BRITISH GUIANA'™*
By CHARLES T. BRUES.
(Figs. 42-43)
In the autumn of 1920, I received from Mr. Alfred
Emerson a most extraordinary termitophilous Hymenopteron
of the parasitic family Braconide. During the course of his
extensive studies on termites and the other insects which
occur with them, carried on in the vicinity of the tropical
station of the New York Zoological Society, at Kartabo,
British Guiana, Mr. Emerson obtained specimens of both sexes
of this strange insect. From observations on its behavior and
the reactions of the termites he assures me that it is un-
doubtedly a true termitophile and his conviction is fully borne
out by its structural characteristics.
The males have the eyes and wings greatly reduced in size
and the neuration of the latter are consequently highly modi-
fied. From observation Mr. Emerson found that they made no
attempts to fly, but moved slowly about the nest among the
termites which displayed no hostility, but appeared rather to
be friendly toward them. A worker termite was once observed
by Mr. Emerson to lick one of the females. The females, in
contrast to the males, have the wings fully developed and are
undoubtedly able to fly readily. This disparity is in itself very
unusual, as there are scarcely any Hymenoptera, or other
insects for that matter, in which the females are fully winged
and the males apterous or partially so. A similar condition
does exist, however, in the fig insects of the family Agaonide
where the females are winged and the males apterous and a
group Idarnine of a related family of Chalcid-flies, the Calli-
1 Contribution from the entomological laboratory of the Bussey Institution,
Harvard University, No. 184.
* Tropical Research Station, Contribution Number 142.
428 Zoologica: N. Y. Zoological Society [TIls 22
momide, parasitic on the fig insects show a similar condition.
Another family of Chalcid flies, the Eulophide, contains
Melittobia, a widely distributed genus parasitic on various
wasps and bees which is subapterous in the male although the
female shows no reduction in wing size, while another Eulo-
phid, Perissopterus, parasitic on scale insects, sometimes has
subapterous males. In a few other Hymenoptera the male is
dimorphic or polymorphic in the development of the wings.
Thus the males of a certain species of Pezomachus (P. flavo-
cinctus Ashm.) belonging to the Ichneumonide have three
types of males, a fully winged one, a subapterous one and an
entirely wingless one, but in this case the female is apterous.
Such is the case also in the Bethylid, Cephalonomia wrichi,
which I have recently shown to have both winged and wingless
males. On the other hand the Trichogrammatid Odphthora has
winged females and both alate and apterous males Still more
recently Picard has shown that a Braconid (Sycosoter lavagnet)
parasitic on Hypoborus ficus has winged and apterous forms,
both sexes being represented by individuals of each type. He
has shown further, in this case, that while the four forms
occur at the same time, the winged ones are most abundant
in warm weather and the wingless ones most numerous in the
spring and autumn, while only apterous ones occur during the
season of hibernation. Whether the present Braconid may also
be dimorphic cannot be stated, but on account of the rarity of
such an occurrence, there seems to be no valid reason for
assuming that it is.
The male is also much lighter in color than the female,
and such is the case also in Melittobia, in at least some of the
species.
Most termitophilous insects are physogastric, having the
abdomen considerably swollen or enlarged and frequently
turned either upwards or downwards, out of the plane in
which it normally rests. Termitobracon shows no distinct
physogastry in either sex, but the abdomen of the male is
perhaps somewhat larger than usual in other male Braconide.
It is, however, distinctly curved downwards, and when the body
is thus partially curled, the aborted wings rest upon its dorsal
1923] Brues: A Termitophilous Braconid 429
surface with their surface bent in conformity to the latter.
This bending takes place almost entirely at two points, the
base and apex of the stigma.
Termitobracon appears to be the first Hymenopteron ever
found as a termite guest’, and is possibly parasitic upon the
termites themselves although it is, of course, quite possible
that it may attack some other insect which occurs regularly
in their nests. I have examined several thousand termites
taken from the nest in which the parasites occured, but have
been unable to find any parasitic larvae either in the bodies
of the termites, attached to them, or free in the alcohol, so that
the host of Termitobracon must remain doubtful. Its size is,
however, just about that of the larger termite workers, as
might very likely be the case if it should be an internal
parasite.
Termitobracon gen. nov.
Female.—Body, including the legs and wings, densely
clothed with very fine yellowish hairs. Head strongly trans-
verse; eyes small, hairy; antennae 14-jointed, filiform, the scape
short, simple at tip and very closely united with the pedicel;
flagellum beyond the third joint marked by fine longitudinal
ridges, the first three joints strongly emarginate at tip; ocelli
minute, in a small triangle; clypeus semicircular, not emar-
ginate, not horned nor toothed; mandibles small, acute, without
teeth near apex. No parapsidal furrows; propodeum simple,
convex. Abdomen short, oval, with seven well developed seg-
ments, first segment with lateral carinae, but otherwise not
sculptured; ovipositor issuing near the middle of abdomen, but
not extending far beyond the apex, sheaths slender, but dilated
near the apex. Legs rather stout, the basal and apical joints
of tarsi elongated, the others very short. Wings rather large,
2 Since this was written Cushman (Proce. Entom. Soe. Washington, vol. 25,
p. 54, 1923), has described a genus of Braconide, Ypsisterecerus, represented
by two species collected in termite nests by Dr. W. M. Mann in Bolivia, Ypsis-
trocerus and Termitobracon are quite closely related, but differ in a number
of good structural characters in spite of the fact that both occur in the nests
of the same species of termite. Cushman has made Ypsistrocerus the type of
a new subfamily (Ypsistrocerinae) to which Termitobracon must now be added.
430 Zoologica: N. Y. Zoological Society [iis 22
stigma broad, dark, but not heavily chitinized, the radial cell
broad, attaining the wing tip; three cubital cells although the
transverse cubiti are in great part hyaline; nervulus strongly
postfurcal; nervellus interstitial; hind wing without nervulus
or marginal vein.
FIG. 42. TERMITOBRACON EMERSONI SP. NOV.
Female.
Male.—Subapterous, the wings greatly reduced in size,
curved over the abdomen at rest and distinctly bent at each end
of the stigma, reaching just beyond middle of abdomen. Eyes
minute, ocelli wanting. Legs stouter than those of the female.
Color of body much lighter.
Type, the following species:
Termitobracon emersoni ¢p. nov.
(Figs. 42-43)
Female (Fig. 42).—Length 2.2-2.8 mm. Fuscous, the head
black, except about the mouth; thorax distinctly darker than the
abdomen, especially in front above; legs brownish yellow, the
tibiae and tarsi lighter than the femora. Antennae yellowish,
1923] Brues: A Termitophilous Braconid 431
the first three joints of flagellum much darkened and the last
seven joints very pale; clypeus and mandibles, except their
black tips, honey-yellow; propleura on anterior edge and spot
on mesopleura below, yellow; propodeum anteriorly and at the
sides stained with yellow; abdomen darker at the sides of the
first segment and along the posterior margins of the second
and fourth segments; ovipositor black, its sheaths pale; wings
brownish-hyaline, a weak cloud in the upper part of the radial
cell; venation dark fuscous. Head two and one half times as
broad as thick antero-posteriorly, rounded and narrowed behind
the eyes, which are broadly oval, quite ‘small, as long as the
FIG. 43. TERMITOBRACON EMERSONI SP. NOV.
Male.
scape of the antenna; antennae 14-jointed; scape short, the
pedicel closely attached to it, rounded; first joint of flagellum
short, second to fourth longer, about twice as long as broad;
following quite distinctly oval the last pointed at tip. Malar
space twice as long as the eye; surface of head smooth; ocelli
forming an equilateral triangle. Mesonotum and _ scutellum
smooth and shining, not very strongly convex. Propodeum
smooth above, without sculpture. First abdominal segment
carinate at the sides, the space between the carinae twice as
broad as long; surface smooth and polished as is the remainder
of the abdomen; second and third segments of equal length,
432 Zoologica: N. Y. Zoological Society (IIT a
together as long as broad, the articulation between them very
faint; fourth and fifth segments equal in length and width,
each slightly shorter and broader than the third; sixth much
longer and narrower; seventh half the length of the sixth;
eighth minute; ovipositor but slightly exserted. First section
of radius very short; one-fourth as long as the second which
is slightly shorter than the third; radial cell attaining the
wing tip; cubitus arising at the middle of the basal nervure,
the transverse cubiti not complete, the first pigmented, except
below, the second indicated by the absence of trichiation;
nervulus entering near the middle of the discoidal cell;
nervellus interstitial; hind wing with only the basal and sub-
median vein and a stump of the anal, nervulus wanting; sub-
discoidal indicated as a faint cloud extending to the wing
margin. |
Male (Fig. 48).—Length 1.9 mm. Almost entirely light
testaceous, the head blackened above, the carinae on the first
abdominal segment black and the hind margins of the third to
fifth segments infuscated. Eyes much smaller, scarcely as long
as the diameter of the pedicel of the antennae.
Four females and three males from Kartabo, British
Guiana, collected by Mr. Alfred Emerson in a nest of Nasuti-
termes (N.) ephratae (Holmgren), July 28, 1920.
Mr. Cushman has kindly compared a specimen of Termito-
bracon with the unique types of his two species of Ypsistro-
cerus and considers the two genera to be undoubtedly closely
related in spite of many obvious differences, some of which
may be tabulated as follows:
Maxillary palpi 2-jointed; labial palpi apparently wanting;
joints of the filiform antennal flagellum all of similar
form; first and second cubital cells fused (female) ;
Stigma narrow (female) __......... Ypsistrocerus Cushman.
Maxillary palpi 3-jointed; labial palpi 2-jointed; basal
three joints of antennal flagellum each with an oblique
emargination at apex, following joints oval; first and
second cubital cells separated (female); Stigma broad
(female). Seee....22 2 pee Termitobracon Brues.
“4
Volume III, Number 23
TWO MYRMECOPHILOUS PHORIDA®
FROM BRITISH GUIANA’*
By CHARLES T. BRUES
(Fig. 44).
In September, 1920, Mr. William Beebe, director of the
Tropical Research Station of the New York Zoological Society
at Kartabo, British Guiana, in company with Mr. Alfred
Emerson, obtained two remarkable species of Phoride along
the trails of the legionary ant, Eciton burchelli. This ant is
abundant in the region of the Station and like the other species
of the genus undoubtedly harbors many myrmecophilous insects
of various kinds.
On examining the specimens, which they kindly sent me
for study, I find that one form represents a new genus, quite
different from any of those heretofore described and that the
second is identical with a species first made known only a few
years ago from Southern Brazil, where it was found with
another species of Eciton having somewhat similar habits.
As is the case with many of the myrmecophilous Phoride,
only the wingless or subapterous female of these two species
has so far been obtained.
Apterophora gen. nov.
Female.—Wingless, but with the eyes large, half as high as
the side of the head; ocelli present; antenne small, round; palpi
simple, with stout bristles at apex; proboscis slender, four
times as long as the head-height, geniculate at the middle, with
the apical half directed forward. Three transverse series of
frontal sete, the lowest two proclinate, close together; a pair
just above these erect or slightly proclinate; upper row of four,
two of which are next to the ocelli. Head, seen from above,
1 Contribution from the entomological laboratory of the Bussey Institution,
Harvard University, No. 185.
* Tropical Research Station, Contribution Number 143.
436 Zoologica: N. Y. Zoological Society [Ill ; 2a
much produced medially in front. Dorsum of thorax somewhat
wider than long, no scutellum; a large humeral bristle on each
side and a posterior row of stout bristles; pleura oblique, fully
twice as high as the length of the dorsum. Abdomen with five
very heavily chitinized black dorsal plates which are only
slightly separated by pale membrane in engorged specimens;
ventral surface membraneous, nearly white, without any chitin-
ous plates. Legs rather slender, the anterior coxe as long as
the femora; all tibiz without preapical spines or bristles.
Type Apterophora caliginosa sp. nov.
This is similar to Enderlein’s genus Crepidopachys (Ender-
lein 712) from Southern Brazil on account of its long proboscis,
but the type of this is a winged insect and it is difficult to
make further comparisons. The sex is not given by Enderlein,
and if his description should apply to a male, the genus might
be related to the present one. I suspect that his examples
were females, however, from the description of the apex of the
abdomen and particularly the long proboscis, in spite of the
fact that the greatly thickened costa suggests that they might
be males. Even if the latter should be the case, I do not believe
that the two could possibly be congeneric or even closely related,
as the long proboscis is the only striking similarity.
Among the genera known to have wingless or subapterous
females, two have a similarly lengthened proboscis. Psyllomyia
Loew (Loew ’57, Wassman ’00; Brues 701; Schmitz 714) a guest
in the nests of Dorylus helvolus in South Africa has a long,
slender, geniculate proboscis which is, however, not much longer
than the head. It has also a dark, heavily chitinized abdomen
like Apterophora, but the wings are present as large broad
pads. The eyes are much smaller, the ocelli absent, and the ~
legs very stout in Psyllomyia. In the absence of males, there-
fore, it seems unwise to regard them as possibly congeneric.
Rhynchomicropteron Annandale (Annandale 712; Schmitz
"14 and 715) known by two species, one from Ceylon as a guest
of Lopopelta ocellifera Rog. and another from Bombay as a
guest of Prenolepis longicornis Latr., is very similar in some
respects to Apterophora; it has a very long, slender, geniculate
1923] Brues: Two Myrmecophilous Phoridae 437
proboscis and a similarly formed head and thorax. It differs
greatly in having well developed digitiform wing pads, in
having the ocelli absent, and is practically blind, as the com-
pound eyes are mere vestiges, each composed of half a dozen
separate ommatidia. The most striking differences are seen
in the abdomen which is entirely membranous, without any
clearly chitinized plates, and in the dorsum of the mesothorax
which bears a longitudinal impression and distinct median
suture, something of very rare occurrence in insects.
Apterophora caliginosa sp. nov.
(Fig. 44).
Female.—Length 1.7-1.9 mm. Head, thorax, abdominal plates,
and four posterior coxe deep, shining black; legs and proboscis
honey-yellow; antenne pale yellow; palpi fuscous; membranous
parts of abdomen white, with a slightly sooty tinge. Head
distinctly wider and longer than the thorax, the front obtusely
triangularly produced between the antenne, frontal bristles
well developed, but not very strong. Eyes oval, contiguous,
with the antennal excavation and the posterior margin of the
head; cheek one-third the height of the eye, each with a tuft
of four or five small bristles anteriorly above the insertion of
the palpus, but without bristles behind; postocular bristles
weak. Antenne round, small, with apical, strongly pubescent
arista which is one-third longer than the head-height. Proboscis
stout at the base, but narrowed and very slender beyond;
geniculate just before the middle, the basal part straight, at
rest bent somewhat beneath the body and extending to the tip
of the front coxe; apical part curved, projecting forward with
the upper margin convex; tip obliquely truncate, with a few
minute bristles. Palpi with a few moderately large bristles
below near apex. Surface of head impunctate. Mesonotum
one-fourth wider than long; anterior margin arcuately ex-
eavated, the humeri rounded; spiracles visible from above, just
behind the humeri; posterior margin slightly convex. Macro-
chaetze not strong, disposed as follows: a weak post-humeral
one, a series of six longer ones along the posterior margin,
one at each extreme lateral angle and four between these, the
438 Zoologica: N. Y. Zoological Society [III; 28
median two farther apart than the others. Surface indistinctly
punctate. Abdomen highly convex above, the plates densely
and finely punctate; first (visible) one the largest, nearly three
times as long and wide as the mesonotum, almost twice as
broad behind as in front, the posterior margin nearly straight
and fringed with long, bristle-like hairs; second plate only half
FIG. 44. APTEROPHORA CALIGINOSA SP. NOV.
as long as the preceding, but of equal width, similarly punctate
and fringed along the posterior margin; third distinctly shorter
and narrower, fringed; fourth (really the fifth) segment
smaller, the gland opening filling a large anterior emargination
of the plate; fifth very small, not fringed like the others; apex
of abdomen of the usual tubular, retractile form. All of the
abdominal plates clothed with fine, pale, glistening pubescence.
Legs slender, although the anterior tibie are slightly thickened;
spurs of four posterior tibiz small, but distinct; hind metatarsi
each with seven transverse rows of dense fine recumbent bristles.
Described from two specimens, both the type and paratype,
as well as several other specimens which I have not examined,
taken at the same time near a trail of the army ant, Eciton
burchelli, at Kartabo, British Guiana. Concerning their rela-
tionship to the ants, Mr. Emerson writes that the first specimen
1923] Brues: Two Myrmecophilous Phoridae 439
was seen by Mr. Beebe in the ant trail and that further careful
search was rewarded by the finding of several others.
ECITOPHORA COMES SCHMITZ.
Zool. Jahrb. Abth. f. Syst., vol. 36, p. 524 (1914).
Brues, Psyche, vol 30, p. 21 (19238).
Three females from Kartabo, British Guiana taken at the
same time that the previous Phorid was obtained, prove to
belong to this species.
The types were found with Eciton predator Sm. at Sao
Leopoldo, Rio Grande do Sul in southern Brazil, but the Guiana
examples agree in all details with Schmitz’s description and
excellent figures.
It is evident, therefore, that this myrmecophile is widely
distributed in tropical South America and that it occurs with
at least two species of Eciton, FE. burchelli and E. predator.
Ecitophora is much like Ecitomyia Brues, with which
Schmitz has compared it, and differs in only a few details. In
view of the numerous monotypic genera in this group and as
Schmitz has already erected the genus Ecitophora for this
species I have used the name although I am by no means
satisfied that the two genera can be maintained. Nevertheless
Ecitophora is readily separable by the presence of ocelli and
the complete absence of the plate on the third abdominal seg-
ment, although the minute fourth and fifth plates are fully
chitinized and fully colored.
BIBLIOGRAPHY
ANNANDALE, N.
1912. A Micropterous Fly of the Family Phoride Associated with Ants.
Spolia Zeylandica, vol. 8, pp. 85-89, pl. 1
BruEs, C. T.
1901. Two New Myrmecophilous Genera of Aberrant Phoride from Texas.
American Natural, vol. 35, pp. 337-356, 11 figs.
ENDERLEIN, G.
1912. Die Phoriden Siid-Brasiliens. Stettiner Entom. Zeitg., pp. 16-45, 2 figs.
440 Zoologica: N. Y. Zoological Society [IIl; 23
Lorw, H. ¥
1857. Psyllomyia testacea, eine neue Gattung der Phoriden. Wiener Entom.
Monatsscher., vol. 1, pp. 54-56, 1 pl., 22-25 figs.
Scumitz, H.
1914. Die myrmecophilen Phoriden der Wasmann’schen Sammlung. Zool.,
Jahrb., Abth. f. Syst., vol. 37, pp. 509-566, 11 figs., 2 pls.
1915. Berichtigung und Erganzungen zu den ‘‘Myrmecophilen Phoriden
der Wassmannschen Sammlung.’’ Schmitz 1914. Deutsch. Entom.
Zeitschr., vol. 5, pp. 503-504.
WASMANN, E.
1900. Neue Dorylinengiste aus dem neotropischen und dem Athiopischen
Faunengebiet. Zool. Jahrb., Abth. f. Syst., vol. 14, pp. 215-289,
2 pls.
ee eee eee
Zoologica: Vol. III. Nos. 1-23.
Index
Acamatus, 29
Acrocinus longimanus (Linné), 29
Acromyrmex, 46
Acropteryx opulenta, 31
Agaonidae, 427
Agelaius imthurni, 9
Agouti, short-tailed, 286
Agouti paca paca Linn., 9, 273
Aix sponsa Linné
tail down of, 32
Akawai Indian Benab, 16
Akawai Indians
plants raised by, 29
Aleocharinae, 323-364
_ new species, 323-364
Alligators, 16, 20
of Georgetown, 16, 20
Allomerus
octoarticulatus Mayr, 162
on Tachigalia tree, 47
Allotinus horsfieldi, 105
Alouatta
seniculus L., 244
(aes macconnelli Elliot, 9, 10, 242,
two fetuses of, 242-262
figures 19-24
twinning, 244
Amazon, the
explorers of, 3, 4
Amazona imperialis, 9, 25
‘Amazona ochrocephala (Gmel.)
eggs and young, 31
Ameiva, genus, 27
Amphibians
check list of 72 amphibians, 26
life histories, 8
living, sent to N. Y. Zoo. Park, 10
new, ye tropical research station, 287-
tadpoles at Kartabo, 28
Amphisbena alba, 10
Amphisbena fuligenosa, 10
Ani, smooth-billed, 21
development of, 21
Annandale, 436 ,
Annual reports trop. research station
for 1917, 24
for 1918, 26
for 1919, 27
Anoplops refigula, 20
Anoplotermes tenebrosus, 384, 388
Antcatcher, rufus-fronted, 20
Anteater, silky, 24
Anthony, H. E.
mammals collected by Beebe at the
tropical research station, 265-286
acknowledgements, 267
appendix A, 284-285
(figures 25 and 26), 271, 273
list of specimens, 267-286
maps, 266
new species, 267
number of specimens, 265
Ants
army, 24, 25, 27, 29
3 new species, 29
attendants of Homoptera, 47
defoliators of Tachigalia tree, 46
forty-two forms (2 new species), 25
gardens of, 29 :
inquilines of Tachigalia, 46-48, 87
“long John,’ 137
new, 25, 32
obligatory, on Tachigalia, 48-50
of Tachigalia, 39, 46, 135-168.
of genus Pseudomyrma, 36
Aotus boliviensis Elliot, 261
Apes, 252
Aphididae,
honey tubes, 61
Aphidoletes, 85
Aphiochaeta scalaris, 80, 103
Appun, 18
Apterophora gen. nov.
description, 435-437
other genera with wingless or
apterous females, 436
caliginosa sp. nov.
description (fig. 44), 437-438
Apterosltigma, 102
Apterygota, termitophilous,
from British Guiana, 383-402
bibliography, 402
description of new species, 385-402
(Plates XVIII-XXIII, 386, 389,
391, 392, 395, 398
source of material, 383
Aristotle, 117
Armadillo, 269, 270
giant, 286
Armitermes
percutiens sp. nov., 392
teevani sp. nov., 390
Army ant, 24, 25, 27
Arthrocnodaz, 85
Arthropods J
misc. of Tachigalia tree, 46
Artibeus
planirostris fallax, 284
planirostris planirostris (Spix), 284
Alteles sp., 286
Ateleus, 261
Atelura
habits of, 383-385
crispula sp. nov. (Pl. XX), 390-392
cucullata sp. nov. (Pl. X XI), 392-
394 :
formicaria Silv., 384
engin sp. nov. (Pl. XVIII), 385-
3
lepismoidea sp. nov. (PY: <RI;
388-390
synoeketa Silv., 384-387
termitobia Silv., 284
Alta cephalotes Fab., 29, 30, 46, 92, 162
biocoenose, 92
defoliators of Tachigalia tree, 46
fungus gardens, 30
habits, nests, etc., 29, 30
Attas, 29, 30
fungus gardens, 30
habits, nests, ete., 29, 30
Altaphila, 30, 93
Attids
in Tachigalia tree, 46
Aublet, 137
Augochlora,
callichlorura, 26
maromana, 26
Auk, 19
Azteca
of Tachigalia, 39, 48, 137
brevicornis Mayr, 138
sub-
[443]
444 Zoologica
foveiceps sp. nov. (fig. 16) 163-166,
on Tachigalia tree, 48, 49, 50
tachigalia, 137
traili Emery
on Tachigalia tree, 48, 50, 163
Bailey, Jow/.,.55.00; 100, 104
Barber, H. S., 35, 64, 68
new species of Coleoptera, 187-194
Barbour, T., 27
Bartica District (of British Guiana), 4, 18
bird life of, 18
birds, list of 350 species, 18
historical, 18
naturalists of, 18
Bates, H. W., 3, 4
Batrachians
new, from tropical research station,
287-299
vampire, 286
Bay of Butterflies, 29
Beach, a jungie
ecology of, 29
Beebe, William, 4, 5, 7, 8, 15, 16, 17, 18, 19,
205 (24,5255 265 27), 28; 20 a0; Sieve:
243, 289, 301
Contributions of tropical
station, 15-32
mammals seen at station, 286
Bees,
Bartica, of, 22, 26
Beetles, 16
ambrosia, 98, 99
fungus of social beetle, 36
larvae and pupae of social beetles, 195-
AEN b
long armed, 29
rhinoceros, grubs, 16
social, bibliography, 119-126
social, of Tachigalia tree, 35-136
Belt, Thomas, 3, 4, 104
Bequaert, J., 36, 46
Berlese, 61
Bingham, 105
Biocoenose, 91
climax in Tachigalia tree, 49-50
concept of, 91
cotingas, of, 28
insects of Tachigalia, 35
Tachigalia tree, of, 41-50, 93-94
parrakeets, 28
toucans, of, 28
weed, of a, 28
Birds
Akawai Indian and colonial names, 19
Bartica, check lists, 18, 27
behavior, 116
Brazil, of Para, 32
development of white ibis, 32
embryos, collection of, 9
eye, 11
herons, habits, 28
jungle, in second growth, 18
life histories, 8
living, sent to N. Y. Zoo. Park, 9
nests from South America, 17
optical fundi, 9
skeletons, collections of, 9
skins, collections of, 9
syrinx, 9
tail feathers of motmots, 31
trogons, 9
voice 9
Blapticoxenus gen. nov.
description, 362
brunneus sp. nov., 363-364
Blatchley, 176
Blepharidatta brasiliensis Wheeler, 32
Blepyrus tachigaliae Brues, 85, 103
research
[Vol. III
description, 229-230
Blisson, 173
Bliintschli, 244
Bodine, J. H., 72-73
Bodkin, G. E., 309, 315
Borecus gen. nov.
description of genus, 397-399
pinnatus sp. nov. (Pl. X XIII), 399-402
Bothrideridae, 199
Boving, Adam, 35, 64, 88, 96, 110
larvae and pupae of social beetles, 195-
221
bibliographic notes, 213
Coccidotrophous, 203-213
Cucujidae, 198-202
Eunausibius, 203-213
(Plates VII-X), 214-221
Silvanidaec, 201-203
Brachymyrmex
on Tachigalia tree, 47
heeri Forel, 166
var. basalis var. nov., 166
Braconidae
dimorphism,
forms, 427-429
Termitobracon gen. nov., 429-430
emersoni sp. nov., (figs. 42-43),
430-432
Ypsistrocerus Cushman, 429, 432
Bradford, H. B., 323
Bradypus
cuculliger Wagler, 268
tridactylus, 9
British Guiana, 17
amphibians, reptiles and mammals, 26
ant gardens of, 29
army ants, 29
our search for a wilderness, 32
tropical wild life, 17, 26
Broek, 261
Brontes, 90, 198
planatus L., 178
Brues, C. T., 36
Blepyrus tachigaliae sp. nov., 229-230
Brues, C. T. ” .
Termitobracon, a termitophilous bra-
conid from British Guiana, 427-432
compared with Ypsistrocerus, 432
description of genus, 429-430
description of emersoni (figs. 42-
43) 480-432
dimorphism, alate and apterous
forms, 427-428
source of material, 427
Brues, C. T. :
two myrmecophilous Phoridae from
British Guiana, 435-440
bibliography; 439-440
source of material, 435-439 ‘ee
Apterophora gen. nov., description,
435-437 z
other genera with wingless or sub-
apterous females, 436 Pie as
caliginosa sp. nov., description
(fig. 44), 437-438
Ecitophora comes Schmitz, 439
winged and apterous
Bufo
marinus, 10 .
typhonius, 10
Bufonidae, 291
Bushbird, cinerecus, 20
Bushmaster, 15
method of capturing, 15
Butorides striata
development of wing, 21
Butterflies,
Gerydine Lycaenid, 105
Heliconid, 25
Ithomid, 25
Caccicus persicus, Linné
<—— oe
Nos. 1-23]
_ nests and eggs, 31
Caicus cela
_ development of wing, 21
Caiman sclerops, 10, 16, 20
of Georgetown, 16, 20
Callandra oryzae, 97
Callimomidae, 428
pay onous
eebei, 25
parabiosis of, 29
femoratus Fabr., 167
___ attendant on membracids, 47, 93
pittieri Forel
poenalis var. noy., 167-168
zoc Forel, 167
on Tachigalia tree, 47, 48
Cannon, 117
Cantharidin, 379
Capnodes albicosta, 31
Caprimulgus nigrescens, 20
embryology, 21
Capritermes
bodkini sp. novy., 312-313 (Plate XIIT)
bodkini Silv. modestior var. noy. (Plate
XIV), 315-318
Carl, 397
Garter, T. D., 5
Casey, 179, 180, 182, 183
Cassique
yellow-backed, 31
Catarrhines, 252
Caterpillar
Lycaenid, 68
on Tachigalia tree, 50
Catfish, giant singing,
habits, 29
Cathartus, 59, 67, 89, 96, 104, 198, 202
advena, 96, 97, 175
cassiae, 176
excisus Reitter, 176
longulus Blatchley, 176
quadricollis Guénin, 176
Catogenus rufus Fabr., 183, 199
Catopsilia |
migration and habits of 5 species, 29
Caudell, A. N., 31
new species of Orthoptera, 31
Cebus, 261
_ apella apella (Linnaeus), 283
apella brunneus, 283
apiculatus, 283
Cecidomyia, 85
caryaecola O. Sxcken, 50
Centipede
of Tachigalia tree, 46
Cephalonomia urichi, 428
Cercropia, 35
Cercropias
dead queens of Azteca in, 48
Cerapachyinae, 106
Ceratorenus gen. nov.
description, 360
tricornis sp. nov. (fig. 39), 360-362
Cercocebeis cynomolgus, 244
twins, 244
Cercopithecus pygerythrus, 244
twins, 244
Cercosaura ocellata Wagler, 301
Cerdocyon thous,9
Chaemepelia talpacoti, 20
Chalcid-flies, 427-428
Chalcididae, 85 .
Chamberlin, Ralph V., 35, 46,
four termitophilous _millipeds
British Guiana, 411-421
description of new genera and
species, 411-421
(Plates XXV, XXVI, XXVII), 412,
416, 418 J
source of material, 411
from
Index
445
Champion, 174
Check-lists
amphibians, 26
birds, 18, 27
birds of Para, 32
mammals, 26
reptiles, 26
Cheliomyrmex megalonyx, Wheeler, 29
Chicks, Trumpter, 16
Chironectes sp., 286
Chittenden, 373, 174, 175, 180, 182
Chlorion neotropicus, 23
Choloepus didactylus, 9, 269
Cicada septemdecim, 85
Claphe laudissima, 31
Calviger, 368
Clavigeridae, 67
Cleroidea, 199
Coccids
cultivated by ants, 35
cultivated by beetles, 35, 90
insects attendants on, 104-106
of Tachigalia tree, 45, 49, 50, 55, 90
ostioles, 61, 62, 67
Coccidomyia, 85
Coccidotrophus gen. nov., 97, 203
description, 189-190
socialis sp. nov.
behavior, 111-118
cannibalism, 72
cocoon (fig. 6), (Pl. V), 73,
74-76, 109-111
copulation, 77-78
description (Pl. III, fig. 2),
(Pl. VI, figs. 1-5), 51-52, 190-
191
distribution, 51
eggs (fig. 1), 52, 56
enemies, 78-88
establishment of colony, 52-57
food, 55, 59, 63, 69-73
fungi, 81
larva (Pl. VII-VIII), 52, 57,
64, 67, 73, 96
description, 203-211
“milking’’ Pseudococcus, 63,
69-72
moults, 73
pupa (Pl. IX), 52, 57,
description, 211-212
synoeketes, 80
Tachigalia tree, on, 51-88
Taxonomic position, 198
traits, 97
tropisms, 112
Coccinellids, 35
of Tachigalia, 35, 83
Cockerell, T. A., 26
Cock-of-the-rock, 15
habits, 15
Cockroach, parasitic, 30
Coendu sp., 286
Coleoptera
catalogue of, 213
Histeridae, 364-366 : e
larvae and pupae of social beetles, 195-
221
new genera and species, 187-194
of termitophilous coleoptera, 323-
366
social life among, 97-103
Staphylinidae, 323-364
Corotocint, 323-338
Oxypodini, 338-362
Gyrophaenae, 362-364
Collembola
new genus and_ species from British
Guiana, 383, 397-402
synoeketes of social beetles, 80
achigalia tree, on, 36
446
Colpocephalum armiferum Kellogg, 31 .
Comstock, 61
Constrictotermes
cavifrons (Holmgren), 325, 328, 367
Convict Trail, 25
Cooper, Isabel, 5, 8
Coquerel, 89, 173
Cordia, 35
Cornitermes pugnax Emerson, 348, 394
Corotoca
guyanaeé sp. nov., 327-329
melantho Schiddte, 329
phyllo Schiddte, 329
Corotocini
new species, 323-338
Corymbogaster gen. nov.
description, 346-347 ;
miranda sp. nov. (fig. 35), 347-349
Corythalinae, 97
Crab-dog, 286
Craig, 116
Craighead, 96, 197, 199
Crake
Cayénne, 20
white-necked, 20
Crandall, L. S., 15, 32
Creciscus viridis, 20
Creeper, blue honey, 20
Crematogaster
on Tachigalia tree, 47, 48
delitescens sp. nov., 152-154
¢« limata F. Smith
subsp. ludio Forel, 151
var. palans Forel, 151
parabiolica
attending membracids, 93, 152
Tachigalia tree, on, 47, 93
ornalipilis, 25
parabiosis of, 29
Crill, 117
Crotophaga ani, 21
development of, 21
Cryptocercus, 48
Cryptodesmidae
four new genera and species from British
Guiana, 411-421
Crypturus
soui soui, 20
variegatus variegatus, 20
Cucujidae, 94-97, 200
families and subfamilies, 200
food, 104
habits, 171-183, 176-179
list of descriptions and figures of larvae,
oui LS}
pupation, 109, 110
taxonomy, 198-203
Cucujids
pronotum, 90
Cucujinae, 198, 200
Cucujus, 90, 96, 198
pupation, 110, 177
cinnaberinus Scop., 177
clavipes Fabr., 176
haematodes, Erich., 177
Curculionids
larvae of. in Tachigalia, 42
Cushman, 429, 432
Cuyuni River, 4
Cyanerpes cyaneus, 20
Cyclopes didactylus, 9, 24, 269
Cynedesmus, 417, 420
Cynipid gall-flies, 104
Cyphodeirus albinos, 80
Cyphoderus, 399
Cyphomyrmex, 102
Dactylopius, 61
Dahlgren, U., 10
Darwin, Charles, 3, 4
Dasyprocta aguti flavescens (Thomas), 272
Zoologica
[Vol. III
Dasypus novemcinctus Linn., 269
Dauber, one-banded, 23
Defoliators
ants of Tachigalia tree, 46
Dendrodates sp., 10
Dendroides canadensis, 96
Dendrophagus crenatus, 177, 198, 213
food, 104, 177
Dentifibula, 85
Deroptyus accipitrinus, 9
Desmodus sp., 286
Diadiplosis, 85, 103
cocci, 225
pseudococci, sp. nov., 225-226
f in Tachigalia tree, 84
Dichocrocopsis maculiferalis, 31
Dicharodiplosis, 85
sen A marsupialis marsupialis, Linnaeus,
Dimmock, 91, 177, 183
Diptera
parasites of coccids, 36
Doflein, 111 e
Dolichoderinae, 162-166
attendant on Homoptera, 106
Dolichoderus attelaboides Fabr.
on Tachigalia tree, 47, 93, 162
Dolphin, fresh-water, 286
Donacia
pupation, 110
Doran, 176
Doras granulosus Valenciennes, 29
Dorylinae, 106
Dorylus helvolus, 436
Dove
Talpacoti ground, 20
red mountain, 20
Drever, 117
Dryocera, 67, 198
Ducks, :
tail down of, 32
Dutch, 18
in Bartica, 18
Diy ar els Geo
new species of Lepidoptera, 31
Dysmerus, 199
Eburniola gen. nov.
description, 333-334
leuccgaster sp. nov. (fig. 31), 334-336
Echimys longirostris Anthony (fig. 26),8273
Eciton
angustinode Emery
subsp. emersoni Wheeler, 29
burchelli, 24, 25, 27, 29, 30, 439
pilosum F. Smith, 29
var. beebei Wheeler, 29
praedator F. Smith
var. guianense Wheeler, 29
Ecitophora comes Schmitz, 439
Ectatomma tuberculatum Oliv.
on Tachigalia tree, 47, 138
Edmonson, Miss J. E., 323
Eels, electric, 10, 11
Eichhoff, 98, 99. 101
Electrophorus electricus, 10
Eleutherodactylus, 291
Emarginea empyra, 31
Emerson, A., 5, 27, 309, 318, 323, 367, 369,
acai 383, 385, 401, 403, 411, 427, 432,
Emerson, Gertrude, 5
Emerson, W. J., 5
Empidonomus varius, 20
embryology, 21
Emporius signatus Frauenf, 178
Encyrtids, 85
Endaphis, 85
Enderlein, 436
Endoastus productus, Osborn
of Tachigalia tree, 47, 233-234
Nos. 1-23]
fig. 17, 234
Entomobrya, 103
dissimilis, 80
myrmecophila, 80
wheeleri sp. nov., 80 (fig. 18), 237-238
Entomobryidae, 399
Eois costalis, 31
Epicharis maculata barticana, 26
Epilachna borealis, 379
gland cells of, 379
Eriococcinae. 61
Erismatura jamaicensis (Gmel.)
tail down of, 32
Escherich, 64, 383, 396
Euglossa,
decorata ruficauda, 26
ignita chlorosoma, 26
Eulophidae, 428
Eulophids
in gall on Tachigalia, tree, 50
Eumenes,
canaliculata, 22
sp., 22
Eumomotus sp., 31
tail feathers, 31
Eu
mops
barbatus (Allen), 281
milleri (Allen), 281
Eunausibius Grouvelle, 191, 203
wheeleri sp. nov. (Plate VI)
activities, 97
behavior, 111-118
cocoon, 109-111
description, 192
larva, 203-211
pronotum, 89
pupa (Pl. IX), 211-213
pee ieota tree, in (Pl. III), 88-91,
Eutermes, 376
holmgreni Banks, 309
microsoma Silv., 309, 384
parvellus, 307-309 (Pl. XI), 308
Explorers of Eastern South America
list of six best known, 3
Exudate organs of insects, 367-381
Falco rufigularis, 9
Fauna of four square feet of jungle, 32
Felt, E. P., 36, 84
- peeves pseudococci sp. nov., 225-226
‘is
living, sent to N. Y. Zoological Park, 10
giant singing catfish, 29
Fiske, 96, 107
Florilegus barticanus, 26
Floyd, J. F. M., 5
Flycatcher, 17
Guiana pigmy, 17
oily, 17, 20
varied, 20
Folsom, J. W., 36
A new Entomobdrya, 237-238 (fig. 18)
Folsom, J. W. A
termitophilous apterygota from British
Guiana, 383-402
bibliography, 402
descriptions of new species, 385-402
(Plates XVIII-X XIII), 386, 389,
391, 392, 395, 398
source of materials, 383
Forbes, 91
Formicidae, 36
Formicinae, 166-168
attendant on Homoptera, 106
Fouilleé, 117
Fowler, 174, 175, 177, 178, 179, 180, 182
Frédéric, 253 ’
Fresh-water dolphin, 286
Index
447
Frog, 16
giant tree, 16
Fungus
of social beetle, 36, 81 (fig. 12)
ambrosia of beetles, 98-101
Furipterus horrens (F. Cuvier), 280
Galbula ruficauda Cuv., 31
nests and eggs, 31
Galeocoptes carolinensis
development of wing, 21
pinon development, 21
Galls
Itonidid on Tachigalia tree, 50
Ganglbaur, 173, 176, 178, 179, 180, 182, 183,
190, 213
Garden, a tropic, 28
Gardens, Ant, 29
Gasatomus gen. nov., 420
description, 417
emersoni sp. nov. (Pl. XX VII), 419
Gernet, 213
Gifford, H., 5
tirault, 173
Glamyromyrmex beebei Wheeler, 32
Glands
insect, abdominal append., 367-381
Glossophaga soricina soricina (Pallas), 279
Glover, 174, 176, 181
Goddard, 117
Gonatodes
beebei sp. nov., 301-303
description of type, 301
diagnosis, 301
measurements, 302
ferrugineus, 303
Goniothoraz, 158-162
Grassiella, 384
Gravely, F. H., 213
Grosbeak,
brown breasted pigmy, 20
Grouvelle, 190, 191
Guerlinguetus
aestuans aestuans (Linn.), 277
Guiana,
explorers of, 3, 4
tropical wild life, 17
hinterland of, 23
Guiana Indians
charms and superstititions, 23
Guillebeau, 173, 174
Guinevere the mysterious, 28
Guira alba,
ontogeny of, 32
Guthrie, 117
Gyrophaenae
new species, 362-364
Hadena niphetodes, 31
Hagedorn, 97
Hamilton, 96, 177
Hamitermes
excellens sp. nov., 309-312
(Plate XII), 309
hamifer Silv., 312
Hammock Nights, 26
Handlirsch, 1908
Hapale, 261
Hartley, G. I., 5, 7, 8, 16, 17, 21, 22
Hartley, R., 5
Heins, 72 i
Hemiderma p. perspicillatum, 16, 280
Hemipectem cleptes, 31
Hemipeplus Latr., 199 ;
marginipennis Lec., 183
microphthalmnus Schwarz, 183
Herpailurus jaguarondi unicolor, 9, 279
Herrick, 116
Heteroptera ,
three new species of Termilaphis (PI.
XXIV), 403-408
Hilhouse, 18
448 Zoologica
Histeridae
new species, 364-366
Hoatzin, see ‘
Opisthocomus hoazin
Holmgren, 379
Homoptera
attendants on, 104-109
attended by larvae of ants, 107
of Tachigalia tree, 47
Hopkins, 98, 99
Hyleborus xylographus, 99-101
Horn, 95
Hoorie, British Guiana,
new species of insects from, 31
Hositea gynaecia, 31
Howard and Marlatt 174
Howes, P. G., 5, 7, 16, i ly (nee a
Hubbard, 98
habits of Platypus compositus, 98
Hudson, W. H., 3. 4
Humming Bird, 17
cayenne hermit, 17
Hunter, G. W., 5, 26
Huntress, blue, 23
Hydrochoerus hydrochoerus (Linn.), 272
Hyla ;
maxima, 16
ornatissima sp. noy., 291-293
description of type, 292
diagnosis, 291
measurements, 293
Hylaea, 39-41
Hylesia indurata, 31
Hyliotinae, 198, 200
Hyloxalus beebei sp. nov., 289-291
description of type, 290
diagnosis, 289
measurements, 291
Hymenoptera
behavior, 111 :
dimorphism, alate and apterous, 427 -
429
in gall on Tachigalia tree, 50
new genus of braconid from B. Guiana,
427-432
Hypoaspis sp., 80
Hypoborus ficus, 428
Ibis, white
ontogeny of, 32
Ichneumonidae, 428
Icterus chrysocephalus, 20
Icticyon sp., 286
Idarninae, 427
Iguana iguana, 10
Iilice biota, 31
Imperial Parrott 25
im Thurn, 18
Incarsha aporalis, 31
Indian charms, 23
Inia sp., 286
Inopeplus praeustus Chevr., 183. 199
Inquilines, :
of Tachigalia tree, 45, 47, 48, 87
Insect behavior, 7, 27, 111-118
bibliography, 119-126
Tnsects,
attendant on coccids, 104-106
bees, 22, 26
butterflies, 25, 105
collections of, 9
glandular abdominal appendages, 367-
1
glandular excretions, 367-381
new species collected in B. Guiana, 31
parasitic, 35
paralyzed by wasp poison, 23
pupation, controlled, 23
social, bibliography, 119-126
synoeketic, 35
Tachigalia tree, of, 32
[Vol. III
termites, new species, 307-321
tyrants, 25
wasps, 16, 22, 23 °
Invertebrates
of four square feet of jungle, 32
Ipidae, 97
Ischnurges bicoloralis, 31
Islands,
West Indies, 25
Itonidids, 85
gall on Tachigalia tree, 50
in Tachigalia tree, 84
Jacamar,
Venezuelan rufous-tailed, 31
Jacana jacana, 21, 28
development of, 21
habits, 28
Janet, 383
Jennings, 112
tropisms, 112
Joseph, 260
Jungle
and its life, 18
beach, ecology, 29
birds, 18
clearing, 28
dangers, 18
edge of, 7, 30
fauna of four square feet, 32
gardens, 29
life zones, 18
mammals, 18
night, 24
night life, 26
peace, 7, 26
seasons, 18
temperature, 18
Kalacoon, 17
wild life near, 20
Kaltenbach, 91, 175, 177, 178, 180
Kartabo, 4, 28
Katydid
eggs in Tachigalia tree, 46
Kellogg, V. L., 31
new species of Mallophaga, 31
Kempf, 117, 118
Kieffer, 85
Knab, 110
Kolbe, 95
Kiister, 85
Label-making in field, 25
Labidus, 29
Lachesis mutus, 10, 15
method of capturing, 15
Laemophloeidae, 96, 97, 179-183, 199, 200
food, 104
pupation, 109, 110
Laemophloeinae, 199, 201
Laemophloeus, 97, 199
abutis, 213
aliernans Erich, 179, 213
ater Oliv., 179
biguttatus Say, 179
bimaculatus Payk, 179
castaneus Erich, 179
clematidis Erich, 180
corticinus Erich, 180
denticulatus Preysl., 180
dufouri Laboulb, 180
duplicatus Waltl., 180
fasciatus Mels., 180
ferrugineus Steph., 180
hypobori Perris, 181
juniperi Grouv., 181
modestus Say, 181
monilis Fabr., 181
perrisi Grouv., 181
pusillus Schon, 182
testaceous Fabr., 182
turcicus Grouy., 182
abe
OS a es aol Mage
Nos. 1-23]
vestitus, 95
Laemotmetus
ferrugineus Gerst., 182
. rhizophagoides Walker, 182
Larval sacrifice, 22
Lasius
attendant on Homoptera, 106, 109
umbratus Nyl., 383
Lathropus, 199
sepicola Miiller, 182
vernalis Erich, 182
LeBaron, 177
Lebedinsky, 397
Leconte, 95
Leguminosae,
Tachigalia, 39-41
Leibnitz, 117
Leng, 95, 179, 183, 213
Leontocebus sp., 286
taeniata sp. nov., 303
description of type, 303
diagnosis, 303
measurements, 304
Leptodactylus
caliginosus, 295, 297
minutus sp. nov., 295-297
description of type, 296
diagnosis, 295
measurements, 297
pentadactylus, 10, 295
pulcher, 295
rhodomystax, 295
rugosus sp. nov., 297-299
description of type, 297
diagnosis, 297
measurements, 298
strictigularis sp. nov., 293-295
description of type, 293
diagnosis, 293
measurements, 294
Leptothorax
on Tachigalia tree, 47,48
echinatinoidis Forel __
subsp. aculeatinodis Emery
xe pleuriticus var. nov., 158-
16
umbratilis sp. nov., 160-162
Lestodiplosis, 85
Leucolepia musica, 20
Leucotermes
crinitus (Emerson), 405
tennis (Hagen), 406
Leuritus gen. nov.
description, 411-413
lermitophilus sp. nov. (Pl. XXV),
413-414
Lipeurus absitus Kellogg, 31
Lithocoryne, 95
Lloyd, 18
Lobodiplosis, 85
Loew, 436
Lugger, 180
Luira mitis Thomas, 277
Lycaenid caterpillars, 68
MacGillivray, 60-62, 167
McConnell, 18
MelIndoo, N. E., 372
glandular structure of the abdominal
appendages of a termite guest (Spirach-
tha), 367-381
abdominal appendages, 368-373
bibliography, 381
discussion, 376-380
interpretation of results, 373-376
Index
449
introduction, 367-368
technique, 367
Macacus, 257
tail of, 257
Macalla pallidomedia, 31
Macrophyllum macrophyllum (Wied), 285
Mallophaga, 31
new species, 31
Mammals
Akawai Indian and colonial names, 19
No list of 119 species of Br. Guiana,
6
collected by Beebe at tropical research
station, 265-286
appendix A, 284-285
collected at Georgetown, 284-285
(figures 25-26), 271, 273
list of specimens, 267-286
maps, 266
number of specimens, 265
seen but not collected, 286
collections of, 10
Guiana howling monkey, 243-262
Manakin, orange-headed, 20
Manatees, 28, 286
Mann, W. M., 403
New genera and species of termito-
philous coleoptera from Northern South
America, 323-366
Histeridae, 364-366
Staphylinidae, 323-364
Aleocharinae, 323-364
Corotocini, 323-338
Gyrophaenae, 362-364
: Oxypodini, 338-362
Margay tigrina vigens (Thomas), 278
Marmosa
chloe Thomas, 267
cinerea demerarae Thomas, 267
murina, 17
Marmoset, 286
Martin, 251
Martin, grey-breasted, 21
nesting habits, 21
Mazama
americana tumatumari Allen, 271
memorivaga (F. Cuvier), 271
Mazaruni River, 4, 17, 29
life of tidal area 29
Meal-worms, 72
Mealy-bug
of Tachigalia tree, 45
devoured by midges, 225
Megasoma actacon, 16
Melittobia, 428
Membracids,
of Tachigalia plant, 36, 47, 233-234
Endoastus productus sp. noy., 233
Microcentrus sp., 234
Merganetta columbiana Des Murs
caudal down of, 32
Mesophylla macconelli Thomas, 280
Metachirus
nudicaudatus nudicaudatus Geoff., 268
Microcentrus
on Tachigalia tree, 47, 234
Microdon, 87
Midge, new species, 225-226
Miller, 280
Miller, G. S., 267
Millipeds
four new genera and species of termito-
philous millipeds from British Guiana,
411-421
description, 411-421
(Plates XXV, XXVI, XXVII),
412, 416, 418
source of material, 411
Minacrgides arnacis, 31
Mirotermes
450
nigritus Silv., 366
tuberosus, 388
Mites
synoeketes of social beetles, 80
Mjéberg, 403
Mobius, 91
Mollison, 250
Molossus
obscurus Geoffroy, 281
rufus Geoffroy, 281
Momotus
tail feathers, 31
Mongoose, 28
Monkey, Beesa, 16
photographs, 16
Monkey
Guiana Howling
fetuses of (figs. 19-24), 243-262
measurements of fetuses, 244-245
twinning, 244
Red Howling
embryology of, 10
Spider, 286
Monodelphis
brevicaudata brevicaudata Schreb., 268
Morrison, Harold, 35, 45
three new species of Termitaphis, 403-
408
description of new species, 405-407
key to species, 408
literature, 403
(Plate XXIV), 404
source of material, 403
Mosso, 117
Motmots
tail feathers, 31
Mungos
birmanicus Thomas, 284
mungo, 28
Mus musculus musculus Linn., 274
Mycodiplosis, 85
Myoprocta sp., 286
Myriopoda
of Tachigalia plant, 36
Myrmecophaga tridactyla Linn., 269
Myrmecophytes, 137
of British Guiana, 35
Myrmelachista, 48
Myrmicinae, 147-162
attendant on Homoptera, 106
Myrmicine parabiosis, 29
Myrmicocrypta, 102
Myrmobrachys, 167
beebei, 25
Myrmothrix, 167
Narthecius, 199
Nasua phoecephala Allen, 278
Nasutitermes
acajutlae (Holmgren), 356, 385, 390, 401
baileyi (Emerson), 419
beebet (Emerson), 363
brevipilus (Emerson), 414, 417
cavifrons (Holmgren), 325, 328, 367
costalis (Holmgren), 331, 340, 342, 345
353, 390, 401
ephratae (Holmgren), 332, 337, 344, 388
390, 432
guayanae (Holmgren), 333, 336, 350
352, 358, 362, 388
intermedius (Banks), 359
nasulississimus, 388
octopilis (Banks), 338, 388, 401
surinamensis (Holmgren), 336, 388, 390
Tachigalia tree, in, 46
Naturalists
of Bartica, 18
Nausibius, 59, 67, 89, 198, 203
clavicornis Kug, 176
food, 104
Neacomys guianoe Thomas, 275
Zoologica
[Vol. IIl
Nectarina, 104-105
Nectomys squamipes melanius, 275
Neger, 98
Neophaenis aedemon, 31
Neoponera
crenata, 138
on Tachigalia tree, 47
unidentata, 138
Neusticurus bicarinatus (Linné), 301
Newman, 254
Newstead, 61
Nicaragua
explorers of, 3, 4
Niceville, 68
Nicoletia
emersoni sp. noy. (Plates XXII, XXIII),
394-396
neotropicalis Silv., 396°
Nighthawk,
dusky, 20
Noble, G. K., 27 :
New Batrachians, 287-299
acknowledgments, 289
new species, 289-299
New Lizards, 301-305
Ocelot, 287
Oecomys
guianae Thomas, 274
nitedulus Thomas, 275
rutilus Anthony, 275
Ohaus, 101, 102, 118
Oncopodura, 397
crassicornis Shoebotham, 397
hamata, 397
Oncopodurinae, 399
Oophthora, 428
Opisthocomus hoazin, 10, 15, 16, 30
distribution, 30
ecology, 30
eggs, 15
hunt for, 23
life history, 19
nest, 15
nestlings, 15, 16
parasites, 30, 31
photographs, 30
pinion development, 21
wing development, 21
Opossum
mouse, 17
water, 286
Oreopelia montana, 20
Oriole, Moriche, 20
Ornithological discoveries, 19
Orthocrema, 151-154
Orthoptera
new species collected by Beebe, 31
Oryzaephilus, 59, 89 (fig. 7), 198, 203
bicornis Erich, habits, 174
food, 104
gossypit Chitt, 174
mercator Fauvel, habits, 174
pupation, 110
surinamensis, 96
habits, 173 ‘ 5
Oryzoborus angolensis breviosiris, 20
Oryzomys, 276
¢aracolus Thomas, 277
laticeps, 276
macconnelli, 276
meridensis, 277
trinitatis, 276
velutinus, 276
Osborn, H. F., 1-11, 28, 36 i
The Tropical Research Station, 1-11
Osborn, Herbert 2
two Tachigalia Membracids, 233-234
Osgood, W. H., 267
Ostioles
of Coccids, 61, 62, 67
Nos. 1-23]
Otostigmus limbatus Meinert
in Tachigalia tree, 46
Oxypodini
new species, 338-362
Pachira aquatica, Aubl., 48
Pachysima, 377, 378
Packard, 91
Palorus subdepressus, 97
Panthera onca (Linn.), 278
Papio porcarius, 244
twins, 244
Para, Brazil
birds of, 32
invertebrates of, 32
Parabiosis, 29
myrmicine. 29
Paracraga amianta, 31
Parrot
Imperial, 25
nestling Pionus, 17
yellow-fronted Amazon, 31
Passalidae, 97, 101
Passalus cornutus, 101
stridulating organs, 102
Passandra, 95
Pecari tajacu macrocephalus Anthony, 270
Peccary, 270
Pediacus, 95, 198
depressus Herbst., 178
fuscus Erich, 178, 213
Penard, T. E., 19
Pelagic life, 25
Perai, 16, 22
Perinthus
dudleyanus Casey, 337
silvestrii, 336
tarsatus sp. nov., 336
vestitus sp. nov., 337-338
wasmanni sp. noy., 337
_ Perissopterus, 428
Berrie, BY, 91, 173, 175, 178, 179, 180, 182
8
Peyerimhoff, de, 95, 183
Pezomachus, 428
Pheidole
cramptoni, 147-148
scrobifera Emery, 150
Tachigalia tree, on, 47
tachigaliae sp. nov., 148-150
Phloeostichus
denticollis W. Redtb., 182, 199
Phoethornis superciliosus, 17
Phoridae
two species from British Guina, 435-440
Apterophora gen. nov., 435-437
caliginosa sp. nov. (fig. 44),
437-438 :
ecitophora comes Schmitz, 439
Phorids,
synoeketes of social beetles, 80
Phrenapates bennetti Kirby, 118
_ Phyllobates, 289, 291
Phyllomedusa bicolor (Bodd), 28
Phyllostomus h. hastatus, 17, 280
Picard, 92, 104, 114, 174, 180, 181, 428
Pierce, 175
Pionus fuscus, 17
Pipra aureola, 20
Pipramorpha oleaginea, 17, 20
Pitangus sulphuratus, 21
embryology, 21
pinion development, 21
wing development, 21
Pithecia pithecia (Linn.), 6, 283
photographs, 16
Plant communities, 18 |
in second growth jungle, 18
Platisus, 198
Platypodidae, 97
Platypus compositus
Index 451
habits of, 98-99
Platyrrhines, 243
Plodia interpunctella, 97
Podium
ruficrus, 46
rufipes, 22
Pomeroon Trail, 17
Ponerinae, 106, 138
Pope, Clifford, 7
Porcupine, tree, 286
Potos flavus flavus (Schreber), 277
Porzana albicollis, 9, 20
Priodontes sp., 286
Procryptocercus, 48
Procyon cancrivorus cancrivorus (Cuv.), 277
Proechimys cayennensis (Desmarest), 273
Progue chalybea, 21
pinion development, 21
wing development, 21
Prostominae, 199, 201
Prostominia, 199
Prostomis mandibularis Fabr., 182, 198
Psammoechus
bipunctatus Fabr., 179
desjardinsi Guérin, 179
Pseudococcus
bromeliae (figs. 9 and 10), 60, 62
devoured by midges, 225
enemies, 79, 83-87, 225
“‘milked"’ by Coccidotrophus, 63,
69, 70-72
ostioles, 61, 67, 68
Tachigalia tree, in, 45, 59, 60
wax, 59-61
farinosus
ostioles, 67
Pseudomyrma
damnosa sp. nov. (fig. 13), 141
on Tachigalia tree, 48, 139-143
exterminated by Solenopsis, 48
on Tachigalia tree, 39, 48, 50
latinoda, 137, 143
var. endophyta, 138
maligna sp. nov. (fig. 14), 144
description, 143-146
var. cholerica var. noy., 48, 146
var. crucians var. nov., 147
on Tachigalia tree, 48, 49, 50
mexicana, 87
queens, 87
picta Stitz, 138, 143
Pseudomyrminae, 106, 137, 138, 139-147
of Tachigalia tree, 139-147
Psittacula passerina, 21
development of wing, 21
Psophia crepitans, 16, 20, 21
development of wing, 21
pinion development, 21
Psyllomyia Loew, 436
Pleroglossus
aracari atricollis. 16, 19
pinion development, 21
wing development, 21
viridis, 19
Puma, 286
Pupation, controlled, 23
Pygocentrus niger, 10
Quadrille Bird, 20
Racheolopha nivetacta, 31
Ramphastos
monilis, 19
vitellinus, 19
Rattus
norvegicus, (Erxleben), 284
rattus alerandrinus (Geolf,), 274
Reese, A. M., 7
Reimarus, 117
Reitter, 174, 175, 176, 177, 178, 179, 180,
181, 182, 183
Reptiles
452 , Zoologica
Ameiva, 27
check list of 112 reptiles, 26
life histories, 8
living, sent to N. Y. Zoological Park, 10
lizards, new from tr opical research
station, 301-305
Research,
methods of, 19
Rhathymus beebei, 26
Rhinotermes marginalis, 401, 402
Rhynchiscus naso (Wied), 279
Rhynchomicropteron Annandale, 436
Rifargia phanerostigma, 31
Rignano, 116
Rodway, James, 23
Roosevelt Theodore, 17
and tropical research station, 24
description of tropical station by, 17
Riibsamen, 85
Rupicola rupicola, 9, 15
habits, 15
Rupununni River, 23
Ruthven, 295
Saalas, 213
Saccopleura lycealis, 31
Saccopteryx bilineata (Temminck), 279
Saimiri
cassaquiarensis (Humboldt), 282
sciureus (Linnaeus), 282
Saissetia nigra nietu, 85, 225
Salientia,
eggs of, 291
Sargossa weed organisms, 25
Satterlee, M., 7, 8
Scalariidae, 96
Scalida, 67, 199
Scalidiidae, 183, 199, 201
Sceliphron fistulare, 23
Schiddte, 368
Schmitz, 436, 439
Schneider-Orelli, 98
Schomburgk, 18
Schultz, A. H.
fetuses of Guiana howling monkey, 242-
262
acknowledgments, 243
bibliography, 262
figures (19-24), 248, 253, 254, 256,
258, 260
indices compared with human,
246-248
measurements, 244-245
twinning in monkeys, 244-254
Schwalbe, 253
Schwarz, E. A., 35, 64, 88
new species of Coleoptera, 187, 194
Sciurillus sp., 286
Scolytidae, 97, 99
Scymnus, Kugelann, 103, 193
xantholeucus sp. NOv., 193
in Tachigalia tree, 83, 86
Sea Wrack, 25
Seedeater,
black-headed, 20
chestnut-bellied, 20
Selenidera culik, 19
Selenka, 244
Sequels, 30
Serrasalmo niger, 16, 22
capture, 16
general, 22
habits, 16
photographs, 16
Sherman, 213
Silvanidae, 59, 67, 96, 198, 200
behavior of social silvanids, 111-118
feeding habits, 103-109
habits, 173-176
new species, 189-192
pronotum, teeth on, 89
[Vol. II]
pupation, 109, 110
Tachigalia tree, in, 51-126
Silvaninae, 200
description of subfamily, 201-202
_ genera of, 202-203
Silvanus, 59, 89, 95, 198, 203
bidentaitus Fabr., 174
fagi Guerin, 175
food, 104
gemellatus Duv., 174
planatus Germ, 175
_ unidentatus Fabr., 175
Silvestri, F., 377, 384, 403, 405
Descriptiones vermitum in Anglorum
Guiana, 307-32
Smith, 117
Smith, J: B., 173, 174, 175, 176, iid, oon
181, 182
Smolucha, T. V., 7
Snodgrass, R. E., 323
Solenopsis, 4
altinodis, on Tachigalia, 48, 79, 82, 83,
88, 154-156
(fig. 15), 155
helena Emery
subsp. nov. hermione, 157
subsp. nov. ultriz, 157-158
Spanish
in Bartica, 18
Sphaerodactylus molei Boettger, 301
Spiders
food of wasps of Tachigalia tree, 46
in Tachigalia tree, 46
Spilographa, 23
Spirachtha
eurymedusa Schiddte, 325, 326, 327
mirabilis sp. nov. (fig. 28), 323-326, 367
glandular structure of abdominal
appendages, 367-381
abdentnd appendages, 368-
basement membrane, 372
bibliography, 367, 368, 381
discussion, 376-380
drawings (fig. 41, Pl. XVI,
XVII), 367
hypodermis, 371-372
interpretations, 373-376
introduction, 367-368
technique, 367
schiodtei sp. nov,. 326-327, 367
Spongiocerinae, 97
Sporophila,
bouvronides, 20
castaneiventris, 20
Spruce, 137
Squirrel, wart, 286
Stach, 399
‘Stagmomantis hoorie Caudell, 31
Staphylinidae
new species, 323-364
Stenitus gen. nov.
description, 414-415
guiananus sp. nov. (PL. XXVI),
415-417
Stitz, 138
Sulc, 61, 62, 68
Sycosoter lavagnei, 428
Symbranchus marmoratus, 10
Synechtrans
of social beetles, 81
Synoeca irina Spinola, 30
sounds made by, 30
Synoeketes
of social beetles, 80
Synoemis Pascoe, 190
Syntermes
molestus Burm.,
parallelus sp. ie "Plate XV), 318-321
Tachigalia
Nos 1-23]
ants, 135-168
biocoenose of, 41-50, 130-131
description, 39-41, 42-45
distribution, 40
formicarum, 39, 137
new midge from, 225-226
paniculata Aublet, 118
social beetles of, 35-118
Tadpoles, tropical, 28
Tail down,
specialization of, in ducks, 32
Tail feathers,
of motmots, 31
Tamandua
tetradactyla tetradactyla Linn., 269
Tapinoma, 48
Tapir, 286
Tapirus sp., 286
Tatu kappleri (Krauss), 270
Tayassu Fee beebei Anthony, 270
Taylor, Anna, 7
Tayra barbara barbara (Linn), 277
Tee-Van, J., 7, 8, 28
Telephanus, 67, 198
velox Haldem, 176
Tenebrio molitor, 72
Termitaphis
three new species, 403-408
description of new species, 405-407
key to species, 408
literature, foot-note, 403
(Plate XXIV), 404
circumvallata Wasm., 408
guianae sp. nov. (figs. 1-14),
405, 408
insularis sp. noy., 406, 408
mexicanus Silv., 405, 407, 408
trinidadensis sp. nov., 406, 408
. sufafra Silv., 406, 407, 408
Termites, 8, 27
behavior, 111
fifty new species, 8
guests, 323-366, 367-381
Kartabo, of, 27
new species from B. Guiana
described by Silvestri, 307-321
Tachigalia tree, of, 46
Termitobracon gen. nov.
Pe pail with Ypsistrocerus Cushman,
description of genus, 429-430
source of material, 427, 432
emersoni sp. nov.
description (figs. 42 and 43),
430-432
Termitogaster
brevis sp. nov., 334-345
emersoni sp. nov. (fig. 34), 342-344
fissipennis, 340
insolens Casey, 345
simopelta sp. nov. (fig. 33), 341-342
simulans sp. nov. (fig. 32), 338-340
Termitomimus, 376, 377 :
Termitophya
amica sp. nov., 351-352
flaviventris sp. nov., 352-353
heyeri Wasm., 351
punctata sp. nov. (fig. 36), 349-351
Tetraponera, 377, 378
Thamnomanes glaucus, 20
Thaumataerius gen. nov.
description, 364-365
emersoni sp. nov. (fig. 40), 365-366
Thaxter, R., 36
Thermesia dorsilinea, 31
Thomann, 68
Thomas, 272
Thyonaea dremma, 31
Thyreozrenus gen. nov.
description, 329-330
Index
453
major sp. nov., 332-333
parviceps sp. noy. (fig. 29), 330-331
pulchellus sp. nov. (fig. 30), 332
Thysamera
five new species from British Guiana,
383-402
Tidopterus gen. nov.
description, 420
sequens sp. nov. (Pl. XVII), 420-
421
Tinamou,
great blue, 31
Guiana great, 20
variegated, 20
Tinamus
major, 20
tao Temm., 31
nest and eggs, 31
Tocaca, 35
Toucan, 16, 19
black-necked Aracari, 19
breeding habits of five species, 19
green Aracari, 19
photographs, 16
red-billed, 19
sulphur- and white-breasted, 19
Toucanet, Guiana, 19
Trachopeplus gen. nov.
description, 353-354
setosus sp. nov. (fig. 37), 354-356
Tragardh, 376, 377, 379
Trichechus
manatus, 28
sp., 286
Tridesmus, 413
Triplaris, 35, 39
Trogons, 9
Trogonurus curucui curucui (Linné), 30
Tropical Research Station
absence of flies and mosquitoes, 4
annual reports (1917-1920), 24, 26, 27,
area, 4
climate, 4
contributions, 7 (1916-1921) 15~32
curator of birds, 4
director, 4, 5
fauna and flora, 4, 26
first season, 16, 17
founding, 4, 15, 17, 24
fourth year, 28
living organisms sent to N. Y. Zoo.
Park, 10
maps, 6, 17, 18, 38
moving picture film, 8
nature of terrain, 4, 18
objects of, 3-11
opportunities for research, 24
paintings by staff artists, 8
photographs, 8, 16, 17
pictures from, 16
re-establishment at Kartaho, 27, 28
Roosevelt, Theodore, and, 24
staff, 5-7
water colors, 8
wild life, 26
work accomplished, 8
Tropisms, of insects, 111
Trosia nigripes, 31
Trumpeters, grey-backed, 20
Tyranniscus acer, 17, 20
Tyrantlet, Guiana, 20
Trypozrylon
cinereohirtum, 22
leucotrichium, 22
Twinning °
human, 255
monkeys,
Alouatta, 244
Cercocebus, 244
454
Cercopithecus, 244
Papio, 244
_ degree of resemblance, 254
Ule, 137, 138
ant-fauna of Tachigalia, 137
Vampyrus, sp., 16
spectrum spectrum (Linn.), 280
Venezuela
birds of Orinoco region, 31
mainland forest, 31
mangrove forest, 31
ornithological reconnaissance, 30
pitch lakes, 31
Vermileo, 111
Vertebrates, higher, 26
amphibia, reptiles, mammalia, 26
Wallace, Alfred Russell, 3, 4
Wasman, 64, 351, 379, 380, 403, 436
Wasps
at tropical research station, 16
attending membracids, 104-105
Bartica, of, 22 $
black reed, 22
blue huntress, 23
buff Eumenes, 22
forest shell, 23
one-banded dauber, 23
poison, paralysis by, 23
red potter, 22
Tachigalia tree, of, 36, 46
tapping, 30
white-footed, 22
Waterton, Charles, 3, 4
Wheeler, W. M., 7, 8, 25, 29, 35, 101, 189,
191, 192, 197, 225, 377, 378, 379
Wheeler, W. M.
habits of European and North American
Cucujidae, 171-183
bibliography, 119-126
social beetles in Brit. Guiana, 35-126
acknowledgments, 35-37
bibliography. 119-126
pear of social Silvanids, 111-
biocoenose of Tachigalia, 41-50
Coccidotrophus socialis, 51-88
Zoologica
[Vol. IlI
cocoon, 109-111
eunausibus wheeleri, 88-91
feeding habits, 103-109
(figs. 4-12, Plates I-V), 43, 44, 54,
57, 58, 60, 62, 66, 81, 128-134
general considerations, 91
introduction, 35-37
maps, 38
social beetles, 51, 118
soci life among Coleoptera, 97-
10
Tachigalia plant, 39-41
Tachigalia ants, the, 135-168
bibliography, 119-126
(figs. 13-16) 141, 144, 155, 164
White, W. G., 23, 104, 177, 178
in British Guiana, 7
Wilderness, laboratory, 24
Wilderness
animal life, 32
British Guiana, 32
our search for, 32
Wing,
development of, 21
Wood, C. A., 7, 11
Xenogaster fossulato sp. noy., 359
Xenopelia gen. nov.
description, 356-357
cornuta sp. nov. (fig. 38), 357-358
Xyleborinae, 97, 99
Xyleborus zylographus, 99-101
Xyloterus, 102
Ypsistrocerinae, 429
Ypsistrocerus, 429, 432
compared with Termitobracon, 432
Zaevius calocore, 31
Zatrephes cardytera, 31
Zethusculus hamotus, 23
Zoologica
plate volumes, 8
Zoological Society, 16
ann. report, 16
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in ZOOLOGICA VOL. I. pe
1—Geographic Variation in Birds (Beebe).............0005 wee ;
2-3—Ecology of the Hoatzin; Birds of Venezuela (Beebe)... .50 er ae -
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7—Juv. Plumage, Yucatan Jay (Bechbe) ..........cc ecu eees 15
8—Northern Elephant Seal (Townsend).............00005: 25 sf
9—Affections of Respiratory and Digestive Organs, Primates «
DEMERS esa SNS ici da: stgld Ai iiayt kN diovan o hn eine a oe View 6 15
- 10—New Blood Pheasant (Beebe) ..-........cccceceeceenes 15
_ 11—Feeding Habits, Serpents (Ditmars) .............0005. 50
3 12-13-14—-Ontogeny, the White Ibis; Specialization, Tail-
down in Ducks; Postponed Moult, Certain Passerines “tary
Ras MERRIE PM aL ita eo 5c abebw Aiea ne WOR Oh 9 evs 25 VS
> 15—Pheasant Studies (Beebe) .........cccccccsecccdcecres 25 Ne hee
16—Porpoise in Captivity CHOUNS ENE) ii 5. ni Vieihiw bie oruse Caio Lea
17 18—The Genus Gennaeus (Beebe) ......... 6 i ae ees ren 25 as
“fe Notes on Costa Rican Birds (Crandall) ween 7
19—Natural History, Whale Shark (Gudger) .............. 30 fs"
20—Ectoparasitic Trematodes (MacCallum) ............065 25 ae:
Postage, Zoologica, 5 cents OES
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Postage-weight, 4 pounds
ZOOLOGICA VOL. II » aS
ie Paper Cloth
_-—s« 1&Two Series of Amphibians (Deckert)..........+++0000 25 a:
Wie 2—Tetrapteryx Stage, Ancestry Birds (Beebe)............. 25 eee
a 3 4—Notes, Birds, Para, Brazil; Fauna of Jungle Debris Ae
_ ERGNE NOME ek. ahve waiatia enables << Se PN en Shee 25 Gig!
a. 5—The Gaff-Topsail Catfish (Gudger)..-.......6eeeceeenees 25 oe.
‘ 6—Mammals of Australia in the Zoological Park (W. H. othe
Die: BOWOL 7 eiv'en da ajc niss dees tecevoes Teeth: > cliincald'e Ste 9 fs a Coa
7, 8, 9—Vertebrates, Bri. Guiana (Beebe).......-+--0++e0+: 25
10—Habits of Sage Grouse (Horsfall).......-...-seeeeeeee 25 ws
11—Eclipse Plumage, Domestic Fowl (Crandall)...........- 25 tA
12—Life History Puffer (Welsh-Breder).......+.++++eeee0% 25 ae
13—Hermaphroditism of Croaker (Bréder) ais de sv csccsyeces 25 ie
ZOOLOGICA VOL. Il—Continued
14--Food of Certain Minnows (Breder-Crawford)...........
15—Fishes of Sandy Hook Bay (Breder)...........22+085-
16—Weaving of Weaver Birds (Friedman).............:45.
17—Scales of Whitefishes (Van QOosten)..........2..0.000-s
18—The Anderson Tree Frog (Noble)...........c-ceceeves
Postage, Zoologica, 5 cents
ZOOLOGICA VOL. III
1-2—Objects, Tropical Research Station (Osborn) ..........
Contributions Tropical Research Station (Beebe)........
3-11—Social Beetles, British Guiana (Wheeler) Sin tye latlebape raretta Ce
Tachigalia® Anta SCW helen) 6 dete Ud sein ss aglakelels alate es es
Habits of ,Cucujidae: (Wheeler). 20h fo acts amie tak
New Coleoptera (Schwarz-Barber) .....:..0cccceewceeecs
Larvae,’ Pupae, Social Beetles (Boving)................
New. Diadiplosis “(Relt) 2 7oa. witne ts a bisis tote bgiatotele AWaTe
New. ‘Blepyrus):(Brues)...op icin aaa ote see
Tachigalia Membracids (Herbert Osborn)...............
Wow. Entomobtya (Folsom) ois. . signs yaw ute ean ek eal
12—Fetuses, Guiana Howling Monkey (Schultz)..........:.
13—Mammals Collected Trop. Res. Sta. (Anthony)..........
14-15—New Batrachians (Noble). New Lizards (Noble)..
16—Descriptions Termitum in Anglorum Sone Reportum
CBHUCStrs) i. ERE SE oes sa eia eee nie ot ode ae eepmigiiats OMe
17—New Genera and Species , Termitophilous Coleoptera
CU CaM) ciae Dl cL UES nua SECRET gl a! oad ne Cn ene shea
18—Glandular Structure, Abdominal Appendages, Termite
Guest CM clndoo) ie eis. ¢ Kato aaie SV iy eyelet ke told Re -
. 19—Termitophilous Apterygota (Folsom) BEIM Pico i ne
20—Three Apparently New Species, Termitaphis (atorrison)
21—Termitophilous Millipeds (Chamberlin) .............+.
22—A Termitophilous Braconid (Brues) ...........¢0..0.
23—Two Myrmecophilous Phoridae (Brues).
(16-23 incl. (Trop. Res. Sta.) in preparation)
_ Postage, Zoologica, 5 cents
ZOOPATHOLOGICA VOL. I
Parasitic Trematodes (MacCallum) ............0:e0eeeeees
New Parasitic Worms (MacOallum) ........0..0cceceweweees
Telorchis and other Trematodes (MacCallum)...............
The Polystomidwe (MacCallum)... 22... lucie oa scale eee ue euihe
Camallanus and other Nematodes (MacCallum).............-
Studies in Helminthology (MacCallwm).........4.....0+02-
Postage, Zoopathologica, 5 cents
Corrected to October: 1923
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