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Title: The Biotic Associations of Cockroaches

Author: Louis M. Roth
Edwin R. Willis

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SMITHSONIAN

MISCELLANEOUS
COLLECTIONS

VOL. 141

"EVERY MAN IS A VALUABLE MEMBER OF SOCIETY WHO, BY HIS OBSERVATIONS,
RESEARCHES,
AND EXPERIMENTS, PROCURES KNOWLEDGE FOR MEN"—JAMES SMITHSON

(Publication 4470)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1961

PORT CITY PRESS, INC.
BALTIMORE, MD., U. S. A.

ADVERTISEMENT

The Smithsonian Miscellaneous Collections series contains, since the
suspension in 1916 of the Smithsonian Contributions to Knowledge, all the
publications issued directly by the Institution except the Annual Report and
occasional publications of a special nature. As the name of the series
implies, its scope is not limited, and the volumes thus far issued relate to
nearly every branch of science. Papers in the fields of biology, geology,
anthropology, and astrophysics have predominated.
Leonard Carmichael,
Secretary, Smithsonian Institution.

SMITHSONIAN MISCELLANEOUS
COLLECTIONS
VOLUME 141 (WHOLE VOLUME)

THE BIOTIC ASSOCIATIONS
OF
COCKROACHES

(With 37 Plates)

By

LOUIS M. ROTH
AND

EDWIN R. WILLIS

Pioneering Research Division, United States Army
Quartermaster Research and Engineering Center
Natick, Mass.

(Publication 4422)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 2, 1960

PL. 1
Blaberus craniifer, c. X 2. 1. (Photograph by Jack Salmon, Philadelphia
Quartermaster Depot.).

SMITHSONIAN MISCELLANEOUS
COLLECTIONS
VOLUME 141 (WHOLE VOLUME)

THE BIOTIC ASSOCIATIONS OF
COCKROACHES

(With 37 Plates)

By

LOUIS M. ROTH
AND

EDWIN R. WILLIS

Pioneering Research Division, United States Army
Quartermaster Research and Engineering Center
Natick, Mass.

(Publication 4422)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 2, 1960

THE LORD BALTIMORE PRESS, INC.
BALTIMORE, MD., U. S. A.

FOREWORD
People having only casual interest in insects usually express amazement
when they learn how much is known about this most numerous group of
animals. However, while entomologists have good reason to take pride in
the accomplishments of their contemporaries and predecessors, they are
more likely to be appalled by how much remains to be learned. We are
indeed ignorant of even the identity of fully half and probably much more
than half the total number of insect species. Of those that have been
described, we have reasonably complete information about the behavior and
basic environmental relationships for only a comparative few. The great
majority of the remainder are known only from specimens found in
museum collections. Such information as we have about these species
usually amounts to no more than date and locality of collection.
This is true of the cockroaches, which now include approximately 3,500
described species. Conservative estimates based on partially studied
museum collections and the percent of new species found in recent
acquisitions, particularly from tropical and subtropical countries, indicate
that at least 4,000 species remain unnamed. Although the group is well
known in general terms to nearly all entomologists, there is an almost
complete void of information about all except the few domestic species and,
to a progressively diminishing degree, some 400 others. Many details about

the lives of even those that share man's habitations are not fully understood.
This then is a rough measure of how little is known about cockroaches.
With the exception of mosquitoes and a few other comparatively small
groups of insects on which work has been concentrated, it is doubtful if any
other comparable segment of the world's insect fauna is better known.
Already an estimated 800,000 kinds of insects have been described, and
since this figure is generally regarded as less than half the actual total, think
what this means in terms of knowledge yet to be assembled. No wonder
entomology is a growing science with a promising future, but the
magnitude of the task also presents a serious obstacle to progress. Progress
can continue only if the scattered literature resulting from the diversified
labors of hundreds of contributors is brought together and summarized in
thorough and well-organized compilations that can serve as a solid basis for
future research.
The present work is such a compilation, for it assembles what has been
gleaned from approximately 1,700 sources, including correspondence with
a large number of other workers. Original observations during some eight
years of concentrated effort in U. S. Army Quartermaster research
laboratories are a valuable supplement to what others have done, and with
this background of experience the authors are especially well qualified to
appraise previous work. Seldom has a compilation been done so thoroughly
or a single large group of insects been the subject of such uninterrupted
effort.
The contents gives the categories of subject matter treated and the
introduction discusses the value of this assembled information and offers
suggestions for future study. No longer are cockroaches regarded only as
disagreeable pests; many species appear to be important, actually or
potentially, as carriers of disease. Recognition of this importance has grown
considerably, even in the period since World War II. Consequently, anything
that increases our knowledge of the basic bionomics of cockroaches will be
consulted widely for factual information and for clues to new approaches.
In spite of this extensive compilation, the limitations of present information
about cockroach bionomics must be kept in mind. The cited observations of
many writers were fragmentary, or their conclusions disagreed. But it is
fundamental to scientific inquiry that we should know and attempt to

evaluate the results of previous study, and that is what Drs. Roth and Willis
have done. Fortunately, their review is readily available. Sometimes, a piece
of work fails to be of maximum value because the results are not generally
accessible to later students. For this reason I am especially glad that the
Smithsonian Institution, by disseminating the results of the authors' labors,
has this opportunity to exercise one of its traditional functions—that of
diffusing knowledge.
Throughout the period of research by Drs. Roth and Willis at Natick, I was
in frequent correspondence with them, and I admire their many
accomplishments. Our warmest commendations should go not only to them
personally but also to those in administration who encouraged their
fundamental research and who aided in the financial support of this
publication.
Ashley B. Gurney
Entomology Research Division
United States Department of Agriculture

CONTENTS
Page
Foreword iii
List of Plates vii
List of Illustrations viii
I. Introduction 1
Historical 2
Methods 4
Future work 5
Illustrations 7
II. Species of cockroaches 7

III. Ecological relationships 14
Cave habitats 16
Cavernicolous cockroaches 17
Cockroaches from burrows 23
Desert habitats 25
Desert cockroaches 27
Aquatic habitats 30
Amphibious cockroaches 31
Outdoor habitats 33
Cockroaches from outdoor habitats 35
Structural habitats 70
Land-based structures 73
Cockroaches associated with land-based
74
structures
Ships 82
Cockroaches associated with ships 85
Aircraft 87
Cockroaches associated with aircraft 88
IV. Classification of the associations 91
V. Mutualism 96
Bacteroids 96
Cockroaches in which bacteroids have been
99
found
Bacteria 100
Protozoa 101
VI. Viruses associated with cockroaches 103
VII. Bacteria associated with cockroaches 104
VIII. Fungi and yeasts 127
Fungi associated with cockroaches 129
IX. Higher plants associated with cockroaches 139
Damage to plants by cockroaches 162
X. Protozoa associated with cockroaches 166
XI. Helminths associated with cockroaches 190

Helminths for which cockroaches serve as
192
primary hosts
Helminths for which cockroaches serve as
203
intermediate hosts
Helminths whose eggs have been carried by
208
cockroaches
XII. Arthropoda associated with cockroaches 210
Arachnida 211
Chilopoda 222
Insecta 224
Hymenoptera 234
Predators and parasites of cockroach eggs 234
Host selection by egg parasites 254
Cockroach-hunting wasps 255
Ants predaceous on cockroaches 266
XIII. Vertebrata associated with cockroaches 268
Pisces 268
Amphibia 269
Reptilia 272
Aves 276
Mammalia 283
Checklist of cockroaches and symbiotic
XIV. 290
associates
XV. Cockroaches as commensals 310
Hosts of commensal cockroaches 310
Checklist of commensal cockroaches with
315
their hosts
Obscure associations 316
XVI. Cockroaches as predators 319
Interspecies predation 319
Intraspecies predation 322
XVII. Associations among cockroaches 324
Familial associations 325
Gregariousness 330

Intraspecies fighting 336
Interspecies compatibility 337
Interspecies antagonism 341
XVIII. Defense of cockroaches against predators 343
XIX. The biological control of cockroaches 348
Invertebrates 349
Vertebrates 353
Acknowledgments 354
References 356
Index 441

LIST OF PLATES
Plate Page
1 Blaberus craniifer, c. X 2. 1. (Photograph by Jack
Salmon, Philadelphia Quartermaster Depot.). a
2 Blaberus craniifer, nymph. (Photograph by Jack
Salmon.) A-3
3 Blaberus giganteus, c. X 2.2. (Photograph by Jack
Salmon.) A-4
4 Blatta orientalis, c. X 3.8. A, Male. B, Female.
(Photographs by Jack Salmon.) A-5
5 A-B, Blattella germanica, c. X 5.2. A, Male. B,
Female.
C-D, Blattella vaga, c. X 5.2. C, Male. D, Female
with oötheca. A-6
6 Byrsotria fumigata, c. X 2. A, Brachypterous male.
B, Macropterous male. C, Female. A-7

7 A and B, Cariblatta lutea minima, X 10. A, Male.
B, Female with partly formed oötheca. C, Ectobius
pallidus, female with completely formed oötheca,
X 8. (C, From Roth and Willis [1957].) A-8
8 A, Cryptocercus punctulatus, c. X 4.6. (Photograph
by Jack Salmon.) B, Panesthia australis, X 2.8. A-9
9 Cutilia sp. near sedilloti, c. X 5. A, Male. B,
Female. A-10
10 Diploptera punctata, c. X 5. A, Male. B, Female. A-11
11 Eurycotis floridana, c. X 2.8. A, Male. B, Female.
(Photographs by Jack Salmon.) A-12
12 A-B, Gromphadorhina portentosa, c. X 1.5. A,
Male nymph. B, Adult female. C, Coleolaelaps (?)
sp., a mite from G. portentosa, c. X 32. (Glycerine
jelly preparation and photograph of C by Dr.
Barbara Stay.) A-13
12A Ischnoptera deropeltiformis, c. X 5.3. A, Male. B,
Female. A-14
13 Leucophaea maderae, c. X 2.2. A, Male. B,
Female. (Photographs by Jack Salmon.) A-15
14 Nauphoeta cinerea, c. X 3.4. A, Male. B, Female. A-16
15 Neostylopyga rhombifolia, c. X 3.4. A, Male. B,
Female with partially formed oötheca. A-17
16 Panchlora nivea, X 4.5. A, Dead individual
showing normal, pale green coloration. B, Dead
individual showing the bright red coloration (very
dark areas) characteristic of infection with Serratia
marcescens. C, Living male. D, Living female. A-18
17 A, Parcoblatta pensylvanica, female with
completely formed oötheca, X 4. B, Parcoblatta
virginica, female with partly formed oötheca, X
7.3. A-19
18 Parcoblatta uhleriana, c. X 5.5. A, Male. B,
Female with oötheca. A-20
19 Periplaneta americana, c. X 3. A, Male. B, Female.
(Photographs by Jack Salmon.) A-21

20 Periplaneta australasiae, c. X 3.2. A, Male. B,
Female. (Photographs by Jack Salmon.) A-22
21 Periplaneta brunnea, c. X 2.9. A, Male. B, Female. A-23
22 Periplaneta fuliginosa, c. X 2.9. A. Male. B.
Female. A-24
23 Platyzosteria novae seelandiae, c. X 2.9. A. Male.
B, Female with oötheca. A-25
24 Pycnoscelus surinamensis, c. X 3.7. A, Male from
Hawaii. B, Macropterous parthenogenetic female
from Florida. C, Brachypterous nonparthenogenetic
female from Hawaii. D, Late instar nymph.
(Photograph of nymph D, by Jack Salmon.) A-26
25 Supella supellectilium, c. X 6.3. A, Male. B,
Female. A-27
26 Bacteroids from Blattella germanica. A, Part of
abdomen showing mycetocytes in fat body, X 225.
B, Lobe of fat body showing 3 mycetocytes, X 750.
C, Single mycetocyte; bacteroids appear hollow as
result of fixation in Carnoy's fluid, X 1725. D,
Smear of fat body showing bacteroids in various
stages, X 1800. (All preparations and photographs
through the courtesy of Dr. Marion A. Brooks.) A-28
27 Fungi parasitic on cockroaches. A, Herpomyces
arietinus growing on antennae, legs, body, and
cerci of a nymph of Parcoblatta virginica, X 7. B,
Herpomyces stylopygae on antenna of Blatta
orientalis, X 35. (Reproduced from Richards and
Smith [1955, 1956].) C, Herpomyces sp. [probably
H. stylopygae] on antenna of B. orientalis, X 132.
(Photographs B and C through the courtesy of Dr.
A. G. Richards.) A-29
28 A-B, Gregarines (Diplocystis sp.?) from Blaberus
craniifer. A, Organisms removed from intestine, X
50. B, Organisms removed from hemocoele, X 32.
C, Gregarine cysts in feces of Leucophaea
maderae, X 12. A-30

29 A, Undetermined mermithid that parasitizes
Ectobius pallidus. X 9. The worm has partly
emerged from the neck region of the cockroach.
(Reproduced from Roth and Willis [1957].) B,
Undetermined gordian worm that parasitized
Eurycotis floridana shown beside its host, X 1.8.
(Specimen courtesy of Dr. T. Eisner.) A-31
30 A, Heteropoda venatoria, a cockroach-hunting
spider, slightly less than natural size, on bananas.
(Reproduced from a Kodachrome transparency
through the courtesy of Dr. B. J. Kaston.) B to E,
Lycosa sp. (avida?) capturing and feeding on a
nymph of Supella supellectilium in the laboratory,
X 1.4. A-32
31 The centipede Scutigera coleoptrata capturing and
feeding on cockroaches in the laboratory. A to E,
Pursuit, capture, and eating of a nymph of Supella
supellectilium, c. X 1.2. F, Centipede feeding on
adult of Blattella germanica, X 1.8. A-33
32 The mantid Hierodula tenuidentata (?) devouring a
nymph of Periplaneta australasiae, c. X 1.5. A-34
33 A, Prosevania punctata ♂ beside an oötheca of
Periplaneta americana, X 5. B, Hyptia harpyoides
with oötheca of Parcoblatta uhleriana from which
it had emerged, X 5. C, Larva of a lampyrid beetle
feeding on Parcoblatta virginica in the laboratory,
X 4. A-35
34 Chalcid parasites of cockroach eggs. A, Anastatus
floridanus ovipositing into an oötheca which is still
being carried by Eurycotis floridana, c. X 4. B,
Comperia merceti ovipositing into an oötheca of
Supella supellectilium, c. X 13. C, Tetrastichus
hagenowii ovipositing into an oötheca of
Periplaneta americana, c. X 10. (C from Roth and
Willis [1954b].) A-36
35 A, The wasp finds a cockroach. B, She stings the A-37
prey in the thorax. C, She then leads the disabled

cockroach (antennae clipped) to her nest. D, The
wasp's egg was placed on the coxa of the
cockroach's right mesothoracic leg where it
hatched. E, Portion of the host's abdomen removed
to show feeding larva. F, New adult wasp emerging
from dead host. (Reproduced from F. X. Williams
[1942] from the color paintings of the late W.
Twigg-Smith, through the courtesy of F. A.
Bianchi.)
36 Chemical defense of Diploptera punctata against
predators; the spray pattern is displayed on KI-
starch indicator paper. A, Spray pattern after right
mesothoracic leg was pinched. B, Cumulative spray
pattern after left mesothoracic leg of the same
insect was pinched. C, The defensive glands of the
cockroach on the left had been excised, and it is
under persistent attack by ants from a laboratory
colony of Pogonomyrmex badius (Latreille). The
intact cockroach on the right was also attacked by
the ants, but it discharged a spray of quinones and
repelled the attackers. (From Eisner [1958],
through the courtesy of Dr. T. Eisner.) A-38

LIST OF FIGURES
Fig. Page
1 Diagram illustrating the relationship between a 128
mature plant of Herpomyces stylopygae and the
integument of Blatta orientalis. (Reproduced from

Richards and Smith [1956], through the courtesy of
Dr. A. G. Richards.)
2 Representative Protozoa associated with
cockroaches. A, Monocercomonoides melolonthae,
X 3094 (after Grassé). B, Coelosporidium
periplanetae, X 1310 (after Sprague); trophozoite
with spores and chromatoid bodies. C, Endamoeba
blattae, X 273 (after Kudo); trophozoite. D,
Lophomonas striata, X 330 (after Kudo). E,
Lophomonas blattarum, X 660 (after Kudo). F,
Retortamonas blattae, X 3094 (after Wenrich). G,
Nyctotherus ovalis, X 175 (after Kudo). H,
Gregarina rhyparobiae, c. X 52: mature
trophozoite attached to intestinal wall of
Leucophaea maderae. (Redrawn from J. M. Watson
[1945].) I, Diplocystis schneideri, c. X 14.4 (after
Kunstler). J, Gregarina blattarum, c. X 57 (after
Kudo). K, Protomagalhaesia serpentula, X 36
(after Pinto). L, Gamocystis tenax, magnification
not known (after Schneider). (All figures except H
redrawn from Kudo [1954] after sources indicated.) 168
3 Protozoa from the gut of the wood-feeding
cockroach Cryptocercus punctulatus. A,
Eucomonympha imla, female above, male below, c.
X 375. (From Cleveland [1950c].) B,
Barbulanympha sp. (From Cleveland [1953].) C,
Urinympha talea, c. X 712. (From Cleveland
[1951a].) D, Rhynchonympha tarda, c. X 450.
(From Cleveland [1952].) E, Trichonympha
okolona or T. algoa, c. X 390. (From Cleveland
[1949].) (All drawings reproduced through the
courtesy of Dr. L. R. Cleveland.) 175
4 The cockroach mite, Pimeliaphilus
podapolipophagus. (From Baker et al. [1956];
reproduced through the courtesy of Dr. E. W. Baker
and the National Pest Control Association.) 221

5 Evania appendigaster. Left, dorsal view, X 8.
Right, side view, X 5. (Reproduced with
permission. British Museum [Natural History],
1951, figs. 1A and 1B.) 238
6 Ampulex canaliculata attacking Parcoblatta
virginica. A, Female wasp stinging her prey, c. X
4.8. B, Wasp's egg attached to the coxa of the
mesothoracic leg of the cockroach. C, Larva of A.
canaliculata (about three-quarters grown) feeding
on the internal organs of the host from the exterior,
c. X 4. (Reproduced from F. X. Williams [1929],
through the courtesy of Dr. F. X. Williams and F. A.
Bianchi, Hawaiian Sugar Planters' Association.) 258
7 Cockroach-hunting wasps. A, Dolichurus stantoni
leading a nymph of Blattella lituricollis to her nest,
c. X 4. (Reproduced from F. N. Williams [1919].)
B, Podium haematogastrum attaching her egg to an
Epilampra sp. while on the side of a termite mound
that contains the wasp's nest, c. X 1.6. C,
Epilampra sp. parasitized by P. haematogastrum
showing the wasp's egg attached to the right fore
coxa, c. X 3.2. (B and C reproduced from Williams
[1928], through the courtesy of Dr. F. X. Williams
and F. A. Bianchi.) 263

THE BIOTIC ASSOCIATIONS OF
[1]
COCKROACHES
[2]
By Louis M. Roth and Edwin R. Willis

Pioneering Research Division, United States Army
Quartermaster Research and Engineering Center
Natick, Mass.

(With 37 Plates)

With most of us collectors the life history of an insect begins in the net
and ends in the bottle.
Hanitsch (1928)

I. INTRODUCTION
Recently we brought together much of the literature linking cockroaches
with the transmission of certain organisms that cause disease in man and
other vertebrates. In that paper (1957a) we concluded that cockroaches,
being potential vectors of pathogenic agents, should not be regarded simply
as minor annoyances. Obviously the associations of cockroaches with
agents of vertebrate diseases are of more immediate importance than their
relations with pathogens of lower animals or with nonpathogens. On the
other hand, cockroaches are of general economic as well as medical
importance, and their control is sought by many who are unaware of their
medical significance. That the control of domiciliary cockroaches is far
from satisfactory may be inferred from current entomological and pest-
control journals in which new insecticides are continually advocated to
replace others found to be inadequate. Possibly new approaches to the
control of cockroaches are needed. Whether these lie in the direction of
increased use of parasites and predators for the biological control of these
insects remains to be seen. In any event, the more we know about any
insect, especially its ecology, the greater the likelihood of achieving
satisfactory control. In order to advance knowledge in any field of science,
new research should proceed from the results of prior investigations when
these exist. We hope that the observations and experiments cited herein may
suggest areas for future research and exploitation.
To the best of our knowledge no previous publication has brought together
the vast literature on the parasites, predators, commensals, and other

symbiotic associates of the Blattaria. For this reason, we have tried to
assemble observations on all such known associations. Undoubtedly we
have overlooked some records, as, for example, those buried in papers
dealing with other phases of cockroach biology. We hope that such
inadvertent omissions will not seriously impair the usefulness of this
compilation. Whatever its defects, this review should be a unified source of
information for all who are interested in the biotic associates of
cockroaches.
In addition to previously published information, this monograph also
contains original records and observations on the associations of
cockroaches that are reported here for the first time. Although some of the
observations were made by us, others were made by colleagues who have
graciously made their knowledge available to us in private communications.

HISTORICAL

Chopard (1938) in his book La Biologie des Orthoptères reviewed much of
the literature on cockroaches, but of the many biotic associations that exist
he discussed only the commensal cockroaches, gregariousness, and familial
associations. Asano (1937), who reviewed the natural enemies of
cockroaches, mentioned about 10 groups of animals that attack
cockroaches. Thompson (1951) in his Parasite Host Catalogue, which was
based mainly on papers abstracted or noted in the Review of Applied
Entomology, listed only 19 insect parasites of cockroaches. Eighteen of
these were Hymenoptera which attack only cockroach eggs; the single
dipteron listed (Sarcophaga lambens Wiedemann, supposedly parasitic on
Pycnoscelus surinamensis) is not a parasite in this case, but deposits its
eggs on the dead insects (see p. 229). Cameron (1955) listed as parasites
and predators of the cockroach 24 species of hymenopterous egg parasites,
7 species of Ampulex which hunt nymphs and adults, 17 Protozoa, 13
nematodes, 5 bacteria, 2 mites, and a few other miscellaneous predators. In
his classified list of the protozoan parasites of the Orthoptera of the world,
Semans (1943) listed about 26 species from cockroaches. Linstow (1878,
1889) recorded 14 species of helminths from cockroaches. Van
Zwaluwenburg (1928) listed 33 names of roundworms which are

commensals or secondary parasites of cockroaches, but some of these
names are synonyms. La Rivers (1949) extended this list with 13 additional
species. Chitwood (1932) recognized 24 species of nematodes which are
primary parasites (probably commensals) of blattids. Steinhaus (1946) gave
many instances of biological relationships between cockroaches and
bacteria, fungi, and yeasts, but the cockroaches were not discussed as an
entity and the information is scattered throughout the book.
In surveying the literature on this subject we have collected a far more
extensive list of animals and plants associated with cockroaches than one
might have expected from an examination of any one of the previous papers
on this subject. In our review of the medically important organisms
associated with the Blattaria, we pointed out that in addition to many
experimental associations cockroaches have been found to harbor, naturally,
4 strains of poliomyelitis virus, about 40 species of pathogenic bacteria, the
eggs of 7 species of pathogenic helminths, and to serve as intermediate
hosts of 12 other species of helminths pathogenic for vertebrates;
cockroaches have also been found to carry, on occasion, 3 species of
Protozoa that are pathogenic to man and 2 species of fungi which are
sometimes found associated with pathological conditions.
In addition to the above organisms of medical importance, we have
compiled records of other organisms, nonpathogenic to vertebrates, which
are naturally associated in some way with cockroaches. None of the
following numbers can be considered absolute because some names may be
synonyms. However, we believe that these figures are very close to the
actual numbers of species that have been isolated because we have
attempted to refer all obvious synonyms to the currently accepted name for
each organism. On this basis there are about 45 species of bacteria, 40
fungi, 6 yeasts, 90 Protozoa, and 45 helminths that have been found
associated naturally with cockroaches. Of the arthropods there are about 2
species of scorpions, 4 spiders, 15 mites, 4 centipedes, and 90 insects. Of
vertebrates there are 4 species of fish, 16 amphibians, 12 reptiles, 20 birds,
and 27 mammals. Besides these there are many records of experimental
associations that have been contrived in the laboratory.
Some idea of the increase in our knowledge of the biotic associations of
cockroaches, during the last 70 years, may be gathered from a comparison
of the above figures with those of Miall and Denny (1886) who presented "

... a long list of parasites which infest the Cockroach." This list included 2
bacteria, 6 Protozoa (some of the names are synonyms), 7 nematodes (some
of these names are also synonyms), 1 mite, 1 wasp, and 1 beetle. In
addition, they mentioned as other foes of the cockroach: monkeys,
hedgehogs, polecats, cats, rats, birds, chameleons, and frogs.

METHODS

We have listed the organisms known to be associated with cockroaches
systematically by phylum, class, order, and family. Within each family the
organisms are listed alphabetically by genus and species. Under each
organism the associated cockroaches are listed as natural or experimental
hosts, vectors, or prey. Identified cockroaches are listed by the currently
accepted name. Unidentified cockroaches are indicated by the word
"Cockroaches." The name of each cockroach is followed by the country in
which the observation was made, the authority for the record, and with a
few exceptions[3] pertinent biological information, where this is known.
Question marks following the names of organisms or countries indicate
tentative or questionable identifications.
Records of predators capturing and feeding on cockroaches in zoos and on
shipboard we consider natural, even though it is very likely that these
particular predators would not normally have access to this prey in nature.
Experimental prey are cockroaches that were fed to predators in the
laboratory. Although these predators may have little, if any, access to these
cockroaches in nature, we have included such records to indicate the
relative acceptability of cockroaches as food by a wide variety of animals.
Records of presumed or known cockroach associates that give no
information about an associated cockroach are not included in this review,
even though certain of these (e.g., species of Ampulex, Evania, Podium)
probably prey upon or parasitize cockroaches exclusively.
The validity of a host-parasite or predator-prey record is dependent upon
the accuracy and knowledge of the observer. In assembling these records
we have had to accept, in most instances, the identifications of species

made by the original authors. However, as a result of our studies on the
biology of various species of cockroaches, including some work on their
hymenopterous parasites, we have questioned certain records in the
literature. Other dubious records which have been perpetuated from one
publication to the next, but which apparently were not based on fact, have
also been questioned or have been clarified with the aid of specialists in
particular groups.
Because the records cover a period of many years, the names of many of the
organisms as well as the names of some of the cockroaches have been
changed. Although it would have been comparatively simple to list the
names as they appeared in the original references, this would have resulted
in misleading redundancy with the same organism being catalogued under
several synonyms. We have attempted to list each organism by its currently
accepted name. However, no attempt was made to prepare complete
taxonomic synonymies; the only synonyms given are those that identify the
organisms by the names used by the authors of the papers cited. The
synonyms under which the cockroaches may have been cited originally are
listed in section II. The synonyms of associated organisms are listed with
each organism. Although authorities for the name changes of the
cockroaches are given, these workers are not necessarily those who were
initially responsible for the synonymies. Various sections have been
checked by specialists in the particular groups. Although we have accepted
name changes suggested by these reviewers, we assume full responsibility
for the names.

FUTURE WORK

After having examined thousands of references on cockroaches, we are
impressed by how little is known about the biology of most species. As a
conservative estimate there are 3,500 described species of Blattaria (J. W.
H. Rehn, 1951). In our literature survey we found records of biotic
associations for about 400 species. Unfortunately, many of these records
contain only a sentence or two of biological information. Our detailed
knowledge of cockroaches is based on studies of the few domiciliary pests
that man attempts to eradicate. Comparable studies of the bionomics of the

less-well-known species should add much valuable information to our
knowledge of this ancient group.
Our understanding of most predator-prey and parasite-host relationships has
barely progressed beyond the taxonomic stage. The total effect of predators
and parasites in limiting natural populations of cockroaches remains to be
determined. It is still not known how, for example, predatory or parasitic
wasps select specific cockroaches from among all other insects. Secretions
produced by certain cockroaches (e.g., 2-hexenal by Eurycotis floridana)
will ward off certain predators. The identities and biological activities of
most cockroach secretions are unknown, but the use of protective chemicals
against predators may be widespread among cockroaches. If so, how
effective are these repellents in protecting the individual or the species? It is
not known whether cockroaches are protected by apparent mimetic
resemblances to other arthropods. There is no experimental proof that insect
parasites can successfully attack the eggs of cockroaches that incubate their
eggs while they are being carried by the female.
It is conceivable that biological control of cockroaches might be achieved in
limited areas such as man-made structures or sewers, but this possibility has
not been thoroughly explored. It would be informative to know what
effects, if any, organisms such as bacteria, Protozoa (e.g., gregarines),
intestinal nematodes, or other helminths have on cockroaches. Possibly
pathogenic microorganisms can be used for biological control of
cockroaches; this approach seems to have been little explored.
Associations of colonial cockroaches (e.g., Cryptocercus spp.) may be truly
familial or they may merely result from gregariousness. Newly hatched
nymphs of species that carry their oöthecae until the eggs hatch cluster near
the mother. This may be a response to the mother as such, a search for
shelter beneath the nearest object (thigmotaxis? or negative phototaxis?), or
there may well be yet another explanation. Tepper in 1893 stated that the
native cockroaches of Australia are almost wholly carnivorous; little
supporting evidence for this claim has been brought forward since that time.
The apparent supersedure of one species of domiciliary cockroach by
another may result from antagonism between different species, or it may
result from more rapid breeding and more effective utilization of available
food and space; but which? Several species of cockroaches are frequently
found associated with certain plants (e.g., bromeliads and bananas); the

ecological relations in these associations remain to be determined. Many of
the obscure associations between cockroaches and other insects, spiders,
birds, and burrowing animals have never been adequately defined. The
factors influencing cannibalism have never been thoroughly investigated
experimentally. These are only a few ideas for future work that have
occurred to us during the preparation of this review. We hope that these
suggestions as well as other questions that may occur to readers will
stimulate further research in areas where it is obviously needed.

ILLUSTRATIONS

Unless otherwise credited, the illustrations were prepared from photographs
taken by the authors. Except where otherwise stated, all photographs were
taken of unposed living specimens.

II. SPECIES OF COCKROACHES
The cockroaches referred to in this paper are listed below. The currently
accepted name for each species is given alphabetically by genus and species
irrespective of its taxonomic affinities. Synonyms used by certain authors
whose work we have quoted are given in brackets under the respective
species; the synonymy is supported by the reference citation that follows
each synonym. References to illustrations of certain species (e.g., Blaberus
craniifer) that appear in the paper follow the names of the describers.

Agis orientalis Chopard

Aglaopteryx absimilis Gurney

diaphana (Fabricius) [Ceratinoptera diaphana Fabricius; Rehn
and Hebard (1927)]
facies (Walker) [Aglaopteryx devia Rehn; Princis (1929). A.
diaphana (Fabricius) in records from Puerto Rico only; Rehn
(1932b); Gurney (1937)]
gemma Hebard [In Florida records = Ceratinoptera diaphana
R. and H.; Hebard (1917)]
vegeta Rehn
ypsilon Princis

Allacta similis (Saussure) [Phyllodromia obtusata Brunner;
Zimmerman (1948)]

Alluaudellina cavernicola (Shelford) [Alluaudella cavernicola
Shelford; Chopard (1932)]

Amazonina emarginata Princis

Anaplecta asema Hebard
azteca Saussure
decipiens Saussure and Zehntner
fallax Saussure
hemiscotia Hebard
lateralis Burmeister
mexicana Saussure

Aneurina tahuata Hebard
viridis Hebard

Apotrogia angolensis Kirby [Acanthogyna deplanata Chopard;
Princis (1957)]

Aptera fusca (Thunberg) [Aptera cingulata (Burmeister); Gurney
(personal communication, 1957)]

Apteroblatta perplexa Shelford

Archiblatta hoevenii Vollenhoven

Archimandrita marmorata (Stoll)
tessellata Rehn

Arenivaga apacha (Saussure)
bolliana (Saussure)
erratica (Rehn)
floridensis Caudell
grata Hebard
roseni (Brancsik) [Heterogamodes roseni; Bei-Bienko (1950).
"Polygamia" roseni is undoubtedly an erroneous citation of
Polyphaga roseni, as there is no genus Polygamia (Gurney,
personal communication, 1957)].
tonkawa Hebard

Aristiger histrio (Burmeister) [Plumiger histrio (Burm.); Bruijning
(1948). Hemithyrsocera histrio Burm.; Hebard (1929)]

Aspiduchus boriquen J. W. H. Rehn [In Puerto Rico records =
Aspiduchus deplanatus R. and H.; Rehn, J. W. H. (1951a)]
cavernicola J. W. H. Rehn
deplanatus (Saussure)

Attaphila aptera Bolívar
bergi Bolívar
flava Gurney
fungicola Wheeler
schuppi Bolívar
sexdentis Bolívar

Atticola mortoni Bolívar

Audreia bromeliadarum Caudell
jamaicana Rehn and Hebard

Balta godeffroyi (Shelford)
patula (Walker)

platysoma (Walker) [Temnopteryx platysoma (Walker); Hebard
(1943)]
quadricaudata Hebard
scripta (Shelford)
torresiana Hebard
verticalis Hebard

Bantua stigmosa (Krauss) [Derocalymma stigmosa Krauss; Princis
(1957)]

Blaberus atropos (Stoll) [Blabera fusca Brunner; Hebard (1917)]
boliviensis Princis
craniifer Burmeister (pls. 1, 2)
discoidalis Serville [Blaberus cubensis Saussure; Hebard
(1916)]
giganteus (Linnaeus) (pl. 3)

Blaptica dubia (Serville) [Blaberus clarazianus Saussure; Rehn, J.
W. H. (1951)]

Blatta orientalis Linnaeus (pl. 4) [Periplaneta orientalis; Hebard
(1917)]
(Shelfordella) lateralis (Walker) [Shelfordella tartara
(Saussure); Princis (1957). Periplaneta tartara Saussure; Bei-
Bienko (1950)]

Blattella germanica (Linnaeus) (pls. 5, A, B; 31, F) [Blatella
germanica; Gurney (1952). Phyllodromia germanica; Hebard
(1917). Ectobius germanicus; Gurney (personal communication,
1957)]
humbertiana (Saussure) [Blatta humbertiana; Phyllodromia
humbertiana; Hebard (1929)]
lituricollis (Walker) (fig. 7, A) [Blattella bisignata (Brunner);
Bei-Bienko (1950)]
schubotzi Shelford
vaga Hebard (pl. 5, C, D)

Buboblatta armata (Caudell) [Latindia armata Caudell; Hebard
(1920)]

Byrsotria cabrerae Rehn and Hebard
fumigata (Guérin) (pl. 6)

Cahita borero Rehn
nahua (Saussure)

Capucinella delicatula Hebard

Cariblatta antiguensis (Saussure and Zehntner)
cuprea Hebard
delicatula (Guérin) [Blattella delicatula Guérin; Cariblatta
punctulata (Beauvois); Rehn and Hebard (1927)]
hylaea Rehn
imitans Hebard
insularis (Walker)
landalei Rehn and Hebard
lutea lutea (Saussure and Zehntner)
lutea minima Hebard (pl. 7, A, B)
nebulicola Rehn and Hebard
orestera Rehn and Hebard
punctipennis Hebard
reticulosa (Walker)
stenophrys Rehn and Hebard

Cariblattoides instigator Rehn and Hebard
suave Rehn and Hebard

Ceratinoptera picta Brunner

Chorisoneura barbadensis Rehn and Hebard
flavipennis Saussure and Zehntner
formosella Rehn and Hebard
parishi Rehn
specilliger Hebard

texensis Saussure and Zehntner [Chorisoneura plocea Rehn;
Rehn and Hebard (1916)]
translucida (Saussure)

Choristima sp.

Choristimodes sp.

Chromatonotus infuscatus (Brunner)
notatus (Brunner)

Compsodes schwarzi (Caudell)

Comptolampra liturata (Serville) [Compsolampra liturata;
Comptolampra is the original spelling, which is followed by Dr.
K. Princis, according to Gurney (personal communication,
1959)]

Cosmozosteria lateralis (Walker)

Cryptocercus punctulatus Scudder (pl. 8, A)
relictus Bei-Bienko

Cutilia nitida (Brunner)
soror (Brunner)
sp. near sedilloti (Bolívar) (pl. 9) [Determined by Dr. A. B.
Gurney from photographs.]

Cyrtotria capucina (Gerstaecker)

Dendroblatta sobrina Rehn

Derocalymma cruralis (Stål) [Homalodemas cruralis (Stål);
Gurney (personal communication, 1957)]
lampyrina Gerstaecker
porcellio Gerstaecker

Deropeltis autraniana Saussure
erythropeza Adelung

melanophila (Walker)
nigrita Saussure

Diploptera punctata (Eschscholtz) (pls. 10, 36) [Diploptera
dytiscoides (Serville); Princis (1950). Eleutheroda dytiscoides
(Serville); Zimmerman (1948)]

Dryadoblatta scotti (Shelford) [Homalopteryx scotti Shelford;
Rehn (1930)]

Ectobius africanus Saussure
albicinctus (Brunner)
duskei Adelung
lapponicus (Linnaeus) [Ectobius perspicillaris Herbst, as used
by Lucas (1920); Blair (1934)]
lucidus Hgb.
nicaeensis (Brisout)
pallidus (Olivier) (pls. 7, C; 29, A) [Ectobius lividus
(Fabricius); Ectobius livens (Turton); Kevan (1952); Princis
(in Roth and Willis, 1957)]
panzeri Stephens [Ectobius ericetorum (Wesmaël); Ramme
(1923)]
panzeri var. nigripes Stephens
semenovi Bei-Bienko
sylvester (Poda) [Ectobius sylvestris (Poda); Ramme (1951)]
tadzihicus Bei-Bienko
vittiventer (Costa) [Ectobius vittiventris (Costa); Ramme
(1951)]

Ellipsidion Saussure [Apolyta Brunner; Hebard (1943)]
affine Hebard
australe Saussure [Ellipsidion pellucidum (Brunner); Hebard
(1943)]
bicolor (Tepper)
simulans Hebard
variegatum (Fabricius) [Ellipsidion aurantium Saussure;
Hebard (1943)]

Epilampra abdomen-nigrum (De Geer)
annandalei Shelford
azteca Saussure
conferta Walker
conspersa Burmeister
grisea (De Geer)
maya Rehn
mexicana Saussure
mona Rehn and Hebard
notabilis Walker
sodalis Walker
tainana Rehn and Hebard
wheeleri Rehn
sp. (fig. 7, B, C)

Eremoblatta subdiaphana (Scudder)

Ergaula capensis (Saussure) [Dyscologamia capensis Saussure;
Dyscologamia wollastoni Kirby; Princis (1957)]
scarabaeoides Walker [Dyscologamia piolosa (Walker); Princis
(1957). Parapolyphaga erectipilis Chopard; Princis (1950).
Dyscologamia chopardi Hanitsch; Bruijning (1948).
Miroblatta silphoides Chopard; Hebard (1929)].

Escala sp.

Euandroblatta palpalis Chopard

Eublaberus posticus (Erichson)

Eudromiella bicolorata Hebard
calcarata Bei-Bienko

Euphyllodromia angustata (Latreille)
liturifera [Euphyllodromia decastigmata Hebard; Princis
(1959)]

Eurycotis bananae Bei-Bienko
biolleyi Rehn [Eurycotis carbonaria Biolley; Rehn (1918)]
caraibea (Bolívar)
decipiens (Kirby)
dimidiata (Bolívar)
ferrum-equinum Rehn and Hebard
floridana (Walker) (pl. II) [Platyzosteria ingens Scudder;
Platyzosteria sabalianus Scudder and hence, by inference,
Eurycotis sabalianus (Scudder); Hebard (1917)]
galeoides Rehn and Hebard
improcera Rehn
kevani Princis
lixa Rehn
manni Rehn
opaca (Brunner)

Euthlastoblatta abortiva (Caudell)

Euthyrrhapha nigra Chopard
pacifica Coquebert

Geoscapheus robustus Tepper

Graptoblatta notulata (Stål) [Blatta notulata Stål; Hebard (1929).
Phyllodromia hieroglyphica Brunner; Kirby (1904)]

Gromphadorhina laevigata S. and Z.
portentosa (Schaum) (pl. 12, A, B)

Gyna kazungulana Giglio-Tos
maculipennis (Schaum) [Gyna vetula Brunner; Shelford
(1909b)]
tristis Hanitsch

Hebardina concinna (Haan) [Blatta concinna Haan; Blattina
concinna (Haan); Bei-Bienko (1950)]

Hemiblabera brunneri (Saussure)

Henicotyle antillarum (Brunner)

Heterogamodes krügeri (Salfi)
rugosa (Schulthess)

Holocompsa azteca (Saussure)
cyanea (Burmeister)
fulva (Burmeister)
metallica Rehn and Hebard
nitidula (Fabricius)
zapoteca Saussure

Hololampra bivittata (Brullé)
chavesi (Bolívar)
maculata (Schreber) [Aphlebia maculata Schreber; Harz
(1957); Gurney (personal communication, 1959)]
marginata (Schreber)
punctata (Charpentier) [Aphlebia punctata Charpentier;
Ramme (1951)]

Hololeptoblatta sp.

Homalopteryx laminata Brunner

Hoplosphoropyga babaulti Chopard

Hormetica apolinari Hebard
laevigata Burmeister
ventralis Burmeister

Ignabolivaria bilobata Chopard

Ischnoptera deropeltiformis (Brunner) (pl. 12A) [Temnopteryx
deropeltiformis Brunner; Hebard (1917)]
panamae Hebard

podoces Rehn and Hebard
rufa occidentalis Saussure
rufa rufa (De Geer)
schenklingi Karney

Karnyia discoidalis (Brunner)

Kuchinga hemerobina (Gerstaecker) [Phyllodromia hemerobina
Gerstaecker; Rehn (1932)]
remota Hebard

Lamproblatta albipalpus Hebard
meridionalis (Brunner)

Latiblattella chichimeca (Saussure and Zehntner) [Blattella
chichimeca S. and Z.; Hebard (1932)]
lucifrons Hebard
rehni Hebard
vitrea (Brunner)
zapoteca (Saussure)

Leucophaea maderae (Fabricius) (pl. 13) [Rhyparobia maderae;
Hebard (1917). Panchlora maderae; Kirby (1904). Very
probably "Blaberus" maderae is a careless reference to this
species; Gurney (personal communication, 1957)]

Leurolestes pallidus (Brunner)

Litopeltis biolleyi (Saussure)
bispinosa (Saussure) [Audreia marginata Caudell; Hebard
(1920)]
deianira Rehn
musarum Rehn

Lobolampra subaptera Rambur

Loboptera decipiens (Germar)
thaxteri Hebard

Lobopterella dimidiatipes (Bolívar) [Loboptera dimidiatipes
(Bolívar); Princis (1957a). Loboptera sakalava (Saussure);
Hebard (1933a). Loboptera extranea Perkins; Hebard (1922).
Princis (1957a) in erecting Lobopterella pointed out that only
the nontypical variety of sakalava is identical with dimidiatipes.]

Lophoblatta arawaka Hebard

Macropanesthia rhinocerus Saussure

Mareta acutiventris Chopard

Maretina uahuka Hebard

Megaloblatta blaberoides (Walker) [Megaloblatta rufipes Dohrn;
Hebard (1920)]

Megamareta verticalis Hebard

Melanosilpha capensis Saussure and Zehntner

Methana canae Pope
curvigera (Walker)
marginalis (Saussure)

Moluchia (?) dahli Princis

Monastria biguttata (Thunberg)

Muzoa madida Rehn

Myrmeblattina longipes (Chopard)

Myrmecoblatta rehni Mann
wheeleri Hebard

Namablatta bitaeniata (Stål)

Nauclidas nigra (Brunner) [Poroblatta nigra Brunner; Rehn
(1930)]

Nauphoeta cinerea (Olivier) (pl. 14) [Nauphoeta bivittata
Burmeister; Zimmerman (1948)]
flexivitta (Walker) [Nauphoeta brazzae (Bolívar); Rehn (1937)]
punctipennis Chopard

Nelipophygus ramsdeni Rehn and Hebard

Neoblattella brunneriana (Saussure) [Blattella brunneriana;
Gurney (personal communication, 1959)]
carcinus Rehn and Hebard
celeripes Rehn and Hebard
detersa (Walker)
dryas Rehn and Hebard
eurydice Rehn and Hebard
fratercula Hebard
fraterna (Saussure and Zehntner)
grossbecki Rehn and Hebard
laodamia Rehn and Hebard
nahua (Saussure) [Blattella nahua Saussure and Zehntner of
Caudell (1914); Hebard (1920)]
proserpina Rehn and Hebard
semota Rehn and Hebard
tridens Rehn and Hebard
vatia Rehn and Hebard

Neostylopyga rhombifolia (Stoll) (pl. 15) [Dorylaea rhombifolia;
Rehn (personal communication, 1956)]

Nesomylacris cubensis Rehn and Hebard
relica Rehn and Hebard

Nocticola bolivari Chopard
caeca Bolívar

decaryi Chopard
simoni Bolívar
sinensis Silvestri
termitophila Silvestri

Nothoblatta wasmanni (Bolívar)

Notolampra antillarum Shelford

Nyctibora azteca Saussure and Zehntner
brunnea (Thunberg)
laevigata (Beauvois)
lutzi Rehn and Hebard
mexicana Saussure
noctivaga Rehn
obscura Saussure
sericea Burmeister
stygia Walker
tomentosa Serville [Nyctibora latipennis Burmeister; Hebard
(1917, p. 263)]

Oniscosoma granicollis (Saussure)

Opisthoplatia maculata Shiraki
orientalis (Burmeister)

Oulopteryx meliponarum Hebard

Oxyhaloa buprestoides (Saussure)
deusta (Thunberg)

Panchlora antillarum Saussure
exoleta Burmeister
fraterna Saussure and Zehntner
nivea (Linnaeus) (pl. 16) [Panchlora cubensis Saussure;
Gurney (1955). Pycnosceloides aporus Hebard; Hebard
(1921c)]

peruana Saussure
sagax Rehn and Hebard
virescens (Thunberg)

Panesthia angustipennis (Illiger) [Panesthia javanica Serville;
Hebard (1929)]
australis Brunner (pl. 8, B)
laevicollis Saussure
lobipennis Brunner
spadica (Shiraki)

Parahormetica bilobata (Saussure)

Parcoblatta americana (Scudder)
bolliana (Saussure and Zehntner) [Kakerlac schaefferi Rehn;
Hebard (1917)]
caudelli Hebard [ ♀ ♀ of Ischnoptera insolita R. and H.;
Ischnoptera uhleriana fulvescens S. and Z. (in part); Hebard
(1917)]
desertae (Rehn and Hebard) [ ♂ ♂ of Ischnoptera insolita R.
and H.; Hebard (1917)]
divisa (Saussure and Zehntner) [Ischnoptera divisa S. and Z.;
Hebard (1917)]
fulvescens (Saussure and Zehntner) [Ischnoptera uhleriana
fulvescens S. and Z. (in part); Hebard (1917)]
lata (Brunner) [Ischnoptera couloniana R. and H. (not
Saussure); Ischnoptera major R. and H. (not S. and Z.);
Hebard (1917)]
notha Rehn and Hebard
pensylvanica (De Geer) (pl. 17, A) [Ischnoptera pennsylvanica
Saussure; Hebard (1917)]
uhleriana (Saussure) (pl. 18) [Ischnoptera uhleriana Saussure;
Hebard (1917)]
virginica (Brunner) (pls. 17, B; 27, A; 33, C; fig. 6)
[Ischnoptera borealis Brunner; Hebard (1917)]
zebra Hebard

Pelmatosilpha coriacea Rehn
kevani Princis
marginalis Brunner
purpurascens (Kirby)
rotundata Scudder
vagabunda Princis

Periplaneta americana (Linnaeus) (pls. 19, 35) [Stylopyga
americana; Blatta americana L.; Hebard (1917)]
australasiae (Fabricius) (pls. 20, 32)
brunnea Burmeister (pl. 21)
cavernicola Chopard
fuliginosa (Serville) (pl. 22)
ignota Shaw
lata (Herbst)

Perisphaerus armadillo Serville
glomeriformis (Lucas)

Phaetalia pallida (Brunner)

Phidon (?) dubius Princis

Phlebonotus pallens (Serville)

Pholadoblatta inusitata (Rehn)

Phorticolea boliviae Caudell
testacea Bolívar

"Phyllodromia" treitliana Werner

Phyllodromica brevipennis (Fischer)
graeca (Brunner)
irinae (Bei-Bienko)
maculata (Schreber)
megerlei (Fieber)
polita (Krauss)

pygmaea (Bei-Bienko)
tartara (Saussure)
tartara nigrescens Bei-Bienko

Platyzosteria analis (Saussure) [Polyzosteria analis Saussure;
Kirby (1904)]
armata Tepper
bifida (Saussure)
castanea (Brunner)
novae seelandiae (Brunner) (pl. 23) [Periplaneta fortipes
Walker; Shelford (1912); Platyzosteria novae-zealandiae]
scabra (Brunner)

Plectoptera dorsalis (Burmeister)
infulata (Rehn and Hebard)
lacerna Rehn and Hebard
perscita Rehn and Hebard
poeyi (Saussure) [Plectoptera floridana Hebard; Rehn and
Hebard (1927)]
porcellana (Saussure)
pygmaea (Saussure)
rhabdota (Rehn and Hebard)
vermiculata Rehn and Hebard

Polyphaga aegyptiaca (Linnaeus) [Blatta aegyptiaca L.; Bei-
Bienko (1950). Heterogamia aegyptiaca (L.); Gurney (personal
communication, 1957). "Polygamia" aegyptiaca; according to
Gurney (p. c.), there is no genus Polygamia and almost surely
the reference is to Polyphaga aegyptiaca.]
indica Walker [Polyphaga pellucida (Redtenbacher); Princis
(1957)]
saussurei (Dohrn)

Polyzosteria limbata Burmeister
melanaria (Erichson)

Pseudoderopeltis aethiopica (Saussure) [Blatta aethiopica
Saussure; Gurney (personal communication, 1957)]

Pseudomops cincta (Burmeister) [Thyrsocera cincta Scudder;
Hebard (1917)]
laticornis Perty
septentrionalis Hebard

Pseudophoraspis nebulosa (Burmeister)

Pycnoscelus niger (Brunner)
striatus (Kirby) [Leucophaea striata Kirby; Gurney (personal
communication, 1957)]
surinamensis (Linnaeus) (pl. 24) [Leucophaea surinamensis
(L.); Hebard (1917). Blatta melanocephala Stoll; Kirby
(1904)]

Rhicnoda natatrix Shelford

Rhytidometopum dissimile Princis

Riatia fulgida (Saussure) [Lissoblatta fulgida (Saussure); Gurney
(personal communication, 1959)]
orientis Hebard

Robshelfordia circumducta (Walker) [Escala circumducta
(Walker); Gurney (personal communication, 1957)]
longiuscula (Walker) [Escala longiuscula (Walker); Gurney
(personal communication, 1957)]

Salganea morio (Burmeister)

Sibylloblatta panesthoides (Walker)

Simblerastes jamaicanus Rehn and Hebard

Spelaeoblatta gestroi Bolívar

Sphecophila polybiarum Shelford
ravana Fernando

termitium Shelford

Steleopyga (?) sinensis Walker [Dr. Gurney (personal
communication, 1957) could not find a reference to this species.
Walker described species named sinensis in three different
genera of cockroaches, and it is uncertain which one this
combination represents.]

Stictolampra buqueti concinula (Walker)

Styphon bakeri Rehn

Supella hottentotta (Saussure)
supellectilium (Serville) (pls. 25; 30, B-E; 31, A-E)
[Phyllodromia supellectilium (Serv.); Bei-Bienko (1950)]

Symploce breviramis (Hanitsch)
cavernicola (Shelford) [Ischnoptera cavernicola (Shelford);
Phyllodromia nigrocincta Chopard; Hebard (1929)]
curta Hanitsch
flagellata Hebard
hospes (Perkins) [Symploce lita Hebard; Hebard (1922)]
jamaicana (Rehn)
kevani Chopard
parenthesis (Gerstaecker) [Phyllodromia parenthesis
Gerstaecker; Rehn (1932)]
remyi (Hanitsch) [Ischnoptera remyi Hanitsch; Chopard
(1938)]
ruficollis (Fabricius) [Symploce bilabiata Rehn and Hebard;
Princis (1949a)]

Tartaroblatta karatavica Bei-Bienko

Temnopteryx obliquetruncata Chopard
phalerata (Saussure)

Theganopteryx straminea Chopard

Therea nuptialis (Gerstaecker) [Corydia nuptialis Gerstaecker;
Princis (1950)]

Tivia australica Princis
brunnea (Chopard)
fulva (Burmeister)
macracantha Chopard
obscura (Chopard)

Typhloblatta caeca (Chopard) [Spelaeoblatta caeca Chopard;
Chopard (1924b)]

Typhloblattodes madecassus Chopard

Xestoblatta festae (Griffini)
immaculata Hebard

III. ECOLOGICAL RELATIONSHIPS
The ecology of extinct cockroaches is necessarily a highly speculative
subject. From the coexistence of fossil cockroaches and fossil plants in the
same geological stratum, one might conclude that there had been intimate
associations between them during prehistoric life. Heer (1864) and
Goldenberg (1877) suggested that Carboniferous cockroaches fed on the
plants with which they have been found as fossils. Scudder (1879)
concurred with this hypothesis. However, Bolton (1911), remarking on the
noticeable associations of blattoid wings with vegetable remains, suggested
that the cockroaches may have been partly carnivorous, feeding on the
snails Spirorbis pusillus, which were attached to the leaves of Cordaites.

Yet the proximity of fossil insects and plants in the same geological
formation is hardly proof of a similar association during life. In fact,
Sellards (1903), Bolton (1921), and Laurentiaux (1951) have all pointed out
that the cockroach remains, particularly the more resistant wings, may have
been washed into streams by heavy rains and transported with drifting plant
material to places where permanent deposits were accumulating.
Some species of fossil cockroaches have long, well-developed ovipositors,
very unlike present-day cockroaches whose ovipositors are small and
nonprotruding. Brongniart (1889) and Zalesskii (1939, 1953) have
suggested that certain Permian and Carboniferous cockroaches with long
ovipositors may have inserted their eggs singly into trees and other plants,
rather than protecting the eggs with an oötheca. However, Laurentiaux
(1951), although conceding the possibility of egg laying in vegetable
material, suggested that oviposition into the earth is more probable because
of the unbending nature of the ovipositor.
Although the ecological associations of modern cockroaches should be well
known from direct observation, actually most species are still little more
than names on museum specimens, and our knowledge of them is
fragmentary. All too frequently ecological observations have been only
incidental to taxonomic or faunistic studies; yet the biological information
that is contained in such papers is all that we know of many species. For
this reason we have cited these observations in some detail, especially when
they were brief; longer accounts of cockroach bionomics, of necessity, have
been abstracted.
Very few exclusively ecological studies of insects have included
cockroaches. The native woodroaches (Parcoblatta pensylvanica, P.
uhleriana, and P. virginica) of the northern United States were included in
ecological studies of the Orthoptera by Hubbell (1922), Strohecker (1937),
and Cantrall (1943). Fifteen species of cockroaches were included in an
ecological study of the Orthoptera of northern Florida by Friauf (1953). The
original papers should be consulted for detailed descriptions of the habitats
and accounts of the associated plants and other Orthoptera.
In this chapter the cockroaches are grouped into those that have been found
in man-made structures and those that occur in other habitats. Certain
species may appear in several categories because they live both indoors and

out. The structural pests are divided into cockroaches that occur in land-
based structures, those on ships, and those in aircraft. The nonstructural
cockroaches are divided into those that occur in quite specific habitats
(caves, water, and deserts) and those that occur generally out of doors.
Nests of various arthropods serve as microhabitats of commensal
cockroaches; these latter associations are discussed on pages 310-318.
In this chapter our discussion is limited to the physical environment and
specific habitats of cockroaches, and only very general references are made
to associated organisms. The relationships of cockroaches to the biota are
examined in detail in subsequent chapters. To show the full extent of the
associations, the associates, from bacteroids to vertebrates, are arranged
phyletically. These associate-centered classifications serve admirably to
relate various species of cockroaches within common bounds, but fail to
give an integrated account of the total biotic relationships in the ecology of
each species. Although physically separated in this monograph, the many
associates of each species of cockroach should all be considered in
appraising the ecology of that species. To assist the reader to achieve this
end, we have included a checklist (p. 290) which serves as a convenient
index to certain organisms associated with particular species of
cockroaches.

CAVE HABITATS

Caves, mines, and animal burrows are somewhat similar habitats that
provide many species of cockroaches with shelter and frequently with food.
The microclimates of these cockroach habitats have not been described in
detail in the papers cited, but it seems rather obvious that natural caves,
man-made caves (mines), and burrows offer relatively stable temperatures
and humidities and protection from adverse climatic conditions. Although
such cavernicolous animals as birds and bats periodically leave caves to
search for food, cockroaches find the accumulated guano and animal and
plant detritus an entirely adequate dietary (Chopard, 1938). Cockroaches in
mines presumably subsist on the food and feces dropped by man and mine
animals (e.g., pit ponies). Food stored in their nests by burrowing animals is
undoubtedly utilized by the associated cockroaches.

Cavernicolous cockroaches show varying degrees of dependence on and
adaptation to these specialized habitats. Some of the common domiciliary
species (Blatta orientalis, Blattella germanica, and Periplaneta americana)
may have accompanied man into caves and remained there after he left
(Chopard, 1929a, 1936, 1938). Other species, from the paucity of records
noting their occurrence in caves, are undoubtedly accidental inhabitants that
may never become established. Besides these, however, many other species
of cockroaches have established large breeding colonies in caves. Although
some of the latter species show very pronounced morphological adaptations
to a cave life, many others resemble their noncavernicolous relatives. The
possible origin of cavernicolous Orthoptera has been discussed by Chopard
(1938).
Cavernicolous cockroaches have been segregated into four groups
according to their ability to adapt to their environment and the degree of
their specialized evolution (Chopard, 1936, 1938): (1) Trogloxenes:
Cockroaches that occur in caves in a sporadic fashion (the domiciliary
cockroaches and accidentals such as Ectobius and Heterogamodes). (2)
Troglophiles: Cockroaches found habitually in caves (Symploce,
Periplaneta cavernicola). (3) Guanobies: Cockroaches that live in the
guano of cavernicolous vertebrates (Gyna, Acanthogyna, Dyscologamia,
Pycnoscelus). (4) Troglobies: Cockroaches that apparently cannot live
outside of caves and which show very marked adaptive characters
(Alluaudellina, Nocticola, Spelaeoblatta, Typhloblatta). For complete
discussions of these groups including descriptions of the adaptive characters
shown by certain genera, the original sources should be consulted.
Although we know very little of the ethology of most of the cavernicolous
cockroaches, it is intriguing that three of the six known species of Nocticola
are cave dwellers, two are inhabitants of termite nests (p. 315), and one (N.
bolivari) was found under stones and cement blocks (Chopard, 1950b). In
the rather extensive list of cavernicolous cockroaches only two (Arenivaga
grata and Parcoblatta sp.) were taken from caves in North America north
of Yucatan. All other records are from Africa, Asia, Central America,
Europe, West Indies, East Indies, and the Philippine Islands. This we find
puzzling. Packard (1888) in his extensive study of the cave fauna of North
America listed no cockroaches. Dearolf (1941) found only the above-
mentioned Parcoblatta in one of 37 caves in Pennsylvania. Kohls and

Jellison (1948) listed no cockroaches among the arthropods from six bat
caves in Texas. We would expect Periplaneta americana to inhabit mines in
North America, but we have found no such records. Have cockroaches been
ignored in fauna collections from North American caves, or has our cave
fauna been less extensively studied than that of other parts of the world?
The two species of cockroaches found in mines (Blattella germanica and
Periplaneta americana) are also found in caves. For this reason we have
included them in the list headed Cavernicolous Cockroaches. On the other
hand, the cockroaches found in animal burrows are generally different
species from those found in caves, so we have grouped these together in a
second list.

CAVERNICOLOUS COCKROACHES

Alluaudellina cavernicola

Tanganyika.—From Kulumusi caves, near Tanga. The eyes of this
cockroach are reduced to a pair of slender streaks (Shelford, 1910a;
Chopard, 1932a).
East Africa.—Chopard (1936).

Apotrogia angolensis

Belgian Congo.—A troglophile without well-marked adaptive characters.
Collected in moist sand on floor of a sandstone grotto inhabited by bats
(Chopard, 1927, 1950a). Taken in many caves in Bas Congo (Leleup,
1956).

Apteroblatta perplexa

East Africa.—Accidental inhabitant of cave (Chopard, 1936).

Arenivaga grata

Arizona.—"A female and many nymphs were taken by Flock in the guano
in a bat cave in the Tucson Mountains" (Ball et al., 1942).

Aspiduchus borinquen

Puerto Rico.—In limestone cavern by thousands in grass and on walls
(Rehn and Hebard, 1927; Rehn, J. W. H., 1951a).

Aspiduchus cavernicola

Puerto Rico.—In limestone cave, in caves inhabited by bats, and apparently
seen in other caves well removed from entrance. "In this latter situation
great numbers were seen on the side walls and roof" (Rehn, J. W. H.,
1951a).

Blaberus atropos

Yucatan.—Found once, in Xmahit cave (Pearse, 1938).

Blaberus craniifer

Yucatan.—Collected within three caves, near the entrances (Pearse, 1938).

Blaberus giganteus

Panama.—Two males and several nymphs were taken under rocks in the
second chamber of the Chilibrillo cave; some also were on the walls
(Caudell, 1924).

Blatta lateralis

Turkmen S.S.R.—All stages, but more often females and nymphs, were
found in the middle and back part of Bakharden cavern, which was
inhabited by tens of thousands of bats (Vlasov, 1929).

Blatta orientalis

Turkmen S.S.R.—All stages found in front part of Bakharden bat cave. This
cave was uninhabited by man but supported a variety of other animals
(Vlasov, 1929).

Blattella germanica

South Africa.—Numerous in a gold mine on the Witwatersrand (Porter,
1930).
Tonkin.—Chopard (1929a); Colani (1952).

Byrsotria fumigata

Cuba.—Cueva de las Cucarachas, La Pantana, Baracoa, Oriente Province:
21 specimens, "It is evident ... that the species is also a cave inhabitant"
(Rehn and Hebard, 1927).

Deropeltis erythropeza

East Africa.—Found at entrance of cave; not a strictly cavernicolous form
according to Chopard (1936).

Ectobius pallidus

France.—Nymph in cave in Basses-Pyrénées, accidental inhabitant
(Chopard, 1936).

Ectobius vittiventer

Italy.—In detritus at base of entrance shaft of Acquaviva cave in the
Venezia Tridentina (Conci, 1951).

Ectobius sp.

Italy.—Found in the heap of saprophytic detritus at the base of the entrance
shaft in the Acquaviva cave (Conci, 1951).

Ergaula scarabaeoides

Sumatra.—West coast (Hebard, 1929).
Malaya.—Found burrowing in bat guano among stones at entrance to caves
in Selangor (Chopard, 1919, 1929).

Euthyrrhapha nigra

Madagascar.—Three males and six females in guano in Antsinomy grotto
(Chopard, 1949a).

Gyna kazungulana

East Africa.—This species is especially found in caves although it shows no
special adaptive characters. It is a typical guanobe (Chopard, 1936).

Gyna maculipennis

Belgian Congo.—Troglophile, guanophile. Found in two caves in Lualaba
(Leleup, 1956).

Gyna tristis

Belgian Congo.—In three caves in Uele (Leleup, 1956).

Heterogamodes krügeri

North Africa.—An accidental inhabitant of caves (Chopard, 1938).

Holocompsa zapoteca

Yucatan.—Common throughout rather dry, dusty caves in southern Yucatan
(Pearse, 1938).

Hoplosphoropyga babaulti

Stated to be a troglophile by Chopard (1938).

Nocticola caeca

Philippine Islands.—Bolívar (1892).

Nocticola decaryi

Madagascar.—A true troglobite according to Chopard (1945).

Nocticola simoni

Philippine Islands.—Bolívar (1892).

Parcoblatta sp.

Pennsylvania.—Found in Merkle cave, Berks County (Dearolf, 1941).

Periplaneta americana

IN CAVES

East Africa.—Its presence in the cave at Shimoni was thought to indicate
that man had sought refuge there and brought the cockroaches in with
baggage or provisions (Chopard, 1936).
India.—Many present in cave at Vengurla, the floor of which was covered
with bird guano (Abdulali, 1942).
Madagascar.—Thought to have been introduced into the cave entrance by
man (Chopard, 1945, 1949a).

IN MINES

Great Britain.—In a coal mine at Pontewydd where they had been
established for some years (Lucas, 1916). In the Pentre Pit mine where they
were abundant (Lucas, 1918). Abundant in a Welch mine 2,166 feet below
the surface (Lucas, 1925). This species was found quite commonly in a
number of South Wales coal mines; in one deep mine a white-eyed mutant
form comprised about 5 percent of the cockroach population for the
preceding 11 years (Jefferson, 1958).
India, western Bengal.—Very numerous in coal mines where the sole food
apparently was human faeces (Chandler, 1926).
South Africa.—Numerous in four deep-level gold mines on the
Witwatersrand.
Sumatra.—Numerous males and females from Sawah Lunto "'from a coal
mine where they lived in great numbers on the faeces of miners'" (Hanitsch,
1929).

Periplaneta australasiae

Sarawak.—Found swarming on walls of caves and in soft bird guano in
company with Symploce cavernicola (Moulton, 1912).
Tonkin.—Chopard (1929a); Colani (1952).

Periplaneta cavernicola

Malaya.—Taken on walls of inner caverns, where they were particularly
abundant (Chopard, 1919).

Periplaneta lata

Tonkin.—Chopard suggested that its presence in caves is probably linked
with man (Chopard, 1929a; Colani, 1952).

Periplaneta sp.

Malaya.—From a cave in Jalor (Annandale et al., 1913).

Perisphaerus sp.

Malaya.—The wingless females and nymphs mined in bats' guano in a
cavern of the Jalor caves (Annandale, 1900).

Polyphaga aegyptiaca

Turkmen S.S.R.—Females found in front part of Bakharden bat cave on
several occasions (Vlasov, 1929).
Turkey.—At Magharadjik and Arab Dede, found in caves with various other
animals (Lindberg, 1954).

Polyphaga sp.

Burma.—Hsin Dawng Cave, S. Shan States, 1 immature male under stone
in complete darkness (Chopard, 1924b).

Pycnoscelus niger

Tonkin.—Apparently not an accidental inhabitant as nymphs were present
(Chopard, 1929a; Colani, 1952).

Pycnoscelus striatus

Malaya.—Found burrowing in bats' guano at entrance to caves in Selangor,
where it was very abundant 50 to 600 feet from entrance; also on walls of
inner cavern (Chopard, 1919, 1929). In the absence of other evidence, the
presence of P. striatus in a cave indicates that bats also inhabit the cave
(Chopard, 1929a).

Pycnoscelus surinamensis

Assam.—Found 300 to 400 feet from entrance of Siju cave in the Garo Hills
(Chopard, 1924b).
South Celebes.—Hanitsch (1932).

Spelaeoblatta gestroi

Burma.—Chopard stated that this species shows marked characteristics of
adaptation to a life in darkness (Bolívar, 1897; Annandale, 1913; Chopard,
1919).

Symploce breviramis

South Celebes.—Hanitsch (1932).

Symploce cavernicola

Sarawak, Borneo.—Swarming on walls of caves and in soft bird guano on
the cave floor (Moulton, 1912). Hanitsch (1931) noted that this species was
first recorded by Shelford from a cave in Sarawak and that there is a series
from a cave in the Oxford University Museum, taken by Banks in 1928.
Malaya.—On the walls of the inner cavern of a cave at Biserat; the insects
covered the walls in places (Chopard, 1919).
Sumatra.—From Baso cavern, on the west coast (Hebard, 1929).

Symploce curta

South Celebes.—Hanitsch (1932).

Symploce remyi

Tonkin.—This seems to be a true cavernicolous species (Chopard, 1929a;
Colani, 1952).

Tivia macracantha

Belgian Congo.—A troglophile without well-marked adaptive characters
(Chopard, 1950a). At Haut-Katanga, troglophile and guanophile (Leleup,
1956).

Tivia sp.

Madagascar.—Last-stage nymphs captured in guano in Antsinomy grotto
(Chopard, 1949a).

Typhloblatta caeca

India, Assam.—An eyeless species with noticeably elongated appendages
(Chopard, 1945).

Typhloblattodes madecassus

Madagascar.—Unpigmented integument and reduced eyes (Chopard,
1945).

Xestoblatta immaculata

Panama.—Found under rocks on guano-covered floor of the Chilibrillo bat
caves (Caudell, 1924).

Unidentified cockroaches

Malaya.—The walls of a cave were covered by dense groups of a species of
"Blatta" (Annandale, 1900).
England.—"The chief insect pests of the mines are cockroaches, which
often swarm in hot mines and those with pit pony stables...." (Hardy, 1941).

COCKROACHES FROM THE BURROWS OF
VERTEBRATES

Arenivaga apacha

Arizona.—In the nests of wood rats, Neotoma sp. (Hebard, 1917).

Arenivaga bolliana

Texas.—In the nests of wood rats, Neotoma sp. (Hebard, 1917; 1943a).

Arenivaga erratica

Arizona.—The wingless females were commonly found in burrows of
Dipodomys spectabilis spectabilis Merriam, the kangaroo rat. The winged
males were never found in the burrows (Vorhies and Taylor, 1922). Found
most commonly in wood-rat and ground-squirrel dens in the desert regions
(Ball et al., 1942).

Arenivaga floridensis

Florida.—Found in a burrow of Peromyscus polionotus rhoadsi (Bangs),
the white-footed mouse (Young, 1949).

Arenivaga roseni

Turkmen S.S.R.—Occasionally found in burrows of Rhombomys opimus
Lichtenstein; in the burrows of the desert turtle, Testudo horsfieldi Gray;
and frequently in burrows of the ground squirrel, Spermophilopsis
leptodactylus Lichtenstein (Vlasov, 1933; Vlasov and Miram, 1937).

Arenivaga tonkawa

Texas.—An immature specimen was found in a prairie-dog hole (Hebard,
1943a).

Cariblatta lutea

Florida.—It has been taken in burrows of the pocket gopher, Geomys sp.
(Hubbell and Goff, 1940).

Euthlastoblatta abortiva

Texas.—In the nests of wood rats, Neotoma sp. (Hebard, 1917).

Parcoblatta fulvescens

Texas.—In the nests of wood rats, Neotoma sp. (Hebard, 1917).

Polyphaga aegyptiaca

Turkmen S.S.R.—Nymphs and adult females were often found in burrows of
the sand mouse, Rhombomys opimus (Vlasov, 1933).

Polyphaga indica

Turkmen S.S.R.—This species prefers sandy soils where it can be found in
burrows of Spermophilopsis leptodactylus and Pallasiomys meridionalis
pennicilliger Heptner (Vlasov and Miram, 1937).

Polyphaga saussurei

Tadzhikistan.—Found in burrows of turtles and rodents (Zmeev, 1936).
Turkmen S.S.R.—Nymphs and adult females are common in burrows of
Rhombomys opimus and in burrows of Testudo horsfieldi. Its principal
habitat is rodent burrows in loess dust, where it is not infrequently found in
the food stores of the host (Vlasov and Miram, 1937).

Pycnoscelus surinamensis

Texas.—In the nests of wood rats, Neotoma sp. (Hebard, 1917).

DESERT HABITATS

There is relatively little ecological information about cockroaches that live
in deserts, even though certain species, notably Polyphaga aegyptiaca, have
long been known to inhabit arid zones. In fact, so little is known about the
ecology of arid-zone insects in general that it is more a subject for research
than for review (Pradhan, 1957). In their account of the cockroaches of
Northern Kenya and Jubaland, Kevan and Chopard (1954) describe in some
detail the vegetational areas of this arid desert or semidesert country, which
averages only about 10 inches of rain per year. The other sources that are
cited below contain very little more biological information than the
abstracted material that is given under each species.
Nearly all the Polyphaginae are said to be marked xerophiles whose
distribution coincides with that of the deserts (Bei-Bienko, 1950). With the
exception of Arenivaga floridana, the species of Polyphaginae in the United
States all occur in the Southwest, where they are (with a few exceptions)
the only cockroaches that inhabit the desert regions proper (Hebard, 1917).
The Polyphaginae reach their greatest diversity in the deserts of Northern
Africa and Anterior and South-Central Asia (Bei-Bienko, 1950). Some of
the desert-inhabiting species have also been found under nondesert
conditions. This only further exemplifies the plasticity of cockroaches in
adapting to different environments.
The ability of desert insects to live under what appear to be extremely
unfavorable conditions has been abundantly illustrated by Pradhan (1957).
Uvarov (1954) has pointed out that a desert "covers a great variety of
landscapes, which provide desert animals with a wide range of habitats,
some of them offering very favorable conditions for life." Pradhan (1957)
stated that many desert animals avoid the extremes of desert climates by
choosing suitable microclimates for diurnal resting places, that a permanent
or temporary underground existence is very common among insects in arid
zones, and that many nocturnal Orthoptera burrow into the soil or hide
under stones where temperatures are lower. For example, the type of
Parcoblatta desertae was found under a boulder on the bare desert (Rehn
and Hebard, 1909).
Symbiosis with burrowing animals is another solution to the problem of
existence in the desert; in fact, symbiosis is a mode of life adopted by

nearly half of the desert cockroaches about which we have any information.
Vlasov and Miram (1937) found Polyphaga indica, Polyphaga saussurei,
and Arenivaga roseni in the burrows of rodents and desert turtles. In the
desert regions of Arizona, females of Arenivaga erratica were found
commonly in burrows of the kangaroo rat (Vorhies and Taylor, 1922) and in
dens of wood rats and ground squirrels (Ball et al., 1942). Arenivaga
apacha and Arenivaga bolliana have also been found inhabiting the nests of
wood rats (Hebard, 1917; 1943a). Bei-Bienko (1950) has suggested that the
adaptation of desert-inhabiting cockroaches to rodent burrows might enable
these insects to survive in the severe climatic conditions of deserts in
summer.
Under desert conditions in southern Arizona, the relative humidity outside
of the burrows of the kangaroo rat is 1 to 15 percent during the day and 15
to 40 percent at night; but inside the burrows the relative humidity is 30 to
50 percent, and the temperature, even during the day, is below 30° C.
(Schmidt-Nielsen, 1949). Thus by living in rodent burrows during the day
and going outside at night, the desert cockroaches could avail themselves of
the most favorable microclimates obtainable. Presumably whatever food
these insects eat provides them with sufficient water to enable them to
survive under desert conditions. Bodenheimer (1953) has suggested that the
extent of utilization of dew, which is sometimes heavy in the desert, should
be investigated; he stated that tenebrionid beetles have been seen in the
early morning eating dry [dead?] herbs that were still wet with dew. It is
obvious that there is a need for additional detailed information without
which we can only guess about the ecology of desert cockroaches.
In the following list we have cited only those species that were stated to
have been found under desert conditions. Undoubtedly, related species that
have been taken in similar localities are also desert-inhabiting forms, as, for
example, other species of Arenivaga that were collected in Texas by Hebard
(1943a). In the absence of specific information linking such other species
with deserts, we have arbitrarily relegated those forms to the section on
outdoor habitats. In addition to the species listed below, desert cockroaches
are said to be found in the following genera: Anisogamia,
Mononychoblatta, and Nymphytria (Chopard, 1938).

DESERT COCKROACHES

Agis orientalis

Northern Kenya.—In desert-grass and thorn-bush country; scattered, dry
tufts of grasses interspersed among acacia bush and scattered trees (Kevan
and Chopard, 1954).

Arenivaga apacha

U.S.A.—Inhabits desert regions of the Southwest, has been found in nests of
wood rats (Hebard, 1917).

Arenivaga bolliana

U.S.A.—On gravelly hillocks, in scattered scrub, and in the nests of wood
rats in Texas. It is a desert inhabitant in the Southwest (Hebard, 1917;
1943a).

Arenivaga erratica

U.S.A.—Inhabits desert regions of the Southwest (Hebard, 1917). In
Arizona it has been found in rodent burrows in the desert (Vorhies and
Taylor, 1922; Ball et al., 1942).

Arenivaga roseni

Turkmen S.S.R.—Predominantly found in burrows in sand; all stages
"swim" in sand and loess dust (Vlasov and Miram, 1937).

Blattella vaga

Arizona.—Found in small numbers on the dry desert (Flock, 1941a).

Compsodes schwarzi

U.S.A.—Occurs in the Southwest where it is confined to the desert and
semidesert mountainous areas, rarely being found on the desert floor
(Hebard, 1917). Taken in an ant nest in mountains of Arizona (Ball et al.,
1942).

Cyrtotria capucina

Eastern Africa.—"Commonly met with under débris, the apterous females
being most frequent." Thorn-bush country (Kevan and Chopard, 1954).

Derocalymma lampyrina

Northern Kenya.—Very abundant; both sexes under débris in desert-grass
and thorn-bush country (Kevan and Chopard, 1954).

Derocalymma porcellio

Northern Kenya.—Taken in upland grassland and bush (Kevan and
Chopard, 1954).

Deropeltis autraniana

Northern Kenya.—In thorn-bush country (Kevan and Chopard, 1954).

Deropeltis melanophila

Northern Kenya.—"Very commonly found at the base of tufts of grass and
other débris, the apterous female particularly in the latter situation"; in
upland grassland near forest; in thorn-bush country (Kevan and Chopard,
1954).

Deropeltis nigrita

Northern Kenya.—Taken in upland grassland and bush (Kevan and
Chopard, 1954).

Eremoblatta subdiaphana

U.S.A.—Apparently found in greatest abundance in the extreme desert
conditions of the southwestern United States (Hebard, 1917). Two small
groups of males were observed in the midst of the sandy desert north of
Yuma, Ariz.; these insects alternately flew and ran over the sand in the hot
sun while headed in a southwesterly direction (Wheeler, 1911).

Euandroblatta palpalis

Northern Kenya.—In desert-grass and thorn-bush country (Kevan and
Chopard, 1954).

Heterogamodes rugosa

Northern Kenya.—"All from desert grass and thorn bush (on sand)." It was
stated (under discussion of Tivia fulva) that Heterogamodes females live
more or less buried in the sand (Kevan and Chopard, 1954).

Namablatta bitaeniata

Southwestern Africa.—Limited in distribution to the more arid portions,
being peculiar to extreme desert conditions (Rehn, 1937).

Nauphoeta punctipennis

Northern Kenya.—In desert grass and thorn bush; "probably the commonest
of all the medium-sized cockroaches occurring in the area under discussion,
coming very freely to light" (Kevan and Chopard, 1954).

Parcoblatta desertae

U.S.A.—In the desert and semidesert mountainous areas of the Southwest; it
is rarely found on the desert floor (Hebard, 1917). Found under boulder on
bare desert (Rehn and Hebard, 1909).

Polyphaga aegyptiaca

Caucasus.—The wingless female was found buried in sand and dust (Burr,
1913).
Turkmen S.S.R.—Although this species is secondarily encountered in
dwellings and courtyards, it is a very characteristic insect of the Trans-
Caspian deserts; the females are encountered fairly frequently as inhabitants
of sand, where they run slowly over the surface, or dig themselves into the
sand to continue their forward motion not far below the surface (Fausek,
1906). Uvarov (in Chopard, 1929b) indicated that females of this genus are
found in various desert localities, particularly where vegetative debris
occurs, but they are not strictly attached to sandy terrain.

Polyphaga indica

Turkmen S.S.R.—This species prefers sandy soils where the nymphs, alate
males, and wingless females "swim" readily through the sand; they can also
be found in the burrows of desert animals (Vlasov and Miram, 1937).

Polyphaga saussurei

Turkmen S.S.R.—Its principal habitats are rodent burrows in loess dust and
burrows of the desert turtle (Vlasov and Miram, 1937).

Supella hottentotta

Northern Kenya.—Taken in bushes by dry river bed and in desert-grass and
thorn-bush country at several stations (Kevan and Chopard, 1954). " ...
taken with light at night running on bark of a large acacia tree" (Rehn,
1947).

Symploce kevani

Northern Kenya.—In desert grass and thorn-bush country (Kevan and
Chopard, 1954).

Theganopteryx straminea

Northern Kenya.—Taken at three stations in desert grass and thorn bush
(Kevan and Chopard, 1954).

Tivia brunnea

Northern Kenya.—In open sandy, riverine bush (scanty ground cover
among acacia trees and doum palms) (Kevan and Chopard, 1954).

Tivia fulva

Northern Kenya.—In desert grass and thorn bush; distributed in semidesert
areas south of Sahara; the apterous females probably live buried in sand
(Kevan and Chopard, 1954).

Tivia obscura

Northern Kenya.—In desert grass and thorn bush (Kevan and Chopard,
1954).

AQUATIC HABITATS

The so-called aquatic or amphibious cockroaches are all members of the
subfamily Epilamprinae (Chopard, 1938). These forms are not nearly as
aquatic as water beetles or aquatic Hemiptera, but in their relations to water
they behave differently from nonamphibious cockroaches, which tend to
avoid water except for drinking. There are apparently no special
morphological characteristics that distinguish amphibious cockroaches
(Shelford, 1907, 1909a; Chopard, 1938), although Takahashi (1926) listed
several characters that he considered made Opisthoplatia maculata adapted
for an aquatic life: (1) Back of body easily wetted; (2) long hairs on
underside of thorax trap air; (3) terminal abdominal spiracles open into
tubes that extend rearward; (4) long hairs on ventral surfaces of cerci

"protect" terminal abdominal spiracles. Annandale (1906) also suggested
that the position of the posterior abdominal spiracles, at the base of tubes
that project rearward from beneath the seventh tergite, are an adaptation to
an aquatic life. However, as Shelford (1907) and Chopard (1938) have
pointed out, this same feature may be observed in many terrestrial
cockroaches. The legs of amphibious cockroaches are similar to those of
nonaquatic species and are not modified for swimming (Shelford, 1909a;
Takahashi, 1926).
Biological observations have been made on relatively few species, but
representatives of at least six genera occur in quasi-aquatic habitats. Strictly
speaking, these cockroaches live on land at the edges of streams or pools
and spend relatively brief periods in the water. A few species are found in
water-filled bromeliads. The behavior of the known amphibious species of
cockroaches in relation to their habitats is discussed below.

AMPHIBIOUS COCKROACHES

Audreia bromeliadarum

Panama.—These insects when disturbed would dive into the water that had
collected in the base of the bromeliad; they would disappear beneath the
surface and remain submerged for some considerable time (Caudell, 1914).

Dryadoblatta scotti

Trinidad.—This species was taken from the leaf bases of Tillandsia sp. at
3,100 feet; water had collected between the leaves and the insect was
presumed to be more or less amphibious (Scott, 1912). Subaquatic in the
bromeliad Glomeropitcairnia erectiflora: "This large and handsome species
[D. scotti] is very common in the larger, water-filled, epiphytic bromeliads
of the rain forest. Within these plants it is usually to be found, often in
considerable numbers, just above the surface of the water or partly
immersed in it. The cockroaches will descend rapidly into the water when
alarmed and probably obtain their nourishment from the accumulated

organic matter in the water. Floating material is probably taken and it seems
less likely that they feed below the surface. They appear to be
ovoviviparous." (Princis and Kevan, 1955.)

Epilampra abdomen-nigrum

Puerto Rico.—Abundant in wet "malojillo" meadows. The nymphs swim
easily and remain under water for long periods, as do the adults (Seín, 1923;
Wolcott, 1950).
Panama.—A swimming nymph, captured in a dipper with mosquito larvae
in a lagoon of the Rio Chilibre, was kept under observations in an
aquarium. If disturbed, the insect dived into the water from floating
vegetation and swam rapidly below the surface for a minute or two. Finally
becoming quiescent, the cockroach would then cling to submerged roots;
twice it remained still for 15 minutes before climbing to the surface, where
it remained for five or more minutes before emerging completely (Crowell,
1946).

Epilampra annandalei

Lower Burma.—One male and three nymphs were collected in the Dawna
Hills by Annandale who made the following observations: "The wingless
specimens were under stones in a jungle stream and behaved just as the one
I obtained in Chota, Nagpur, did [Annandale, 1906]. The winged specimen
was under a stone at the edge of the stream, but swam readily. It did not
seem so much at home in the water, however, and apparently could not,
owing to the wings, raise the tip of its abdomen above the surface."
(Shelford, 1909a.)

Epilampra sp.

Siamese Malay States.—Wingless females rested on floating logs from
which they would dive into the water upon the least disturbance; they
remained under water for several minutes, then surfaced beneath the shelter
of the log. In the jungle all females were taken either in the water or among

matted roots on the sides of the stream. Winged males were seen rising
from the surface of the water (Annandale, 1900).
Sarawak.—All specimens were immature; they swam and dived well, but
were soon drowned if prevented from rising to the surface to breathe.
"When at rest the body of the cockroach is almost entirely submerged, the
tip of the abdomen alone projecting above the surface of the water; the
abdomen moves gently up and down and every 30-40 seconds a bubble of
air issues from the prothoracic spiracle on each side." (Shelford, 1901,
1916.)
India.—A nymphal female, found in a jungle stream at Chota Nagpur, could
swim with belly or back upward. When held under water it drowned in a
few minutes. The tip of the abdomen was held out of water (Annandale,
1906).
Shelford (1907) has suggested that the immature stages of terrestrial species
of Epilampra may well be amphibious. This is an area that could profit by
more field observations.

Opisthoplatia maculata

Formosa.—Invariably found under or between rocks near mountain
streams. The wingless adult and the nymph have similar habits. Normally
the cockroach lives on land, and when it goes into the water it returns to
land within a few minutes. This cockroach rarely swims, but when it does, it
maintains its body in a horizontal position just below the surface of the
water. Ordinarily, it walks on the river bottom or on water-covered rocks.
This insect feeds on decayed leaves and, according to Shikano, it will eat
human feces. (Takahashi, 1926.)
This species has a large number of long hydrophobic hairs on the ventral
sides of the thorax and anterior abdominal segments. When the insect
submerges, air is trapped in these hairs. The thoracic and one pair of
abdominal spiracles open into the bubble of trapped air. However, the insect
apparently does not use this plastron of air to replenish its tracheal air
supply, but, like Rhicnoda natatrix (see below), it inspires air while at the
surface through its posterior abdominal spiracles and expires air into the
bubble under the thorax. While the insect is submerged, the air bubble

increases in volume until part of it breaks away and floats to the surface.
(Takahashi, 1926.)

Opisthoplatia orientalis

Formosa.—Lives on or in swampy ground (Takahashi, 1924).

Rhicnoda natatrix

Sarawak.—Immature cockroaches were found in sodden leaves at the edge
of a pool, where they rested for hours at a time. Generally the fore part of
the body was in the water but the tip of the abdomen was always in air.
When disturbed the insects dived into the water and hid under sticks and
stones on the bottom. Air is inspired through the posterior abdominal
spiracles, when they projected above the water surface, and expired through
the thoracic spiracles. In experiments in which the insects' abdomens were
held immersed in water, with the thorax exposed, the insects died in 6 to 12
hours or less. (Shelford, 1907.)

Stictolampra buqueti concinula

Westsumba.—Found under moist fallen leaves on gravelly shore of Melolo
River. The nymphs distinguished themselves through their amphibious
mode of life and were often good swimmers (Princis, 1957a).

Unidentified epilamprines

Brazil.—These cockroaches were found under stones at the side of a rocky
stream at Ouro Preto. When disturbed they ran down under the surface of
the water and hid under stones at the bottom. When thrown on the water
surface, they were helpless, and to get beneath the water surface they had to
walk down some object. When they had penetrated the surface film they
could swim freely. Specimens kept in jars lived several days with only a
portion of their abdomens exposed to the air. (Bristowe, 1925.)

OUTDOOR HABITATS

This category is a catchall for all cockroaches that are not limited to the
more circumscribed habitats that have been previously considered. Some
cockroaches in this section select specific microhabitats (e.g., Cryptocercus
spp., which live exclusively in rotten logs; and Neoblattella dryas, N.
eurydice, and N. grossbecki in bromeliads). Others are found in a wide
variety of habitats (e.g., Ischnoptera deropeltiformis and Parcoblatta spp.).
But some species are so little known that their actual habitats are barely
suggested in the collection data.
Williams (1941) made an ecological study of the floor fauna of the Panama
rain forest. He found Orthoptera (nearly all were unidentified nymphal
cockroaches) in the litter of dead leaves, twigs, and other plant products in
over 90 percent of the quadrats he examined. These insects represented
about 0.25 percent of the total animal population.
Delamare Deboutteville (1948) made a quantitative study of the animal
population in suspended soil that had accumulated between the roots of
forest epiphytes of the lower Ivory Coast. He analyzed 2 dm.3 samples of
soil from an epiphyte located 45 meters above ground on a main branch of
Parinarium, with these results: Horizon A.—Superficial zone of large
rootlets, 6 cm. deep: 2 cockroaches, 4 arachnids, and 4 beetles. Horizon B.
—Zone of fine rootlets, 6 cm. deep: 6 cockroaches and numerous other
arthropods. Horizon C.—Humid zone, 8 cm. deep: 7 cockroaches and
numerous other arthropods. Plants, such as Palissota, were also living in
this very original biotype.
The species of cockroaches listed below have been found in the following
kinds of outdoor microhabitats: In jungle, forest, and woodlands they have
been found in rotten wood; under bark of living, dead, and fallen trees; in
decay cavities in trees; burrowing in living bark; on foliage of trees, shrubs,
bushes, and low herbage; on vines and in bromeliads and epiphytic ferns;
under signs on trees and stumps; in piles of logs and firewood; under dead
leaves and debris; in and under decaying fruit on the ground. Cockroaches
have been found between the leaves and under leaf sheaths of sugarcane,
corn, and other grasses; under dry fibers and fronds of coconut trees; in
hollow stems and bases of tree-fern fronds; under bracts of banana

blossoms and in bunches of bananas (p. 146). Cockroaches also inhabit
abandoned cocoons and larval tents, wasp nests, ant nests, termite nests,
bird nests, rat nests, and burrows of other rodents (pp. 23-25, 310-319).
Cockroaches have been found in rock crevices and under rocks; under
boards and other objects on ground; under seaweed, drift, and other debris
on beaches; burrowing in soil and under clods of earth; in marshes and
swamps; in dumps and rubbish heaps.
The above list does not exhaust the available outdoor microhabitats that
cockroaches find suitable for their continued existence, but it is fairly
representative. Although we have no measurements to substantiate this
conclusion, we suggest that the microhabitats cited above have a more
constant temperature and a relatively higher humidity than is provided by
the surrounding macrohabitats. We would expect insects such as
cockroaches, whose water balance is dependent on a continuous supply of
fluid water or moist food, to seek moist environments or to avoid situations
in which their transpiration might increase. Deviations, presumably brief,
from this expected behavior must occur to account for the cockroaches that
are found under relatively unfavorable environmental conditions. Despite
the apparent preference for cryptic habitats, some cockroaches are found in
hot sunlight (Ellipsidion spp.; Tepper, 1893); Rehn (1945) has stated that
many kinds are diurnal rather than nocturnal. Movement of cockroaches
between habitats may be assumed to occur; but movement from an
unfavorable environment to a more favorable one, following a shift in water
balance, has not been observed in nature; however, laboratory experiments
suggest that the mechanism for mediating such behavior is present in some
species of cockroaches (Gunn and Cosway, 1938; Roth and Willis, 1952a).
Obviously, additional research is needed on the bionomics of all species.
Further conclusions based on current limited knowledge can only be
speculative and possibly misleading.

COCKROACHES FROM OUTDOOR HABITATS

(Except Amphibious, Desert, and Cavernicolous Forms)

Aglaopteryx absimilis

Puerto Rico.—Living in rotten, wooden fence; living between leaves of
Samanea saman and in abandoned cocoons of Megalopyge krugii on bucare
trees (Wolcott, 1950).
Leeward Islands.—On coconut tree (Princis and Kevan, 1955).

Aglaopteryx facies

Puerto Rico.—As diaphana, in dead branch 10 feet above the ground on
Mona Island (Hebard, 1917). In trunks of trees under bark and very often in
abandoned cocoons of the "plumilla" (Seín, 1923). On rotten, wooden
fence; in empty cocoons of Megalopyge krugii on trunks of bucare trees,
Erythrina glauca; on trunk of Inga laurina; in larval tents of Tetralopha
scabridella on Inga vera (Wolcott, 1936). In large numbers in nests of the
gray kingbird (Wolcott, 1950).

Aglaopteryx diaphana

West Indies.—In Cuba, under corky bark of large tree in open; Jamaica,
under loose bark of shade trees and in bracts of banana blossoms; in
bromeliads and hollow bases of dead tree-fern fronds (Rehn and Hebard,
1927).

Aglaopteryx gemma

Florida.—On Long Key, under coquina boulder in heavy scrub; under
loose, dry fibers near head of standing coconut palm (Rehn and Hebard,
1912). Climbing on roots of red mangrove, Rhizophora mangle, in swamp;
under loose bark on trunk of Exothea paniculata in dense jungle; under
limestone boulder in keys scrub; under signs on oaks, sweet gum, and
longleaf pines in southeastern and southern States (Hebard, 1917).
Infrequent in the shrub growth of the Sandhills habitat (Friauf, 1953).
Texas.—In undergrowth of pine forest; under sign on oak near river; in
Tillandsia sp. (Hebard, 1917). Usually in hiding places on trees; only once

found under a stone on ground (Hebard, 1943a).

Allacta similis

Hawaii.—Common in hollow stems and under bark (Swezey and Williams,
1932).

Amazonina emarginata

Trinidad.—On low herbage, on hibiscus at night, and in banana bunch
(Princis and Kevan, 1955).

Anaplecta asema

Panama.—Under dead leaves in jungle (Hebard, 1920).

Anaplecta decipiens

Costa Rica.—In decayed leaves (Rehn, 1906).

Anaplecta fallax

Costa Rica.—Under stones on borders of Surubres River (Rehn, 1906).

Anaplecta hemiscotia

Panama.—Under rubbish at edge of jungle and in overgrowth of heavy
vines on low bushes (Hebard, 1920).

Anaplecta lateralis

Panama.—Under drift on edge of coral-sand beach (Hebard, 1920).

Arenivaga bolliana

Texas.—In dense jungle brush of the river plain; on gravelly hillocks in
scattered scrub; under debris and leaf mold under mesquite trees; in rat's
nests, Neotoma sp. (Hebard, 1917). In dry earth under bush; inhabits litter
on ground and nests of rats (Hebard, 1943a).

Arenivaga floridensis

Florida.—Male on ground under leaves of cabbage palmetto (Blatchley,
1920). Females in sand under boards and debris along lake shore (Friauf in
Cantrall, 1941). Infrequent on bare soil and ground under vegetation in the
longleaf-pine flatwoods habitat (Friauf, 1953). In rodent burrow (Young,
1949).

Arenivaga grata

Texas.—Under stones in upper canyon; under rocks in pine-oak forest; from
oak-manzanita forest along dry stream bed (Hebard, 1943a).

Aristiger histrio

Malaya.—Lives freely on bushes and flowers of Passiflora sp. (Karny,
1924).

Aspiduchus boriquen

Puerto Rico.—"Apparently the species [as deplanatus] is locally numerous
in suitable locations, such as caves, rock crevices and the shelter of large
stones." (Rehn and Hebard, 1927).

Audreia bromeliadarum

Panama.—Perfectly at home in bromeliads (see p. 31) (Caudell, 1914).

Audreia jamaicana

Jamaica.—In bromeliads; under dead wood in dense forest (Rehn and
Hebard, 1927).

Balta godeffroyi

Australia.—Under bark (Hebard, 1943).

Balta quadricaudata

Australia.—From sugarcane (Hebard, 1943).

Balta scripta

Australia, Queensland.—On leaves, grass, and sugarcane (Hebard, 1943).

Balta torresiana

Australia.—From leaves, under bark, from sugarcane (Hebard, 1943).

Balta verticalis

Australia.—In leaves, from tree, from sugarcane (Hebard, 1943).

Blaberus atropus

Trinidad.—Female in rotting log (Princis and Kevan, 1955).

Blaberus discoidalis

Jamaica.—Under dead coconut petioles in open spot. Gundlach found it
under stones in a field in Cuba (Rehn and Hebard, 1927).

Blaberus giganteus

Trinidad.—Nymph in rotten palm tree (Princis and Kevan, 1955).

Blaberus spp.

Venezuela.—Only taken in the forests of the Orinoco near the trunks of
rotten trees at night (Doumerc in Blanchard, 1837).
Panama.—Among dead leaves and debris on floor of rain forest (E. C.
Williams, Jr., 1941).

Blatta lateralis

U.S.S.R.—Found among rocks at 2,000 or more meters elevation. It is
found in cultivated areas as well as in mountainous landscapes and in
semideserts (Bei-Bienko, 1950).

Blatta orientalis

Great Britain.—One female nymph under bark of tree 10 feet above the
ground (Burr, 1900). Swarming within a rubbish heap in February (Lucas,
1912). In refuse tip under old sacks and sheets of linoleum (Hallett in
Lucas, 1922). Male under bark of oak far from houses (Donisthorpe, 1918).
One adult female and nymph in prone dead elm 50 yards from house (Burr,
1937). An immature male at the roots of Ballota nigra (Buck in Gardner,
1954). Four additional records of this species outdoors away from houses
(Lucas, 1920).
Southern Crimea.—Under stones, dead leaves, and detritus in small copses
of Quercus pubescens, Carpinus orientalis, Cornus mas, Paliurus
aculeatus, and Dictamnus fraxinella; 19 specimens, apparently breeding
outdoors (Adelung, 1907).
North-central U.S.—Observations since 1950 indicate a marked increase in
frequency and duration of infestations outdoors; observed in bare soil,
vegetation, debris, alongside foundations in sodded areas, along sidewalks,
and at edge of parking areas throughout the year; in some urban residential
areas, the yards of whole blocks of houses were "alive" with this species on
warm summer nights; in winter they have been found under stones, leaf
debris, and soil near structures (Shuyler, 1956).

Blattella germanica

Algeria.—Under moist leaves in woods (Lucas, 1849).
California.—Under rubbish and on date palms (Herms, 1926).
Connecticut.—In city dump under loose material, very numerous (Walden,
1922). Additional infestations of dumps by this species have been reported
in New York (Felt, 1926, 1928) and New Jersey (Hansens, 1949, 1950)
England.—Swarming within a rubbish heap in February (Lucas, 1912).
Formosa.—Lives among fallen leaves on the ground (Takahashi, 1924).
North-central U.S.—Reported living outdoors near buildings and in soil
under basementless buildings from early summer to late fall (Shuyler,
1956).

Blattella humbertiana

India.—Common among decaying vegetation and on trees (Chopard and
Chatterjee, 1937).
Formosa.—Normally found in sugarcane fields, pineapple fields, and
grasslands where it feeds on decayed leaves and other decayed vegetable
matter and dead insects. It lies concealed among and under fallen leaves and
clods of earth on or close to ground and never on the upper parts of plants,
except pineapple where it is found among the leaves (Takahashi, 1940).

Blattella vaga

Arizona.—Typically an inhabitant of irrigated fields and yards, it is found in
fewer numbers on the dry desert. It is found under stones, plant debris, and
clumps of earth; found in greatest numbers around decaying dates on
ground (Flock, 1941a).
Texas.—Beneath duff under athel trees; rather abundant in clumps of
Rhodes grass (Riherd, 1953).

Byrsotria cabrerae

Cuba.—In sea-coast woods: "The species [this and Byrsotria fumigata] are
ground-dwelling, hiding under stones and other shelter" (Rehn and Hebard,
1927).

Byrsotria fumigata

Cuba.—Ground dwelling, hiding under stones, etc.; also a cave inhabitant
(Rehn and Hebard, 1927).

Cahita borero

Brazil, Matto Grosso.—Beaten from tree foliage in dry scrub, from tree
foliage at edge of dry riverine tangle, and from undergrowth in a dry forest
area (Rehn, 1937a).

Cahita nahua

Honduras.—All beaten from foliage along roads or in thickets, during rainy
season (Rehn, 1937a).

Cariblatta antiguensis

Virgin Islands, St. Croix.—Common under heaps of rubbish (Beatty, 1944).
Trinidad.—On herbage below bananas; all stages on Hibiscus at night; in
grass at dusk; on low herbage under old coconut (Princis and Kevan, 1955).

Cariblatta cuprea

Jamaica.—In leaves on leaf mold in hillside forest (Hebard, 1916a).

Cariblatta delicatula

West Indies.—In debris in short grass in open, Cuba. Under dead petioles of
coconut palms, San Domingo. In leaves on leaf mold in hillside forest,
Jamaica (Hebard, 1916a).

Cariblatta hylaea

Honduras.—Found at foot and on lower slopes of first ridges of the Sierra
Pija, from 75 to at least 800 feet above sea level, where vegetation ranged
from abandoned banana patches overgrown with Heliconia and Cecropia
and interspersed with forest trees, at the foot of the hills, to primeval
lowland forest (ceibas, figs, palms, etc.) on the slopes. In the banana
patches C. hylaea was found on hanging dead banana and Cecropia leaves;
on the slopes it was found on undergrowth foliage, hanging dead leaves,
and in dead leaves on ground (Rehn, 1945a).

Cariblatta imitans

Panama.—Among loose leaves on leaf mold in heavy jungle (Hebard,
1916a).

Cariblatta insularis

Jamaica.—One of the most frequently encountered orthopterous insects in
bromeliads on trees (Hebard, 1916a, 1917; Rehn and Hebard, 1927).

Cariblatta jamaicensis

Jamaica.—In decaying herbage (Rehn and Hebard, 1927).

Cariblatta landalei

Jamaica.—All specimens taken from under drying bracts of banana
blossoms (Rehn and Hebard, 1927).

Cariblatta lutea lutea

North Carolina.—Under pine straw on ground in woods (Brimley, 1908).
Southeastern U.S.—Under dead oak leaves; under dead needles in longleaf-
pine woods; in wire grass; under refuse; beaten from undergrowth in pine

and oak woods (Rehn and Hebard, 1916). In undergrowth of shortleaf-pine,
longleaf-pine, and oak woods; in heavy scrub in damp spot of sand dune
area; from high bushes, Ilex coriacea [= lucida] along inland swampy area
(Hebard, 1916a). "The species is in large part terrestrial, being usually
found among dead leaves and litter on the ground. Occasional specimens
are, however, sometimes beaten from bushes. Individuals are decidedly
active and are usually to be found in the greatest numbers in sandy
situations" (Hebard, 1917).
Florida.—Throughout winter and spring they are frequent beneath leaves
and other debris on ground, especially in dry, sandy locations (Blatchley,
1920). Friauf (1953) found this species under debris, fallen leaves, leaf
mold, or decaying wood in these habitats: Dry, ruderal grassland
(infrequent), scrub (frequent), sandhills (dominant), xeric hammock
(infrequent), mesic hammock (dominant), pond margin (infrequent),
longleaf-pine flatwoods (frequent), bayhead (occasional), low hammock
(frequent), and alluvial hammock (occasional). In the shrub stratum in these
habitats: Scrub (frequent), sandhills (dominant), and xeric hammock
(infrequent). In herbaceous stratum in these habitats: Sandhills (dominant),
mesic hammock (dominant), and black-pine flatwoods (infrequent). On
bare soil or bare sand under vegetation in these habitats: Sandhills
(dominant), pond margin (infrequent), longleaf-pine flatwoods (frequent),
and slash-pine flatwoods (frequent) (Friauf, 1953).

Cariblatta lutea minima

Florida.—Series of specimens captured on Long Key under dead petioles
of coconut palm on moist ground at edges of pools of brackish water.
Specimens from Key West were in dry dead grass under boards (Rehn and
Hebard, 1912). Nymphs frequent under bark on decaying pine logs in pine
woods; occasional in leaf mold in heavy junglelike scrub (Rehn and
Hebard, 1914). In water-soaked leaves in heavy red-mangrove swamp
(Hebard, 1915). Under dead petioles of coconut palm on sandy soil in
grapefruit grove (Hebard, 1916a). Numerous at bases of tufts of coarse
grass growing just back of sea beach (Blatchley, 1920). Friauf (1953) found
this species in leaf duff, leaf mold, debris, or decaying wood in these
habitats: Dry, ruderal grassland (occasional), scrub (infrequent), sandhills

(infrequent), mesic hammock (infrequent), pond margin (occasional),
longleaf-pine flatwoods (occasional), and low hammock (infrequent). On
bare soil or bare sand under vegetation in these habitats: Longleaf-pine
flatwoods (occasional) and slash-pine flatwoods (occasional). Dominant in
the spartina marsh habitat in the grass stratum and duff around clumps.
Frequent in the saw-grass marsh habitat in the grass stratum and, during the
dry season, in decaying vegetation on the marsh floor.

Cariblatta nebulicola

Jamaica.—Adults in dead leaf litter alongside the trail in dense forest of
tree ferns, Podocarpus, Cyrilla, and other trees; the forest was bathed in fog
much of the time (Rehn and Hebard, 1927).

Cariblatta reticulosa

Jamaica.—In leaves on leaf mold in hillside forest (Hebard, 1916a).
Moderately numerous in leaf litter in mangrove swamp; in decaying
herbage (Rehn and Hebard, 1927).

Cariblatta stenophrys

Puerto Rico.—Between the leaves and under the leaf sheaths of corn (Sein,
1923; Wolcott, 1936).

Cariblatta spp.

West Indies.—The tropical species of this genus inhabit heavy forest, living
among the fallen leaves resting on the leaf mold, in epiphytic bromeliads,
and in dead agaves (Hebard, 1916a; Rehn and Hebard, 1927).

Cariblattoides instigator

Cuba.—In siftings from under sea grapes, other shrubs, and low trees (Rehn
and Hebard, 1927).

Cariblattoides suave

Puerto Rico.—On dry limestone hills (Rehn and Hebard, 1927).

Ceratinoptera picta

Trinidad.—Under bark of old cacao tree (Princis and Kevan, 1955).

Chorisoneura flavipennis

Costa Rica.—Under stones on borders of Surubres River (Rehn, 1906).

Chorisoneura formosella

Jamaica.—Swept from huckleberry trees (Vaccinium meridionale) (Rehn
and Hebard, 1927).

Chorisoneura parishi

Panama.—From jungle undergrowth (Hebard, 1920).

Chorisoneura specilliger

Panama.—In grass (Hebard, 1920).

Chorisoneura texensis

Florida.—"The almost impenetrable jungle on Key Largo was examined,
and in its depths the two specimens of this species were secured by beating
the lower branches of gumbo limbo, other trees and the lower bushes and
shrubs, among which latter are to be found such tropical forms as Ocotea
catesbyana [= Nectandra coriacea] and Citharexylum villosum" (Rehn and
Hebard, 1912). In nests of webworm and beaten from bushes of bayberry,
Myrica cerifera, along edge of pine woods (Rehn and Hebard, 1916).
Beneath dead leaves in oak woods and beaten from foliage of oak and

bayberry (Blatchley, 1920). Infrequent in the tall shrub stratum of the xeric
hammock habitat (Friauf, 1953).
Texas.—The great majority of specimens were beaten from foliage of
bushes (Hebard, 1943a).
Southeastern and southern U.S.—In undergrowth in pine woods; beaten
from shrubbery, from bayberry bushes, from lower branches of gumbo
limbo and other trees, from lower bushes and shrubs in jungle, and from
low oaks on hills. In Texas, beaten from tall weeds in opening in river-plain
jungle scrub (Hebard, 1917).

Chorisoneura translucida

Panama.—In jungle vegetation, including vines covering low bushes
(Hebard, 1920).

Chromatonotus infuscatus

Trinidad.—Males on low herbage under old cacao tree (Princis and Kevan,
1955).

Chromatonotus notatus

Trinidad.—Males in orchard on low herbage at night; females under refuse
and in grass (Princis and Kevan, 1955).

Comptolampra liturata

Malaya.—Often found between dry foliage in the beakers of the epiphytic
fern, Asplenium nidus, although the species lives mainly in bamboo bushes
(Karny, 1924).

Cryptocercus punctulatus

North Carolina.—"They were never found except in parts of the logs
[chestnut] where the decayed wood was soft, punky and wet" (Rehn and

Hebard, 1910).
Oregon.—In fir logs where sap wood was soggy (Hebard, 1917).
Virginia.—In decaying chestnut and pine logs; taken six times in chestnut
and once in pine (Hebard, 1917). In rotten logs in deep ravines of moist
woods (Davis, 1926).
Appalachian Mountains, U.S.—In southern Virginia and eastern Tennessee,
it is usually quite abundant in well-forested areas at elevations from 3,000
to 5,000 feet; "sometimes even a majority of the dead logs on a mountain
side have roaches in them" (Cleveland et al., 1934). This cockroach not
only lives in rotten, dead logs but also in sound logs that have been down
only a few years. In Virginia it is found more often in chestnut and
hemlock. "It occurs fairly often in oak, and has been found in pine, spruce,
and arbor vitae.... There is little evidence that they ever leave the log and
enter the ground" (Cleveland et al., 1934).

Cryptocercus relictus

Eastern Manchuria.—In great numbers under rotting fallen trees and in
rotten dead wood (Bei-Bienko, 1950).

Cutilia soror

Marquesas Islands.—Males under stones and dead log (Hebard, 1933a).
Hawaii.—In soil about roots of pineapple (Illingworth, 1927). Often found
about roots of grasses and weeds and other debris (Williams et al., 1931).
Under stones and pineapple mulching paper (Fullaway and Krauss, 1945).
Wake Island.—Numerous, some from rotten logs. Found in bunch grass on
Ocean Island (Bryan, 1926).

Cutilia spp.

Australia.—Frequent woods where they leave shelter soon after sunset and
run actively on ground or ascend shrubs and trees in quest of prey (Tepper,
1893).

Dendroblatta sobrina

Panama.—Colony on tree trunk; on surface of trunk of fallen tree (Hebard,
1920).

Diploptera punctata

Hawaii.—"Crowds of these insects in various stages of development
sometimes gather in cypress trees, in suitable chinks, in old flowerhead
sheaths of palms, etc., and even more or less openly on leafy twigs, in
bunch grass, and the species is at times locally abundant behind the older
leaf bases of sugar cane" (Williams et al., 1931). Williams also lists the
following as food plants: Cryptomeria, algaroba, lime trees, ripening
mangoes, papayas, and oranges. However, Bianchi (personal
communication, 1954) doubted that any of the above are the main dietary,
because the largest populations he had seen "were found in the fairly dry
litter of Star Jasmine (Jasminum pubescens Willd.), well removed from any
of the plants mentioned by Williams."
Raiatea, Society Islands.—Beaten out of bracken (Cheesman, 1927).
Uahuka, Marquesas Islands.—Under bark (Hebard, 1933a).

Dryadoblatta scotti

Trinidad.—Very common in water-filled, epiphytic bromeliads in the rain
forest (see p. 31) (Princis and Kevan, 1955).

Ectobius africanus

Belgian Congo.—Females in forest margin and in forest undergrowth
(Rehn, 1931).

Ectobius albicinctus

South France.—Females and young beneath stones (Blair, 1922).

Ectobius duskei

U.S.S.R.—In the steppe belt, it is a very characteristic member of feather-
grass steppes, where it is found in associations of typically steppe
vegetation, with feather grasses at the head (Stipa lessingiana and others),
and on rocky slopes; it occurs frequently in cultivated fields of young crops
and also in young geological strata in sections with virgin soil. The
populations of this steppe cockroach average 6 to 8 individuals per square
meter from the middle to the end of July. By the end of summer most
individuals were observed at the bases of straw stacks with a canopy,
having their south sides sheltered. This is the only species of Ectobius
adapted to a purely steppe biocenose. (Bei-Bienko, 1950.)

Ectobius lapponicus

Southeastern Europe.—Numerous under stones on Trebovic (Burr, 1898).
U.S.S.R.—Found in wooded communities and peat bogs (in northern part of
its range); males occur predominantly on herbaceous plants and bushes, but
females hide under fallen leaves, moss, etc. (Bei-Bienko, 1950). It
populated about 25 percent of the aspen trees in an experimental plot,
feeding in galleries in the bark of young branches; there were 25 or more
individuals per tree (Stark in Bei-Bienko, 1950).
Germany.—Abundant in woods; in pine woods in company with
Stenobothrus vagans and Tettix kraussi. Numerous in low aspen bushes in
forest. Numerous in deciduous and coniferous forests on trees and
underbrush; under fallen leaves and moss; on oaks (Zacher, 1917). In
foliage of young oak on top of mountain (Ramme, 1923).
Great Britain.—Under moss and dry leaves, among woodland undergrowth,
and, generally, on vegetation close to the ground; occasional on bushes and
trees (Lucas, 1920). Nymphs in heather in February and later; adults among
rushes fringing pond in July (Lucas, 1925). Nymphs and males on rushy
vegetation; unusually abundant on low herbage in dried-up swamp (Lucas,
1930).

Ectobius nicaeensis

France.—In dry woods, on bushes, and at the base of trees (Chopard,
1947).

Ectobius pallidus

Algeria.—Under stones; in moist places that are shaded and covered with
plants (Lucas, 1849).
England.—Very abundant on sand dunes and among bracken in July
(Buxton, 1914).
Germany.—In deciduous and coniferous forests; at edge of forest, from
bare woods and bushes; numerous under leaves in oak woods and under
moss (Zacher, 1917). In forest well lighted by the sun (Ramme, 1923).
Massachusetts.—Under loose lichens and bark on oak trees; under boxes,
baskets, paper, etc., near houses; on Swiss chard (Flint, 1951). On roofs of
houses, in shrubbery (Gurney, 1953). We have collected this species for
several summers in a fairly dense, wooded area near dwellings, among
fallen leaves and climbing on the erect stems and undersides of the leaves
of periwinkle. Oöthecae were found on the ground under leaves and debris.

Ectobius panzeri

England.—Abundant on sandhills along shoreline among roots of grass
(Burr, 1908). Under dead seaweed and other rubbish a few yards from shore
on ground that would be washed by the sea (Lucas, 1896). Nymphs found
among marram grass (Buxton, 1914). On sandhills near coast and covered
with marram grass; often found on heather and low herbage; under old bark
and rotten wood on posts; in decayed stump (Lucas, 1920). Swarming on
Beta maritima and other plants in July (Lucas, 1920a). Very common in all
stages in August, being frequently found under stones (Lucas, 1925).
Common on sand dunes especially under stems of dead marram grass.
Viable oöthecae found buried in sand (Brown, 1952).
Germany.—In beech woods and in pine woods (Zacher, 1917).

Ectobius semenovi

Kazakhstan.—Along the shores of the Syr-daria it is found on and around
living willows and on Populus euphratica; under loose bark of dying and
dead trees (Bei-Bienko, 1950).

Ectobius sylvester

U.S.S.R.—In wooded steppe zones; probably only occurs in association
with forests (Bei-Bienko, 1950).

Ectobius tadzhicus

Tadzhikistan.—Great numbers at the roots of Eleagnus shrubs on the banks
of reservoirs and frequently under the bark of old trees (Bei-Bienko, 1950).

Ectobius vittiventer

South France.—One male beneath stone (Blair, 1922).

Ellipsidion affine

Australia.—From leaves, from scrub (Hebard, 1943). Collected in trees
(Pope, 1953a).

Ellipsidion australe

Australia.—On eucalyptus leaves, on wattle, under bark (Hebard, 1943).
Collected in trees (Pope, 1953a).

Ellipsidion bicolor

Australia.—In corn and from tree (Hebard, 1943).

Ellipsidion simulans

Australia.—From sugarcane (Hebard, 1943).

Ellipsidion spp.

Australia.—All stages are diurnal moving about the foliage of shrubs and
small trees in bright sunlight on hottest summer days (Tepper, 1893).

Epilampra abdomen-nigrum

Trinidad.—In dried-up drain; among grass; in debris under old cacao tree;
under old leaves (Princis and Kevan, 1955).
Puerto Rico.—Abundant in damp lowlands (Seín, 1923). Under dead leaves
in wet malojillo meadow (Wolcott, 1936).
This species is amphibious (p. 31). Shelford (1907) suggested that
immature stages of other species of the genus may be aquatic, which would
place them in moist situations on the shores of rivers and other bodies of
water.

Epilampra azteca

Panama.—Very scarce, under palm trees in decaying leaf mold and litter;
one found under decaying bark of a log (Hebard, 1921a).

Epilampra mona

Mona Island, Puerto Rico.—One specimen under bark of dead tree (Ramos,
1946).

Epilampra tainana

Cuba.—Under dead leaves on stream bank (Rehn and Hebard, 1927).

Epilampra wheeleri

Puerto Rico.—In siftings from high-altitude primeval forest (Rehn and
Hebard, 1927).

Epilampra spp.

Australia.—By day the insects live under bark, stones, logs, dead vegetable
debris, or buried in loose dust or soil. After sunset females wander in grass
or ascend low objects (Tepper, 1893).

Ergaula capensis

Uganda.—In open bush and short grass (Princis, 1955).

Eudromiella bicolorata

Panama.—Under rubbish on edge of jungle (Hebard, 1920).

Euphyllodromia liturifera

Colombia.—In brushwood (Princis, 1946).

Eurycotis biolleyi

Costa Rica.—Numbers of individuals were found in the large bromeliads of
the temperate localities (Picado, 1913).

Eurycotis decipiens

Trinidad.—In old, rotten coconut stump (Princis and Kevan, 1955).

Eurycotis dimidiata

Cuba.—"This species was recorded from under stones in the fields ... by
Gundlach" (Rehn and Hebard, 1927).

Eurycotis ferrum-equinum

Cuba.—Under stones in woods (Rehn and Hebard, 1927).

Eurycotis floridana

Florida.—Moderately common under bark of dead pine stumps and logs; at
Key West it fairly swarmed under coquina boulders in the woods (Rehn and
Hebard, 1905). Many specimens under palmetto leaves on ground (Caudell,
1905). In pine woods under dry bark of dead logs; on Long Key in dry
fibers at the base of the heads of coconut palms; "at Key West, a large
colony was discovered among boards lying on dry grass in a field, and
several were captured upon turning over coquina boulders in the dense
bush" (Rehn and Hebard, 1912). Particularly numerous in tree cavities and
under bark along the edge of hammock areas (Hebard, 1915). Abundant
between basal leaves of Tillandsia utriculata; beneath loose bark of logs
and stumps; in and beneath decaying palmetto trunks and leaves; under
rubbish (Blatchley, 1920). On ground in heavy tangle after dark; in
decaying log of Sabal palmetto; in bromeliads; common under debris and
bark in jungle; under signs on Pinus caribaea; in almost every sheltered
outdoor place (Hebard, 1917). It moves about at night and hides under bark
of logs and in other recesses during the day; where pines are present it
almost invariably hides under bark of dead logs and stumps (Rehn and
Hebard, 1914). Friauf (1953) found this species in leaf duff, leaf mold, or
decaying wood in these habitats: Sandhills (infrequent), xeric hammock
(dominant), mesic hammock (frequent), and low hammock (dominant); on
tree trunks in sandhills habitat (infrequent) and mesic hammock (frequent);
infrequent in saw-grass marsh habitat in the grass stratum and, during the
dry season, in decaying vegetation on floor of marsh. Under the bark of logs
and beneath logs in the woodpile habitat (Friauf, 1953).

Eurycotis galeoides

Cuba.—Under stones in deep woods (Rehn and Hebard, 1927).

Eurycotis kevani

Trinidad.—Under debris, trash, and vegetable refuse (Princis and Kevan,
1955).

Eurycotis opaca

Cuba.—In pine and palmetto region (Rehn and Hebard, 1927).

Euthlastoblatta abortiva

Texas.—Under dense tangle of bushy vegetation, palms, and vines near Rio
Grande; in leaves and dry litter on ground; on dead petiole hanging from
palm tree (Hebard, 1917). Under bark of dead hackberry; abundant in dead
leaves, dry litter, and rats' (Neotoma sp.) nests in heavy scrub (Hebard,
1943a).

Graptoblatta notulata

Tahiti.—On foliage in sun or concealed among dead leaves that collect
between the fronds of tree ferns (Cheesman, 1927).
Hawaii.—Quite active during the day, occurring on sugarcane, etc., in the
wetter districts; it is also a household insect (Williams et al., 1931).

Hemiblabera brunneri

Puerto Rico.—Under bark of tamarind tree (Rehn and Hebard, 1927).
Under the bark on a fence post (Wolcott, 1950).

Henicotyle antillarum

Dominica.—From rotting wood and wood soil (Rehn and Hebard, 1927).

Holocompsa metallica

Dominican Republic.—Along railroad through jungle and swamp (Rehn
and Hebard, 1927).

Hololampra bivittata

Canary Islands.—Found in numbers among pine needles; nymphs were in
the majority, adults rare (Burr, 1911).

Hololampra chavesi

Azores.—Very common in the hedges, particularly in brambles. Contrary to
most species of this genus, which live on the ground under stones, this
species is exclusively dendricolous and is only captured by beating the
bushes on which it abounds (Chopard, 1932).

Hololampra maculata

Germany.—Abundant in deciduous forest in grass and under fallen leaves;
in pine forests under lichens and between fallen needles; in edge of
coniferous forest; under stones (Zacher, 1917).

Hololampra marginata

Macedonia.—Usually found crawling on the flowers and stems of giant
thistles in May; common on thistles in June (Burr, 1923).

Hololampra sp.

Caucasus.—Numerous beneath dry leaves in a garden (Burr, 1913).

Hololeptoblatta sp.

Seychelles.—Apparently only inhabits Pandanus between the leaf bases
(Scott, 1910, 1912).

Homalopteryx laminata

St. Vincent.—In decaying leaves in forest (Rehn and Hebard, 1927).
Trinidad.—In forest debris and debris under old cacao trees; it is not
uncommon under dry leaves; it feigns death when disturbed (Princis and

Kevan, 1955).

Hormetica laevigata

Brazil.—From crown of palm between leaf bases (Hancock, 1926).

Ignabolivaria bilobata

U.S.S.R.—Under rocks and on the edges of woods in the lowlands in the
north and in the mountains in the south (Bei-Bienko, 1950).

Ischnoptera deropeltiformis

North Carolina.—Under pine straw on ground in woods (Brimley, 1908).
Georgia.—Under dead oak leaves; under debris in garden; running on
ground in pine and oak woods (Rehn and Hebard, 1916).
Indiana.—It is "a ground-frequenting, forest-loving insect, hiding beneath
cover or about the edges of deep woodland, more frequently in damp
places, and rarely taken beneath bark, signs, or at lights" (Blatchley, 1920).
Missouri.—Twenty to 30 males found resting on heads of wild oats on
successive evenings (Rau, 1947).
Texas.—It preferred damp, open woodlands (Hebard, 1943a).
Eastern and southeastern U.S.—Under stone in heavy deciduous forest;
under damp, dead leaves on edges of forests; under bark of pine log; in wire
grass and sphagnum bordering stream thicket; in leaf mold and rubbish
about pothole in pine woods, Pinus caribaea; under debris and leaf mold in
hammock; under dead oak leaves in heavy deciduous forest (Hebard, 1917).
Florida.—"This species is distinctly geophilous and appears to prefer damp
surroundings" (Rehn and Hebard, 1912). Under boards on very wet ground
in everglades; in debris and leaf mold in heavy, junglelike areas of trees,
bushes, and vines (Rehn and Hebard, 1914). Adults and numerous nymphs
beneath weeds, grass, and other debris washed up on beach of Lake
Okeechobee (Blatchley, 1920). Friauf (1953) found this species in leaf duff,

leaf mold, and/or decaying wood on ground in these habitats: Dry, ruderal
grassland (infrequent), scrub (infrequent), sandhills (occasional), xeric
hammock (frequent), mesic hammock (dominant), shrubby, longleaf-pine
flatwoods (infrequent), bayhead (dominant), and low hammock (dominant).
On open bare soil or bare sand under vegetation in these habitats: Dry,
ruderal grassland (infrequent), mesic hammock (dominant), moist, ruderal
grassland (infrequent), pond margin (occasional), longleaf-pine flatwoods
(infrequent), slash-pine flatwoods (infrequent), and low hammock
(dominant). Infrequent in the herbaceous stratum of these habitats: Dry,
ruderal grassland, moist, ruderal grassland, and longleaf-pine flatwoods.
Infrequent in the shrub stratum of the dry, ruderal grassland habitat. (Friauf,
1953.)
Tennessee.—Taken in traps baited with cantaloupe in a parklike stand of
oak, gum, hickory, and tulip trees in a creek bottom, and in a stand of oak
on a dry ridge (Walker, 1957).

Ischnoptera panamae

Panama.—Under rubbish at edge of jungle and under drift on edge of coral-
sand beach (Hebard, 1920).

Ischnoptera podoces

Jamaica.—In dead leaf litter along side trail through mountain forest (Rehn
and Hebard, 1927).

Ischnoptera rufa rufa

Virgin Islands, St. Croix.—Common under rubbish and on shrubbery at
night (Beatty, 1944).
Barbados.—Occasionally found in cane fields (Tucker, 1952).
West Indies.—In Puerto Rico, under stones in cultivated area, under debris
on alkalie flat. In Jamaica, under dry petioles of coconut palm in grassy
area; under logs, logwood on docks, and litter on limestone and near beach.

In Panama, under drift on edge of coral-sand beach; under rubbish at edge
of jungle (Hebard, 1916c).
Jamaica.—Under limbs and leaf litter in mangrove swamp (Rehn and
Hebard, 1927).

Lamproblatta albipalpus

Panama.—Under drift on edge of coral-sand beach. Several under decayed
banana stem (Hebard, 1920).

Lamproblatta meridionalis

Trinidad.—Under debris in forest and debris under old cacao trees (Princis
and Kevan, 1955).

Latiblattella chichimeca

Costa Rica.—Very common in the bromeliads of all Costa Rica (Picado,
1913).

Latiblattella lucifrons

Arizona.—"Most commonly seen feeding on pollen and dead insects on the
flower stalks of Yucca elata in June in the Santa Rita Mountains" (Ball et
al., 1942).

Latiblattella rehni

Florida.—Widely distributed throughout pine woods (Pinus caribaea);
under signs on Pinus clausa and Pinus caribaea (Hebard, 1917). Beneath
bark of dead pine tree; beating Spanish moss; they seldom attempt flight
when disturbed, but hide in crevices or drop to ground (Blatchley, 1920).

Latiblattella zapoteca

Costa Rica.—Under stones on borders of Surubres River (Rehn, 1906).

Leucophaea maderae

Barbados.—In cane fields (Tucker, 1952).
Dominica.—In vegetation of royal palms, guava, etc.; under loose bark and
banana sheaths. In Jamaica, on logwood docks (Rehn and Hebard, 1927).

Litopeltis biolleyi

Costa Rica.—Under bark of tree in forest; in epiphytic bromeliads (Rehn,
1928).

Litopeltis bispinosa

Panama Canal Zone.—About 80 specimens from rotting banana stalks at
bases of leaves; boring in decaying banana stem (Hebard, 1920).

Litopeltis deianira

Costa Rica.—In tree stump on edge of mountain forest; in dead wood on
ground (Rehn, 1928).

Litopeltis musarum

Costa Rica.—Shaken from dead banana leaves. Footnote to specific name:
"In relation to the liking of species of this genus for bananas (Musa) as
shelter and possibly food" (Rehn, 1928).

Lobolampra subaptera

France.—Under stones and dead leaves, always rare (Chopard, 1947).

Loboptera decipiens

France.—All stages common beneath stones (Blair, 1922). Under stones
and dead leaves (Chopard, 1947).
Maltese Islands.—Quite common in open country under stones (Valletta,
1955).
Dalmatia.—On seashores under rocks and seaweed cast up on shore (Bei-
Bienko, 1950).

Loboptera thaxteri

Argentina.—Common in rubbish and leaf litter in small woodlot (Hebard,
1932).

Lobopterella dimidiatipes

Hawaii.—Abundant in wet districts, both in lowlands and to a considerable
altitude in the forests, under trash, stones, boards, etc. (Williams et al.,
1931). Often it is found with nymphs of Periplaneta australasiae (Fullaway
and Krauss, 1945).

Lophoblatta arawaka

Trinidad.—On grass, maize, and cut sugarcane fodder; under vegetable and
garden refuse; under old cacao (Princis and Kevan, 1955).

Macropanesthia rhinocerus

Australia.—Infrequently seen during dry season from March to October.
"They burrow quite deeply, about two feet below the surface of the sandy
soil in stands of cypress pine (Callitris sp.). They make a nest of dead
leaves, grass roots, etc., frequently among the pine roots. The young
nymphs rarely appear above ground, but following rain the adults burrow to
the surface, especially at night.... This species is also found in the brigalow
(Acacia harpophylla) scrub about 70 miles west of Rockhampton,
Queensland, and on Fraser Island off the Coast of Queensland" (Henson in
Day, 1950).

Megaloblatta blaberoides

Panama.—Under bark on tree (Hebard, 1920).
Ecuador.—Under a dense pile of dead leaves around base of tree (Campos
R., 1926).

Megamareta verticalis

Australia.—In sugarcane (Hebard, 1943).

Methana canae

Australia.—Under loose bark on dead upright tree (Pope, 1953a).

Methana curvigera

Australia.—Under loose bark on trees and logs; many specimens on wattle
trees where in strong sunlight they hid in curled-up leaves; oöthecae
attached to underside of loose bark and leaves (Pope, 1953a).

Methana marginalis

Australia.—Under loose bark of trees and logs (Pope, 1953a).

Moluchia (?) dahli

Chile.—Collected from lichens and mosses on tree trunks (Princis, 1952).

Muzoa madida

Costa Rica.—Under dead wood in dense second-growth forest; in thick mat
of hanging dead vegetation in dense forest; under leaves in forest (Rehn,
1930).

Nauclidas nigra

St. Vincent.—Under rotten fruit (Rehn and Hebard, 1927).

Nelipophygus ramsdeni

Cuba.—Under rotten bark (Rehn and Hebard, 1927).

Neoblattella detersa

Jamaica.—Under dried leaves of coconut palm; in dry leaves under acacia
on hillside; in debris on beach; under stones on coral rock; in leaf mold
under dense brush on hillside; under bracts of banana blossoms (Rehn and
Hebard, 1927).

Neoblattella dryas

Jamaica.—In bases of dead tree-fern fronds; numerous in bromeliads;
nearly all collected specimens were taken in these plants (Rehn and Hebard,
1927).

Neoblattella eurydice

Jamaica.—Nearly all collected specimens taken in bromeliads (Rehn and
Hebard, 1927).

Neoblattella grossbecki

Jamaica.—In epiphytic bromeliads and hollow bases of dead tree-fern
fronds; nearly all collected specimens taken in bromeliads (Rehn and
Hebard, 1927).

Neoblattella proserpina

Jamaica.—Under bark of huckleberry; nearly all collected specimens taken
in bromeliads (Rehn and Hebard, 1927).

Neoblattella semota

Jamaica.—All specimens collected from under drying bracts of banana
blossoms (Rehn and Hebard, 1927).

Nesomylacris cubensis

Cuba.—In dry region of palmettos and pines (Rehn and Hebard, 1927).

Nesomylacris relica

Jamaica.—Widely distributed from sea level to 5,700 feet elevation; in
bromeliads in mountain forest; among dead leaves in heavy leaf mold under
dense hillside scrub; under stones and in ground litter about banana trees;
under bark of tree in dense ridge-type forest; in dead agave in scrub forest
(Rehn and Hebard, 1927).

Nocticola bolivari

Ethiopia.—Always found under stones or cement blocks, but not
necessarily deeply buried in the ground (Chopard, 1950b).

Nyctibora laevigata

Jamaica.—In cracks in dead stump of mimosa; in bromeliads (Rehn and
Hebard, 1927).

Nyctibora lutzi

Puerto Rico.—Possibly to be found most often in rotten tree trunks in the
highest mountains; found in rotten stump with termites, ants, and beetle
grubs (Wolcott, 1950).

Nyctibora obscura

Trinidad.—Under pile of cornstalks (Princis and Kevan, 1955)

Nyctibora stygia

Haiti.—Under loose dead bark of mesquite tree, 52 specimens (Rehn and
Hebard, 1927).

Oniscosoma spp.

Australia.—The females bury themselves in loose soil or dust (Tepper,
1893).

Opisthoplatia orientalis

Formosa.—On or in swampy ground or under rotten trees on the ground
(Takahashi, 1924).

Panchlora antillarum

Dominican Republic.—In cultivated grounds, palms, fruits, etc. (Rehn and
Hebard, 1927).

Panchlora nivea

Panama.—As Pycnosceloides aporus, in jungle under decaying banana
stem in which were boring individuals of Litopeltis bispinosa (Hebard,
1920).

Texas.—Lives in foliage and in the green sheaths of plants (Hebard, 1943a).
Cuba.—On cane leaves; according to Gundlach this genus lives under the
loose bark of trees (Rehn and Hebard, 1927).
Puerto Rico.—In rotting trunks of coconut palms (Seín, 1923). Most
specimens have been collected from the very rotten interior of coconut
palms (Wolcott, 1950).
Trinidad.—On corn; under old log; flies readily to lights (Princis and
Kevan, 1955).

Panchlora sagax

Dominica.—In decaying stump in banana patch and in rotting wood. In
Puerto Rico, in rotten coconut palm (Rehn and Hebard, 1927).

Panesthia australis

Australia.—In burrows under the thick bark of fallen and rotting trees
(Shaw, 1914). In loose detritus, beneath clods of earth, and in fissures at
foot of cliffs along the seashore beyond direct action of the waves (Tepper,
1893).

Panesthia laevicollis

Australia.—Under decayed logs in coastal scrub. It burrows into the soft
part of the log (Froggatt, 1906).

Parcoblatta bolliana

North Carolina.—Under pine straw on ground in pine woods (Brimley,
1908).
Texas.—Under dry cow dung in pine woods (Hebard, 1917).
Nebraska.—Under pile of old boards (Hauke, 1949).

Parcoblatta caudelli

North Carolina.—From under the bark of dead trees (Rehn and Hebard,
1910).
Virginia.—At night on shrubbery. In South Carolina, under sign on tree
(Hebard, 1917).
Tennessee.—In traps baited with cornmeal, cantaloupe, or fish in a stand of
oak on dry ridge, and in abandoned rocky field on a south-facing slope
(Walker, 1957).

Parcoblatta desertae

Texas.—From mountains, arid, and semi-arid regions; under small boulder
on desert (Hebard, 1917). On ground in dry-creek bed through scrub oak,
pine, and juniper forest (Hebard, 1943a).

Parcoblatta divisa

Eastern and southeastern U.S.—All specimens taken from under signs on
red oaks and longleaf and shortleaf pines in Georgia and Virginia (Rehn and
Hebard, 1916). Trapped in molasses-baited jar in oak forest in New Jersey;
under signs on red and white oaks, sweet gum, and other deciduous trees;
under signs on shortleaf and longleaf pines and pine stumps (Hebard, 1917).
Widespread in southeastern U.S. in habitats as diverse as dry pine lands,
oak scrub, moist hammocks in northern Florida, and deep, cool ravines
along Apalachicola River (Hebard, 1943a).

Parcoblatta fulvescens

Eastern and southeastern U.S.—Trapped in molasses jars: in heavy, barrier-
beach forest; in typical pine-barrens undergrowth; in pine barrens with
heavy, grassy undergrowth; on border of pine barrens and on edge of
swamp; in heavy deciduous forest; in heavy oak woods. Found under debris
in dead, shortleaf-pine needles; under dead leaves on edge of oak and

shortleaf-pine woods; under bark of pine log; among dead leaves under live
oaks; under sign on Pinus caribaea (Hebard, 1917).
Georgia.—From under bark of pine log, among dead leaves under live
oaks, and under leaves on edge of oak and shortleaf-pine woods (Rehn and
Hebard, 1916).
Florida.—Very common among dead leaves, under logs, beneath loose
bark, and wanders about at night in pinelands, hammock, turkey oak, and
sand-scrub habitats (Hubbell and Goff, 1940). Beneath drift, cow dung,
leaves, boards, bark of logs, and other debris, usually in open pine woods in
sandy areas; frequent at the base of thistle leaves (Blatchley, 1920). Friauf
(1953) found this species in leaf duff, debris, or decaying wood in these
habitats: Scrub (dominant), sandhills (dominant), xeric hammock
(dominant), mesic hammock, longleaf-pine flatwoods (infrequent), low
hammock (infrequent), and alluvial hammock (infrequent). In the shrub
stratum in these habitats: Scrub (dominant), sandhills (dominant), xeric
hammock (dominant), and longleaf-pine flatwoods (infrequent). In the
herbaceous stratum of the longleaf-pine flatwoods habitat, and under bark
and beneath logs in the woodpile habitat.

Parcoblatta lata

Southeastern and southern U.S.—Under bark of pine logs and stumps; in
sweet-gum logs and stumps; moderately numerous under bark of dead
shortleaf pines; under bark of longleaf-pine stumps; under signs on red oak
and longleaf pines; in dead oak. In Texas, under bark of pine stumps
(Hebard, 1917).
North Carolina.—All stages under loose bark of dead pines, both prostrate
and upright, and stumps. "It seems to prefer the space under the bark to be
rather damp" (Brimley, 1908). Under bark of dead pine trees (Rehn and
Hebard, 1910).
Florida.—Infrequent in leaf duff and decayed wood of low hammock
habitat (Friauf, 1953).
Indiana.—Beneath rocks on sides and tops of high hills, in limestone glades
where cedar abounds (Blatchley, 1920).

Missouri.—In leaf stratum of oak-hickory forest (Dowdy, 1951). Earlier,
Dowdy (1947) reported finding numerous immature Pseudomopinae
[presumably Parcoblatta sp.] in soil and leaf strata of oak-hickory forest.
Texas.—Captured in molasses-baited traps in low, wet, oak woods and in
dry woodlot on hillside (Hebard, 1943a).

Parcoblatta pensylvanica

Eastern and southeastern U.S.—Trapped in molasses-baited jars; in oak
and in chestnut forests, and on knoll with high deciduous trees. Found in
oak and pine woods, under bark of decaying chestnut log and dead chestnut
stump, and under signs on trees including oaks (Hebard, 1917).
North Carolina.—In all stages under loose bark of upright, dead pines,
when the space under the bark was dry (Brimley, 1908).
Virginia, North and South Carolina.—Under signs on trees (white and red
oaks); under bark of dead shortleaf-pine and sweet-gum logs and stumps
(Rehn and Hebard, 1916).
Indiana.—Beneath bark of logs and stumps; empty oöthecae common
beneath loose bark of logs, especially shellbark hickory (Blatchley, 1920).
Under loose bark on logs in January (Blatchley, 1895).
Illinois.—In pine forest associes, in black oak forest on sand, in oak-
hickory forest on clay, and in climax forest; it evidently moved into the pine
associes nightly, great numbers of oöthecae were found under bark of pine
logs, where, in October and November, hibernating nymphs were found
(Strohecker, 1937). In nests of Vespula maculata (Balduf, 1936; McClure,
1936).
Missouri.—Usually in hollow trees, under loose bark, in woodpiles, and in
cracks in rural buildings (Rau, 1940).
Michigan.—Common in oak-dune and beech-maple forests, under loose
bark on dead trees and fallen logs, and under debris on forest floor
(Hubbell, 1922). "A characteristic inhabitant of the low shrub-terrestrial and
probably the terrestrial-hypogeic stratum." It occurred throughout the
upland forests; groups were found established in and under logs 100 to 200
feet from the nearest forest (Cantrall, 1943).

Ontario.—Very abundant in rocky, sparsely-wooded country, where it
occurred in rotten logs and under loose bark; on tree trunk at night on rocky
island in lake (Walker, 1912).

Parcoblatta uhleriana

North Carolina.—Under pine straw on ground in woods (Brimley, 1908).
Under bark of dead trees; 92 males attracted to lights (Rehn and Hebard,
1910).
Virginia.—Resting on woods foliage; at night on road (Rehn and Hebard,
1916).
Eastern and southeastern U.S.—Trapped in molasses-baited jars: in oak
and pine woods, in heavy barrier-beach forest, in both scant and typical
undergrowth on pine barrens, in heavy grassy undergrowth on pine barrens,
on border of pine barrens, on edge of swamp, in heavy deciduous forest, in
heavy oak woods, in upland oak and chestnut forest, in chestnut forest, in
forested ravine, and on ridge with heavy oak, chestnut, and maple forest.
Found under damp leaves on edge of forest, under bark of decayed chestnut
log, inside decaying chestnut log with Cryptocercus punctulatus, under
palmetto roots, under bark of pine stump, and in dry leaves under live oaks
(Hebard, 1917).
Tennessee.—In traps baited with cornmeal or cantaloupe in maple-gum-oak
forest in a mesic valley, and in a stand of oak on a dry ridge (Walker, 1957).
Indiana.—Beneath cover on slopes of high wooded hills. "This is
essentially a forest-loving species; usually occurring beneath leaves and
other debris on or along the borders of heavy hardwood timber." (Blatchley,
1920.)
Illinois.—In oak-hickory forest on clay and in climax forest (Strohecker,
1937).
Michigan.—In oak-dune woods (Hubbell, 1922). Restricted to woodlands,
where it inhabited piles of moist dead leaves and rotten logs in oak-hickory
forest (Cantrall, 1943).
U.S.A.—This species, P. uhleriana, and P. virginica were attracted at night
to honeydew secreted by aphids on Pyrus sp. (Davis, 1918).

Parcoblatta virginica

New England.—Females under loose stones, boards, and other debris on
ground; beneath loose bark (Morse, 1920).
North Carolina.—Under debris in dead shortleaf-pine needles (Rehn and
Hebard, 1916).
Florida.—Infrequent in the shrub stratum of the scrub habitat. This was the
only habitat of 25 studied in which this species was found (Friauf, 1953).
Eastern and southeastern U.S.—Trapped in molasses-baited jars: in pine
and oak woods, in pine barrens, in pine woods with heavy grass
undergrowth, in oak forest, in heavily forested ravine, on rocky slope with
few deciduous trees, on knoll with high deciduous trees, in lofty chestnut
forest, and in heavy low chestnut and oak forest on high ridge; under bark
of decaying chestnut log and stump; under stones in chestnut forest; under
bark of pine stumps (Hebard, 1917).
Indiana.—Frequents borders of open woods and fields; under debris, loose
bark, and half-buried logs (Blatchley, 1920).
Illinois.—In black-oak forest on sand, in oak-hickory forest on clay, and in
climax forest (Strohecker, 1937).
Michigan.—Common in oak-dune and beech-maple forests; under loose
bark on dead trees and fallen logs and under debris on forest floor (Hubbell,
1922). Restricted to woodlands, where it inhabited piles of moist dead
leaves and rotten logs in oak-hickory forest (Cantrall, 1943).
Texas.—Captured in molasses traps in moist woods of maple, oak, and pine
with much undergrowth and a heavy layer of duff; in open, rather dry
woodlot of Spanish oak and other trees; and in low wet woods of willow
and oak along creek (Hebard, 1943a).

Parcoblatta zebra

Indiana.—Beneath log in cypress swamp (Blatchley, 1920).
Louisiana and Mississippi.—In decay cavity in sweet gum; under sign on
shortleaf pine (Hebard, 1917).

Parcoblatta spp.

Alabama.—In the dry wall of a sweet-gum stump together with serropalpid
and tenebrionid beetles (Snow, 1958).
Ohio.—Oöthecae under loose bark of fallen trees, where as many as 184
oöthecae were found within a few feet of each other; others found under
boards and in piles of firewood (Edmunds, 1952).

Pelmatosilpha coriacea

Puerto Rico.—Mona Island, under bark of dead trees and under guava
leaves (Ramos, 1946). Under bark of Sideroxylon foetidissimum (Wolcott,
1941). Common along the coast and in mountains. "Very much at home"
under the loose bark of Sideroxylon foetidissimum (Wolcott, 1950).

Pelmatosilpha kevani

Trinidad.—Under debris in bush (Princis and Kevan, 1955).

Pelmatosilpha purpurascens

Dominica.—In decaying logs in forest (Rehn and Hebard, 1927).

Periplaneta americana

Bermuda.—Among and under decaying debris, just above high-tide line
(Verrill, 1902).
Johnson Island.—Nocturnal, coming out at night in great numbers about
Tribulus blossoms. Under timbers on French Frigate Shoals (Bryan, 1926).
United States.—Alleyways and yards may be overrun during the summer;
adults and hundreds of nymphs found in decaying maple trees along
residential street (Gould and Deay, 1938, 1940). Around fumaroles where a
railroad fill was burning internally (Davis, 1927). Common in palm trees

along the gulf coast of Texas, where they often fly around street lights at
night (Zimmern in Gould and Deay, 1940).

Periplaneta australasiae

Bermuda.—Very abundant under stones (Rehn, 1910).
Jamaica.—Under bark of dead tree and under bases of leaves of coconut
palms (Rehn and Hebard, 1927).
Virgin Islands, St. Croix.—Common in sugarcane fields and in woodlands
(Beatty, 1944).
Florida.—Juveniles under bark of dead logs of Pinus caribaea (Hebard,
1915). Frequently found under signs on trees near borders of towns; under
bases of dead petioles of cabbage palmetto (Hebard, 1917). Beneath logs,
burlap bags, and other cover in old orange orchards (Blatchley, 1920).
Marquesas Islands.—Under coconut fronds and grass (Hebard, 1935).
Nihoa Island.—Nymphs only, on Sida, Pritchardia, bunch grass, and about
camp (Bryan, 1926).

Periplaneta brunnea

Georgia.—Under signs on oaks (Rehn and Hebard, 1916).
Florida.—Beneath bark of stump (Blatchley, 1920).

Periplaneta fuliginosa

Southeastern and southern U.S.—"This species is usually encountered out
of doors, in or near towns. Over its range it is frequently found under signs
on trees" (Hebard, 1917).

Phidon (?) dubius

Chile.—Collected from mosses and lichens on tree trunks (Princis, 1952).

Phoraspis spp.

Brazil and Guiana.—In grasslands, plantations of maize, sugarcane, and
other plants on the borders of forests; the cockroaches were always found
between the leaves which form the branches of the plants (Doumerc in
Blanchard, 1837).

Phyllodromica brevipennis

Asia Minor and western Europe.—On ground among grasses; under moss
and brushwood in mountain meadows (Bei-Bienko, 1950).

Phyllodromica graeca

U.S.S.R., western Georgia.—In pine forest mixed with deciduous trees
(Bei-Bienko, 1950).

Phyllodromica irinae

U.S.S.R., Turan Lowland.—Along margins of "tugas" under half-fallen
bushes of Salsola kali that overhang the ground (Bei-Bienko, 1950).

Phyllodromica maculata

Central Europe and western U.S.S.R.—On the edges of forests of the
central-European type that are lighted by the sun; under fallen leaves; on
bushes and conifers (Bei-Bienko, 1950).

Phyllodromica meglerei

U.S.S.R.—Among fallen leaves under bushes; on oak branches; under
mown hay (Bei-Bienko, 1950).

Phyllodromica polita

Caucasus.—Under fallen leaves on slopes of mountains covered by forest
or brushwood (Bei-Bienko, 1950).

Phyllodromica pygmaea

U.S.S.R.—In the sands of Un-dzhal-kum and Zhety-konur it is found in the
dense turf of Aristida pennata (Bei-Bienko, 1950).

Phyllodromica tartara

Central Asia.—In lowlands and in mountains up to 2,500 meters; in fruit
orchards under trap rings fastened to trees to combat lesser apple worm
(Bei-Bienko, 1950).

Phyllodromica tartara nigrescens

Southern Uzbekistan.—Under bark of Juniperus sp., under stones and on
flowers of Scorzonera acanthoclada (Bei-Bienko, 1950).

Platyzosteria castanea

Australia.—Under loose wood or bark (Shaw, 1914).

Platyzosteria novae seelandiae

New Zealand.—Swarms under loose dry bark and logs (Walker, 1904).

Plectoptera dominicae

Dominica.—On moss-covered lime trees. "The species of the genus
Plectoptera are all foliage and flower frequenters, generally secured by
beating low arborescent vegetation, or are attracted to light" (Rehn and
Hebard, 1927).

Plectoptera dorsalis

Puerto Rico.—In caladium, grass, weeds, coffee, and bananas; in flowers of
Ipomoea tiliasea (Rehn and Hebard, 1927). "... living in trees between
leaves, or in 'butterfly-nests' of Tetralopha scabridella in leaves of Inga
vera, or of Pilocrocis secernalis in the leaves of 'capá blanco' (Petitia
domingensis) in the mountains. Along the coast they have been found under
the bracts of cotton squares or bolls, and under the leaf-sheaths of sugar
cane, in curled-up leaves of grapefruit, or in the dry flower clusters of
'espino rubial' (Zanthoxylum caribaeum)." These observations apply also to
Plectoptera infulata and P. rhabdota (Wolcott, 1950).

Plectoptera floridana

Florida.—On fringe of tall bushes at edge of mangrove swamp (Rehn and
Hebard, 1914). Rehn and Hebard (1927) stated that on the Keys it
frequented dry scrubby vegetation, particularly Ilex cassine.

Plectoptera infulata

Puerto Rico.—See Wolcott's (1950) comments under Plectoptera dorsalis
above.

Plectoptera lacerna

Cuba.—In grasses, sedges, etc., about a waterhole; on grass, pines and oak
(Rehn and Hebard, 1927).

Plectoptera perscita

Dominica.—On moss-covered lime trees (Rehn and Hebard, 1927).

Plectoptera porcellana

Cuba.—Taken on flowers of "Júcaro" (Gundlach in Rehn and Hebard,
1927).

Plectoptera pygmaea

Jamaica.—In relatively dense forest foliage; in shrubbery (Rehn and
Hebard, 1927).

Plectoptera rhabdota

Puerto Rico.—In mixed vegetation; on grapefruit tree and guava (Psidium
guajava); on bushes and shrubs (Rehn and Hebard, 1927). On coffee trees;
on Spondias; on sugarcane; in caterpillar nests of Tetralopha scabridella on
Inga vera; in old cotton bolls; on grapefruit (Wolcott, 1936). See also
Wolcott's (1950) comments under Plectoptera dorsalis.

Plectoptera vermiculata

Cuba.—On pine in palmetto region (Rehn and Hebard, 1927).

Polyphaga aegyptiaca

Algeria.—Nymphal females under decaying leaves at the end of November
(Lucas, 1849).
Transcaucasia.—In burrows in argillaceous cliffs along ravines. Females
often covered by attached clay particles, an indication, according to Bei-
Bienko, that this species is ecologically connected to compact clay soils or
at least does not avoid them (Bei-Bienko, 1950).
See also the section on desert habitats (p. 29).

Polyphaga saussurei

South-central Asia.—Occupies compact clay soils; distributed in drier
regions than P. aegyptiaca; frequently found near dwellings, in yards,
stables, and houses (Bei-Bienko, 1950).

Polyzosteria limbata

Australia.—Common, usually "resting among the foliage or sunning itself
on a fence or stumps, seldom or never hiding under bark or logs like most
of the species" (Froggatt, 1906).

Poroblatta spp.

Tropical America.—"The species of Poroblatta apparently live as borers in
stumps and logs in a manner similar to those of Cryptocercus Scudder in
the United States" (Gurney, 1937).

Pseudomops septentrionalis

Texas.—In dead-brush pile; not scarce in heavy weeds, sunflowers, etc., in
openings of river-plain jungle scrub (Hebard, 1917). It lives largely in
herbage (Hebard, 1943a).

Pycnoscelus surinamensis

Florida.—Under stones and rubbish; very abundant under coquina boulders
in woods at Key West (Rehn and Hebard, 1905). "This species is common
under planks, stones, and other debris on the ground ... also found at Long
Key in the dry fibres at the base of the petioles of a coconut palm" (Rehn
and Hebard, 1912). At Musa Isle, found burrowing in sand (Hebard, 1915).
In fallen leaves and decaying wood in xeric and mesic hammock habitats
(Friauf, 1953).
Hawaii.—The soil swarmed with young of various stages during the
summer (Illingworth, 1915). In soil about roots of pineapple under
mulching paper; feeding on pineapple roots (Illingworth, 1927, 1929).
Fakarava, Tuamoto Archipelago.—Numerous among dead leaves in tree
holes (Cheesman, 1927).
West Indies.—Under decayed stalks of sugarcane and in siftings from
mangrove swamps, Cuba. Under manure, bases of leaves of coconut palm,
litter, logs, and stones on coral rock and in bromeliads, Jamaica. Under
wood, tiles, and boards in stable yards; immature individuals bored into the
soil, Puerto Rico. (Rehn and Hebard, 1927.)

Barbados.—Frequents cane fields (Tucker, 1952).
Puerto Rico.—"Altho primarily a xerophytic species: collected among dry
stones on Mona Island, under dry cow dung at Boquerón, and under boxes
at Guánica, it is reasonably common in the more humid parts of Puerto
Rico" (Wolcott, 1950).
Virgin Islands, St. Croix.—Common under rubbish; frequently seen feeding
on chicken feces around chicken roosts (Beatty, 1944). By feeding on
chicken feces it may become the vector of the chicken eyeworm,
Oxyspirura mansoni, as described in the references cited on page 204.
Egypt.—Large numbers were found in moist soil at the site of a manure pile
(Chakour, 1942).
Germany.—Under greenhouse conditions the depth to which P.
surinamensis penetrated the soil was determined; 21 dug down to a depth of
8 to 10 cm., 3 dug down 10 to 12 cm., but only one dug 13 cm. below the
surface. Often the tubes in the soil ended in a chamber which the cockroach
might not leave for several days; nymphs molted in such chambers and
females bore their young there (Roeser, 1940).

Rhytidometopum dissimile

Trinidad.—Male on low herbage in orchard at night; under sacking; on
Hibiscus at night (Princis and Kevan, 1955).

Riatia orientis

Trinidad.—Numerous specimens of both sexes at night on roadside
Hibiscus rosa-sinensis or low herbage in orchard (Princis and Kevan,
1955).

Simblerastes jamaicanus

Jamaica.—Numerous in fragmentary debris of an abandoned termite nest
on ground in the dry Liguanea Plain; a specimen was also taken under a
stone in a field of short grass (Rehn and Hebard, 1927).

Styphon bakeri

Costa Rica.—Among humus and rubble in crevices and large cavities in
rocks of the Tertiary limestone rim and the metamorphosed and igneous
rocks of the interior of the islands (Baker in Rehn, 1930).

Supella supellectilium

Virgin Islands, St. Croix.—Under rubbish heaps; in sugarcane straw (Beatty,
1944).
Africa.—"A cosmotropical species which occurs both out of doors and as a
household pest in many warmer parts of the world. It is apparently endemic
to non-forested areas in much of Africa north of the Equator." (Kevan and
Chopard, 1954.)

Symploce flagellata

Puerto Rico.—Under low trees on hillside and dead leaves in thicket of sea
grape (Hebard, 1916c).

Symploce hospes

Hawaii.—Under stones and rubbish (Illingworth, 1915).
Virgin Islands, St. Croix.—Under rubbish and on shrubbery at night (Beatty,
1944).

Symploce jamaicana

Jamaica.—In dead leaves under acacia and other shrubs in desert tract;
under log and rubbish in open on limestone sand near beach (Hebard,
1916c). Very common in short dry grass in roadside gutter at night, often
clustered together; under beach trash in stony wash of Hope River (Rehn
and Hebard, 1927).

Symploce ruficollis

Virgin Islands, St. Croix.—Under rubbish and on shrubbery at night (Beatty,
1944).
Puerto Rico.—In siftings from sea-grape thicket on sandy soil (Rehn and
Hebard, 1927). Often living under leaf-sheaths of sugarcane (Wolcott,
1950).

Tartaroblatta karatavica

Asia, Kara-tau Mountains.—Many hundreds of individuals found only
under stones on moist earth and not where ground seemed dry; found on
very stony slopes with sparse vegetation, often with undergrowth present
(Bei-Bienko, 1950).

STRUCTURAL HABITATS

In this category we include all man-made structures, whether inhabited by
man or not, that may become infested with cockroaches. A nonexhaustive
list of such structures would include dwellings, restaurants, mess halls,
barracks, groceries, markets, bakeries, dairies, drug stores, department
stores, hotels, hospitals, warehouses, mills, factories, packing houses,
animal houses, breweries, incinerators, privies, sewers, sewage treatment
plants, ships, aircraft, etc. Although dwellings are only one of the many
kinds of structures that are colonized by cockroaches, the several species
that have adopted this mode of life are generally referred to as domiciliary
cockroaches. This term is adequate only if we remember that these
cockroaches are not restricted to domiciles but are pests in other structures
as well.
Associations between man and certain species of cockroaches possibly
started as casually as the short-lived association that Beebe (1953) observed
when he discovered three cockroaches in the newly built couch of an orang-
utan. Obviously, when man came down from the trees, his fellow travelers
found his cave dwellings and other abodes particularly favorable habitats.
From such primitive beginnings, domiciliary cockroaches have spread into
every kind of structure that man has since devised. We predict that when

man develops a suitable vehicle, cockroaches will someday accompany him
into space. Yet despite the apparent predilection of certain species of
cockroaches for man, man is only incidental to these associations. Only the
shelter and food that man unwittingly provides for these unwelcome guests
attract cockroaches to him; man's physical presence is unnecessary.
Most, if not all, of the common domiciliary cockroaches apparently
originated in the Tropics or sub-Tropics from whence they have spread,
through normal commercial channels, into most of the inhabited world. At
least eight domiciliary cockroaches originated in Africa (Rehn, 1945):
Blatta orientalis, Blattella germanica, Leucophaea maderae, Nauphoeta
cinerea, Oxyhaloa buprestoides, Periplaneta americana, P. australasiae,
and Supella supellectilium; and, perhaps, Periplaneta brunnea as well;
Neostylopyga rhombifolia was probably of Indo-Malayan origin;
Pycnoscelus surinamensis was of oriental origin; and Leurolestes pallidus
was endemic in the West Indies (Rehn, 1945). Princis (1954a) rejected
Africa as the original home of Blatta orientalis and advanced reasons for
placing its origin in Central Asia.
Several domiciliary species have become well established in temperate
zones and some even in the Arctic. Bei-Bienko (1950) listed the following
10 species as sinanthropes in the Palearctic zone: Blatta lateralis, B.
orientalis, Blattella germanica, Leucophaea maderae, Periplaneta
americana, P. australasiae, Polyphaga saussurei, Pycnoscelus
surinamensis, and Supella supellectilium. In the warmer parts of the
temperate regions, as in their native Tropics, certain domiciliary species
breed outdoors as well as indoors. In the less temperate extensions of their
ranges most domiciliary species are nearly always found indoors. In regions
with low winter temperatures these cockroaches do not survive in unheated
structures; but in heated buildings Blattella germanica, for example, has
been able to withstand the rigorous climate of Alaska, where it has caused
severe infestations (Chamberlin, 1949).
The limiting factors that determine whether man-made structures will
provide suitable habitats for cockroaches are favorable temperature and
availability of water and food. The range of temperatures that man provides
for his own comfort and protection fosters the rapid increase of cockroach
populations indoors. Gunn (1934, 1935) has demonstrated that the preferred
temperature range (zone of indifference) of Blatta orientalis is 20-29° C.

The upper limit of the preferred temperature of Blattella germanica and
Periplaneta americana is 33° C. (Gunn, 1935). The lower limits of
temperature tolerance were not sharply defined in Gunn's work. However,
less than optimum temperatures, if they last for only short periods, are not
necessarily lethal. The 24-hour mortality for P. americana that had been
held for one hour at 0° C. was only 2±2 percent (Knipling and Sullivan,
1957). Gunn (1934) observed that Blatta orientalis would not settle at
temperatures above 33° C. and would react violently against higher
temperatures (e.g., 39° C.) by running away; thus the thermotactic behavior
of cockroaches might be presumed to bring them into favorable
environments within structures. Thermal death points have been determined
for the above three species by Gunn and Notley (1936).
It is common knowledge among those who rear cockroaches experimentally
that, unless the water content of the food is high, fluid water is essential in
the insects' dietary. Ten species of domiciliary cockroaches have been
shown to be unable to survive as long on dry food alone as they could on
food and water at 36-40 percent relative humidity (Willis and Lewis, 1957).
Blatta orientalis, when in a state of normal water balance, usually spent
more time in the drier part of a humidity gradient; but desiccated insects
tended to become hygropositive (Gunn and Cosway, 1938). We presume
that other domiciliary species behave similarly. If water is available nearby,
it may be presumed that partially desiccated cockroaches could locate a
source through the mediation of a humidity sense. Hygroreceptors have
been demonstrated on the antennae of Blattella germanica (Roth and Willis,
1952a) and suggested for Blatta orientalis (Gunn and Cosway, 1938).
Drinking water is available to cockroaches in the traps of sinks, wash
basins, tubs, and toilet bowls; in flush tanks; as condensation on cold pipes,
flush tanks, and windows; around leaking pipes and faucets; as spillage; in
miscellaneous water-filled containers, such as pet drinking dishes, aquaria,
vases; empty beverage bottles; and drainage from ice boxes. Soft, juicy
fruits and vegetables can provide both moisture and food. There seems to be
a tendency for certain species (Blatta orientalis and Blattella germanica) to
become established in the more humid parts of structures, such as
basements, around sinks, and in bathrooms. Whether this is a reaction to a
preferred humidity or merely a fortuitous aggregation near sources of
drinking water and food has never been clearly demonstrated. The rather

widespread dissemination of these species into zones of low as well as high
humidity suggests that detailed studies of the microclimatic conditions of
structural microhabitats will be needed before meaningful conclusions can
be drawn about the stratification of cockroaches within structures according
to species.
In nearly all structures infested by cockroaches, food of some kind is
available, either in the structure itself or nearby. This may be the food
stored by man for his own use or the use of kept animals; it may be crumbs,
food spillage, garbage, or excreta; glues and pastes on cartons, boxes,
stamps, envelopes, labels, and wall paper; sizing on cloth and book covers;
various dried animal and plant products; dead insects; living plants; etc. In
fact, it is almost impossible, despite good housekeeping, to keep any
structure used by man free of all food suitable for cockroaches.
That the requisite temperature, water, and food are provided, more or less
adequately, by a variety of structures is attested by the innumerable
infestations of cockroaches that develop when control measures are relaxed.
Within structures the accessibility of certain harborages to cockroaches
probably depends on the habits of the species and to some extent on their
size. Similar types of harborages in different structures may be used by the
same species, although there seems to be some overlapping by different
species into the same kinds of daytime shelters. The comparative ecology of
domiciliary cockroaches has not been thoroughly investigated, so any
interpretation of observational data is necessarily speculative and
inconclusive at this time. Our discussion on pages 324 to 343 is also
pertinent to this section.

LAND-BASED STRUCTURES

Dwellings provide a variety of microhabitats that are acceptable to
cockroaches. It has been stated that old houses, or houses that have many
cracks and crevices, or have basement kitchens that are not kept clean and
in good repair are particularly liable to invasion by cockroaches (Laing,
1946; British Museum [Nat. Hist.], 1951). Although this statement is
undoubtedly true, it has been our personal experience, as well as the

experience of others, that new, clean, and well-planned houses and
apartments are also easily and sometimes quickly invaded by cockroaches.
Mallis (1954) has cited the following places that are frequently infested by
cockroaches in homes. In the kitchen, cockroaches are found in and around
sinks, in cupboards above and below sinks, under tables and chairs, in
stoves, around breadboards, in utility cabinets, in kitchen closets, under
linoleum, behind, under, and inside refrigerators and iceboxes. In living
rooms cockroaches are found in furniture, studio couches, sewing
machines, closets, and bookshelves; behind picture frames, pennants,
calendars, and other wall ornaments. In bathrooms cockroaches are found in
and behind utility cabinets and toilets; they may be found in wicker clothes
hampers, in brooms and mops, and in door hinges. Ordinarily, cockroaches
are not found in bedrooms unless they are abundant elsewhere in the
dwelling. Additional harborages are cited under specific cockroaches in the
list below.
In markets DeLong (1948) found the German cockroach in bags of potatoes
and onions, in crates of citrus fruits, in pads and shredded papers in banana
boxes, and in cases of bottled beverages. The insects were attracted by
coffee and crawled into the folds of coffee bags. They were found in cartons
of canned goods; in bread and baked goods; in cartons of packaged cookies,
cakes, and crackers. Packaged cereals were attractive, and cockroaches
were sometimes found in packages of cigarettes. The insects occurred in
scales (by the hundreds) and in cash registers. Rather heavy infestations
were found under stainless-steel cappings that covered wooden arms on the
fish cleaning stand. The insects were numerous in display cases where they
were warm and sheltered. They were also found behind mirrors above
produce racks, in electrical switch boxes and conduits, and in telephone
boxes, as well as generally in cracks and behind loose moldings or loose
wall boards. Enclosed boxlike tables were frequently heavily infested.
In restaurants cockroaches may be found in the following places: Crevices
in wood, plaster, concrete, and metal; in the bar; in the kitchen and in the
associated equipment; in cupboards, lavatories, and garbage storage areas;
and on the undersides of chairs and tables (Mallis, 1954).
In drug stores Frings (1948) found cockroaches behind the mirror and
between the sink and the cooler. Thousands were found in hollow

ornamental shelf edging. The hollow bases of malted-milk dispensers and
drink mixers were cockroach havens.
In a hospital Frings (1948) found cockroaches in decorative trim around
doorways, by the thousands in wicker laundry baskets, and in incubators for
premature babies. In military hospitals we have seen cockroaches (Blattella
germanica) in kitchens and dining halls in the usual hiding places
mentioned above and on the undersides of stainless-steel serving tables.
In department stores cockroaches have been found in food departments,
beauty salons, rest rooms, dressing rooms, linen departments, and stationery
departments (Anonymous, 1952). The infestation in the linen department
was traced to clean towels which, when returned from the laundry,
contained at least 500 cockroaches per bundle. The insects were carried into
the rest rooms and beauty salon when the towels were distributed.
The microhabitats of cockroaches in privies and sewers have not been
studied. These habitats are particularly important in view of the
demonstrated migrations of cockroaches from sewers into dwellings and the
possible dissemination of pathogenic microorganisms from feces to food.
The reader is referred to our 1957(a) paper for a summary of the known
information on cockroach dispersal from sewers.

COCKROACHES ASSOCIATED WITH LAND-BASED
STRUCTURES

Most of the cockroaches listed below are either known domiciliary species
or they have been found one or more times in houses or other man-made
structures. The known structural pests breed within the building. Certain
other species, which have been observed only infrequently in structures and
are not known to breed there, may possibly be incipient pests; these latter
species may attain future economic importance if they establish breeding
colonies within a structure. A few species have undoubtedly wandered
indoors by accident. It is difficult to decide whether a particular species was
an accidental invader or whether it was attracted indoors in response to
some stimulus. Only additional information will provide the desired
answers.

Aglaopteryx ypsilon

Trinidad.—Male found indoors (Princis and Kevan, 1955).

Allacta similis

Hawaii.—Found only indoors at Nauhi. Otherwise this is apparently an
outdoor species (Swezey and Williams, 1932).

Blaberus craniifer

Cuba.—Household pest (Deschapelles, 1939). Particularly abundant in
houses in Santiago and Havana (Rehn and Hebard, 1927).
Florida.—Under boards in woodshed (Rehn and Hebard, 1912, 1914).

Blaberus discoidalis

Ecuador.—In eating places (Campos R., 1926).
Hispaniola.—In houses (Rehn and Hebard, 1927).
New Jersey.—In greenhouse (Weiss, 1917).
Puerto Rico.—In homes (Seín, 1923). In fruit stores (Wolcott, 1950).

Blatta lateralis

Central Asia.—Household pest, often found in homes with clay floors (Bei-
Bienko, 1950).
Turkmen S.S.R.—Males and females occurred in dwellings (Vlasov, 1929).

Blatta orientalis

This species is a cosmopolitan domiciliary pest (Hebard, 1917; Rehn,
1945). It is reported to occur particularly in basements and crawl spaces
under basementless houses (Mallis, 1954). In damp basements where food

is available large colonies are not unusual, but it also may infest offices and
apartments several floors off the ground (Gould and Deay, 1940). The
number encountered on upper floors is seldom large, but the frequency of
occurrence may reach 30 percent of the observations (Spear et al., in
Shuyler, 1956). In supermarkets this species hides during the day inside
concrete blocks or cracks in the foundation, under furniture, or behind
cartons; it is conspicuous on the floors of the markets at night (De Long,
1948). In Great Britain the kitchen is preferred by this pest (and by Blattella
germanica); they shelter beneath steam radiators and gas stoves, behind
hot-water pipes, underneath furniture and floor coverings, sinks and baths;
basements and underground kitchens are especially likely to be infested
(Laing, 1946; British Museum [Nat. Hist.], 1951). Goodliffe (1958) noted
that B. orientalis may travel long distances to find food.

Blattella germanica

This species is a cosmopolitan domiciliary pest (Hebard, 1917; Rehn,
1945). It is one of the commonest insects in homes and restaurants (Gould
and Deay, 1940). It is found in kitchens, larders, bathrooms, furnace rooms,
and storage rooms of bakeries, breweries, hospitals, barracks, as well as
dwellings, where, during daylight, it hides behind cupboards, furniture,
hanging pictures, panels and skirting boards, in cracks around drains, water
pipes, electric wires, and hot-water and steam heating units (Wille, 1920).
The German cockroach may be found in cracks around baseboards, pipes,
conduits, sinks, and drawers; behind cabinets; inside switch boxes and
refrigerators; on under surfaces of tables, chairs, and shelves; between
stacks of stored goods, and in almost every place that is not readily
observed (Kruse, 1948). We have also seen this species packed in electric-
clock cases and loud-speaker baffles, in cash registers, and clinging to the
undersurface of stainless-steel steam tables. The infestation of markets by
this species has been described above. Very narrow cracks provide refuges
for the German cockroach. Wille (1920) found first-instar nymphs in cracks
0.5 mm. wide and adult males and females without oöthecae in cracks 1.6
mm. wide.
Shuyler (1956) has observed extensions into relatively new structural
habitats by Blattella germanica in the north-central area of the United

States. A few German cockroaches are now being encountered in living
rooms, bedrooms, clothes closets, bedroom furniture, lobbies, entrance
halls, checkrooms, nonfood storerooms, nonfood warehouses, and coin-
vending machine repair shops. In these situations this species is behaving
much like the brown-banded cockroach, Supella supellectilium.

Blattella schubotzi

Cameroon.—Five specimens in a house (Princis, 1955).

Blattella vaga

U.S.A.—Although this is mostly an outdoor species, during dry seasons it
may temporarily enter houses in great numbers, occasionally breeding
indoors in Arizona (Flock, 1941a). Two adults were collected indoors in
Texas (Riherd, 1953).

Chromatonotus notatus

Trinidad.—A male was found indoors (Princis and Kevan, 1955).

Cutilia soror

Hawaii.—Almost as common in houses as Neostylopyga rhombifolia
(Hebard, 1922).

Ectobius duskei

U.S.S.R.—Frequently occurs in living apartments in farming localities as an
accidental inhabitant (Bei-Bienko, 1950).

Ectobius pallidus

Massachusetts.—A summertime pest in houses along coast (Gurney, 1953;
E. R. Willis, personal observation). Generally an outdoor species.

Epilampra abdomen-nigrum

Trinidad.—Male, indoors (Princis and Kevan, 1955). An outdoor species
generally.

Ergaula capensis

Cameroon.—A male taken in a house (Princis, 1955).

Eublaberus posticus

Trinidad.—Indoors, feeding on bat feces (Princis and Kevan, 1955).

Eurycotis floridana

Florida.—Occasionally found in homes (Creighton, 1954; Roth and Willis,
1954a).

Euthyrrhapha pacifica

Hawaii.—Found outdoors and indoors where it breeds in neglected
cupboards and in rubbish (Fullaway and Krauss, 1945).

Holocompsa azteca

Mexico.—Household pest (Ball et al., 1942).

Holocompsa cyanea

Costa Rica.—One specimen in house (Rehn, 1906).

Holocompsa nitidula

Apparently domiciliary in American Tropics (Hebard, 1917). In houses
under chests, etc., Cuba (Gundlach, 1890-1891); Puerto Rico (Gundlach,

1887). In folds of burlap bag, Florida (Rehn and Hebard, 1914).

Ischnoptera rufa occidentalis

Panama.—Thrives about human habitations under litter, though not
domiciliary (Hebard, 1920).

Ischnoptera rufa rufa

Jamaica.—In hotel. "While hardly a domiciliary form it would seem to
frequent environments where man has considerably disturbed natural
conditions, as under debris, docks, under logs and stones in cultivated
areas" (Rehn and Hebard, 1927).

Leucophaea maderae

West Indies.—In habitations, warehouses, and other structures; "At times it
is a very abundant and serious pest" (Rehn, 1945). In Puerto Rico it was
also found in fruit stores, markets, and inns (Seín, 1923; Wolcott, 1950).
Reported as a domiciliary pest in Madeira (Heer, 1864); Windward Islands
(Marshall, 1878); Tropics and sub-Tropics (Rehn, 1937); Philippine Islands
(Uichanco, 1953); New York City (Anonymous, 1953; Gurney, 1953);
Trinidad (Princis and Kevan, 1955). This species is also established in
coastal Brazil, Central America, all the Greater Antilles, several other
tropical islands, and tropical Africa, where it probably originated (Rehn,
1945).

Leurolestes circumvagans

Hispaniola, Grenada.—Largely domiciliary (Rehn and Hebard, 1927).

Leurolestes pallidus

Cuba.—All over island, in houses, under lockers, etc. (Rehn, 1945;
Gundlach, 1890-1891).

Florida.—Rehn and Hebard (1914).
This species has been recorded from various islands in the West Indies,
from Mexico, Guatemala, and Brazil (Rehn, 1945).

Methana marginalis

Australia.—Reported entering houses (Pope, 1953a).

Nauphoeta cinerea

Australia.—In hospital (Mackerras and Mackerras, 1948). In dwellings,
grain stores, and fowl-feeding pens (Pope, 1953).
Sudan.—Domiciliary in huts of the Shilluk natives; fairly widely distributed
in eastern Africa (Rehn, 1945).
Hawaii.—In feed rooms of poultry plants (Illingworth, 1942).
Florida.—Established in feed mills around Tampa (Gresham, 1952;
Gurney, 1953).
The wide distribution of this species from East Africa, where it originated,
to the Orient and the New World was undoubtedly mediated by shipping
(Rehn, 1945).

Neoblattella sp.

Puerto Rico.—Observed [as Blatta caraibea, Rehn and Hebard (1927)] in
houses (Gundlach, 1887).

Neostylopyga rhombifolia

Domiciliary in Indo-Malaya and New World Tropics (Rehn, 1945);
Philippine Islands (Uichanco, 1953); and Hawaii (Hebard, 1922).

Oxyhaloa buprestoides

Presumably to some extent domiciliary, as it evidently spread from Africa
to the New World via slave ships (Rehn and Hebard, 1927; Rehn, 1945).

Panchlora nivea

Colombia.—A male and a female taken in a dwelling (Princis, 1946). This
is primarily an outdoor species which is frequently taken indoors as an
adventive on bananas (see p. 150 for references).

Parcoblatta fulvescens

Florida.—Males found in laboratory and dormitory buildings, ostensibly
attracted by lights (Friauf, 1953).

Parcoblatta lata

Connecticut.—Domiciliary pest (Moore, 1957). Generally an outdoor
species.

Parcoblatta notha

Arizona.—It may occasionally be a nuisance in houses (Ball et al., 1942).

Parcoblatta pensylvanica

U.S.A.—Country houses often badly infested, Indiana (Blatchley, 1920).
Frequently taken in houses in wooded areas, Michigan (Hubbell, 1922).
Infestation by males, females, and nymphs on fourth floor of building,
South Dakota (Severin, 1952). Houses in wooded areas infested by nymphs
and occasionally by adults (Gould and Deay, 1940).
Canada.—Pest in summer cottages in Ontario (Walker, 1912).

Periplaneta americana

This is a cosmopolitan domiciliary pest (Hebard, 1917; Rehn, 1945). It is
common in restaurants, grocery stores, bakeries, and where food is prepared
or stored; it was trapped regularly in the basement and upper floors of store
buildings, and it was also found in all heated parts of an old meat-packing
plant (Gould and Deay, 1938). P. americana was numerous in latrines in
Iran (Bei-Bienko, 1950) and in privies in Texas (Dow, 1955) and Georgia
(Haines and Palmer, 1955). Large numbers of this species also occur in
sewers adjacent to human habitations (Roth and Willis, 1957a).

Periplaneta australasiae

Generally domiciliary, but also occurs outdoors in the West Indies (Rehn
and Hebard, 1927). It is a very abundant domiciliary pest in tropical Africa
and tropical America (Rehn, 1945); Ecuador (Campos R., 1926); Puerto
Rico (Sein, 1923); Philippine Islands (Uichanco, 1953); Australia (Pope,
1953).
Also occurs as a pest in greenhouses in Pennsylvania (Thilow and Riley,
1891); France (Giard, 1900); Italy (Boettger, 1930); Great Britain (Laing,
1946; British Museum [Nat. Hist.], 1951).

Periplaneta brunnea

Circumtropical domiciliary pest which is apparently more nearly peculiar to
the Tropics and adjacent regions than P. americana (Hebard, 1917). This
species has been trapped in significant numbers in privies and dwellings in
Georgia (Haines and Palmer, 1955). It is the species of Periplaneta present
in homes in San Antonio, Texas (Mallis, 1954).

Periplaneta fuliginosa

U.S.A.—Frequently encountered out of doors, but has been reported
common after dark about a hotel in Alabama and was captured in a house in
Louisiana; it was also extremely abundant on the wharves at night in
Jacksonville, Florida (Hebard, 1917). As a domiciliary pest it was, next to
Blattella germanica, the most common cockroach inside homes in
southwest Georgia, where it was also the most common cockroach in

privies (Haines and Palmer, 1955). This species has become a very common
domiciliary pest in Texas (Eads, personal communication, 1955). It infested
a greenhouse for five years in Indiana (Gould and Deay, 1940).

Periplaneta ignota

Australia.—It occurs in dwellings occasionally (Pope, 1953).

Phaetalia pallida

Colombia.—Three specimens from three dwellings (Princis, 1946).
Trinidad.—Male indoors; male and female at light (Princis and Kevan,
1955).

Plectoptera dorsalis

Puerto Rico.—According to Gundlach (1887) it enters houses at night
attracted by light (Rehn and Hebard, 1927).

Polyphaga aegyptiaca

Iraq.—Common in houses (Weber, 1954).
Caucasus.—Winged male in kitchen (Burr, 1913).
U.S.S.R.—Listed as a sinanthrope (Bei-Bienko, 1950).

Polyphaga saussurei

South-central Asia.—One of the commonest domiciliary species (Bei-
Bienko, 1950).

Pseudophoraspis nebulosa

East Indies.—This species is sometimes difficult indoors (Karny, 1925).

Pycnoscelus surinamensis

A household pest in the East Indies (Karny, 1925); Philippine Islands
(Uichanco, 1953); Tanganyika (Smith, 1955); Trinidad, eight records
indoors (Princis and Kevan, 1955). It is also a greenhouse pest (Hebard,
1917; Zappe, 1918; Doucette and Smith, 1926; Saupe, 1928; Roeser, 1940).
Common in or around chicken batteries and yards in Hawaii (Schwabe,
1949).

Supella supellectilium

Domiciliary wherever distributed (Rehn, 1945), this species is especially
difficult to control because of its apparently nonselective dispersal
throughout dwellings. For example, Mallis (1954) observed in Texas that it
was widely distributed throughout the apartment and was probably the most
common cockroach seen in the bedroom; its favorite harborages were
beneath and behind corner braces on kitchen chairs, underneath tables,
behind pictures and other objects on walls, and in shower stalls; its oöthecae
were commonly fastened on walls and ceilings throughout the house. Gould
and Deay (1940) reported that this species prefers high locations, such as
shelves in closets, behind pictures, and picture molding; oöthecae were
found about kitchen sink, desks, tables, and other furniture, and even in
bedding. Hafez and Afifi (1956) stated that the adult wanders in nearly all
rooms of the house and only visits the kitchen when searching for food; it
hides in cupboards, pantries, closets, bookshelves, drawers, and behind
picture frames; the nymphs normally hide in the corners of drawers, behind
frames, and in similar situations.

Symploce bicolor

Puerto Rico.—In houses, Sardinera Beach, Mona Island (Ramos, 1946).

Symploce hospes

North American Tropics.—Domiciliary, but not exclusively so, and
apparently widely distributed (Hebard, 1917). In Florida, as Ischnoptera

rufescens, found in a greasy cupboard (Rehn and Hebard, 1914).
Hawaii.—Illingworth (1915).

SHIPS

Sailing ships have long been notorious for their unwelcome hordes of
cockroaches, and it was by ship that at least 11 domiciliary species migrated
from their centers of origin to other parts of the world (Rehn, 1945). Over
40 nondomiciliary species have been carried by ship from the American
Tropics to other parts of the world in cargoes of bananas (p. 146). In
addition to these, other adventive cockroaches appear from time to time in
ports to which they have been carried by ships. Yet by far the most
numerous cockroaches on shipboard are the breeding populations of a few
common domiciliary pests. Except in the most rigorously disinsectized
ships, this commerce in cockroaches has continued to the present day.
Cockroaches undoubtedly infested the first ships that sailed the
Mediterranean; of these we have no records. The earliest recognizable
record of cockroaches on shipboard is Moffett's (1634) statement that when
Drake captured the ship Philip, he found it overrun with cockroaches
[Blattarum alatarum]. Bligh (1792) described disinfesting H.M.S. Bounty
with boiling water to kill cockroaches. Chamisso (1829) reported that he
had seen ships casks, in which rice or grain had been stored, that were
found to be filled with Blattella germanica when opened. During a voyage
from England to Van Diemen's Land, Lewis (1836) was greatly annoyed by
hundreds of cockroaches flying about his cabin at night; the most numerous
resembled Periplaneta americana and another was similar to Ectobius
lapponicus. This latter was undoubtedly B. germanica, which is the only
ship-infesting species that resembles the feral E. lapponicus. Lewis
continued, P. americana "were in immense profusion, and had
communication with every part of the ship, between the timbers or skin.
The ravages they committed on everything edible were very extensive; not
a biscuit but was more or less polluted by them, and amongst the cargo 300
cases of cheeses, which had holes in them to prevent their sweating, were

considerably damaged, some of them being half devoured and not one
without some marks of their residence."
Kingsley (1870), Kellogg (1908), Gates (1912), Heiser (1936), and Bronson
(1943) have all reported that cockroaches were so numerous on ships that
they gnawed the skin and nails of the men on board. These are all
independent observations of what may well have been a common
occurrence on ships. We have discussed in detail the subject of cockroaches
biting man in our 1957(a) paper.
Mosely (1892) reported, "At the time that England was left the ship [H.M.S.
Challenger] seemed nearly free of animals, other than men, dogs, and
livestock required for food. The first cockroaches apparently came on board
at St. Vincent, Cape Verdes.... Cockroaches soon became plentiful on board,
and showed themselves whenever the ship was in a warm climate.
"At one period of the voyage, a number of these insects established
themselves in my cabin, and devoured parts of my boots, nibbling off all the
margins of leather projecting beyond the seams on the upper leathers."
Sir Edmund Freemantle (1904) recalled some of his experiences in the
British Navy. "Cockroaches in the tropics were also terrible scourges. One
saw little of them in fine, dry weather, but in damp, wet weather they
seemed to come from every hidden corner ... our remedy in the 'Spartan'
was to make the boys catch them—on pain of being caned.... One brig, the
'Lily,' was so overrun by cockroaches that the officers' clothes smelt of
them."
The quotation from Sonan (1924) on page 348 describes similar conditions
in the Japanese Navy.
On modern cargo ships cockroaches are reported to be extremely numerous
in the galley, the crew's quarters, and sometimes in the holds; they dwell in
hot, humid environments such as the casing around steam pipes (Monro,
1951). Williams (1931) reported that Blattella germanica was the most
important cockroach pest on ships seen at New York. Although often
numerous in the holds, the cockroaches as a rule congregated in living
quarters. They were also frequently found between tarpaulins covering the
hatches. It was not unusual to kill 20,000 to 50,000 in the forecastle, and
more than 20,000 have been taken from a single stateroom. Simanton

(1946) inspected the S.S. William Kieth when it berthed at San Francisco
from a 10-month voyage to the South Pacific. The holds were infested with
thousands of Periplaneta americana, but in the crew's quarters, mess halls,
and storerooms B. germanica predominated. After insecticidal treatment
about 2,000 P. americana were seen in each hold and as many as 24 B.
germanica in each cabin. Richardson (1947) reported that in Army
transports inspected between 1943 and 1946 at New York, P. americana
was found in the galleys and messes, and occasionally heavy infestations
were found deep in the holds; B. germanica was found in the galleys and
messes; Blatta orientalis was found only in the hold.
Additional references indicating the presence of Blattella germanica on
ships may be found in the account of its parasite Ripidius pectinicornis (p.
232). Although Rice (1925) and Williams (1931) cite B. germanica as the
most numerous cockroach on ships, Brooke (1920) stated that the great
majority of ship cockroaches were Periplaneta americana. In addition to
citing cockroach infestations on ships, the following authors reported
various methods for disinfecting ships: Canalis (1916), Pryor (1918),
Brooke (1920), Rice (1925), Williams (1931), Simanton (1946), Richardson
(1947), and Anonymous (1951, 1954).

COCKROACHES ASSOCIATED WITH SHIPS

In the following list we include some previously unpublished data on
cockroaches that were recovered from ships at the Miami, Fla., Quarantine
Station for the periods November 1945 through May 1946; May, June,
August, and September 1950; and 17 July 1957 (Porter, personal
communication, 1958). These data were lumped, without breakdown to
species, under the entry Orthoptera in Porter (1958).
Certain of the species listed below occur only accidentally on shipboard and
will probably never establish breeding colonies on ships or become pests on
shipboard or elsewhere; some were merely passengers between one land-
based colony and another. Others, the truly domiciliary pests, are as likely
to be pests on shipboard as they are in land-based structures.

Blaberus discoidalis

Hispaniola.—On board ship (Rehn and Hebard, 1927).

Blatta orientalis

U.S.A.—At Port of New York (Richardson, 1947).

Blattella germanica

At sea?—In ships casks (Chamisso, 1829).
U.S.A.—Port of New York (Williams, 1931; Richardson, 1947). San
Francisco (Simanton, 1946). At Miami, 7,852 live specimens recovered
from ships (Porter, personal communication, 1958). Most numerous species
on ships (Rice, 1925).

Epilampra maya

At sea.—One male and one female found dead on S.S. Tenadores (Hebard,
1917).

Epilampra sp.

Florida.—One dead specimen, Miami (Porter, personal communication,
1958).

Ischnoptera sp.

Florida.—Five live and one dead specimen, Miami (Porter, personal
communication, 1958).

Latiblattella sp.

At sea.—One female alive in hold of S.S. Tenadores (Hebard, 1917).

Leucophaea maderae

Brought from West Africa to West Indies and Brazil by slave ships (Rehn,
1945).

Nauphoeta cinerea

Widely disseminated by sailing ships (Rehn, 1945).

Neoblattella fratercula

At sea.—Two females found dead on S.S. Tenadores (Hebard, 1917).

Neoblattella fraterna

At sea.—One male found dead in hold of S.S. Tenadores (Hebard, 1917).

Neoblattella nahua

At sea.—One female dead in hold of S.S. Tenadores (Hebard, 1917).

Neoblattella sp.

Florida.—Five dead specimens recovered from ships, Miami (Porter,
personal communication, 1958).

Neostylopyga rhombifolia

Widely distributed by sailing ships (Rehn, 1945).
England.—Captured on a sugar vessel from Java (Lucas, 1920).

Nyctibora noctivaga

At sea.—One male and one female nymph found dead on S. S. Tenadores
(Hebard, 1917).

Nyctibora sp.

Florida.—Two dead specimens recovered from ships at Miami (Porter,
personal communication, 1958).

Oxyhaloa buprestoides

Spread from Africa to New World by ships (Rehn and Hebard, 1927; Rehn,
1945).

Panchlora nivea

At sea.—One female dead in hold of S.S. Tenadores (Hebard, 1917).
Florida.—Fifteen dead specimens taken from ships, Miami (Porter,
personal communication, 1958).
From the numerous records of this species as an adventive taken on bananas
(p. 150), it may be presumed to be a frequent traveler on banana boats.

Periplaneta americana

At sea.—Lewis (1836). Hebard (1933a) stated that this is "often a serious
pest on the smaller ships sailing the South Seas."
U.S.A.—San Francisco (Simanton, 1946). Port of New York (Richardson,
1947). At Miami, 62 live and 123 dead specimens (Porter, personal
communication, 1958).

Periplaneta australasiae

Migrated from West Africa to America in slave ships (Rehn, 1945).

Pycnoscelus surinamensis

Probably in part reached the New World by way of Africa in slave ships
(Rehn, 1945).

Supella supellectilium

Reached America from West Africa by slave ship (Rehn, 1945).

Xestoblatta festae

At sea.—One female found dead in hold of S.S. Tenadores (Hebard, 1917).

AIRCRAFT

Michel (1935) stated that the development of air transportation brought the
same insect dispersal problems that exist in land and water transportation;
in addition, the problem of cockroach infestation had become a very serious
one, quite aside from the hygienic point of view, because it had been
discovered that these insects seek out the wings of airplanes, where they
subsisted on the glue and dope used in airplane construction. However,
Dethier (1945) found no cockroaches in dismantled or wrecked wing and
tail structures of metal aircraft in central Africa. In fact, all-metal aircraft
would seem to provide little in the way of food or water for stowaway
cockroaches.
Laird (1951, 1952, 1956a) found living specimens of Blattella germanica,
Periplaneta americana, and Periplaneta australasiae in baggage
compartments and/or kitchens in aircraft. Other species which have been
recovered from undisclosed spaces in aircraft are listed below. Some of the
cockroaches that were reported as dead may not have died from exposure
during flight but may have been killed by insecticide applied by inspecting
personnel at the airports.

COCKROACHES ASSOCIATED WITH AIRCRAFT

In the following list we include some previously unpublished data on
cockroaches that were recovered from aircraft in Miami, Fla., International
Airport from 1 July 1956 through 30 June 1957 (Porter, personal
communication, 1958). These data were lumped under the entry Orthoptera
without breakdown to species in Porter (1958).
Species reported by Hughes (1949), and cited below as from southern
United States, were recovered from aircraft that arrived at Brownsville, Fort
Worth, Miami, New Orleans, and San Juan. There was no way of linking a
specific record with any particular city.
The comments we made above about species that are infrequently
encountered on ships apply with equal validity to similar species found on
aircraft.

Anaplecta sp.

U.S.A.—One live and 15 dead specimens recovered from 16 aircraft at
Miami (Denning et al., 1947).

Blatta orientalis

U.S.A.—Six live and four dead specimens recovered from six aircraft at
Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949).

Blattella germanica

Hawaii.—Williams (1946a).
Khartoum.—Whitfield (1940).
New Zealand.—Laird (1951, 1952, 1956a).
U.S.A.—At Miami 193 live and 184 dead specimens were recovered from
141 aircraft (Denning et al., 1947). Recovered at airports in southern U.S.
(Hughes, 1949). Recovered at Miami, 51 live and 24 dead specimens
(Porter, personal communication, 1958). Exposed experimentally in jet
aircraft (Sullivan et al., 1958).

Blattella sp.

Khartoum.—Whitfield (1940).
Southern U.S.—Hughes (1949).

Cariblatta ssp.

U.S.A.—One live and three dead specimens recovered from three aircraft at
Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949).

Epilampra sp.

U.S.A.—One dead specimen, Miami (Denning et al., 1947). Southern U.S.
(Hughes, 1949).

Eublaberus posticus

Southern U.S.—Hughes (1949).

Ischnoptera rufa rufa

U.S.A.—Two dead specimens recovered from two aircraft at Miami
(Denning et al., 1947).

Ischnoptera sp.

U.S.A.—Two live and one dead specimen recovered from three aircraft,
Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949). One dead
specimen, Miami (Porter, personal communication, 1958).

Leucophaea maderae

U.S.A.—Three dead specimens recovered from three aircraft at Miami
(Denning et al., 1947). Southern U.S. (Hughes, 1949).

Nauphoeta cinerea

U.S.A.—One live and one dead specimen, Miami (Denning et al., 1947).

Neoblattella sp.

U.S.A.—One dead specimen recovered at Miami (Denning et al., 1947).

Panchlora nivea

U.S.A.—Two dead specimens recovered from two aircraft, Miami (Denning
et al., 1947).

Periplaneta americana

U.S.A.—Five live and three dead specimens recovered from seven aircraft,
Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949). Three live
and five dead, Miami (Porter, personal communication, 1958).
Experimentally exposed in jet aircraft, U.S. (Sullivan et al., 1958).
New Zealand.—Laird (1951, 1952).

Periplaneta australasiae

U.S.A.—Two dead specimens recovered from two aircraft, Miami (Welch,
1939). Five live and three dead from five aircraft, Miami (Denning et al.,
1947). Southern U.S. (Hughes, 1949). Three live and two dead, Miami
(Porter, personal communication, 1958).
New Zealand.—Laird (1952).

Periplaneta spp.

U.S.A.—One live and three dead specimens from four aircraft, Miami
(Denning et al., 1947). Southern U.S. (Hughes, 1949). One live and six
dead, Miami (Porter, personal communication, 1958).

Pycnoscelus surinamensis

U.S.A.—Two live and three dead specimens from five aircraft, Miami
(Denning et al., 1947). Southern U.S. (Hughes, 1949).
New Zealand.—Laird (1956a).

Supella supellectilium

U.S.A.—Two live and one dead specimen from one aircraft, Miami
(Denning et al., 1947). Southern U.S. (Hughes, 1949).
Khartoum.—Whitfield (1940).

Supella sp.

U.S.A.—Two live specimens from two aircraft, Miami (Welch, 1939).
Southern U.S. (Hughes, 1949).

Symploce sp.

U.S.A.—Two live and one dead specimen from three aircraft, Miami
(Denning et al., 1947). Southern U.S. (Hughes, 1949).

Unidentified cockroaches

Anglo-Egyptian Sudan.—At Khartoum (Whitfield, 1940).
Brazil.—From flying boats, 62 specimens; from land planes, 45 specimens
(Carneiro de Mendonça and Cerqueira, 1947).
Central Africa.—Dethier (1945).
Kenya.—At Kisumu (Symes in Whitfield, 1940).
New Zealand.—Laird (1956a).
U.S.A.—Four live and 14 dead specimens from 4 aircraft, Miami (Welch,
1939). One live and 15 dead specimens (adults?) from 16 aircraft; 8
oöthecae from 7 planes; 147 live and 83 dead nymphs from 108 planes,

Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949). Five live and
7 dead specimens, Miami (Porter, personal communication, 1958).

IV. CLASSIFICATION OF THE
ASSOCIATIONS
Asano (1937) classified the natural enemies of cockroaches into two types
as follows:

1. Enemies that feed mainly on cockroaches (certain ripiphorid
beetles and certain chalcid, evaniid, and ampulicid wasps).
2. Organisms which, in their search for food, devour cockroaches
that may be encountered (certain species of scorpions, spiders,
ticks, centipedes, Strepsiptera, ants, birds, rats, and "parasitic
bacteria").

Cameron (1955) arranged the associates of cockroaches in two groups as
follows:

Group A. Parasites and predators.
1. Parasites: Hymenoptera (Evaniidae, Eulophidae, Eupelmidae,
Encyrtidae, Pteromalidae, Cleonymidae) and Coleoptera
(Ripiphoridae).
2. Predators: Hymenoptera (Ampulicidae), Hemiptera
(Reduviidae), Coleoptera (Dermestidae), Arachnida (Araneae,
Acarina).

Group B. Parasites and symbionts.
1. Protozoa (including examples of both parasites and
commensals).

2. Nematoda (including both primary "parasites" and secondary
parasites).
3. Bacteria (including the mutualistic bacteroids).
4. Algae [Arthromitis (= Hygrocrocis) intestinalis; see p. 124].

Asano's arrangement, although essentially true, is limited; Cameron's
system is divided into arthropods (group A) and lower forms (group B), but
does not include higher animals. Both attempts at classification need
amplification; this we have endeavored to do below.
In classifying the biotic associates of cockroaches, we were immediately
confronted with a problem in semantics. The concepts parasitism,
predatism, and symbiosis have all been used with various shades of
meaning by different authors. The problem is not solved merely by
accepting as authoritative specific definitions, however apt they seem to be,
because, unfortunately, these concepts are not mutually exclusive. For
example, among the entomophagous insects, as Sweetman (1936) has
pointed out, there can be no definite line of separation between parasitism
and predatism: the two intergrade, only the extremes being quite distinct. In
fact, Andrewartha and Birch (1954) generalized these relationships by
calling both categories predatism. These authors divided natural populations
of associated organisms into nonpredators and predators. Although this
simplifies their presentation of the general principles of ecology, for our
purpose more narrowly defined terms have proved useful.
In the main we have followed Sweetman (1936) and Allee et al. (1949) in
arriving at the following definitions:
Symbiosis is the living together in more or less intimate association of
organisms of different species; it includes virtually all relationships between
cockroaches and other organisms, such as parasitism, predatism,
commensalism, and mutualism. Allee et al. (1949) apparently do not
include predatism in symbiosis.
Mutualism is symbiosis in which both members benefit by the association.
The smaller partner has commonly been called a symbiont or symbiote by
authors.
Commensalism includes associations in which neither party appears to
benefit or be harmed. One partner may live on the surplus food or wastes of

the other; shelter and transport may be involved.
Parasitism is the state of symbiosis in which one of the members feeds
upon the other during the whole of either the immature or mature feeding
stage; the host is harmed in some way and may be killed.
Predatism is an association in which one member attacks and feeds upon,
or stores as food for its progeny, one or more other organisms; the predator
spends less than the immature or mature feeding period on the prey. This
category includes a few invertebrates and all the vertebrates that capture,
kill, and feed on cockroaches. This association may be divided into
interspecies predatism, in which the predator preys upon a different species,
and intraspecies predatism (cannibalism) in which the predator preys upon
its own species.
Although we have attempted to adhere to these definitions throughout this
discussion, we realize that in doing so we may have tended to oversimplify
complex relations. Some questionable interpretations stem from insufficient
knowledge of the basic relationships between cockroaches and their
associates. Only further study will clarify these relationships. Some of the
problems are discussed below.
Probably many of the so-called parasites (e.g., Protozoa like Nyctotherus,
and intestinal nematodes of the family Thelastomatidae), which do not
invade the host's tissues and seem to have no effect on the activity and
vitality of the host, are commensals. Although we consider these forms to
be commensals, we realize that they might actually affect the host in some
way even though this has not been shown. It is possible that Rothschild and
Clay's (1957) statement about bird parasites may well apply to the
apparently harmless organisms found in the cockroach. These authors
wrote, "It cannot be too strongly emphasized that the effect of all types of
parasites on the host is detrimental. If we find that a bird seems little, if at
all, inconvenienced by the presence of Protozoa or worms or lice, or a
cuckoo in the nest, we can nevertheless assume that it would be better off
without them.... Small effects such as lack of vitality, loss of voice,
excessive blinking, or perverted habits like dirt eating are extremely
difficult to gauge. Nevertheless, it is only a question of degree. Potentially
all parasites are harmful." It should also be pointed out that some workers
would consider certain of our commensals of cockroaches to be parasites.

Thus Faust (1955) stated that "A truly successful parasite is one which has
developed a state of equilibrium with its host, so that no detectable damage
is produced which endangers the health or life of the host. In a suitable host
the parasite may obtain food and shelter without any evidence of trauma or
toxicity. The damage produced may be so slight that repair and functional
readjustment keep pace with the injury." Faust's successful parasite would
be indistinguishable from a commensal, but there is undeniably a difference
between an organism causing slight, and undetectable, damage to a host and
one causing none. Certain of the organisms we list as commensals may
eventually be shown to be parasites.
Certain organisms which live in cockroaches appear to have no effect on the
vitality of the host even though the tissues of the host are invaded.
Gregarines may penetrate the intestinal wall of the cockroach without
seeming to injure the host. Fungi of the genus Herpomyces invade the
cuticle of cockroaches producing pathological changes; yet the insects'
behavior is apparently unaffected (see p. 129). We consider these organisms
to be parasites because the host's tissues are invaded and, as far as we know,
no benefit to the host results.
In the literature certain insects have been considered to be either parasites
or predators or both. Among these are the ensign wasps (Evaniidae), whose
larvae feed on the eggs of cockroaches within the oötheca, and the
ampulicid wasps, which capture, paralyze, and store in their nests (as food
for their larvae) nymphs and adults of cockroaches. Clausen (1940) claimed
that the evaniid Zeuxevania splendidula is a true egg parasite when it
destroys the first egg in a cockroach oötheca; but after the wasp larva molts
and proceeds to devour the other eggs, he considered it to be a predator.
Clausen's definition of an entomophagous parasite is an insect that in its
larval stage develops either internally or externally upon a single host which
is eventually killed; with few exceptions the adults are free-living and their
food is usually different from that of the larvae. A predatory insect, by
Clausen's definition, is principally free-living in the larval as well as adult
stage, kills the host immediately by direct attack, and requires a number of
victims to reach maturity; the predator is of greater size than the prey, and
the food sources of the adults and immature stages are frequently the same.
It is apparent, as Clausen and other writers have pointed out, that there are
instances of a particular species showing characteristics which fit both the

definitions for predator and parasite. Thus, among the evaniids one wasp
larva destroys all the eggs in an oötheca, but in spite of this the larva has
more of the characteristics of a parasite than of a predator; the adult wasp
does not utilize the same food as the larva (adults have been taken on
flowers and on honeydew from scale insects). It is questionable whether the
evaniid larva kills the cockroach egg outright. The wasp larva, being
restricted to the inside of the oötheca, is not free-living. Probably the only
criterion by which the evaniid could be judged to be a predator, by
Clausen's definition, is that more than a single egg is devoured by the
maturing wasp larva.
Among the other wasp parasites (Encyrtidae, Eulophidae, Eupelmidae) of
cockroach eggs many individuals develop in a single oötheca. When a
hundred or more wasps emerge from an oötheca which contained less than
20 eggs, it is obvious that a single cockroach egg supported more than one
wasp, yet it is possible that one particular wasp larva may have fed upon
more than one cockroach egg before becoming an adult. We consider all
entomophagous wasps that develop in cockroach oöthecae to be parasites
rather than predators.
On the other hand, even though Anastatus floridanus, A. tenuipes, and
Tetrastichus hagenowii are egg parasites as larvae, the adult females are, in
a sense, predators when they sometimes eat part of the cockroach egg that
oozes through the oviposition puncture (Roth and Willis, 1954a, 1954b).
Williams (1929) has seen the female of Ampulex canaliculata imbibe blood
that oozed from the cut ends of the cockroach's antennae after she had
clipped them off before leading the prey to her nest. Yet despite this evident
predatism on the part of the adult, the larva feeds as a parasite on the stored
cockroaches in accordance with Sweetman's (1936) (though not Clausen's
1940) definition of parasitism, which is "that form of symbiosis in which
one symbiont lives in or on the host organism and feeds at its expense
during the whole of either the immature or mature feeding stage." The
ampulicid larva, as the evaniid, is not free-living and does not kill the host
immediately by direct attack, even though it may require more than one
victim to reach maturity. Thus, within one individual both parasitic and
predatory behavior are operant during different stages of its life history.
With the above discussion in mind we have summarized below the various
biotic associations of cockroaches. Only a few examples are given for each

section, but all organisms with similar habits presumably would be
classified in the same categories.

Class A. Associations in which cockroaches serve as hosts, vectors, or
prey for other organisms.

Type I. Obligate associates. Animals and plants that normally develop
only on or in the cockroach; in general, these organisms depend
entirely upon the cockroach for survival.

Group 1. Mutuals (symbiotes or symbionts of authors).
(a) Bacteria-like organisms (bacteroids which are found in the
fat body of all cockroaches that have been examined; p. 96).
(b) Bacteria (wood-digesting forms in Panesthia, and possibly
certain bacteria in the intestines, of other cockroaches; p.
100).
(c) Protozoa (several genera and species found in Cryptocercus;
p. 101).

Group 2. Commensals.
(a) Protozoa (Nyctotherus, Herpetomonas, Lophomonas, etc.; p.
172).
(b) Nematodes (Thelastomatidae; p. 193).

Group 3. Parasites.
(a) Fungi (Laboulbeniales; p. 134).
(b) Protozoa (gregarines, Plistophora, etc.; p. 181).
(c) Helminths.
(1) Primary parasites (mermithids and gordian worms; p.
201).
(2) Secondary parasites (Gongylonema neoplasticum,
Oxyuris mansoni, Moniliformis spp.; p. 206).
(d) Arthropods.
(1) Mites (Pimeliaphilus podapolipophagus; p. 219).
(2) Insects (larvae of ripiphorids, evaniids, and ampulicids;
p. 231).

Type II. Facultative associates. Animals and plants that prey on
cockroaches or are incidentally or accidentally picked up by the
cockroach, but which can survive or propagate readily on some
other host or prey. Steinhaus (1946) emphasized the importance of
the environment in determining the type of microbial flora
associated with the cockroach which may carry one type of flora in
an area which is exposed to filth and a different type in other areas.
Because many of these organisms survive passage through or on
the cockroach, the blattid may act as a vector of these animals and
plants.

Group 1. Commensals.
(a) Viruses (strains of poliomyelitis virus; p. 103).
(b) Bacteria (Enterobacteriaceae, Pseudomonadaceae,
Micrococcaceae, etc.; p. 111).
(c) Fungi (Aspergillus; p. 130).
(d) Protozoa (Iodamoeba, Dobellina, and cysts of Entamoeba
coli and Entamoeba histolytica; p. 179).
(e) Helminths (cysts of various helminths parasitic in
vertebrates; p. 208).
(f) Arthropods.
(1) Mites (Tyrophagus lintneri; p. 218).

Group 2. Parasites.
(a) Bacteria (Serratia marcescens; p. 117).
(b) Helminths (Protospirura spp.; p. 206).
(c) Arthropods.
(1) Mites (Locustacarus sp.; p. 219).
(2) Insects (Melittobia chalybii; p. 248).

Group 3. Predators, active.
(a) Arthropods.
(1) Spiders (p. 214).
(2) Scorpions (p. 212).
(3) Centipedes (p. 222).

(4) Mites (Rhizoglyphus tarsalus; p. 218).
(5) Insects (dermestids, reduviids, and on occasion adult
females of Tetrastichus, Anastatus, Ampulex; p. 234).
(b) Vertebrates.
(1) Amphibia (p. 269).
(2) Reptilia (p. 272).
(3) Aves (p. 276).
(4) Mammalia (p. 283).

Group 4. Predators, passive: Pitcher plants (p. 154).

Class B. Associations in which cockroaches serve as commensals or
predators.

Group 1. Commensal cockroaches.
(a) Associates of social insects (Attaphila spp., etc.; p. 315).
(b) Obscure associates (p. 316).

Group 2. Predatory cockroaches.
(a) Interspecies predators (p. 319).
(b) Intraspecies predators (p. 322).

Class C. Associations of cockroaches with other cockroaches.

Group 1. Intraspecies associations.
(a) Familial associations (p. 325).
(b) Other conspecific associations (aggregations and fighting)
(p. 336).

Group 2. Interspecies associations.
(a) Compatible associations (p. 337).
(b) Antagonistic associations (p. 329).

Class D. Ecological associations of cockroaches with higher plants.

Group 1. Benign associations (p. 139).

Group 2. Associations detrimental to plants (p. 162).

V. MUTUALISM
BACTEROIDS

Blochmann (1887, 1888) discovered intracellular particles (the bacteroids
or symbiotes of authors) that resembled bacteria in the fat body of males
and females of Blatta orientalis and Blattella germanica (pl. 26), in the ova
of these insects, and in their embryos. Bacteroids have since been found in
at least 25 species and 19 genera of cockroaches. Presumably such
microorganisms are universally distributed throughout the Blattaria.
General reviews of the bacteroids of cockroaches and other insects have
been published by Glaser (1930b), Schwartz (1935), Steinhaus (1946,
1949), Buchner (1952, 1953), Brooks (1954), and Richards and Brooks
(1958). The reader is referred to these papers, and those of authors cited in
the list at the end of this section, for discussions of the morphology of the
bacteroids, their distribution within the host, and attempts to culture them in
vitro.
It has long been assumed, without proof, that cockroaches and their
bacteroids form a mutually beneficial association. As it has not been
possible to cultivate bacteroids apart from their cockroach hosts, it may be
assumed that the host is essential to the continued existence of the
microorganism, which also derives from the association other obvious
benefits as well. Experiments to show that the host also benefits from the
association have centered around rendering cockroaches bacteroid free.
Starvation, parasites, electromagnetic radiation, heat or cold, or chemicals
have all been used in attempts to eliminate the bacteroids. Of these, only
chemical treatment has provided a satisfactory technique.

Few chemicals other than antibiotics have proved to be useful in the
elimination or reduction of bacteroids. Yetwin (1932) injected various
dilutions of 22 compounds into Blattella germanica. He observed decreases
in the bacteroids of the fat body only following injection of methylene blue,
but did not pursue this lead further. Gier (in Steinhaus, 1946) observed
reduction in the numbers of bacteroids after cockroaches were injected with
crystal violet, hexylresorcinol, or metaphen. Bode (1936) reported that
injection of irritants such as lithium salts or quinine hydrochloride had no
apparent effect on the symbiotes.
Brooks (1957) reared Blattella germanica on diets containing different
concentrations of inorganic ions. On a manganese-deficient diet the
cockroaches grew poorly and some of their progeny lacked normal
bacteroids; about 10 percent of the aposymbiotic generation grew and
reproduced on a diet fortified with yeast. Varying the concentrations of
other salts in the diets gave results in which the progeny were either
aposymbiotic or the fat body was abnormal but the mycetocytes were
abundant; all these cockroaches soon died even on fortified diets.
The administration of certain antibiotic drugs has produced cockroaches
very nearly free of bacteroids. Brues and Dunn (1945) found that although
sulfa drugs had no effect on the bacteroids, penicillin in large doses reduced
the number of bacteroids in Blaberus craniifer, but the cockroaches died
within a few days. Death was attributed to lack of bacteroids rather than
effect of the drug; this is perhaps an unwarranted conclusion in view of the
survival of aposymbiotic cockroaches that have been produced by other
drugs (Brooks, 1954; Brooks and Richards, 1955). Glaser (1946) found that
the bacteroids of Periplaneta americana could be adversely affected or
destroyed by sulfathiazole, or sodium or calcium penicillin. Fraenkel (1952)
questioned the conclusions of Brues and Dunn (1945) and of Glaser (1946)
because of the high mortality in their experiments, and he suggested that the
described phenomena "were due rather to direct toxic effects on the host
than to loss of the symbionts." Noland (in Brooks, 1954; Brooks and
Richards, 1955) confirmed Glaser's results with penicillin and extended
sulfa treatments to include Blattella germanica. Every female whose
bacteroids were reduced to the vanishing point resorbed her ovaries and was
incapable of reproduction. Brooks (1954; Brooks and Richards, 1955)
found that administration of several antibiotics did not eliminate the

bacteroids from the fat body of B. germanica unless the dose was so high
that it caused excessive mortality. Frank (1955, 1956) was able to eliminate
bacteroids from Blatta orientalis by injecting or feeding chlortetracycline,
oxytetracycline, or penicillin; survival of treated insects was not good and
reproduction was poor; the aposymbiotic individuals were smaller than
normal. As Richards and Brooks (1958) have pointed out, it is uncertain
how much of this difference was the result of loss of bacteroids and how
much the effect of the drug. It is obvious that in all these experiments the
action of the drugs on the bacteroids was accompanied by equivocal side
effects which confused interpretation of the results. The effect on the
cockroach of a loss of bacteroids cannot be separated from a possible toxic
effect of the drug.
Fortunately Brooks (1954; Brooks and Richards, 1955) obtained completely
aposymbiotic offspring from Blattella germanica that had been reared on
aureomycin. These bacteroid-free nymphs were practically incapable of
growth on a natural diet that was adequate for nymphs with symbiotes.
However, the addition of large amounts of dried yeast to the diet enabled
aposymbiotic nymphs to mature in two to three times the period required by
normal nymphs. Final proof of the function of the bacteroids was obtained
by reestablishing them in aposymbiotic cockroaches. The insects that
received implants of normal fat body of B. germanica showed a slow,
steady gain in weight over the controls (Brooks, 1954; Brooks and
Richards, 1956). Obviously, the intracellular symbiotes subserve the normal
nutrition of the cockroach. Whether the bacteroids produce only vitaminlike
substances, as suggested by Keller (1950), or function in some other way is
still to be determined. Brooks (1954) concluded that the amount of vitamin-
containing food required for increased growth by aposymbiotic cockroaches
is much greater than the known vitamin requirements; hence the factor(s)
needed is unknown and present in low concentration, or it serves as a
precursor of a second factor(s) whose synthesis is aided by the bacteroids.
Brooks (in Richards and Brooks, 1958) has since found that the bacteroids
of Blattella germanica "can supply the insect with B vitamins, amino acids
and some larger protein fragment."
Gier (1947) stated that the symbiotes of cockroaches are generically all the
same. However, as the symbiotes are presumed to have been associated
with cockroaches for over 300,000,000 years (Buchner, 1952) they may be

assumed to have developed specific differences that link them inseparably
to their respective hosts. Ries (1932) transplanted symbiote-containing fat
body from Blatta orientalis into the mealworm and larva of Ephestia
kühniella, and from Blattella germanica and Stegobium paniceum
(=Sitodrepa panicea) into B. orientalis. The implants did not become
established in the new host, although most of the transplantations were
successful in that the hosts survived and the implants remained intact for
some time before they were encapsulated by host tissue. Brooks (1954;
Brooks and Richards, 1956) transplanted fat body of Periplaneta americana
and B. orientalis into aposymbiotic B. germanica. The growth of the
cockroaches injected with foreign tissue was not different from that of
aposymbiotic controls. Sections of host insects did not contain mycetocytes
and no bacteroids were found. Haller (1955a) injected bacteroids or
implanted mycetocytes of B. germanica into gryllids, acridids, and
locustids. These implants and innoculations were rapidly destroyed by the
hosts. But as Richards and Brooks (1958) have pointed out, none of these
experiments provide information about the specificity of the bacteroids
themselves.

COCKROACHES IN WHICH BACTEROIDS HAVE BEEN
FOUND

Bantua stigmosa. Fraenkel (1921)
Blaberus craniifer. Brooks (1954); Brues and Dunn (1945);
Hoover (1945).
Blaberus giganteus. Blochmann (1892).
Blatta orientalis. Blochmann (1887, 1888, 1892); Bode (1936);
Brooks (1954); Buchner (1912); Cuénot (1896); Fraenkel
(1921); Frank (1955, 1956); Gier (1936, 1947); Glaser (1920);
Gropengiesser (1925); Gubler (1948); Heymons (1895);
Hollande and Favre (1931); Hoover (1945); Hovasse (1930);
Javelly (1914); Keller (1950); Koch (1949); Menel (1907)?;
Mercier (1906a, 1907, 1907b, 1907c); Ries (1932); Ronzoni
(1949); Tacchini (1946); Wolf (1924, 1924a).

Blattella germanica. Blochmann (1887, 1888, 1892); Bode
(1936); Borghese (1946, 1948, 1948a); Brooks (1954); Brooks
and Richards (1954, 1955, 1955a, 1956); Fraenkel (1921); Gier
(1936, 1947); Glaser (1920, 1930a); Gropengiesser (1925);
Haller (1955, 1955a); Heymons (1892, 1895); Hoover (1945);
Koch (1949); Lwoff (1923); Milovidov (1928); Neukomm
(1927, 1927a, 1932); Pérez Silva (1954, 1954a); Ries (1932);
Rizki (1954); Ronzoni (1949); Tacchini (1946); Wollman
(1926); Yetwin (1932, 1953).
Cryptocercus punctulatus. Gier (1936, 1947); Hoover (1945).
Derocalymma cruralis. Fraenkel (1921).
Ectobius lapponicus. Heymons (1892); Cuénot (1896); Koch
(1949).
Ectobius pallidus. Heymons (1892, 1895); Cuénot (1896).
Epilampra grisea. Fraenkel (1921).
Eurycotis floridana. Gier (1936, 1947).
Loboptera decipiens. Buchner (1930).
Nauphoeta cinerea. Fraenkel (1921).
Nyctibora noctivaga. Gier (1947).
Parcoblatta lata. Gier (1936, 1947).
Parcoblatta pensylvanica. Brooks (1954); Gier (1936, 1947).
Parcoblatta uhleriana. Gier (1936, 1947); Hoover (1945).
Parcoblatta virginica. Glaser (1920); Gier (1947).
Periplaneta americana. Baudisch (1956); Bode (1936); Cuénot
(1896); Gier (1936, 1937, 1947); Glaser (1920, 1930, 1946);
Gubler (1948); Hertig (1921); Hoover (1945); Ketchel and
Williams (1953).
Periplaneta australasiae. Gier (1936, 1947); Koch (1949).
Platyzosteria armata. Fraenkel (1921).
Polyphaga aegyptiaca. Fraenkel (1921).
Pseudoderopeltis aethiopica. Fraenkel (1921).
Pycnoscelus surinamensis. Bode (1936); Koch (1949).
Supella supellectilium. Brooks (1954).

BACTERIA

Evidence showing that intestinal bacteria contribute to the nutrition of
cockroaches is meager. Cleveland et al. (1934) isolated a bacterial organism
from the foregut of the wood-feeding cockroach Panesthia angustipennis.
The bacterium digested cellulose rapidly in vitro and these workers believe
that this cockroach and other related wood-feeding species are dependent
on symbiotic bacteria for the digestion of their food.
Mencl (1907) described cell nuclei in "symbiotic," not closely defined types
of bacilli that he found in abundance in the digestive tract of the
Küchenschabe, Periplaneta (presumably Blatta orientalis). Unfortunately,
he was more concerned about the morphology of the bacteria than the stated
mutualistic relationship, so nothing is known of their physiology.
The growth rates of Periplaneta americana and Blattella germanica were
retarded when the insects were reared aseptically, which suggests that
microorganisms normally found in the digestive tract supply certain
necessary dietary constituents (Gier, 1947a; House, 1949). Noland et al.
(1949) suggested that microorganisms in the digestive tract of B. germanica
synthesized riboflavin since the nymphs reared on a low riboflavin diet
accumulated more of the vitamin than could have been ingested in the diet.
However, Metcalf and Patton (1942) found little or no bacterial synthesis of
riboflavin in P. americana. Noland and Baumann (1951) suggested that
methionine, one of the amino acids essential for rapid growth of B.
germanica, was synthesized by intestinal microorganisms in the insects.

PROTOZOA

It is probable that with few exceptions protozoa found in the digestive tract
are not necessary for survival of the cockroach. However, very few
experiments have been performed to determine the importance, if any, of
these microorganisms to the host. Cleveland (1925) removed the protozoa
from the cockroach (possibly Periplaneta americana) by oxygenation at 3.5
atmospheres. The ciliates Nyctotherus and Balantidium, flagellates
Lophomonas and Polymastix, the amoeba Endamoeba blattae, and three

unidentified protozoa were killed by this treatment, yet the insects lived
normally after defaunation.
Armer (1944) studied the effects of high-carbohydrate, high-fat, and high-
protein diets, as well as starvation, on the intestinal protozoa (Nyctotherus
ovalis, Endamoeba blattae, Endolimax blattae, Lophomonas striata, and
Lophomonas blattarum) in Periplaneta americana. Starvation of the host
lowered the incidence or eliminated most of the protozoa, but a high-
carbohydrate diet maintained them at a relatively high level. Lophomonas
blattarum was eliminated by a high-protein diet, and practically eliminated
by a high-fat diet. Lophomonas striata was eliminated from some hosts that
were kept on high-fat and high-protein diets. Endamoeba blattae showed a
decrease in infection rate when the cockroaches were maintained on high-
fat and high-protein diets. The effects of diets on Endolimax blattae were
not uniform.
It has been shown by Cleveland (1930, 1948) and Cleveland et al. (1931,
1934) that the wood-feeding cockroach Cryptocercus punctulatus depends
upon certain intestinal protozoa for survival; these protozoa utilize as food
the wood ingested by this cockroach. The wood is broken down into
compounds the cockroach can utilize by the protozoa which elaborate a
cellulase and possibly a cellobiase (Trager, 1932). Only molting nymphs of
Cryptocercus can pass the protozoa on to the newly hatched young, so that
molting and hatching must happen concurrently each year or the young die.
The sexual cycles in species of protozoa in the genera Trichonympha,
Saccinobaculus, Oxymonas, Monocercomonoides, Hexamita,
Eucomonympha, Leptospironympha, Urinympha, Rhynchonympha,
Macrospironympha, and Barbulanympha (fig. 3, B) are induced by
hormones produced by Cryptocercus only during its molting period
(Cleveland, 1931, 1947, 1947a, 1949-1956a). Perhaps the prothoracic gland
hormone of the host may be responsible for initiation of the flagellate
sexual cycles (Cleveland and Nutting, 1955). The protozoan sexual cycles
may be used as indicators of the onset of molting in Cryptocercus; thus
different species of protozoa begin their sexual cycles from 35 days before
to 2 days after molting of the cockroach (Cleveland and Nutting, 1954).
Hollande (1952) and Grassé (1952) have reviewed the roles and the
evolution of the flagellates in Cryptocercus and in termites.

The protozoa of cockroaches and termites are clues to the relationship
between these two groups of insects. Kirby (1927) pointed out similarities
between Endamoeba blattae of Periplaneta and the amoebae of the termite
Mirotermes, suggesting that these protozoans were probably derived from
an amoeba in an ancestor common to both blattid and termite. Kirby (1932,
in Kidder, 1937) found a species of Nyctotherus in Amitermes that
resembles Nyctotherus ovalis from domestic cockroaches. The belief that
the termites and cockroaches had a common origin is also strengthened by
the similarities between the hypermastigotes of both Cryptocercus and
termites (Cleveland et al., 1934).
The cockroaches Cryptocercus and Panesthia both feed on wood, but the
protozoa found in Panesthia resemble more closely the species in domestic
cockroaches than those in Cryptocercus. The Clevelandellidae (from
Panesthia) are closely related to Nyctotherus and have probably evolved
from common ancestors. However, the separation of the Clevelandellidae
from Nyctotherus must have taken place at a later date than the divergence
of their hosts, otherwise representatives of that family would probably also
be found in Periplaneta and Blatta (Kidder, 1937).
The protozoa of Cryptocercus can be transferred from one individual to
another (Nutting and Cleveland, 1954). They can also be transferred to the
termite Zootermopsis where they survive only until the host molts; the
reverse is also true, Zootermopsis Protozoa can survive in Cryptocercus
until the cockroach molts (Nutting and Cleveland, 1954a).

VI. VIRUSES ASSOCIATED WITH
COCKROACHES
Annotations on some of the following observations may be found in Roth
and Willis (1957a). Use of asterisk is explained in footnote 3, page 4.

POLIOMYELITIS VIRUSES

* Lansing strain

Experimental vectors.—Blattella germanica, U.S.A. (Hurlbutt, 1949,
1950).
Periplaneta americana, U.S.A. (Hsiang et al., 1952).

* Brunhilde type, Minnesota and Mahoney strains

Experimental vectors.—Periplaneta americana, U.S.A. (Fischer and
Syverton, 1951; Syverton et al., 1952).

* Columbia SK virus

Experimental vectors.—Periplaneta americana, Great Britain? (Findlay
and Howard, 1951): Results with Blattella germanica were negative.

* Four unspecified strains

Natural vectors.—Blattella germanica and/or Blattella vaga, Periplaneta
americana and/or Periplaneta brunnea, and Supella supellectilium, U.S.A.
(Syverton et al., 1952; Dow, 1955; Dow in Roth and Willis, 1957a).

OTHER VIRUSES

* Coxsackie viruses

Experimental vectors.—Periplaneta americana, U.S.A. (Fischer and
Syverton, 1951a, 1957): Recently Fischer and Syverton (1957) found that
after feeding a single meal of Coxsackie virus to Periplaneta americana,
the gastrointestinal tracts of the insects, which were removed at 5-day
intervals up to 20 days, contained sufficient virus to paralyze and kill test

mice. Cockroach salivary glands, removed 5 days after the insects had fed,
contained the virus which caused paralysis and death in test mice; mice
were also infected by virus obtained from salivary glands removed from the
insects 10 and 20 days after the cockroaches had fed once on the virus. The
virus was also isolated from the cockroaches' feces and rarely from the fat
bodies and reproductive organs. Fischer and Syverton concluded that it is
possible that cockroaches could acquire the virus, by feeding on
mammalian excreta, maintain it for a period of time, and transmit it by
contamination of food. The virus could also be transmitted through the
feces of wild mice if the mice happened to feed on virus-infected
cockroaches.

* Mouse encephalomyelitis virus

Experimental vectors.—Periplaneta americana, U.S.A. (Syverton and
Fischer, 1950).

* Yellow-fever virus

Experimental vectors.—Blattella germanica, Great Britain? (Findlay and
MacCallum, 1939).

VII. BACTERIA ASSOCIATED WITH
COCKROACHES
Classification of the bacteria follows Breed et al. (1948). Synonymy in most
cases was taken from the same source. Names of bacteria preceded by the
symbol † are either not listed by Breed et al. or are stated by them to be

insufficiently characterized for definite classification. Use of asterisk is
explained in footnote 3, page 4.

Phylum SCHIZOPHYTA
Class SCHIZOMYCETES

Order EUBACTERIALES

Family PSEUDOMONADACEAE

* Pseudomonas aeruginosa (Schroeter) Migula

Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949).
Blatta orientalis, U.S.A. (Olson and Rueger, 1950).
Blattella germanica, U.S.A. (Olson and Rueger, 1950; Janssen and
Wedberg, 1952).
Periplaneta americana, U.S.A. (Bitter and Williams, 1949, 1949a; Olson
and Rueger, 1950).
Experimental vectors.—Blattella germanica, U.S.A. (Herms and Nelson,
1913).
Cockroaches, U.S.A. (Longfellow, 1913).

* Pseudomonas eisenbergii Migula

Natural and experimental vectors.—Blatta orientalis, Italy (Cao, 1906).

* Pseudomonas fluorescens Migula

Natural and experimental vectors.—Blatta orientalis, Italy (Cao, 1898,
1906; Spinelli and Reitano, 1932).
Periplaneta americana, U.S.A. (Gier, 1947): The organism was pathogenic
to the cockroach when injected.

† Spirillum periplaneticum Kunstler and Gineste

Habitat.—Periplaneta americana, France? (Kunstler and Gineste, 1906):
From intestinal tract.

† Spirillum α, β, and γ Dobell

Habitat.—Blatta orientalis, England (Dobell, 1911, 1912): From hind gut.

Spirillum sp.

Habitat.—Blatta orientalis, U.S.S.R. (Zasukhin, 1930): From intestinal
tract.
Cockroaches, Venezuela (Tejera, 1926): From digestive tract.

* Vibrio comma (Schroeter) Winslow et al.

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898; Spinelli and
Reitano, 1932).
Blattella germanica, Orient (Toda, 1923); Germany (Jettmar, 1927).
Periplaneta americana, Philippine Islands (Barber, 1914); Netherlands
(Akkerman, 1933); Formosa (Morischita and Tsuchimochi, 1926).
Periplaneta australasiae, Formosa (Morischita and Tsuchimochi, 1926).

* Vibrio metschnikovii Gamaléia

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898, 1906).

Vibrio tyrogenus (Flügge) Holland

Synonymy.—Vibrio of Deneke.
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898): The organism
passed through the intestinal tract unchanged.

† Vibrio Types I and II Heiberg

Habitat.—Water.
Natural vectors.—Cockroaches, India (Pasricha et al., 1938): The vibrios
were found in 16 or 17 percent of 94 cockroaches and resembled Vibrio
comma in their morphology and their main biochemical reactions; however,
serum-agglutination reactions differed.

Vibrio sp.

Habitat.—Blatta orientalis, U.S.A. (Leidy, 1853): From intestine.

Family RHIZOBIACEAE

Chromobacterium violaceum (Schroeter) Bergonzini

Synonymy.—B. violaceus.
Habitat.—Water.
Experimental vectors.—Cockroach, U.S.A. (Longfellow, 1913): Recovered
from outer part of body and intestinal tract.

Family MICROCOCCACEAE

* Micrococcus aurantiacus (Schroeter) Cohn

Natural vectors.—Blattella germanica, U.S.A. (Janssen and
Wedberg, 1952).

* Micrococcus citreus Migula

Natural vectors.—Cockroaches, U.S.A. (Longfellow, 1913).
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

* Micrococcus epidermidis (Winslow and Winslow) Hucker

Natural vectors.—Blattella germanica, U.S.A. (Janssen and Wedberg,
1952).

† Micrococcus nigrofaciens Northrup

Source.—Diseased June beetle larvae.
Experimental infection.—Periplaneta americana, U.S.A. (Northrup, 1914):
Three of four adults were infected by feeding them bread saturated with a
broth culture of the Micrococcus. After 11 days the tarsi of the cockroaches
became infected, and the hind legs split and broke off. Antennae and setae
also were affected and micrococci were recovered from the feces.

* Micrococcus pyogenes var. albus (Rosenbach) Schroeter

Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949).
Blatta orientalis, U.S.A. (Tauber, 1940; Tauber and Griffiths, 1942).
Blattella germanica, U.S.A. (Herms and Nelson, 1913; Herms, 1939;
Janssen and Wedberg, 1952).
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898); U.S.A. (Tauber
and Griffiths, 1942).
Micrococcus pyogenes var. albus (=Staphylococcus albus) and an
unidentified short rod form were found by Tauber (1940) in the hemolymph
of B. orientalis. These microorganisms were never found together in the

same insect and caused loss of appetite, sluggishness, irregular respiratory
movements, and paralysis in the cockroach; in the final stages of the disease
the legs were folded under the body, the head was tucked beneath the
forelegs, the whole insect became arched and maintained this position until
death. In some cockroaches infected with the rod pathogen, conjunctival
folds, particularly those between the dorsal abdominal sclerites, and the
joints of the metathoracic legs ruptured liberating thick white hemolymph
filled with bacteria. Tauber suggested that the infection might be spread by
contact, especially to newly molted individuals or by actual ingestion of the
bacteria by the cockroaches feeding on dead or dying individuals. All the
roaches died after successful inoculation with the Micrococcus. The
bacterial infection was associated with high total hemocyte counts and high
percentages of mitotically dividing hemolymph cells (Tauber, 1940); these
responses of the insect were interpreted as a mechanism whereby the
number of hemocytes increases resulting in an increase in the number of
phagocytes for combating the bacteria (Tauber and Griffiths, 1942).

* Micrococcus pyogenes var. aureus (Rosenbach) Zopf

Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949).
Blatta orientalis, Italy (Cao, 1906).
Blattella germanica, U.S.A. (Herms, 1939).
Cockroaches, U.S.A. (Longfellow, 1913).

Micrococcus ureae Cohn

Habitat.—Stale urine and soil containing urine.
Natural vectors.—Blattella germanica, U.S.A. (Janssen and Wedberg,
1952): From intestinal tract and feces.

* Micrococcus spp.

These organisms were obtained from pus or were designated as
staphylococci [i.e., pathogenic micrococci (Blair in Dubos, 1948)].

Natural vectors.—Blatta orientalis, Italy (Spinelli and Reitano, 1932);
Germany (Jettmar, 1935).
Blattella germanica, Germany (Jettmar, 1935).
Experimental vectors.—Blattella germanica, on shipboard (Morrell, 1911);
Germany (Vollbrechtshausen, 1953).

Micrococcus sp.

Natural vectors.—Periplaneta americana? ("Blatella americana"), England
(Shrewsbury and Barson, 1948): From intestinal tract.

† Sarcina alba Zimmermann

Habitat.—Water.
Natural and experimental vectors.—Blatta orientalis, Italy (Cao 1898,
1906): From intestinal contents.

Sarcina aurantiaca Flügge

Habitat.—Air and water.
Experimental vectors.—Blatta orientalis, Italy (Cao, 1906): Intestinal
contents.

Sarcina lutea Schroeter

Habitat.—Air, soil, water, skin surfaces.
Experimental vectors.—Blatta orientalis, Italy (Cao, 1906): From intestinal
contents.

† Sarcina symbiotica Pribram

Habitat.—Oöthecae of Blatta orientalis and/or Blattella germanica,
Germany (Gropengiesser, 1925): It was described as "eine gelbe Sarcina";

Pribram (1933) named the organism.

Sarcina ventriculi Goodsir

Habitat.—Garden soil, dust, mud; isolated from a diseased stomach.
Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946): From
intestinal tract.

Sarcina sp.

Natural and experimental vectors.—Periplaneta americana, U.S.A. (Gier,
1947): Organism not pathogenic to the cockroach when injected.

† Sarcina sp.

Synonymy.—Sarcina "blanche" of Sartory and Clerc.
Natural vectors.—Blatta orientalis, France (Sartory and Clerc, 1908):
Isolated from intestinal tract.

*† Sarcina sp.

Synonymy.—Sarcina alba "patogena" of Cao.
Natural vectors.—Blatta orientalis, Italy (Cao, 1898, 1906).

*† Sarcina sp.

Synonymy.—Sarcina "bianca" and "gialla" of Cao.
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

Family NEISSERIACEAE

* Neisseria meningitidis (Albrecht and Ghon) Holland

Experimental vectors.—Cockroaches, U.S.A. (Longfellow, 1913).

* Veillonella parvula (Veillon and Zuber) Prévot

Natural vectors.—Periplaneta americana, U.S.A. (Hatcher, 1939).

Family LACTOBACTERIACEAE

* Diplococcus pneumoniae Weichselbaum

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).
Cockroaches, U.S.A. (Longfellow, 1913).

† Enterococcus sp.

Natural vectors.—Blatta orientalis, Italy (Spinelli and Reitano, 1932):
From intestinal tract.

Lactobacillus fermenti Beijerinck

Habitat.—Fermenting plant or animal products.
Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946): From
intestinal canal.

† Pneumococcus Type I, No. 1231

Experimental vectors.—Blattella germanica, Germany (Vollbrechtshausen,
1953).

* Streptococcus faecalis Andrewes and Horder

Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946).
Blattella germanica, U.S.A. (Steinhaus, 1941).

Periplaneta americana? ("Blatella americana"), England (Shrewsbury and
Barson, 1948).
Cockroaches [presumably any or all of the above three species], Egypt (El-
Kholy and Gohar, 1945).

* Streptococcus liquefaciens Sternberg emend. Orla-Jensen

Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946).

*† Streptococcus microapoika Cooper, Keller, and Johnson

Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946).

*† Streptococcus non-hemolyticus II Holman

Natural vectors.—Periplaneta americana? ("Blatella americana"), England
(Shrewsbury and Barson, 1948).

* Streptococcus pyogenes Rosenbach

Natural vectors.—Blatta orientalis, Italy (Cao, 1906).
Experimental vectors.—Cockroaches, U.S.A. (Longfellow, 1913).

* Streptococcus sp. (pyogenic group)

Experimental vectors.—Blatta orientalis, Germany (Jettmar, 1935).

* Streptococcus sp. (viridans group)

Experimental vectors.—Blatta orientalis, Germany (Jettmar, 1935).

* Streptococcus spp.

Natural vectors.—Blatta orientalis, Germany (Jettmar, 1935).

Blattella germanica, Germany (Jettmar, 1935); U.S.A. (Janssen and
Wedberg, 1952).
Cockroaches, U.S.A. (Longfellow, 1913).
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

Family CORYNEBACTERIACEAE

* Corynebacterium diphtheriae (Flügge) Lehmann and Neumann

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).
Cockroaches, U.S.A. (Longfellow, 1913).

Family ACHROMOBACTERIACEAE

† Achromobacter hyalinum (Jordan) Bergey et al.

Natural vectors.—Periplaneta americana, U.S.A. (Hatcher, 1939): Isolated
from feces.

Achromobacter sp.

Natural vectors.—Blattella germanica, U.S.A. (Janssen and Wedberg,
1952).

Alcaligenes faecalis Castellani and Chalmers

Synonymy.—Bacillus faecalis alkaligenes; Bacillus alcaligenes faecalis; B.
alcaligenes faecalis.
Habitat.—Intestinal canal of man. Has been isolated from feces, abscesses
related to intestinal canal, and occasionally in the bloodstream. However, it
is generally considered nonpathogenic.

Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949):
Isolated from feces.
Blatta orientalis, Poland (Nicewicz et al., 1946): Isolated from intestinal
tract.
Blattella germanica, U.S.A. (Janssen and Wedberg, 1952): From intestinal
tract and feces.
Periplaneta americana, U.S.A. (Bitter and Williams, 1949): Isolated from
intestinal tract.
Cockroaches [presumably Blatta orientalis, Blattella germanica and/or
Periplaneta americana], Egypt (El-Kholy and Gohar, 1945): From
suspensions of macerated whole insects.

Alcaligenes recti (Ford) Bergey et al.

Synonymy.—B. alcaligenes recti.
Habitat.—Intestinal canal.
Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946): Isolated
from intestinal tract.

Alcaligenes viscosus (Weldin and Levine) Weldin

Habitat.—Water, dairy utensils; produces ropiness in milk.
Natural vectors.—Blattella germanica, U.S.A. (Janssen and Wedberg,
1952): Isolated from intestine and feces.

Family ENTEROBACTERIACEAE

Aerobacter aerogenes (Kruse) Beijerinck

Synonymy.—Bacillus lactis aerogenes.
Habitat.—Grains, plants, intestinal tract of man and other animals.

Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949):
Isolated from feces.
Blatta orientalis, Poland (Nicewicz et al., 1946): Isolated from intestinal
tract.
Blattella germanica, on shipboard (Morrell, 1911): Isolated from feces.
Periplaneta americana, U.S.A. (Bitter and Williams, 1949): Isolated from
intestinal tract.
Cockroaches [presumably Blatta orientalis, Blattella germanica and/or
Periplaneta americana], Egypt (El-Kholy and Gohar, 1945): Isolated from
outer surface of body, intestinal tract, and suspensions of macerated whole
insects.

Aerobacter cloacae (Jordan) Bergey et al.

Synonymy.—Bacillus cloacae.
Habitat.—Sewage, soil, water, human and other animal feces.
Natural vectors.—Blattella germanica, on shipboard (Morrell, 1911): From
feces. U.S.A. (Janssen and Wedberg, 1952; Steinhaus, 1941): From
intestinal tract, feces, and oötheca.
Periplaneta americana, U.S.A. (Bitter and Williams, 1949): From intestinal
tract.

Aerobacter sp.

Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949): Isolated from intestines.

† Eberthella oedematiens Assis

Habitat.—Intestinal canal.
Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949): Intestinal tract.

* Escherichia coli (Migula) Castellani and Chalmers

Natural vectors.—Blatta orientalis, Italy (Cao 1898, 1906; Spinelli and
Reitano, 1932); France (Sartory and Clerc, 1908); Europe (Jettmar, 1935);
Poland (Nicewicz et al., 1946).
Blattella germanica, U.S.A. (Steinhaus, 1941).
Periplaneta americana, U.S.A. (Bitter and Williams, 1949, 1949a).
Cockroaches [presumably one or all of the above three species], Egypt (El-
Kholy and Gohar, 1945).
Cockroaches, U.S.A. (Longfellow, 1913).
Experimental vectors.—Blattella germanica, Germany (Vollbrechtshausen,
1953): When injected anally or orally the bacteria invaded the intestinal
cells and in heavy infections killed the cockroaches.

Escherichia coli var. acidilactici (Topley and Wilson) Yale

Synonymy.—Bacillus acidi lactici.
Source.—Diseased nun moth larvae.
Experimental vectors.—Blatta orientalis, Europe (Filatoff, 1904):
Organism pathogenic to cockroach when injected but not when fed.

Escherichia coli var. communior (Topley and Wilson) Yale

Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): From
feces.

Escherichia freundii (Braak) Yale

Habitat.—Soil, water, intestinal canal of man and other animals.
Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): From
feces.

Blattella germanica, U.S.A. (Janssen and Wedberg, 1952): From intestinal
canal and feces.
Periplaneta americana, U.S.A. (Bitter and Williams, 1949): From intestinal
tract.

Escherichia intermedium (Werkman and Gillen) Vaughn and Levine

Habitat.—Soil, water, intestinal canal of man and other animals.
Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949): From intestinal canal.

* Klebsiella pneumoniae (Schroeter) Trevisan

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

* Paracolobactrum aerogenoides Borman, Stuart and Wheeler

Natural vectors.—Blattella germanica, U.S.A. (Janssen and Wedberg,
1952).
Periplaneta americana, U.S.A. (Bitter and Williams, 1949).

* Paracolobactrum coliforme Borman, Stuart and Wheeler

Natural vectors.—Blattella germanica, U.S.A. (Janssen and Wedberg,
1952).
Periplaneta americana, U.S.A. (Bitter and Williams, 1949).

* Paracolobactrum spp.

Natural vectors.—Blatta orientalis, U.S.A. (Olson and Rueger, 1950).
Blattella germanica, U.S.A. (Olson and Rueger, 1950).
Periplaneta americana, U.S.A. (Bitter and Williams, 1949; Olson and
Rueger, 1950).

* Paracolon bacilli

Natural vectors.—Cockroaches [presumably Blatta orientalis, Blattella
germanica and/or Periplaneta americana], Egypt (El-Kholy and Gohar,
1945).

* Proteus mirabilis Hauser

Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949, 1949a).

* Proteus morganii (Winslow et al.) Rauss

Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949, 1949a).
Cockroaches [presumably Blatta orientalis, Blattella germanica, and/or
Periplaneta americana], Egypt (El-Kholy and Gohar, 1945).

* Proteus rettgeri (Hadley et al.) Rustigian and Stuart

Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949).

* Proteus vulgaris Hauser

Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949).
Blatta orientalis, Italy (Spinelli and Reitano, 1932).
Periplaneta americana, U.S.A. (Bitter and Williams, 1949, 1949a).
Cockroaches, U.S.A. (Longfellow, 1913).

* Proteus spp.

Natural vectors.—Blatta orientalis, U.S.A. (Olson and Rueger, 1950).

Periplaneta americana, U.S.A. (Bitter and Williams, 1949; Olson and
Rueger, 1950).

* Salmonella anatis (Rettger and Scoville) Bergey et al.

Natural vectors.—Periplaneta americana, U.S.A. (Eads et al., 1954).

* Salmonella choleraesuis (Smith) Weldin

Experimental vectors.—Polyphaga saussurei, U.S.S.R. (Zmeev in
Pavlovskii, 1948).

* Salmonella enteritidis (Gaertner) Castellani and Chalmers

Experimental vectors.—Blatta orientalis, U.S.S.R. (Rozengolts and I͡ udina
in Pavlovskii, 1948).

Blattella germanica, U.S.A. (Olson and Rueger, 1950); U.S.S.R.
(Rozengolts and I͡ udina in Pavlovskii, 1948).
Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948).

* Salmonella morbificans (Migula) Haupt

Natural vectors.—Periplaneta americana, Australia (Mackerras and
Mackerras, 1948, 1949).
Experimental vectors.—Nauphoeta cinerea, Periplaneta australasiae,
Periplaneta ignota, and Supella supellectilium, Australia (Mackerras and
Pope, 1948).

* Salmonella paratyphi (Kayser) Castellani and Chalmers

Experimental vectors.—Polyphaga saussurei, U.S.S.R. (Zmeev in
Pavlovskii, 1948).

* Salmonella schottmuelleri (Winslow et al.) Bergey et al.

Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949, 1949a).
Experimental vectors.—Periplaneta americana, Gold Coast Colony
(Macfie, 1922).
Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948).

* Salmonella sp. (Type Adelaidae)

Experimental vectors.—Nauphoeta cinerea, Periplaneta australasiae,
Periplaneta ignota, and Supella supellectilium, Australia (Mackerras and
Pope, 1948).

* Salmonella sp. (Type Bareilly)

Natural vectors.—Periplaneta americana, U.S.A. (Eads et al., 1954).

* Salmonella sp. (Type Bredeny)

Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949, 1949a).

* Salmonella sp. (Type Derby)

Experimental vectors.—Nauphoeta cinerea, Periplaneta australasiae, and
Supella supellectilium, Australia (Mackerras and Pope, 1948).

* Salmonella sp. (Type Kentucky)

Natural vectors.—Periplaneta americana, U.S.A. (Eads et al., 1954).

* Salmonella sp. (Type Kottbus)

Experimental vectors.—Periplaneta australasiae, Australia (Mackerras and
Pope, 1948).

* Salmonella sp. (Type Meleagris)

Natural vectors.—Periplaneta americana, U.S.A. (Eads et al., 1954).

* Salmonella sp. (Type Montevideo)

Experimental vectors.—Periplaneta americana, U.S.A. (Jung and Shaffer,
1952).

* Salmonella sp. (Type Newport)

Natural vectors.—Periplaneta americana, U.S.A. (Eads et al., 1954).

* Salmonella sp. (Type Oranienburg)

Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949, 1949a; Eads et al., 1954).
Experimental vectors.—Blatta orientalis, Blattella germanica, and
Periplaneta americana, U.S.A. (Olson and Rueger, 1950).

* Salmonella sp. (Type Panama)

Natural vectors.—Periplaneta americana, U.S.A. (Eads et al., 1954).

* Salmonella sp. (Type Rubislaw)

Natural vectors.—Periplaneta americana, U.S.A. (Eads et al., 1954).

* Salmonella sp. (Type Tennessee)

Natural vectors.—Periplaneta americana, U.S.A. (Eads et al., 1954).

* Salmonella typhimurium (Loeffler) Castellani and Chalmers

Natural vectors.—Blattella germanica, Belgium (Graffar and Mertens,
1950).
Nauphoeta cinerea, Australia (Mackerras and Mackerras, 1948).
Experimental vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949).
Blatta orientalis, U.S.S.R. (Rozengolts and I͡ udina in Pavlovskii, 1948).
Blattella germanica, Belgium (Graffar and Mertens, 1950); U.S.A. (Olson
and Rueger, 1950; Janssen and Wedberg, 1952; Beck and Coffee, 1943);
U.S.S.R. (Rozengolts and I͡ udina in Pavlovskii, 1948).
Nauphoeta cinerea, Australia (Mackerras and Pope, 1948).
Periplaneta americana, U.S.A. (Beck and Coffee, 1943; Jung and Shaffer,
1952).
Periplaneta australasiae and Supella supellectilium, Australia (Mackerras
and Pope, 1948).
Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948).

* Salmonella typhosa (Zopf) White

Natural vectors.—Blatta orientalis, Italy (Antonelli, 1930, 1943).
Experimental vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949).
Blatta orientalis, Italy (Spinelli and Reitano, 1932); U.S.A. (McBurney and
Davis, 1930); U.S.S.R. (Rozengolts and I͡ udina in Pavlovskii, 1948).
Blattella germanica, U.S.A. (Janssen and Wedberg, 1952); Germany
(Jettmar, 1927); U.S.S.R. (Rozengolts and I͡ udina in Pavlovskii, 1948).
Periplaneta americana, Gold Coast Colony (Macfie, 1922); Netherlands
(Akkerman, 1933); Formosa (Morischita and Tsuchimochi, 1926); U.S.A.
(Olson in Roth and Willis, 1957a).
Periplaneta australasiae, Formosa (Morischita and Tsuchimochi, 1926).
Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948).
Cockroaches [presumably Blatta orientalis, Blattella germanica, and/or
Periplaneta americana], Egypt (El-Kholy and Gohar, 1945).

Serratia marcescens Bizio

Synonymy.—Bacillus prodigiosus, Bacterium prodigiosum.
Habitat.—Water, soil, milk, foods, and various insects.
Natural hosts.—Blatta orientalis, Poland (Nicewicz et al., 1946): From
intestinal tract. Italy (Spinelli and Reitano, 1932).
Blattella germanica, Canada (Heimpel and West, 1959).
Diploptera punctata, Nauphoeta cinerea, Neostylopyga rhombifolia,
Panchlora nivea, Pycnoscelus surinamensis, and Supella supellectilium,
U.S.A. (Roth and Willis, unpublished data, 1958): The organism was
isolated and identified by Dr. Hillel Levinson, Quartermaster bacteriologist,
from dead specimens found in our laboratory colonies which showed the
red coloration characteristic of insects that have died with infections of S.
marcescens (pl. 16, A, B).
Leucophaea maderae, U.S.A. (Levinson, personal communication, 1958):
The organism was isolated from the hemolymph of living insects while
attempting to determine the cause of unexplained mortality in our
laboratory colony of this insect.
Leucophaea maderae or Periplaneta americana, Philippine Islands (Barber,
1912): From hemolymph.
Periplaneta americana, U.S.A. (Gier, 1947; Steinhaus, 1959).
Periplaneta australasiae and Periplaneta brunnea (Roth and Willis,
unpublished data [1954]): In laboratory colonies. Isolated from suspensions
of ground insects. In 1954 we received a culture of Periplaneta brunnea
from the Department of Public Health, University of Minnesota. These
insects began to die off rapidly and the normally lightly pigmented parts of
the body became red. Dr. Hillel Levinson, Quartermaster bacteriologist,
cultured Serratia marcescens from several moribund individuals. The
Department of Public Health of Minnesota had at times in the past cultured
S. marcescens but had discarded the cultures and was unaware that it might
be surviving in the cockroach colonies (Richards, personal communication,
1954).

Periplaneta sp., U.S.A. (Olson in Roth and Willis, 1957a): Isolated from an
undetermined species of Periplaneta, received in a shipment from the
South, a strain of S. marcescens which was toxic to mice when administered
intraperitoneally.
Experimental hosts.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949):
When fed in small numbers, S. marcescens increased to such an extent that
the insect's extremities and upper halves of their bodies turned deep red.
The insects died after this color appeared and practically pure cultures of
Serratia were recovered from the reddened areas.
Blatta orientalis, Italy (Cao, 1898): Isolated from intestinal contents. Passed
unchanged through the gut.
Blattella germanica, Canada (Heimpel and West, 1959): Not normally
pathogenic per os; LD50 by injection, is approximately 38,000 bacteria per
insect.
Periplaneta americana, U.S.A. (Gier, 1947): Organism toxic to the
cockroach when injected.
Cockroaches, U.S.A. (Longfellow, 1913): Isolated from legs and viscera
after feeding experiments.

* Shigella alkalescens (Andrewes) Weldin

Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Williams,
1949, 1949a).

* Shigella dysenteriae (Shiga) Castellani and Chalmers

Experimental vectors.—Blatta orientalis, Italy (Spinelli and Reitano, 1932).
Periplaneta americana, Formosa (Morischita and Tsuchimochi, 1926).
Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948).

* Shigella paradysenteriae (Collins) Weldin

Natural vectors.—Blatta lateralis, Tadzhikistan (Zmeev, 1940).

Experimental vectors.—Periplaneta americana, Gold Coast Colony
(Macfie, 1922).
Polyphaga saussurei, U.S.S.R. (Zmeev in Pavlovskii, 1948).
Cockroaches, Venezuela (Tejera, 1926).

Family PARVOBACTERIACEAE

Bacteroides uncatus Eggerth and Gagnon

Habitat.—Probably intestinal canal of mammals; from human feces.
Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946): From
intestinal canal.

* Brucella abortus (Schmidt and Weis) Meyer and Shaw

Experimental vectors.—Periplaneta americana, U.S.A. (Ruhland and
Huddleson, 1941).

Fusiformis lophomonadis Grassé

Habitat.—Surface of a flagellate (Lophomonas striata) which lives in the
intestine of cockroaches (Breed et al., 1948; Grassé 1926, 1926a).

* Malleomyces mallei (Zopf) Pribram

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

* Pasteurella multocida (Lehmann and Neumann) Rosenbusch and
Merchant

Experimental vectors.—Blatta orientalis, Germany (Küster, 1902, 1903);
Italy (Cao, 1906).

* Pasteurella pestis (Lehmann and Neumann) Holland

Natural vectors.—Blatta orientalis, Hongkong (Hunter, 1906).
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898); Germany
(Küster, 1903).
Blattella germanica, Germany (Jettmar, 1927).
Leucophaea maderae and Periplaneta americana, Philippine Islands
(Barber, 1912).

Family BACTERIACEAE

† Bacterium alkaligenes Nyberg

Natural vectors.—Periplaneta americana? ("Blatella americana"), England
(Shrewsbury and Barson, 1948): From intestinal tract.

† Bacterium delendae-muscae Roubaud and Descazeaux

Source.—Diseased fly larvae.
Experimental infection.—Cockroach, France (Roubaud and Descazeaux,
1923): Organism pathogenic to cockroach when injected.

Bacterium haemophosphoreum Pfeiffer and Stammer

Habitat.—Diseased larvae of Mamestra oleracea.
Experimental infection.—Blatta orientalis and Blattella germanica,
Germany (Pfeiffer and Stammer, 1931): Organism pathogenic, when
injected, to eight B. orientalis and two B. germanica.

Coccobacillus cajae Picard and Blanc

Experimental host.—Blatta orientalis, France (Picard and Blanc, 1913):
The organism was pathogenic to B. orientalis when injected.

Family BACILLACEAE

* Bacillus anthracis Cohen emend. Koch

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898, 1906); Germany
(Küster, 1903).

† Bacillus bütschlii Schaudinn

Habitat.—Blatta orientalis, Germany (Schaudinn, 1902): Isolated from
intestinal tract. Three percent of the cockroaches from Berlin bakeries were
infected.

Bacillus cereus Frankland and Frankland

Synonymy.—Bacillus albolactis.
Habitat.—Soil, dust, milk, plants.
Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): From
feces.
Periplaneta americana, U.S.A. (Hatcher, 1939): In feces.
Experimental host.—Periplaneta americana, U.S.A. (Babers, 1938): The
cockroaches died within 96 hours after being injected with 10-3 ml. of a 24-
hour broth culture.

Bacillus circulans Jordan

Habitat.—Soil, water, dust.
Natural vectors.—Blattella germanica, U.S.A. (Janssen and Wedberg,
1952): From intestine and feces.

† Bacillus flacheriae (Hoffman)

Source.—Diseased nun moth larvae.

Experimental infection.—Blatta orientalis, Europe (Filatoff, 1904): The
organism was not pathogenic when fed to the cockroach, but killed the
insects when injected into the body cavity; after the insects died Filatoff
reisolated this pathogen together with another bacillus from the cadavers.
He succeeded in culturing the new microorganism and found it to be
pathogenic when injected into, but not when fed to, the cockroaches. The
diseased insects became sluggish, failed to eat or drink, turned over on their
backs, their extremities became totally paralyzed, and they finally died.

Bacillus megaterium De Bary

Habitat.—Soil, water, decomposing materials.
Natural vectors.—Periplaneta americana? ("Blatella americana"), England
(Shrewsbury and Barson, 1948): From intestinal tract.
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898): Organism
recovered, apparently unchanged, from intestinal contents.

† Bacillus monachae (von Tubeuf) Eckstein

Synonymy.—Bacterium monache.
Source.—Diseased larvae of nun moth, Lymantria monacha.
Experimental infection.—Blatta orientalis, Europe (Filatoff, 1904):
Organism pathogenic to the cockroach when injected but not when fed.

† Bacillus periplanetae Tichomiroff

Habitat.—Blatta orientalis, U.S.S.R.? (Tichomiroff, 1870[?], in Filatoff,
1904): The infected insects suffered from a diarrhea and the liquid feces
were yellow-brown.

† Bacillus stellatus Hollande

Natural infection.—Blatta orientalis, France (Hollande, 1934): Organism
observed regularly in the intestine (especially rectum). Extensive

description given.

† Bacillus radiciformis

Experimental vectors.—Blatta orientalis, Italy (Cao 1898): Organism
recovered, apparently unchanged, from intestinal contents.

* Bacillus subtilis Cohn emend. Prazmowski

Natural vectors.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949).
Blatta orientalis, Italy (Cao, 1898, 1906; Spinelli and Reitano, 1932);
France (Sartory and Clerc, 1908); Poland (Nicewicz et al., 1946).
Cryptocercus punctulatus, U.S.A. (Hatcher, 1939).
Periplaneta americana? ("Blatella americana"), England (Shrewsbury and
Barson, 1948).
Cockroaches, U.S.A. (Longfellow, 1913).
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898, 1906).

"Bacillus subtilis group"

Natural infection.—Blatta orientalis, Italy (Ronzoni, 1949): Isolated from
oöthecae.

† Bacillus tritus Batchelor

Habitat.—Isolated from feces (man?).
Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946): Isolated
from intestinal tract.

* Clostridium feseri Trevisan

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

Clostridium lentoputrescens Hartsell and Rettger

Habitat.—Soil, intestinal tract of man.
Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946): Isolated
from intestinal tract.

* Clostridium novyi (Migula) Bergey et al. or

* Clostridium sporogenes (Metchnikoff) Bergey et al.

Natural and experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

* Clostridium perfringens (Veillon and Zuber) Holland

Natural vectors.—Cockroaches [presumably Blatta orientalis, Blattella
germanica, and/or Periplaneta americana], Egypt (El-Kholy and Gohar,
1945).

* Clostridium tetani (Flügge) Holland

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

* Clostridium spp.

Natural vectors.—Periplaneta americana? ("Blatella americana"), England
(Shrewsbury and Barson, 1948).

Order ACTINOMYCETALES

Family MYCOBACTERIACEAE

* Mycobacterium avium Chester

Experimental vectors.—Blatta orientalis, U.S.S.R. (Ekzempliarskaia in
Pavlovskii, 1948).

Mycobacterium friedmannii Holland

Habitat.—Parasitic in turtles and possibly sparingly distributed in soils.
Natural vectors.—Periplaneta americana, U.S.A., Texas (Micks, in Roth
and Willis, 1957a): Organism isolated from batches of intestinal tracts of
cockroaches collected at random.

* Mycobacterium lacticola Lehmann and Neumann?

Natural vectors.—Periplaneta americana, U.S.A. (Leibovitz, 1951).

* Mycobacterium leprae (Armauer-Hansen) Lehmann and Neumann

Natural vectors.—Blattella germanica, Southern Rhodesia and Kenya
(Moiser, 1945, 1946, 1946a; Anonymous, 1946).
Periplaneta americana and Periplaneta australasiae, Formosa (Arizumi,
1934, 1934a).
Cockroaches, Venezuela (Tejera, 1926); Belgian Congo (Radna, 1939).
Experimental vectors.—Blatta orientalis, Europe (Paldrock in Klingmüller,
1930); Nyasaland (Lamborn, 1940).
Blattella germanica, Europe (Paldrock in Klingmüller, 1930); Southern
Rhodesia and Kenya (Moiser, 1945, 1946, 1946a, 1947; Anonymous,
1946).
Nauphoeta cinerea, Nyasaland (Lamborn, 1940).
Periplaneta americana, Gold Coast Colony (Macfie, 1922); Formosa
(Arizumi, 1934, 1934a).
Periplaneta australasiae, Formosa (Arizumi, 1934, 1934a).
Cockroaches, Belgian Congo (Radna, 1939); Venezuela (Tejera, 1926).

* Mycobacterium lepraemurium Marchoux and Sorel

Experimental vectors.—Cockroaches, Belgian Congo (Radna, 1939).

* Mycobacterium phlei Lehmann and Neumann

Natural vectors.—Periplaneta americana, U.S.A. (Leibovitz, 1951; Micks
in Roth and Willis, 1957a).

* Mycobacterium piscium Bergey et al.

Natural vectors.—Periplaneta americana, U.S.A. (Leibovitz, 1951).
Experimental vectors.—Blatta orientalis, U.S.S.R. (Ekzempliarskaia in
Pavlovskii, 1948).

* Mycobacterium tuberculosis (Schroeter) Lehmann and Neumann

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898); Germany
(Küster, 1903); U.S.S.R. (Ekzempliarskaia in Pavlovskii, 1948).
Blattella germanica, on shipboard (Morrell, 1911).
Periplaneta americana, Gold Coast Colony (Macfie, 1922).
Cockroaches, Venezuela (Tejera, 1926); U.S.A. (Read, 1933).

* Mycobacterium spp.

Natural vectors.—Periplaneta americana, U.S.A. (Leibovitz, 1951; Micks
in Roth and Willis, 1957a).

Family ACTINOMYCETACEAE

* Nocardia sp.?

Natural vectors.—Periplaneta americana, U.S.A. (Leibovitz, 1951).

Family STREPTOMYCETACEAE

Streptomyces leidynematis Hoffman

Habitat.—Surface of the nematodes Hammerschmidtiella diesingi and
Leidynema appendiculata in Periplaneta americana, U.S.A. (Hoffman,
1952, 1953): Eighteen percent of 192 nematodes found in 52 adult
cockroaches were infected with the bacterium.

Order CARYOPHANALES

Family ARTHROMITACEAE

Arthromitus intestinalis (Valentin) Peshkoff

Synonymy.—Hygrocrocis intestinalis.
Habitat.—Blatta orientalis, Europe (Valentin, 1836; Robin, 1847, 1853;
Peshkoff, 1940): Isolated from intestinal tract. The organism appears as
fragments in fecal masses or as fibers adhering to the mucous membrane of
the large intestine (Robin, 1853).
Cockroach, France? (Chatton and Pérard, 1913).

Order SPIROCHAETALES

Family SPIROCHAETACEAE

† Spirochaeta blattae Tejera

Habitat.—Blaberus atropos, Venezuela (Tejera, 1926): Isolated from
intestinal tract.

* † Spirochaeta periplanetae Laveran and Franchini

Habitat.—Blatta orientalis, France (Laveran and Franchini, 1920a).
Cockroaches, Venezuela (Tejera, 1926): Tejera reported finding
"Spirochaeta blatarum Laveran et Franchini" which may have been a
lapsus.

Family TREPONEMATACEAE

† Treponema parvum Dobell

Habitat.—Blatta orientalis, England (Dobell, 1912); U.S.S.R.? Zasukhin
(1930): From intestinal tract.

† Treponema stylopygae Dobell

Synonymy.—Spirochaeta stylopygae Zuelzer.
Habitat.—Blatta orientalis, England (Dobell, 1912); U.S.S.R.? Zasukhin
(1930): From intestinal tract.

Unidentified spirochaetes

Habitat.—Blatta orientalis, U.S.S.R. (Yakimov and Miller, 1922):
Spirochaetes and spirilla were found in the intestines of 70 percent of 124
specimens collected in Petrograd.
Periplaneta americana, Gold Coast Colony (Macfie, 1922).

ADDITIONAL BACTERIA WHOSE TAXONOMIC
POSITION IS UNKNOWN

* "B. aerobio del pseudoedema maligno" of Cao

Natural vectors.—Blatta orientalis, Italy (Cao, 1906).

B. alcaligenes beckeri

Natural vectors.—Blatta orientalis, Poland (Nicewicz et al., 1946): Isolated
from intestinal tract.

* "B. del pseudoedema maligno" of Cao

Natural vectors.—Blatta orientalis, Italy (Cao, 1906).

* "Bacillo proteisimile>" of Cao

Natural and experimental vectors.—Blatta orientalis, Italy (Cao, 1898,
1906).

* "Bacillo del barbone dei bufali" of Cao

Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

* "Bacillo similcarbonchio" of Cao

Natural and experimental vectors.—Blatta orientalis, Italy (Cao, 1898,
1906).

* "Bacillo similtifo" or "Bacillo tifosimile" of Cao

Natural and experimental vectors.—Blatta orientalis, Italy (Cao, 1898,
1906).

">Bacillus"

Natural infection.—Blatta orientalis, Germany (Heinecke, 1956): Disease
organism found in the hemolymph of infected cockroaches. It can be spread
by mouth and through wound infection. The animals died with symptoms of
paralysis in 85-90 days. The organism has been isolated and is in the culture
collection of the Institute for Microbiology and Experimental Therapy,

Jena, under the numbers SG 896, Strain A; SG 897, Strain B; SG 898,
Strain C.
Experimental infection.—Blattella germanica, Germany (Heinecke, 1956):
Infected animals died in 26-30 days.
Periplaneta americana was unaffected even by heavy inoculations of the
pathogen.

"Bacterium"

Source.—(I) Diseased silkworm larvae. (II) Diseased Ocneria dispar larvae
and blood of Blatta orientalis.
Experimental infection.—(I)(II) Blatta orientalis, Europe (Filatoff, 1904):
Organism pathogenic when injected, nonpathogenic when ingested.
(I) Cockroach, U.S.A. (Glaser, 1925): Organism pathogenic to cockroach
when injected.

"Coccobacillus"

Natural infection.—Blatta orientalis, France (Hollande, 1934): Organism
described morphologically.

"Colon bacilli"

Natural vectors.—Cockroaches [presumably Blatta orientalis, Blattella
germanica, and/or Periplaneta americana], Egypt (El-Kholy and Gohar,
1945): From the outer surface, intestinal tract, and suspensions of
macerated insects.

"Diplococci"

Natural vectors.—Blatta orientalis, Germany (Jettmar, 1935): From
intestinal tract.
Blattella germanica, Germany (Jettmar, 1935): From outer surface of body.

"Diphtheroid I and II"

Source.—Periplaneta americana.
Natural and experimental infections.—Periplaneta americana, U.S.A.
(Gier, 1947): Pathogenicity to the cockroach variable when organism
injected.

"Gram positive rods"

Source.—Feces of Blattella germanica.
Experimental vector.—Blattella germanica, Germany (Vollbrechtshausen,
1953): Nonpathogenic to the insect when injected into the mouth or anus.

"Silkworm disease bacillus"

Cockroaches that were inoculated with living cultures succumbed in a few
days (Glaser, 1925).

† Spirillochaeta blattae Hollande

Habitat.—Blatta orientalis, France (Hollande, 1934; Hollande and
Hollande, 1946): Organism found in hind intestine. This spirillum was
stated to be related in external morphology to Spirillum periplaneticum
Kunstler and Gineste, but it was believed that S. blattae should be in the
Spirochaetaceae rather than the Spirillaceae.

"Spirochaetoid bacteria"

Habitat.—Blatta orientalis, France (Hollande, 1934): Two kinds described
but not named.

† Tetragenous sp.

Natural and experimental infections.—Periplaneta americana, U.S.A.
(Gier, 1947): Pathogenicity to the cockroach variable when organism
injected.

VIII. FUNGI AND YEASTS
By far the greatest number of fungi known to be associated with
cockroaches belong to the Laboulbeniaceae, genus Herpomyces, the species
of which are restricted to parasitizing cockroaches (Thaxter, 1908). Most
species are hyaline, small and inconspicuous (Thaxter, 1931) and are
usually, but not exclusively, found on the insects' antennae. Species of
Herpomyces are highly, but not completely, host specific (Richards and
Smith, 1954). While attached to the host, these fungi appear like minute
dark-colored, yellow, or white (e.g., H. arietinus) bristles or bushy hairs (pl.
27, A).

Fig. 1.—Diagram illustrating the relationship between a mature plant of Herpomyces stylopygae
and the integument of Blatta orientalis. (Reproduced from Richards and Smith [1956], through the
courtesy of Dr. A. G. Richards.)

Richards and Smith (1955, 1955a) have studied the life history of
Herpomyces stylopygae on the oriental cockroach. The plants grow only on
living cockroaches, and the infection is disseminated by contact. The
mature plants are found mostly on the antennae (pl. 27, B), either on setae
or on hard or soft cuticle. Spores are ejected from perithecia singly or in
groups of 2 to 4 spores, although groups as large as 12 spores have been
found. The presence of single, paired, or multiple spore groups on the
surface of the host was correlated with the presence of single, paired, or
multiple plants on infected cockroaches. Development from spore to mature
perithecia takes about two weeks. The plant obtains nutriment from the host
by means of a tubular haustorium that extends through the cockroach's
cuticle and expands into a large bulb in the underlying epidermal cells (fig.
1). Infections on nymphs are lost when the nymph moults, but infections on
adults persist throughout life. However, nymphs which have lost the fungus
upon moulting are readily reinfected. Collart's (1947) statement that
nymphs are never infected with Herpomyces is not true.
Richards and Smith (1956) concluded that there is no evidence of
pathogenicity in Herpomyces infections because heavily infected
cockroaches appear fully active in laboratory colonies; they can run at the
same speed as uninfected cockroaches; they reproduce normally and do not
appear to die prematurely. These workers stated that the infections cause a
dermatitis which is neither pathogenic nor debilitant. So far as we know
there are no comparative data on longevity and reproductive performance of
fungus-infected versus normal cockroaches. However, Gunn and Cosway
(1938) have shown that the presence of these fungi (identified as
Stigmatomyces sp.; see p. 138) on the antennae seemed to interfere with the
humidity reactions of Blatta orientalis. Although Richards and Smith
(1956) admit that humidity receptors and other sense organs on the
antennae may be destroyed by the fungus, they state that "insects possess
such a large number of sensilla that the result may well be more distressing
to the sensory physiologist than to the insect." Yet it seems to us that the
loss of sense organs from fungal infection and concomitant shortening of

the antennae (pl. 27, A) might be considerably more of a handicap to free-
living cockroaches than those in laboratory colonies.
Bode (1936) studied the flora of Periplaneta americana and cultured
Aspergillaceae and Mucorinae from the insect's body surface and intestinal
contents; he also found nonsporulating yeasts in P. americana. To prevent
fungal growth on oöthecae of P. americana, Griffiths and Tauber (1942a)
autoclaved their rearing containers and dipped the oöthecae in 70-percent
alcohol for 10 seconds.
Mercier (1906) isolated and cultured a pathogenic yeastlike parasite which
had invaded the fat body and blood of Blatta orientalis. The abdomens of
the infected insects became swollen, distended, and soft. McShan
(unpublished MS., 1953) consistently isolated Saccharomycetes from the
feces of Periplaneta americana.

FUNGI ASSOCIATED WITH COCKROACHES

The use of the asterisk (*) is explained in footnote 3, page 4.

Phylum THALLOPHYTA
Class FUNGI IMPERFECTI

Order MONILIALES

Family PSEUDOSACCHAROMYCETACEAE

Candida zeylanoides (Castellani) Langeron and Guerra

Natural host.—Oötheca of Blatta orientalis, Italy (Ronzoni, 1949).

Torulopsis sp.

Natural host.—Oötheca of Blatta orientalis, Italy (Ronzoni, 1949).

Family MONILIACEAE

Spicaria prasina (Maublanc) Sawada

Natural host.—Ischnoptera rufa rufa, Puerto Rico (Wolcott, 1950): A dead
specimen of this cockroach was found stuck to a leaf and covered with this
fungus.

Aspergillus flavus Link

Natural hosts.—Oöthecae of Blattella germanica and Eurycotis floridana,
U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1952): On outer
surface. Determination by Miss Mary Downing.
Oöthecae of Periplaneta americana, U.S.A., Pennsylvania (Roth and
Willis, unpublished data, 1952): Inside oöthecae. Determination by Miss
Mary Downing.

* Aspergillus fumigatus Fresenius

Natural vector.—Blatta orientalis, France (Sartory and Clerc, 1908): From
intestine.

* Aspergillus niger van Tieghem

Natural vector.—Periplaneta americana, U.S.A., Texas (McShan in Roth
and Willis, 1957a): From feces.
Experimental vector.—Blatta orientalis, Italy (Cao, 1898): Organism passed
unchanged through the gut of the insects.

Aspergillus sydowi (Bainier and Sartory) Thom and Church

Natural host.—Oötheca of Eurycotis floridana, U.S.A., Pennsylvania (Roth
and Willis, unpublished data, 1952): On outer surface. Determination by
Miss Mary Downing.

Aspergillus tamarii Kita

Natural host.—Oöthecae of Blattella germanica, U.S.A., Pennsylvania
(Roth and Willis, unpublished data, 1952): On exterior surface.
Determination by Miss Mary Downing.

Aspergillus sp.?

Natural and experimental vector.—Blattella germanica, on shipboard
(Morrell, 1911): Isolated from feces. Experimentally Morrell also showed
that the spores of the fungus could be recovered from feces of cockroaches
that had fed on them.

Aspergillus sp.

Natural vector.—Periplaneta americana, England (Bunting, 1956): The
fungus was isolated mostly from imperfectly excreted feces.

Beauveria bassiana (Balsamo) Vuillemin

Experimental host.—Blattella germanica and Periplaneta americana,
U.S.A. (Dresner, 1949, 1950): The nymphs of American cockroaches
became infected when they (1) were injected with a 1-percent suspension of
spores, (2) ate rat pellets sprayed with the spore suspension, or (3) were
dusted with the fungus spores. The symptoms of the fungus infection were
paralysis followed by death; some of the infected insects liquefied, others
dried up after the appearance of a subcuticular blackening.

Cephalosporium sp.

Natural vector.—Periplaneta americana, U.S.A., Texas (McShan,
unpublished MS., 1953): From feces of cockroaches collected in the

basement of a grain elevator at the docks in Galveston.

* Geotrichum candidum Link

Experimental vector.—Blatta orientalis, Italy (Cao, 1898): Organism
retained its pathogenicity after passing through the insect's gut.

Penicillium sp.

Natural vector.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): From
feces.
Periplaneta americana, England (Bunting, 1956): Mostly from imperfectly
excreted feces.

Metarrhizium anisopliae (Metschnikoff) Sorokin

Natural hosts.—Blattidae, Seymour (1929); Charles (1941).
Panesthia australis, U.S.A., Massachusetts (Roth and Willis, unpublished
data, 1957): Growing on adult specimens that were found dead in a
laboratory colony. Determination by Miss Dorothy Fennell.
Periplaneta americana, England (Bunting, 1956): Growing on genitalia of
females where it prevented oöthecal formation.
Cockroach, Puerto Rico (Johnston, 1915): From a "small roach" in the
pathological collection at Rio Piedras (no data).

Family DEMATIACEAE

Memnoniella echinata (Rivolta) Galloway

Natural host.—Oötheca of Blattella germanica, U.S.A., Pennsylvania (Roth
and Willis, unpublished data, 1952): On material that had oozed from a
damaged oötheca. Determination by Miss Mary Downing.

Torula acidophila Owen and Mobley

Natural host.—Periplaneta americana, U.S.A. (Owen and Mobley, 1948):
The digestive tract of this cockroach is the normal habitat of this yeast
which was transmitted to sirup by the insects. The yeast superimposed a
foreign taste, suggestive of malic acid, upon the original flavor of the sirup.

Torula gropengiesseri Lodder

Natural host.—Blatta orientalis, Germany (Gropengiesser, 1925; Lodder,
1934): Isolated from fat body and oöthecae. Gier (1947) is of the opinion
that the so-called yeasts that supposedly may displace the bacteroids in the
fat body (Mercier, 1907b; Gropengiesser, 1925) may actually represent
poorly fixed and insufficiently stained bacteroids.

Torula rosea Preuss

Experimental host.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949):
Upon repeated feeding of massive doses of this yeast to the cockroach,
these workers were able to isolate the organism from the feces up to six
days thereafter. There was no evidence that T. rosea was pathogenic for B.
craniifer.

Class PHYCOMYCETES

Order MUCORALES

Family MUCORACEAE

Mucor guilliermondii Nadson and Filippov

Natural host.—Periplaneta americana, U.S.S.R. (Nadson and Filippov,
1925; Filippov, 1926): Isolated and cultured from intestine.

Mucor sp.

Natural host.—Oötheca of Periplaneta americana, U.S.A., Pennsylvania
(Roth and Willis, unpublished data, 1952): Inside oötheca. Determination
by Miss Mary Downing.
Pycnoscelus surinamensis, Germany (Bode, 1936): Isolated from fat body
which it had stained red.

Rhizopus nigricans Ehrenberg

Natural vector.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): From
feces.

Rhizopus sp.

Natural vector.—Periplaneta americana, U.S.A., Texas (McShan,
unpublished MS., 1953): From feces.

Syncephalastrum sp.

Natural vector.—Periplaneta americana, U.S.A., Texas (McShan,
unpublished MS., 1953): From feces.

Order ENTOMOPHTHORALES

Family BLASTOCYSTIDACEAE

Blastocystis hominis Brumpt

Natural vector.—Blatta orientalis, U.S.S.R. (Zasukhin, 1930): In hind gut
in 40 percent of over 3,000 cockroaches.

Blastocystis sp.

Natural vectors.—Blatta orientalis, U.S.S.R. (Yakimov and Miller, 1922):
Found in the intestinal contents of 29 percent of 124 B. orientalis.
Cockroaches, Venezuela (Tejera, 1926).

The placement of the following fungus is problematic.

Coccidioides periplanetae Avrech

Natural host.—Blatta orientalis, Germany (Avrech, 1931): Found in cells
lining the lumen of midgut and caeca. The whole upper part of the
epithelium was filled with sporangia and spores.

Class ASCOMYCETES

Order ENDOMYCETALES

Family SACCHAROMYCETACEAE

Saccharomyces cerevisiae Hansen

Natural vector.—Blaberus craniifer, U.S.A. (Wedberg et al., 1949): In
feces.

Saccharomyces sp.

Natural vector.—Blattella germanica, U.S.A. (Janssen and Wedberg, 1952):
Found consistently in alimentary tract of B. germanica fed sucrose
solutions.

Order HYPOCREALES

Family HYPOCREACEAE

Cordyceps amazonica Hennings

Natural host.—Cockroaches, British Honduras (Mains, 1940).

Cordyceps blattae Petch

Natural host.—Blattella germanica, Ceylon (Petch, 1924): Collected at
Hakgala twice. A slight covering of brown mycelium overran the insect and
fastened it to the underside of a living leaf.

Order LABOULBENIALES

Family LABOULBENIACEAE

Herpomyces amazonicus Thaxter

Natural host.—Nyctibora obscura, Brazil, Natal (Thaxter, 1931): On
antennae.

Herpomyces anaplectae Thaxter

Natural hosts.—Anaplecta sp., Venezuela, Caracas (Thaxter, 1905, 1908);
Trinidad (Thaxter, 1931): On antennae.
Cockroach, Sumatra (Thaxter, 1931).

Herpomyces appendiculatus Thaxter

Natural host.—Platyzosteria scabra, Australia, N.S.W. (Thaxter, 1931): On
antennae.

Herpomyces arietinus Thaxter

Natural hosts.—Ischnoptera sp., U.S.A., Georgia (Thaxter, 1908).
Parcoblatta uhleriana, U.S.A., Massachusetts (Roth, unpublished data,
1957): The nymphs were in a culture of Parcoblatta virginica which was
infected with this fungus; it is possible that these P. uhleriana became
infected by contact with P. virginica. Fungus identified by Dr. R. K.
Benjamin.
Parcoblatta virginica, U.S.A., Massachusetts (Roth, unpublished data,
1957): Fungus determined by Dr. R. K. Benjamin. Fungus found on
antennae, palpi, legs, body surface (pl. 27, A).
Parcoblatta sp., U.S.A., Kentucky, Massachusetts (Thaxter, 1902, 1908):
On antennae.
It is likely that Thaxter's host records (certainly those assigned to
Temnopteryx and possibly those assigned to Ischnoptera) were species of
Parcoblatta. Hebard (1917) has shown that all the species referred to
Ischnoptera in the United States, except I. deropeltiformis, now belong in
the genus Parcoblatta. All species originally referred to the genus
Temnopteryx in the United States are now synonymized with species of
Parcoblatta.

Herpomyces chaetophilus Thaxter

Natural hosts.—Periplaneta americana, Brazil (Thaxter, 1931).
Periplaneta sp., Zanzibar and Mauritius (Thaxter, 1902, 1908): On spines
of legs, antennae, and cerci.

Herpomyces chilensis Thaxter

Natural host.—Cockroach, Chile (Thaxter, 1918): On antennae.

Herpomyces diplopterae Thaxter

Natural hosts.—Diploptera punctata, Ascension Island (Thaxter, 1902,
1908): On antennae. This species also was infected experimentally
(Richards and Smith, 1954).
Cockroach, Fiji (Thaxter, 1931).

Herpomyces ectobiae Thaxter

Natural hosts.—Blattella germanica, U.S.A., Massachusetts (Thaxter,
1902, 1908); Burma, Tenasserim (Spegazzini, 1915); Argentina, Buenos
Aires (Spegazzini, 1917): On antennae. U.S.A., Minnesota (Richards and
Smith, 1955): Scattered over entire body, wings. France? (Picard, 1913): On
tibial spines. Chile and Philippine Islands (Thaxter, 1931).
"Ectobia" spp., Zanzibar and Saint Kitts, B.W.I. (Thaxter, 1902, 1908):
Possibly on species that are now in the genus Blattella rather than in the
genus Ectobius as it is known today, because Thaxter also used the
synonym Ectobia germanica for the German cockroach, Blattella
germanica.
Experimental hosts.—Blattella germanica and Blattella vaga, U.S.A.
(Richards and Smith, 1954).

Herpomyces forficularis Thaxter

Natural hosts.—Cockroaches, Mauritius? and Fiji (Thaxter, 1902, 1908,
1931): On antennae.

Herpomyces gracilis Thaxter

Natural host.—Blattella humbertiana, Philippine Islands, Luzon (Thaxter,
1931): On antennae.

Herpomyces grenadinus Thaxter

Natural host.—Cockroach, Grenada, B.W.I. (Thaxter, 1931): On antennae
of a "brown wingless blattid."

Herpomyces leurolestis Thaxter

Natural host.—Leurolestes pallidus, British Guiana and Trinidad (Thaxter,
1931): On antennae.

Herpomyces lobopterae Thaxter

Natural host.—Loboptera sp., Argentina (Thaxter, 1931): On antennae.

Herpomyces macropus Spegazzini

Natural host.—Blaberus sp.?, Argentina (Spegazzini, 1917).
Cockroaches, Peru, Puerto Rico, Ecuador, and Haiti (Spegazzini, 1915,
1917): Material previously assigned by Spegazzini (1915) to H. paranensis
was also placed by him in this new species. However, Thaxter (1931)
believed that H. macropus may be synonymous with H. paranensis, but he
provisionally retained H. macropus because he had not seen Spegazzini's
material.

Herpomyces nyctoborae Thaxter

Natural hosts.—Nyctibora tomentosa, U.S.A., Texas (Thaxter, 1905, 1908):
On antennae. This cockroach is not established in Texas, and the specimen
may have been misidentified (Gurney, personal communication, 1958).
Nyctibora sp., Argentina (Spegazzini, 1917): On antennae.

Herpomyces panchlorae Thaxter

Natural hosts.—Panchlora nivea, Trinidad (Thaxter, 1931): On antennae.

Herpomyces panesthiae Thaxter

Natural host.—Panesthia lobipennis, Ceylon (Thaxter, 1915): On antennae.

Herpomyces paranensis Thaxter

Natural hosts.—Blaberus sp.? Brazil (Thaxter, 1902, 1908): On antennae.
Blaberus sp., Brazil and Argentina (Spegazzini, 1917): On antennae.
Cockroaches, Trinidad and Argentina (Thaxter, 1931).

Herpomyces periplanetae Thaxter

Natural hosts.—Blaberus sp.?, Argentina (Spegazzini, 1917).
Blatta orientalis, U.S.A., Massachusetts (Thaxter, 1902, 1908); Locality?
(Spegazzini, 1915); France? (Picard, 1913).
Periplaneta americana, Bermuda and U.S.A., Massachusetts (Thaxter,
1902, 1908); Plains of Biajar, Italian Somaliland, and Argentina
(Spegazzini, 1915, 1917).
Periplaneta australasiae, Bermuda (Thaxter, 1902, 1908).
Periplaneta brunnea, Brazil (Thaxter, 1931).
Periplaneta sp., Mexico, West Indies, Panama, Brazil, Africa, South Seas,
and China (Thaxter, 1902, 1908).
Cockroaches, Belgium (Collart, 1947).
Additional locality records: Grenada, Trinidad, B.W.I., and Tangier
(Thaxter, 1931).
The fungus was found growing on spines, tegmina, integument, and
antennae, at times abundantly.
Experimental hosts.—All the following data are from Richards and Smith
(1954):
Blatta orientalis: A few plants matured.
Neostylopyga rhombifolia: Some development but no mature plants.

Periplaneta americana: Fungus developed prolifically with a density equal
to that on original host.
Periplaneta australasiae: Some development but no mature plants.
Periplaneta brunnea: Fungus developed prolifically with a density equal to
that on original host.

Herpomyces phyllodromiae Thaxter

Natural host.—"Phyllodromia" sp., Abyssinia (Thaxter, 1905, 1908): On
antennae.

Herpomyces platyzosteriae Thaxter

Natural host.—"Eurycotis floridana," Mexico (Thaxter, 1905, 1908): On
antennal setae.
Since this cockroach is not found in Mexico (J. A. G. Rehn, personal
communication, 1957), E. floridana is undoubtedly not the host for this
fungus. W. B. Brown (personal communication, 1957) searched the
cockroach collection at the Museum of Comparative Zoology but was
unable to find Thaxter's insect for reidentification.

Herpomyces stylopygae Spegazzini

Natural hosts.—Blatta orientalis, Argentina (Spegazzini, 1917); U.S.A.
(Richards and Smith, 1955a).
Experimental hosts.—Neostylopyga rhombifolia (Richards and Smith,
1954): A few plants matured.
Pycnoscelus surinamensis (Richards and Smith, 1954): Some development
but no mature plants.
The fungus (fig. 1) is found on antennae (pl. 27, B, C), palpi, cerci, and
femurs. Thaxter (1931) believed H. stylopygae to be synonymous with H.
periplanetae. However, Richards and Smith (1954) concluded that H.
stylopygae would not grow on P. americana under their laboratory

conditions although H. periplanetae would grow on B. orientalis. This
indicated a strain or species difference between the two fungi. Gunn and
Cosway (1938) reported a species of Stigmatomyces on the antennae of B.
orientalis; this fungus was probably H. stylopygae (Richards and Smith,
1956).

Herpomyces supellae (Thaxter)

Natural host.—Supella supellectilium, Trinidad (Thaxter, 1931): On
antennal spines.

Herpomyces tricuspidatus Thaxter

Natural hosts.—Blaberus craniifer, U.S.A., Key West (Richards and Smith,
1955).
Blaberus sp. and Epilampra? sp., Panama (Thaxter, 1902, 1908).
Epilampra sp., Saint Kitts, B.W.I., and Haiti (Thaxter, 1902, 1908).
Leucophaea maderae, Fernando Po (Spegazzini, 1915).
Nauphoeta cinerea, Brazil (Thaxter, 1931).
Cockroaches, China? (Thaxter, 1902); Philippine Islands, Mindanao
(Thaxter, 1931).
Experimental hosts.—Blaberus craniifer, U.S.A. (Richards and Smith,
1955).
Infections on the antennae. Richards and Smith (1954) were unable to
secure experimental infections in L. maderae with H. tricuspidatus.
Experiments with N. cinerea showed some development but no mature
plants although identification of the growing fungus was uncertain because
of simultaneous exposure to H. ectobiae, H. stylopygae, and H.
tricuspidatus.

Herpomyces zanzibarinus Thaxter

Natural hosts.—Eurycotis manni, Brazil (Thaxter, 1931): On antennae.

Gyna sp.?, Isle of Nias (Spegazzini, 1915): On antennae.
Cockroach, Zanzibar (Thaxter, 1902): On antennae.

INCERTAE SEDIS

According to Dr. R. K. Benjamin (personal communication, 1957) and Dr.
E. G. Simmons (personal communication, 1957), the phylogenetic position
of the following genus is uncertain.

Amphoromorpha blattina Thaxter

Natural hosts.—Cockroaches, Grenada, B.W.I. (Thaxter, 1920): On the axis
of the antennae of a dark wingless and a pale winged blattid.

Amphoromorpha sp.

Natural host.—Cockroach, Grenada, B.W.I. (Thaxter, 1920): On antennal
setae.

IX. HIGHER PLANTS
The significance of many observed associations between cockroaches and
the higher plants is still obscure. Undoubtedly many associations are
ecological, but lack of adequate supporting evidence makes this conclusion
somewhat tentative. The ecological aspects are covered in Section III (p.
14). Other associations may be accidental (e.g., certain unique observations
that have never again been confirmed). In the absence of contrary evidence,
most associations are presumed to be benign; exceptions to this conclusion

are found among the cockroaches that feed on living plants (p. 162) and
those allegedly captured as prey by the carnivorous pitcher plants
(Sarracenia and Nepenthes). In all the records cited below the cockroaches
were stated to have been on, in, or feeding on the plant.
The plants are listed below by family according to the taxonomic
arrangement of Lawrence (1951). Botanical nomenclature follows Bailey
(1925), Fernald (1950), or Dr. R. A. Howard (personal communications,
1958, 1959). We take full responsibility for referring to appropriate taxa
certain plants that were reported by common name only in the cited
literature.

Division PTERIDOPHYTA

Family CYATHEACEAE

Alsophila sp.

Associate.—Pycnoscelus surinamensis, Louisiana (Anonymous, 1893):
Feeding on heart of tree fern.

Family POLYPODIACEAE

Asplenium nidus Linnaeus

Associate.—Comptolampra liturata, Malaya (Karny, 1924): Often found
between dry foliage of the beakers of this fern.

Division EMBRYOPHYTA SIPHONOGAMA

Family PINACEAE

Pinus australis Michaux

Associates.—Aglaopteryx gemma and Parcoblatta lata, Alabama (Hebard,
1917): The former species was common under signs on longleaf pines, and
P. lata was occasional.
Parcoblatta divisa, Georgia (Rehn and Hebard, 1916): Under signs.

Pinus caribaea Morelet

Associates.—Eurycotis floridana, Latiblattella rehni, and Parcoblatta
fulvescens, Florida (Hebard, 1917): Many records under signs on the tree
trunks.

Pinus clausa Vasey

Associate.—Latiblattella rehni, Florida (Hebard, 1917): Under sign on tree.

Pinus echinata Mill.

Associates.—Parcoblatta divisa, Virginia (Rehn and Hebard, 1916): Under
signs on shortleaf pine.
Parcoblatta zebra, Mississippi (Hebard, 1917): Under sign.

Pinus sylvestris Linnaeus

Associate.—Ectobius pallidus, England (Milton, 1899; Burr, 1899b): On
Scotch fir.

Pinus spp.

Associates.—Plectoptera lacerna and Plectoptera vermiculata, Cuba (Rehn
and Hebard, 1927).
Latiblattella rehni, Florida (Rehn and Hebard, 1905): Under signs. Cuba
(Rehn and Hebard, 1927).

Family TAXODIACEAE

Cryptomeria sp.

Associate.—Diploptera punctata, Hawaii (Pemberton and Williams, 1938;
Zimmerman, 1948).

Family CUPRESSACEAE

Cupressus macrocarpa Hartweg

Associate.—Diploptera punctata, Hawaii (Hebard, 1922): "The species is
common and injurious in the territory infesting particularly the Monterey
cypress trees ... and doing particular damage by gnawing away the bark."
Similar injury has been cited by Pemberton (1934), Fullaway and Krauss
(1945), and Zimmerman (1948).

Juniperus sp.

Associate.—Phyllodromica tartara nigrescens, Southern Uzbekistan (Bei-
Bienko, 1950): Under bark.

Family PANDANACEAE

Freycinetia sp.

Associate.—Graptoblatta notulata and Kuchinga remota, Tahiti (Hebard,
1933).

Pandanus sp.

Associate.—Hololeptoblatta sp., Seychelles (Scott, 1910, 1912).

Family GRAMINEAE

Aristida pennata Trin.

Associate.—Phyllodromica pygmaea, U.S.S.R. (Bei-Bienko, 1950): Found
in the dense turf.

Bamboo

Associate.—Comptolampra liturata, Malaya (Karny, 1925).

Chloris gayana Kunth

Associate.—Blattella vaga, Texas (Riherd, 1953): This field cockroach was
rather abundant in clumps of Rhodes grass.

Panicum purpurascens Raddi

Synonymy.—Panicum barbinode [Hitchcock, 1936].
Associate.—Epilampra abdomen-nigrum, Puerto Rico (Seín, 1923; Wolcott,
1936): Abundant in "malojillo" meadow.

Saccharum officinarum Linnaeus

Associates.—Balta quadricaudata, Balta scripta, Balta torresiana, Balta
verticalis, Ellipsidion simulans, and Megamareta verticalis, Australia,
Queensland (Hebard, 1943): All collected by J. F. Illingworth on sugarcane.
Blattella humbertiana, Ischnoptera schenklingi, and Pycnoscelus
surinamensis, Formosa (Box, 1953).
Cariblatta stenophrys, Puerto Rico (Seín, 1923; Wolcott, 1936): Between
the leaves and under the leaf sheaths.
Panchlora nivea, Cuba (Rehn and Hebard, 1927): On the leaves.
Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1936).

Phoraspis spp., Brazil and Guiana (Doumerc in Blanchard, 1837).
Plectoptera dorsalis, Plectoptora infulata, and Plectoptera rhabdota,
Puerto Rico (Wolcott, 1950): Under the leaf sheaths.
Symploce ruficollis, Puerto Rico (Wolcott, 1950): Often found living under
the leaf sheaths.
Cockroaches, Philippine Islands (Uichanco in Williams et al., 1931):
Between cane leaf sheaths.

Setaria verticillata (Linnaeus) Beauv.

Synonymy.—Chaetochloa verticillata (Linnaeus) [Howard, personal
communication, 1958].
Associate.—Diploptera punctata, Hawaii (Severin, 1911): The cockroach
was caught on the barbed awns of this grass.

Wild oats

Associate.—Ischnoptera deropeltiformis, Missouri (Rau, 1937).

Zea mays Linnaeus

Associates.—Cariblatta stenophrys, Puerto Rico (Seín, 1923; Wolcott,
1936).
Ellipsidion bicolor, Australia, Queensland (Hebard, 1943).
Lophoblatta arawaka, Trinidad (Princis and Kevan, 1955).
Phoraspis sp., Brazil and Guiana (Doumerc in Blanchard, 1837).
Supella supellectilium, New Caledonia (Cohic, 1956).

Family CYPERACEAE

Cyperus sp.

Associate.—Maretina uahuka, Marquesas Islands, Uahuka (Hebard,
1933a).

Family PALMAE

Acrocomia aculeata (Jacq.) Lodd.

Associate.—Pycnoscelus surinamensis, Trinidad (Princis and Kevan, 1955):
On "gru-gru" fruits.

Cocos nucifera Linnaeus

Associates.—Aglaopteryx gemma, Florida (Rehn and Hebard, 1912).
Cariblatta lutea minima, Florida, and Cariblatta delicatula, San Domingo
(Hebard, 1916a).
Eurycotis floridana, Florida (Rehn and Hebard, 1912; Hebard, 1917).
Periplaneta australasiae, Jamaica (Rehn and Hebard, 1927).
Pycnoscelus surinamensis, Florida (Rehn and Hebard, 1912; Hebard, 1917).
Jamaica (Rehn and Hebard, 1927).

Phoenix dactylifera Linnaeus

Associate.—Blattella germanica, California (Herms, 1926): On date palms.

Pritchardia sp.

Associate.—Periplaneta australasiae, Nihoa Island (Bryan, 1926). Hawaii
(Zimmerman, 1948).

Roystonea regia O. F. Cook

Associate.—Cariblatta punctulata, San Domingo (Hebard, 1916a).

Sabal palmetto Lodd.

Associate.—Eurycotis floridana, Florida (Scudder, 1879).
Periplaneta australasiae, Florida (Hebard, 1917).

Undetermined palms

Associates.—Euthlastoblatta abortiva, Texas (Hebard, 1917).
Hormetica laevigata, Brazil (Hancock, 1926).
Panchlora antillarum, Dominican Republic (Rehn and Hebard, 1927).
Periplaneta americana, Texas (Zimmern in Gould and Deay, 1940).

Family ARACEAE

Arum sp.

Associate.—Latiblattella vitrea, Mexico (Hebard, 1921b): In flower shaft.

Caladium sp.

Associate.—Plectoptera dorsalis, Puerto Rico (Rehn and Hebard, 1927).

Family BROMELIACEAE

Aechmaea porteoides Britton

Associate.—Dryadoblatta scotti, Trinidad (Princis and Kevan, 1955).

Ananas comosus Merr.

Associates.—Pycnoscelus surinamensis, Hawaii (Illingworth, 1927, 1929):
Feeding on roots of pineapple.
Blattella humbertiana, Formosa (Takahashi, 1940): Imago and grown
nymphs occasionally lie concealed in the leaves.

Catopsis fulgens Griseb.

Associates.—Cockroaches, Costa Rica (Calvert and Calvert, 1917).

Glomeropitcairnia erectiflora Mez

Associate.—Dryadoblatta scotti, Trinidad (Princis and Kevan, 1955).

Grevisia sp.

Associate.—Notolampra antillarum, Trinidad (Princis and Kevan, 1955):
One male only.

Tillandsia fasciculata Swartz

Associate.—Eurycotis floridana, Florida (Rehn and Hebard, 1914; Hebard,
1917).

Tillandsia usneoides Linnaeus

Associates.—Parcoblatta sp., Louisiana (Rainwater, 1941).
Latiblattella rehni, Florida (Blatchley, 1920): By beating.
Cockroaches, Louisiana (Rosenfeld, 1911, 1912): One mature and 39
immature blattids were collected from 8 of 12 samples of Spanish moss.

Tillandsia uttriculata Linnaeus

Associate.—Epilampra mona, Mona Island, West Indies (Rehn and Hebard,
1927): The type and one paratypic female of E. mona were collected in this

bromeliad.
Eurycotis floridana, Florida (Blatchley, 1920).

Tillandsia sp.

Associates.—Aglaopteryx gemma, Texas (Hebard, 1917).
Dryadoblatta scotti, Trinidad (Scott, 1912): Found in the leaf bases.

Undetermined bromeliads

Associates.—Aglaopteryx diaphana, Jamaica (Hebard, 1917; Rehn and
Hebard, 1927).
Anaplecta azteca and Anaplecta sp., Costa Rica (Picado, 1913).
Anaplecta mexicana, Costa Rica (Calvert and Calvert, 1917).
Audreia bromeliadarum, Panama (Caudell, 1914).
Audreia jamaicana, Jamaica (Rehn and Hebard, 1927).
Blattella sp., Costa Rica (Picado, 1913).
Buboblatta armata, Panama (Caudell, 1914): "Probably not a typical
bromeliadicolous species."
Cariblatta insularis, Jamaica (Hebard, 1916a, 1917; Rehn and Hebard,
1927).
Cariblatta nebulicola, Jamaica (Rehn and Hebard, 1927); One
immature male.
Dryadoblatta scotti, Trinidad (Princis and Kevan, 1955).
Epilampra conspersa, Dominica (Scott, 1912).
Epilampra maya, Panama (Hebard, 1920).
Epilampra sodalis, Panama (Caudell, 1914).
Epilampra sp. and Hormetica laevigata, Brazil (Hancock, 1926).
Eurycotis biolleyi, Costa Rica (Picado, 1913).
Ischnoptera rufa occidentalis, Mexico (Caudell, 1914).
Latiblattella chichimeca, Costa Rica (Picado, 1913).
Litopeltis biolleyi, Costa Rica (Rehn, 1928).

Litopeltis bispinosa, Panama (Caudell, 1914).
Neoblattella brunneriana, Costa Rica (Calvert and Calvert, 1917).
Neoblattella dryas, Neoblattella eurydice, Neoblattella grossbecki, and
Neoblattella proserpina, Jamaica (Rehn and Hebard, 1927).
Neoblattella fratercula, Mexico (Hebard, 1921b).
Neoblattella nahua, Mexico (Caudell, 1914).
Nesomylacris relica, Jamaica (Rehn and Hebard, 1927).
Nyctibora brunnea(?), Panama (Caudell, 1914): According to Hebard
(1920) Caudell's specimen was almost certainly incorrectly identified.
It may have been Nyctibora noctivaga or a smaller species of the
genus. Brazil (Hancock, 1926).
Nyctibora laevigata, Jamaica (Hebard, 1917; Rehn and Hebard, 1927).
Nyctibora lutzi, Puerto Rico (Rehn and Hebard, 1927): "in epiphytes
with pencil-like leaves."
Pelmatosilpha rotundata, Panama (Caudell, 1914).
Pseudomops laticornis, Costa Rica (Picado, 1913).
Pycnoscelus surinamensis, Costa Rica (Picado, 1913). Mexico
(Caudell, 1914). Jamaica (Rehn and Hebard, 1927).
"Rhicnoda" sp., Costa Rica (Picado, 1913). This genus is now
recognized as not being in the New World fauna. Probably the
specimen was a species of Epilampra or Hyporhicnoda as suggested
by Gurney (personal communication, 1959) and confirmed by Rehn
(p.c., 1959).
Cockroaches, Costa Rica (Calvert, 1910): Cockroaches were said to be
common in bromeliads on the moist Atlantic slope.

Family LILIACEAE

Yucca elata Engelman

Associate.—Latiblattella lucifrons, Arizona (Ball et al., 1942).

Easter lilies

Associate.—Pycnoscelus surinamensis, Connecticut (Zappe, 1918).

Family MUSACEAE

Bananas

Cockroaches have been captured in bunches of bananas, in bracts of banana
flowers, under banana leaves, and burrowing in rotten banana stalks.
Although many of the species associated with bananas are indigenous to the
banana-growing areas of the American Tropics, most of the specimens cited
below were captured elsewhere as adventitious insects that had been
imported with the fruit. It is obvious that many of these insects must have
been closely associated with bananas on the plantations, where,
undoubtedly, the growing plants provided attractive ecological niches.
Bunting (1956) deduced, from the presence of healthy cockroaches on
bananas allegedly sprayed with copper arsenate, that the insects did not feed
on stems or fruit but hid among the bananas and foraged elsewhere;
however, certain reports are of cockroaches actually feeding on bananas.
Some of the records cited by Hebard (1917) were compiled from earlier
reports not all of which we have seen. Numbers in parentheses following
certain citations indicate the number of times the association had been
observed. Known or suspected adventive material is so indicated.
Aglaopteryx diaphana, Jamaica (Rehn and Hebard, 1927): Found in bracts
of banana blossoms. England (Bunting, 1955): Adventive, on bananas from
Dominica.
Aglaopteryx vegeta, Finland (Princis, 1947): Adventive, in banana box.
Amazonina emarginata, Trinidad (Princis and Kevan, 1955): In banana
bunch.
Archimandrita marmorata, Denmark (Henriksen, 1939): Adventive (2), in
bananas from Jamaica(?). As Princis (1947) and Gurney (personal

communication, 1959) point out, this is a Central American species, so
Jamaica may be an error.
Archimandrita tessellate, Sweden (Princis, 1947): Adventive, from
Honduras.
Blaberus atropos(?), Denmark (Henriksen, 1939): Adventive, from
Jamaica. Princis (1947) pointed out that this species was more likely to
have been Blaberus craniifer or Blaberus discoidalis, which are West
Indian species, than B. atropos which is a South American species.
Blaberus boliviensis, Ecuador (Princis, 1952): In a shipment of bananas
from near Puna.
Blaberus discoidalis, Puerto Rico (Rehn and Hebard, 1927): From banana
ripening room. Great Britain (Pearce, 1929): Adventive. England (Bunting,
1955, 1956): Adventive, from Dominica.
Capucinella delicatula, California (Caudell, 1931): Adventive.
Cariblatta delicatula, Cuba (Rehn and Hebard, 1927).
Cariblatta hylaea, Honduras (Rehn, 1945a): Shaken from hanging dead
banana leaves.
Cariblatta insularis, Finland (Frey, 1948): Adventive.
Cariblatta landalei, Jamaica (Rehn and Hebard, 1927): All specimens taken
from under drying bracts of banana blossoms.
Cariblatta punctipennis and Chorisoneura barbadensis, England (Bunting,
1956): Adventive, from Dominica.
Epilampra abdomen-nigrum and Epilampra sp., England (Bunting, 1955):
Adventive, from Dominica.
Epilampra maya, Massachusetts (Hebard, 1917): Adventive.
Epilampra mexicana(?), Denmark (Henriksen, 1939): Adventive (2), from
Danish West Indies. Princis (1947) suggested that this should be Epilampra
sp., because E. mexicana is not a West Indian species.
Eudromiella calcarata and Eurycotis bananae, U.S.S.R., Leningrad (Bei-
Bienko, 1947): Adventive, from Colombia.

Euphyllodromia angustata, Sweden (Princis, 1947): Adventive.
Eurycotis caraibea, New York (Hebard, 1917): Adventive.
Eurycotis dimidiata, Washington, D. C. (Caudell, 1931): Adventive.
Eurycotis lixa, New York (Rehn, 1930): Adventive, on banana ship from
Jamaica.
Graptoblatta notulata, Marquesas Islands, Uahuka (Hebard, 1933a): In
banana leaves.
Holocompsa nitidula, Trinidad (Princis and Kevan, 1955): Eating banana
pulp.
Hormetica laevigata, Wales (Sandemann, 1934): Adventive, in pile of
banana sacks.
Hormetica ventralis, Sweden (Princis, 1947): Adventive, in local warehouse
of banana company.
Hormetica spp., Europe and North America (Bei-Bienko, 1950): Adventive,
introduced with bananas and other tropical fruits.
Ischnoptera rufa rufa, Puerto Rico (Wolcott, 1950): Brought into houses on
bunches of bananas.
Kuchinga remota, Society Islands, Moorea (Hebard, 1933a): In dead banana
leaves.
Lamproblatta albipalpus, Panama Canal Zone (Hebard, 1920): Several
under decayed banana stem.
Latiblattella sp., Finland (Frey, 1948): Adventive.
Leucophaea maderae, New York (Hebard, 1917): Adventive. Dominica
(Rehn and Hebard, 1927): Under banana sheaths. England (Palmer, 1928):
Adventive, captured at railroad station after bananas had been unloaded.
England (Bunting, 1955): Adventive, from Dominica. Trinidad (Princis and
Kevan, 1955): Nymph, eating bananas in cupboard. Puerto Rico (Seín,
1923): Seín stated that bananas are the favorite food of L. maderae.
Litopeltis bispinosa, Panama Canal Zone (Hebard, 1920): From rotting
banana stalks at bases of leaves.

Litopeltis musarum, Costa Rica (Rehn, 1928): Shaken from dead banana
leaves.
Nauclidas nigra, England (Bunting, 1955, 1956): Adventive, from
Dominica.
Nauphoeta flexivitta, Denmark (Vestergaard, 1958): Adventive.
Neoblattella carcinus, Neoblattella celeripes, and Neoblattella laodamia,
England (Bunting, 1956): Adventive, from Dominica. Bunting (1955) first
reported these as Neoblattella spp. and stated that they were common.
Neoblattella detersa, Jamaica (Rehn and Hebard, 1927): From under the
bracts of banana blossoms. Sweden (Princis, 1947): Adventive.
Neoblattella detersa and Neoblattella tridens, Finland (Frey, 1948):
Adventive.
Neoblattella fratercula, Nebraska (Hebard, 1916b): Adventive.
Neoblattella semota, Jamaica (Rehn and Hebard, 1927): From under drying
bracts of banana blossoms.
Neoblattella vatia, Cuba (Rehn and Hebard, 1927).
Neoblattella sp., Finland (Princis, 1947): Adventive, from Jamaica.
Nyctibora azteca, England (Bunting, 1955): Adventive, from Dominica.
Bunting reported this species as Nocticola azteca. Dr. A. B. Gurney called
our attention to the fact that Nocticola is an Old World genus, presumably
combined in error with the New World species azteca. The true identity of
the specimen was confirmed by Dr. D. Ragge (personal communication,
1958), who examined it at the British Museum (Natural History).
Nyctibora holoserica, Canada (Walker, 1912): Adventive.
Nyctibora laevigata, Canada, Maine, Massachusetts, Pennsylvania (2)
(Hebard, 1917): Adventive. Taken from banana boat Annetta at Philadelphia
(Rehn and Hebard, 1927). England (Bunting, 1956): Adventive, from
Dominica. Sweden, Denmark (Princis, 1947): Adventive.
Nyctibora mexicana(?), Denmark (Henriksen, 1939): Adventive (5), from
Jamaica and West Indies. Princis (1947) suggested that these specimens

were probably the West Indian Nyctibora noctivaga, because N. mexicana is
not a West Indian insect.
Nyctibora noctivaga, Canada, Idaho, Illinois, Massachusetts, Nebraska (4),
Virginia (Hebard, 1917): Adventive. Nebraska (Hauke, 1949): Adventive
(2). Panama Canal Zone (Hebard, 1920): From banana stalks. England
(Blair in Turner, 1930): Adventive, from Costa Rica. Washington (Hatch,
1938): Adventive. Sweden (Princis, 1947): Adventive (2). Finland (Princis,
1947): Adventive, from Jamaica.
Nyctibora obscura, Trinidad (Princis and Kevan, 1955): In banana bunch.
Nyctibora sericea, Canada (Stevenson, 1905; Walker, 1912): Adventive;
Hebard (1917) synonymized Walker's specimen under N. laevigata. Isle of
Wight (Meade-Waldo, 1910): Adventive, from Jamaica. England (Tulloch,
1939): Adventive, in banana crates from Brazil.
Nyctibora sp., England (Welch, 1935): Adventive, in railway truck that had
carried bananas. England (Tulloch, 1939): Adventive, from Brazil.
Oxyhaloa deusta, U.S.S.R., Leningrad (Bei-Bienko, 1947): Adventive, from
Colombia.
Panchlora antillarum, England (Bunting, 1955): Adventive, from
Dominica.
Panchlora exoleta, Scotland (Distant, 1902): Adventive. Great Britain
(Shaw, 1902): Adventive. England (Coney, 1918): Adventive. Sweden,
Norway (Princis, 1947): Adventive, Norwegian specimen from Brazil.
Germany (Zacher, 1917): Adventive, from Jamaica.
Panchlora nivea, Colorado, Nebraska, New Jersey, New York (2), Utah
(Hebard, 1917): Adventive. Nebraska (Hauke, 1949): Adventive.
Washington (Hatch, 1938): Adventive. Massachusetts (Roth and Willis,
1958): Adventive. England (Bunting, 1955): Adventive, from Dominica.
U.S.S.R. (Bei-Bienko, 1947): Adventive, from Colombia. Sweden (13),
Norway (3), Finland (3) (Princis, 1947): Adventive, mostly females; origin
(where known) Jamaica.
Panchlora fraterna(?) and Panchlora peruana(?), Denmark (Henriksen,
1939): Adventive; origin (where known) Danish West Indies and Jamaica;
Princis (1947) suggested that both species were probably Panchlora nivea.