The Biotic Associations of Cockroaches
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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
[Illustration: Smithsonian Institution Logo]
"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.
[Illustration: Smithsonian Institution Logo]
(PUBLICATION 4422)
CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
DECEMBER 2, 1960
[Illustration: Plate 1
_Blaberus craniifer_, c. × 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.
[Illustration: Smithsonian Institution Logo]
(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
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 structures 74
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 found 99
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 primary hosts 192
Helminths for which cockroaches serve as intermediate
hosts 203
Helminths whose eggs have been carried by cockroaches 208
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
XIV. Checklist of cockroaches and symbiotic associates 290
XV. Cockroaches as commensals 310
Hosts of commensal cockroaches 310
Checklist of commensal cockroaches with their hosts 315
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
THE BIOTIC ASSOCIATIONS OF
COCKROACHES[1]
BY LOUIS M. ROTH AND EDWIN R. WILLIS[2]
_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 [[F][F] of _Ischnoptera insolita_ R. and H.;
_Ischnoptera uhleriana fulvescens_ S. and Z. (in part); Hebard
(1917)]
_desertae_ (Rehn and Hebard) [[M][M] 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
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