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MISCELLANEOUS COLLECTIONS
VOL. 141
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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 dead specimens (adults?) from 16 aircraft; 8
oöthecae from 7 planes; 147 live and 83 dead nymphs from 108 planes,
Miami (Denning et al., 1947). Southern U.S. (Hughes, 1949). Five live
and 7 dead specimens, Miami (Porter, personal communication, 1958).
IV. CLASSIFICATION OF THE ASSOCIATIONS
Asano (1937) classified the natural enemies of cockroaches into two
types as follows:
1. Enemies that feed mainly on cockroaches (certain ripiphorid
beetles and certain chalcid, evaniid, and ampulicid wasps).
2. Organisms which, in their search for food, devour cockroaches
that may be encountered (certain species of scorpions, spiders,
ticks, centipedes, Strepsiptera, ants, birds, rats, and
"parasitic bacteria").
Cameron (1955) arranged the associates of cockroaches in two groups as
follows:
Group A. Parasites and predators.
1. Parasites: Hymenoptera (Evaniidae, Eulophidae, Eupelmidae,
Encyrtidae, Pteromalidae, Cleonymidae) and Coleoptera
(Ripiphoridae).
2. Predators: Hymenoptera (Ampulicidae), Hemiptera (Reduviidae),
Coleoptera (Dermestidae), Arachnida (Araneae, Acarina).
Group B. Parasites and symbionts.
1. Protozoa (including examples of both parasites and
commensals).
2. Nematoda (including both primary "parasites" and secondary
parasites).
3. Bacteria (including the mutualistic bacteroids).
4. Algae [_Arthromitis_ (= _Hygrocrocis_) _intestinalis_; see p.
124].
Asano's arrangement, although essentially true, is limited; Cameron's
system is divided into arthropods (group A) and lower forms (group B),
but does not include higher animals. Both attempts at classification
need amplification; this we have endeavored to do below.
In classifying the biotic associates of cockroaches, we were immediately
confronted with a problem in semantics. The concepts parasitism,
predatism, and symbiosis have all been used with various shades of
meaning by different authors. The problem is not solved merely by
accepting as authoritative specific definitions, however apt they seem
to be, because, unfortunately, these concepts are not mutually
exclusive. For example, among the entomophagous insects, as Sweetman
(1936) has pointed out, there can be no definite line of separation
between parasitism and predatism: the two intergrade, only the extremes
being quite distinct. In fact, Andrewartha and Birch (1954) generalized
these relationships by calling both categories predatism. These authors
divided natural populations of associated organisms into nonpredators
and predators. Although this simplifies their presentation of the
general principles of ecology, for our purpose more narrowly defined
terms have proved useful.
In the main we have followed Sweetman (1936) and Allee et al. (1949) in
arriving at the following definitions:
=Symbiosis= is the living together in more or less intimate association
of organisms of different species; it includes virtually all
relationships between cockroaches and other organisms, such as
parasitism, predatism, commensalism, and mutualism. Allee et al. (1949)
apparently do not include predatism in symbiosis.
=Mutualism= is symbiosis in which both members benefit by the
association. The smaller partner has commonly been called a symbiont or
symbiote by authors.
=Commensalism= includes associations in which neither party appears to
benefit or be harmed. One partner may live on the surplus food or wastes
of the other; shelter and transport may be involved.
=Parasitism= is the state of symbiosis in which one of the members feeds
upon the other during the whole of either the immature or mature feeding
stage; the host is harmed in some way and may be killed.
=Predatism= is an association in which one member attacks and feeds
upon, or stores as food for its progeny, one or more other organisms;
the predator spends less than the immature or mature feeding period on
the prey. This category includes a few invertebrates and all the
vertebrates that capture, kill, and feed on cockroaches. This
association may be divided into interspecies predatism, in which the
predator preys upon a different species, and intraspecies predatism
(cannibalism) in which the predator preys upon its own species.
Although we have attempted to adhere to these definitions throughout
this discussion, we realize that in doing so we may have tended to
oversimplify complex relations. Some questionable interpretations stem
from insufficient knowledge of the basic relationships between
cockroaches and their associates. Only further study will clarify these
relationships. Some of the problems are discussed below.
Probably many of the so-called parasites (e.g., Protozoa like
_Nyctotherus_, and intestinal nematodes of the family Thelastomatidae),
which do not invade the host's tissues and seem to have no effect on the
activity and vitality of the host, are commensals. Although we consider
these forms to be commensals, we realize that they might actually affect
the host in some way even though this has not been shown. It is possible
that Rothschild and Clay's (1957) statement about bird parasites may
well apply to the apparently harmless organisms found in the cockroach.
These authors wrote, "It cannot be too strongly emphasized that the
effect of all types of parasites on the host is detrimental. If we find
that a bird seems little, if at all, inconvenienced by the presence of
Protozoa or worms or lice, or a cuckoo in the nest, we can nevertheless
assume that it would be better off without them.... Small effects such
as lack of vitality, loss of voice, excessive blinking, or perverted
habits like dirt eating are extremely difficult to gauge. Nevertheless,
it is only a question of degree. Potentially all parasites are harmful."
It should also be pointed out that some workers would consider certain
of our commensals of cockroaches to be parasites. Thus Faust (1955)
stated that "A truly successful parasite is one which has developed a
state of equilibrium with its host, so that no detectable damage is
produced which endangers the health or life of the host. In a suitable
host the parasite may obtain food and shelter without any evidence of
trauma or toxicity. The damage produced may be so slight that repair and
functional readjustment keep pace with the injury." Faust's successful
parasite would be indistinguishable from a commensal, but there is
undeniably a difference between an organism causing slight, and
undetectable, damage to a host and one causing none. Certain of the
organisms we list as commensals may eventually be shown to be parasites.
Certain organisms which live in cockroaches appear to have no effect on
the vitality of the host even though the tissues of the host are
invaded. Gregarines may penetrate the intestinal wall of the cockroach
without seeming to injure the host. Fungi of the genus _Herpomyces_
invade the cuticle of cockroaches producing pathological changes; yet
the insects' behavior is apparently unaffected (see p. 129). We consider
these organisms to be parasites because the host's tissues are invaded
and, as far as we know, no benefit to the host results.
In the literature certain insects have been considered to be either
parasites or predators or both. Among these are the ensign wasps
(Evaniidae), whose larvae feed on the eggs of cockroaches within the
oötheca, and the ampulicid wasps, which capture, paralyze, and store in
their nests (as food for their larvae) nymphs and adults of cockroaches.
Clausen (1940) claimed that the evaniid _Zeuxevania splendidula_ is a
true egg parasite when it destroys the first egg in a cockroach oötheca;
but after the wasp larva molts and proceeds to devour the other eggs, he
considered it to be a predator. Clausen's definition of an entomophagous
parasite is an insect that in its larval stage develops either
internally or externally upon a _single_ host which is eventually
killed; with few exceptions the adults are free-living and their food is
usually different from that of the larvae. A predatory insect, by
Clausen's definition, is principally free-living in the larval as well
as adult stage, kills the host immediately by direct attack, and
requires a number of victims to reach maturity; the predator is of
greater size than the prey, and the food sources of the adults and
immature stages are frequently the same.
It is apparent, as Clausen and other writers have pointed out, that
there are instances of a particular species showing characteristics
which fit both the definitions for predator and parasite. Thus, among
the evaniids one wasp larva destroys all the eggs in an oötheca, but in
spite of this the larva has more of the characteristics of a parasite
than of a predator; the adult wasp does not utilize the same food as the
larva (adults have been taken on flowers and on honeydew from scale
insects). It is questionable whether the evaniid larva kills the
cockroach egg outright. The wasp larva, being restricted to the inside
of the oötheca, is not free-living. Probably the only criterion by which
the evaniid could be judged to be a predator, by Clausen's definition,
is that more than a single egg is devoured by the maturing wasp larva.
Among the other wasp parasites (Encyrtidae, Eulophidae, Eupelmidae) of
cockroach eggs many individuals develop in a single oötheca. When a
hundred or more wasps emerge from an oötheca which contained less than
20 eggs, it is obvious that a single cockroach egg supported more than
one wasp, yet it is possible that one particular wasp larva may have fed
upon more than one cockroach egg before becoming an adult. We consider
all entomophagous wasps that develop in cockroach oöthecae to be
parasites rather than predators.
On the other hand, even though _Anastatus floridanus_, _A. tenuipes_,
and _Tetrastichus hagenowii_ are egg parasites as larvae, the adult
females are, in a sense, predators when they sometimes eat part of the
cockroach egg that oozes through the oviposition puncture (Roth and
Willis, 1954a, 1954b). Williams (1929) has seen the female of _Ampulex
canaliculata_ imbibe blood that oozed from the cut ends of the
cockroach's antennae after she had clipped them off before leading the
prey to her nest. Yet despite this evident predatism on the part of the
adult, the larva feeds as a parasite on the stored cockroaches in
accordance with Sweetman's (1936) (though not Clausen's 1940) definition
of parasitism, which is "that form of symbiosis in which one symbiont
lives in or on the host organism and feeds at its expense during the
whole of either the immature or mature feeding stage." The ampulicid
larva, as the evaniid, is not free-living and does not kill the host
immediately by direct attack, even though it may require more than one
victim to reach maturity. Thus, within one individual both parasitic and
predatory behavior are operant during different stages of its life
history.
With the above discussion in mind we have summarized below the various
biotic associations of cockroaches. Only a few examples are given for
each section, but all organisms with similar habits presumably would be
classified in the same categories.
Class A. Associations in which cockroaches serve as hosts, vectors,
or prey for other organisms.
Type I. Obligate associates. Animals and plants that normally
develop only on or in the cockroach; in general, these
organisms depend entirely upon the cockroach for survival.
Group 1. Mutuals (symbiotes or symbionts of authors).
(a) Bacteria-like organisms (bacteroids which are found in the
fat body of all cockroaches that have been examined; p. 96).
(b) Bacteria (wood-digesting forms in _Panesthia_, and possibly
certain bacteria in the intestines, of other cockroaches;
p. 100).
(c) Protozoa (several genera and species found in
_Cryptocercus_; p. 101).
Group 2. Commensals.
(a) Protozoa (_Nyctotherus_, _Herpetomonas_, _Lophomonas_, etc.;
p. 172).
(b) Nematodes (Thelastomatidae; p. 193).
Group 3. Parasites.
(a) Fungi (Laboulbeniales; p. 134).
(b) Protozoa (gregarines, _Plistophora_, etc.; p. 181).
(c) Helminths.
(1) Primary parasites (mermithids and gordian worms;
p. 201).
(2) Secondary parasites (_Gongylonema neoplasticum_,
_Oxyuris mansoni_, _Moniliformis_ spp.; p. 206).
(d) Arthropods.
(1) Mites (_Pimeliaphilus podapolipophagus_; p. 219).
(2) Insects (larvae of ripiphorids, evaniids, and
ampulicids; p. 231).
Type II. Facultative associates. Animals and plants that prey on
cockroaches or are incidentally or accidentally picked up by the
cockroach, but which can survive or propagate readily on some
other host or prey. Steinhaus (1946) emphasized the importance of
the environment in determining the type of microbial flora
associated with the cockroach which may carry one type of flora
in an area which is exposed to filth and a different type in
other areas. Because many of these organisms survive passage
through or on the cockroach, the blattid may act as a vector of
these animals and plants.
Group 1. Commensals.
(a) Viruses (strains of poliomyelitis virus; p. 103).
(b) Bacteria (Enterobacteriaceae, Pseudomonadaceae,
Micrococcaceae, etc.; p. 111).
(c) Fungi (_Aspergillus_; p. 130).
(d) Protozoa (_Iodamoeba_, _Dobellina_, and cysts of
_Entamoeba coli_ and _Entamoeba histolytica_; p. 179).
(e) Helminths (cysts of various helminths parasitic in
vertebrates; p. 208).
(f) Arthropods.
(1) Mites (_Tyrophagus lintneri_; p. 218).
Group 2. Parasites.
(a) Bacteria (_Serratia marcescens_; p. 117).
(b) Helminths (_Protospirura_ spp.; p. 206).
(c) Arthropods.
(1) Mites (_Locustacarus_ sp.; p. 219).
(2) Insects (_Melittobia chalybii_; p. 248).
Group 3. Predators, active.
(a) Arthropods.
(1) Spiders (p. 214).
(2) Scorpions (p. 212).
(3) Centipedes (p. 222).
(4) Mites (_Rhizoglyphus tarsalus_; p. 218).
(5) Insects (dermestids, reduviids, and on occasion adult
females of _Tetrastichus_, _Anastatus_, _Ampulex_;
p. 234).
(b) Vertebrates.
(1) Amphibia (p. 269).
(2) Reptilia (p. 272).
(3) Aves (p. 276).
(4) Mammalia (p. 283).
Group 4. Predators, passive: Pitcher plants (p. 154).
Class B. Associations in which cockroaches serve as commensals or
predators.
Group 1. Commensal cockroaches.
(a) Associates of social insects (_Attaphila_ spp., etc.;
p. 315).
(b) Obscure associates (p. 316).
Group 2. Predatory cockroaches.
(a) Interspecies predators (p. 319).
(b) Intraspecies predators (p. 322).
Class C. Associations of cockroaches with other cockroaches.
Group 1. Intraspecies associations.
(a) Familial associations (p. 325).
(b) Other conspecific associations (aggregations and fighting)
(p. 336).
Group 2. Interspecies associations.
(a) Compatible associations (p. 337).
(b) Antagonistic associations (p. 329).
Class D. Ecological associations of cockroaches with higher
plants.
Group 1. Benign associations (p. 139).
Group 2. Associations detrimental to plants (p. 162).
V. MUTUALISM
BACTEROIDS
Blochmann (1887, 1888) discovered intracellular particles (the
bacteroids or symbiotes of authors) that resembled bacteria in the fat
body of males and females of _Blatta orientalis_ and _Blattella
germanica_ (pl. 26), in the ova of these insects, and in their embryos.
Bacteroids have since been found in at least 25 species and 19 genera of
cockroaches. Presumably such microorganisms are universally distributed
throughout the Blattaria. General reviews of the bacteroids of
cockroaches and other insects have been published by Glaser (1930b),
Schwartz (1935), Steinhaus (1946, 1949), Buchner (1952, 1953), Brooks
(1954), and Richards and Brooks (1958). The reader is referred to these
papers, and those of authors cited in the list at the end of this
section, for discussions of the morphology of the bacteroids, their
distribution within the host, and attempts to culture them in vitro.
It has long been assumed, without proof, that cockroaches and their
bacteroids form a mutually beneficial association. As it has not been
possible to cultivate bacteroids apart from their cockroach hosts, it
may be assumed that the host is essential to the continued existence of
the microorganism, which also derives from the association other obvious
benefits as well. Experiments to show that the host also benefits from
the association have centered around rendering cockroaches bacteroid
free. Starvation, parasites, electromagnetic radiation, heat or cold, or
chemicals have all been used in attempts to eliminate the bacteroids. Of
these, only chemical treatment has provided a satisfactory technique.
Few chemicals other than antibiotics have proved to be useful in the
elimination or reduction of bacteroids. Yetwin (1932) injected various
dilutions of 22 compounds into _Blattella germanica_. He observed
decreases in the bacteroids of the fat body only following injection of
methylene blue, but did not pursue this lead further. Gier (_in_
Steinhaus, 1946) observed reduction in the numbers of bacteroids after
cockroaches were injected with crystal violet, hexylresorcinol, or
metaphen. Bode (1936) reported that injection of irritants such as
lithium salts or quinine hydrochloride had no apparent effect on the
symbiotes.
Brooks (1957) reared _Blattella germanica_ on diets containing different
concentrations of inorganic ions. On a manganese-deficient diet the
cockroaches grew poorly and some of their progeny lacked normal
bacteroids; about 10 percent of the aposymbiotic generation grew and
reproduced on a diet fortified with yeast. Varying the concentrations of
other salts in the diets gave results in which the progeny were either
aposymbiotic or the fat body was abnormal but the mycetocytes were
abundant; all these cockroaches soon died even on fortified diets.
The administration of certain antibiotic drugs has produced cockroaches
very nearly free of bacteroids. Brues and Dunn (1945) found that
although sulfa drugs had no effect on the bacteroids, penicillin in
large doses reduced the number of bacteroids in _Blaberus craniifer_,
but the cockroaches died within a few days. Death was attributed to lack
of bacteroids rather than effect of the drug; this is perhaps an
unwarranted conclusion in view of the survival of aposymbiotic
cockroaches that have been produced by other drugs (Brooks, 1954; Brooks
and Richards, 1955). Glaser (1946) found that the bacteroids of
_Periplaneta americana_ could be adversely affected or destroyed by
sulfathiazole, or sodium or calcium penicillin. Fraenkel (1952)
questioned the conclusions of Brues and Dunn (1945) and of Glaser (1946)
because of the high mortality in their experiments, and he suggested
that the described phenomena "were due rather to direct toxic effects on
the host than to loss of the symbionts." Noland (_in_ Brooks, 1954;
Brooks and Richards, 1955) confirmed Glaser's results with penicillin
and extended sulfa treatments to include _Blattella germanica_. Every
female whose bacteroids were reduced to the vanishing point resorbed her
ovaries and was incapable of reproduction. Brooks (1954; Brooks and
Richards, 1955) found that administration of several antibiotics did not
eliminate the bacteroids from the fat body of _B. germanica_ unless the
dose was so high that it caused excessive mortality. Frank (1955, 1956)
was able to eliminate bacteroids from _Blatta orientalis_ by injecting
or feeding chlortetracycline, oxytetracycline, or penicillin; survival
of treated insects was not good and reproduction was poor; the
aposymbiotic individuals were smaller than normal. As Richards and
Brooks (1958) have pointed out, it is uncertain how much of this
difference was the result of loss of bacteroids and how much the effect
of the drug. It is obvious that in all these experiments the action of
the drugs on the bacteroids was accompanied by equivocal side effects
which confused interpretation of the results. The effect on the
cockroach of a loss of bacteroids cannot be separated from a possible
toxic effect of the drug.
Fortunately Brooks (1954; Brooks and Richards, 1955) obtained completely
aposymbiotic offspring from _Blattella germanica_ that had been reared
on aureomycin. These bacteroid-free nymphs were practically incapable of
growth on a natural diet that was adequate for nymphs with symbiotes.
However, the addition of large amounts of dried yeast to the diet
enabled aposymbiotic nymphs to mature in two to three times the period
required by normal nymphs. Final proof of the function of the bacteroids
was obtained by reestablishing them in aposymbiotic cockroaches. The
insects that received implants of normal fat body of _B. germanica_
showed a slow, steady gain in weight over the controls (Brooks, 1954;
Brooks and Richards, 1956). Obviously, the intracellular symbiotes
subserve the normal nutrition of the cockroach. Whether the bacteroids
produce only vitaminlike substances, as suggested by Keller (1950), or
function in some other way is still to be determined. Brooks (1954)
concluded that the amount of vitamin-containing food required for
increased growth by aposymbiotic cockroaches is much greater than the
known vitamin requirements; hence the factor(s) needed is unknown and
present in low concentration, or it serves as a precursor of a second
factor(s) whose synthesis is aided by the bacteroids. Brooks (_in_
Richards and Brooks, 1958) has since found that the bacteroids of
_Blattella germanica_ "can supply the insect with B vitamins, amino
acids and some larger protein fragment."
Gier (1947) stated that the symbiotes of cockroaches are generically all
the same. However, as the symbiotes are presumed to have been associated
with cockroaches for over 300,000,000 years (Buchner, 1952) they may be
assumed to have developed specific differences that link them
inseparably to their respective hosts. Ries (1932) transplanted
symbiote-containing fat body from _Blatta orientalis_ into the mealworm
and larva of _Ephestia kühniella_, and from _Blattella germanica_ and
_Stegobium paniceum_ (=_Sitodrepa panicea_) into _B. orientalis_. The
implants did not become established in the new host, although most of
the transplantations were successful in that the hosts survived and the
implants remained intact for some time before they were encapsulated by
host tissue. Brooks (1954; Brooks and Richards, 1956) transplanted fat
body of _Periplaneta americana_ and _B. orientalis_ into aposymbiotic
_B. germanica_. The growth of the cockroaches injected with foreign
tissue was not different from that of aposymbiotic controls. Sections of
host insects did not contain mycetocytes and no bacteroids were found.
Haller (1955a) injected bacteroids or implanted mycetocytes of _B.
germanica_ into gryllids, acridids, and locustids. These implants and
innoculations were rapidly destroyed by the hosts. But as Richards and
Brooks (1958) have pointed out, none of these experiments provide
information about the specificity of the bacteroids themselves.
COCKROACHES IN WHICH BACTEROIDS HAVE BEEN FOUND
_Bantua stigmosa._ Fraenkel (1921)
_Blaberus craniifer._ Brooks (1954); Brues and Dunn (1945); Hoover
(1945).
_Blaberus giganteus._ Blochmann (1892).
_Blatta orientalis._ Blochmann (1887, 1888, 1892); Bode (1936);
Brooks (1954); Buchner (1912); Cuénot (1896); Fraenkel (1921);
Frank (1955, 1956); Gier (1936, 1947); Glaser (1920);
Gropengiesser (1925); Gubler (1948); Heymons (1895); Hollande
and Favre (1931); Hoover (1945); Hovasse (1930); Javelly (1914);
Keller (1950); Koch (1949); Menel (1907)?; Mercier (1906a, 1907,
1907b, 1907c); Ries (1932); Ronzoni (1949); Tacchini (1946); Wolf
(1924, 1924a).
_Blattella germanica._ Blochmann (1887, 1888, 1892); Bode (1936);
Borghese (1946, 1948, 1948a); Brooks (1954); Brooks and Richards
(1954, 1955, 1955a, 1956); Fraenkel (1921); Gier (1936, 1947);
Glaser (1920, 1930a); Gropengiesser (1925); Haller (1955, 1955a);
Heymons (1892, 1895); Hoover (1945); Koch (1949); Lwoff (1923);
Milovidov (1928); Neukomm (1927, 1927a, 1932); Pérez Silva (1954,
1954a); Ries (1932); Rizki (1954); Ronzoni (1949); Tacchini
(1946); Wollman (1926); Yetwin (1932, 1953).
_Cryptocercus punctulatus._ Gier (1936, 1947); Hoover (1945).
_Derocalymma cruralis._ Fraenkel (1921).
_Ectobius lapponicus._ Heymons (1892); Cuénot (1896); Koch (1949).
_Ectobius pallidus._ Heymons (1892, 1895); Cuénot (1896).
_Epilampra grisea._ Fraenkel (1921).
_Eurycotis floridana._ Gier (1936, 1947).
_Loboptera decipiens._ Buchner (1930).
_Nauphoeta cinerea._ Fraenkel (1921).
_Nyctibora noctivaga._ Gier (1947).
_Parcoblatta lata._ Gier (1936, 1947).
_Parcoblatta pensylvanica._ Brooks (1954); Gier (1936, 1947).
_Parcoblatta uhleriana._ Gier (1936, 1947); Hoover (1945).
_Parcoblatta virginica._ Glaser (1920); Gier (1947).
_Periplaneta americana._ Baudisch (1956); Bode (1936); Cuénot
(1896); Gier (1936, 1937, 1947); Glaser (1920, 1930, 1946);
Gubler (1948); Hertig (1921); Hoover (1945); Ketchel and
Williams (1953).
_Periplaneta australasiae._ Gier (1936, 1947); Koch (1949).
_Platyzosteria armata._ Fraenkel (1921).
_Polyphaga aegyptiaca._ Fraenkel (1921).
_Pseudoderopeltis aethiopica._ Fraenkel (1921).
_Pycnoscelus surinamensis._ Bode (1936); Koch (1949).
_Supella supellectilium._ Brooks (1954).
BACTERIA
Evidence showing that intestinal bacteria contribute to the nutrition of
cockroaches is meager. Cleveland et al. (1934) isolated a bacterial
organism from the foregut of the wood-feeding cockroach _Panesthia
angustipennis_. The bacterium digested cellulose rapidly in vitro and
these workers believe that this cockroach and other related wood-feeding
species are dependent on symbiotic bacteria for the digestion of their
food.
Mencl (1907) described cell nuclei in "symbiotic," not closely defined
types of bacilli that he found in abundance in the digestive tract of
the Küchenschabe, _Periplaneta_ (presumably _Blatta orientalis_).
Unfortunately, he was more concerned about the morphology of the
bacteria than the stated mutualistic relationship, so nothing is known
of their physiology.
The growth rates of _Periplaneta americana_ and _Blattella germanica_
were retarded when the insects were reared aseptically, which suggests
that microorganisms normally found in the digestive tract supply certain
necessary dietary constituents (Gier, 1947a; House, 1949). Noland et al.
(1949) suggested that microorganisms in the digestive tract of _B.
germanica_ synthesized riboflavin since the nymphs reared on a low
riboflavin diet accumulated more of the vitamin than could have been
ingested in the diet. However, Metcalf and Patton (1942) found little or
no bacterial synthesis of riboflavin in _P. americana_. Noland and
Baumann (1951) suggested that methionine, one of the amino acids
essential for rapid growth of _B. germanica_, was synthesized by
intestinal microorganisms in the insects.
PROTOZOA
It is probable that with few exceptions protozoa found in the digestive
tract are not necessary for survival of the cockroach. However, very few
experiments have been performed to determine the importance, if any, of
these microorganisms to the host. Cleveland (1925) removed the protozoa
from the cockroach (possibly _Periplaneta americana_) by oxygenation at
3.5 atmospheres. The ciliates _Nyctotherus_ and _Balantidium_,
flagellates _Lophomonas_ and _Polymastix_, the amoeba _Endamoeba
blattae_, and three unidentified protozoa were killed by this treatment,
yet the insects lived normally after defaunation.
Armer (1944) studied the effects of high-carbohydrate, high-fat, and
high-protein diets, as well as starvation, on the intestinal protozoa
(_Nyctotherus ovalis_, _Endamoeba blattae_, _Endolimax blattae_,
_Lophomonas striata_, and _Lophomonas blattarum_) in _Periplaneta
americana_. Starvation of the host lowered the incidence or eliminated
most of the protozoa, but a high-carbohydrate diet maintained them at a
relatively high level. _Lophomonas blattarum_ was eliminated by a
high-protein diet, and practically eliminated by a high-fat diet.
_Lophomonas striata_ was eliminated from some hosts that were kept on
high-fat and high-protein diets. _Endamoeba blattae_ showed a decrease
in infection rate when the cockroaches were maintained on high-fat and
high-protein diets. The effects of diets on _Endolimax blattae_ were not
uniform.
It has been shown by Cleveland (1930, 1948) and Cleveland et al. (1931,
1934) that the wood-feeding cockroach _Cryptocercus punctulatus_ depends
upon certain intestinal protozoa for survival; these protozoa utilize as
food the wood ingested by this cockroach. The wood is broken down into
compounds the cockroach can utilize by the protozoa which elaborate a
cellulase and possibly a cellobiase (Trager, 1932). Only molting nymphs
of _Cryptocercus_ can pass the protozoa on to the newly hatched young,
so that molting and hatching must happen concurrently each year or the
young die.
The sexual cycles in species of protozoa in the genera _Trichonympha_,
_Saccinobaculus_, _Oxymonas_, _Monocercomonoides_, _Hexamita_,
_Eucomonympha_, _Leptospironympha_, _Urinympha_, _Rhynchonympha_,
_Macrospironympha_, and _Barbulanympha_ (fig. 3, B) are induced by
hormones produced by _Cryptocercus_ only during its molting period
(Cleveland, 1931, 1947, 1947a, 1949-1956a). Perhaps the prothoracic
gland hormone of the host may be responsible for initiation of the
flagellate sexual cycles (Cleveland and Nutting, 1955). The protozoan
sexual cycles may be used as indicators of the onset of molting in
_Cryptocercus_; thus different species of protozoa begin their sexual
cycles from 35 days before to 2 days after molting of the cockroach
(Cleveland and Nutting, 1954). Hollande (1952) and Grassé (1952) have
reviewed the roles and the evolution of the flagellates in
_Cryptocercus_ and in termites.
The protozoa of cockroaches and termites are clues to the relationship
between these two groups of insects. Kirby (1927) pointed out
similarities between _Endamoeba blattae_ of _Periplaneta_ and the
amoebae of the termite _Mirotermes_, suggesting that these protozoans
were probably derived from an amoeba in an ancestor common to both
blattid and termite. Kirby (1932, _in_ Kidder, 1937) found a species of
_Nyctotherus_ in _Amitermes_ that resembles _Nyctotherus ovalis_ from
domestic cockroaches. The belief that the termites and cockroaches had a
common origin is also strengthened by the similarities between the
hypermastigotes of both _Cryptocercus_ and termites (Cleveland et al.,
1934).
The cockroaches _Cryptocercus_ and _Panesthia_ both feed on wood, but
the protozoa found in _Panesthia_ resemble more closely the species in
domestic cockroaches than those in _Cryptocercus_. The Clevelandellidae
(from _Panesthia_) are closely related to _Nyctotherus_ and have
probably evolved from common ancestors. However, the separation of the
Clevelandellidae from _Nyctotherus_ must have taken place at a later
date than the divergence of their hosts, otherwise representatives of
that family would probably also be found in _Periplaneta_ and _Blatta_
(Kidder, 1937).
The protozoa of _Cryptocercus_ can be transferred from one individual to
another (Nutting and Cleveland, 1954). They can also be transferred to
the termite _Zootermopsis_ where they survive only until the host molts;
the reverse is also true, _Zootermopsis_ Protozoa can survive in
_Cryptocercus_ until the cockroach molts (Nutting and Cleveland, 1954a).
VI. VIRUSES ASSOCIATED WITH COCKROACHES
Annotations on some of the following observations may be found in Roth
and Willis (1957a). Use of asterisk is explained in footnote 3, page 4.
POLIOMYELITIS VIRUSES
* =Lansing strain=
_Experimental vectors._--_Blattella germanica_, U.S.A. (Hurlbutt, 1949,
1950).
_Periplaneta americana_, U.S.A. (Hsiang et al., 1952).
* =Brunhilde type, Minnesota and Mahoney strains=
_Experimental vectors._--_Periplaneta americana_, U.S.A. (Fischer and
Syverton, 1951; Syverton et al., 1952).
* =Columbia SK virus=
_Experimental vectors._--_Periplaneta americana_, Great Britain?
(Findlay and Howard, 1951): Results with _Blattella germanica_ were
negative.
* =Four unspecified strains=
_Natural vectors._--_Blattella germanica_ and/or _Blattella vaga_,
_Periplaneta americana_ and/or _Periplaneta brunnea_, and _Supella
supellectilium_, U.S.A. (Syverton et al., 1952; Dow, 1955; Dow _in_ Roth
and Willis, 1957a).
OTHER VIRUSES
* =Coxsackie viruses=
_Experimental vectors._--_Periplaneta americana_, U.S.A. (Fischer and
Syverton, 1951a, 1957): Recently Fischer and Syverton (1957) found that
after feeding a single meal of Coxsackie virus to _Periplaneta
americana_, the gastrointestinal tracts of the insects, which were
removed at 5-day intervals up to 20 days, contained sufficient virus to
paralyze and kill test mice. Cockroach salivary glands, removed 5 days
after the insects had fed, contained the virus which caused paralysis
and death in test mice; mice were also infected by virus obtained from
salivary glands removed from the insects 10 and 20 days after the
cockroaches had fed once on the virus. The virus was also isolated from
the cockroaches' feces and rarely from the fat bodies and reproductive
organs. Fischer and Syverton concluded that it is possible that
cockroaches could acquire the virus, by feeding on mammalian excreta,
maintain it for a period of time, and transmit it by contamination of
food. The virus could also be transmitted through the feces of wild mice
if the mice happened to feed on virus-infected cockroaches.
* =Mouse encephalomyelitis virus=
_Experimental vectors._--_Periplaneta americana_, U.S.A. (Syverton and
Fischer, 1950).
* =Yellow-fever virus=
_Experimental vectors._--_Blattella germanica_, Great Britain? (Findlay
and MacCallum, 1939).
VII. BACTERIA ASSOCIATED WITH COCKROACHES
Classification of the bacteria follows Breed et al. (1948). Synonymy in
most cases was taken from the same source. Names of bacteria preceded by
the symbol [cross] are either not listed by Breed et al. or are stated
by them to be insufficiently characterized for definite classification.
Use of asterisk is explained in footnote 3, page 4.
Phylum SCHIZOPHYTA
Class SCHIZOMYCETES
Order EUBACTERIALES
Family PSEUDOMONADACEAE
* =Pseudomonas aeruginosa= (Schroeter) Migula
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949).
_Blatta orientalis_, U.S.A. (Olson and Rueger, 1950).
_Blattella germanica_, U.S.A. (Olson and Rueger, 1950; Janssen and
Wedberg, 1952).
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949, 1949a; Olson
and Rueger, 1950).
_Experimental vectors._--_Blattella germanica_, U.S.A. (Herms and
Nelson, 1913).
Cockroaches, U.S.A. (Longfellow, 1913).
* =Pseudomonas eisenbergii= Migula
_Natural and experimental vectors._--_Blatta orientalis_, Italy (Cao,
1906).
* =Pseudomonas fluorescens= Migula
_Natural and experimental vectors._--_Blatta orientalis_, Italy (Cao,
1898, 1906; Spinelli and Reitano, 1932).
_Periplaneta americana_, U.S.A. (Gier, 1947): The organism was
pathogenic to the cockroach when injected.
[cross] =Spirillum periplaneticum= Kunstler and Gineste
_Habitat._--_Periplaneta americana_, France? (Kunstler and Gineste,
1906): From intestinal tract.
[cross] =Spirillum α, β, and γ= Dobell
_Habitat._--_Blatta orientalis_, England (Dobell, 1911, 1912): From hind
gut.
=Spirillum= sp.
_Habitat._--_Blatta orientalis_, U.S.S.R. (Zasukhin, 1930): From
intestinal tract.
Cockroaches, Venezuela (Tejera, 1926): From digestive tract.
* =Vibrio comma= (Schroeter) Winslow et al.
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898; Spinelli
and Reitano, 1932).
_Blattella germanica_, Orient (Toda, 1923); Germany (Jettmar, 1927).
_Periplaneta americana_, Philippine Islands (Barber, 1914); Netherlands
(Akkerman, 1933); Formosa (Morischita and Tsuchimochi, 1926).
_Periplaneta australasiae_, Formosa (Morischita and Tsuchimochi, 1926).
* =Vibrio metschnikovii= Gamaléia
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898, 1906).
=Vibrio tyrogenus= (Flügge) Holland
_Synonymy._--_Vibrio_ of Deneke.
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898): The
organism passed through the intestinal tract unchanged.
[cross] =Vibrio Types I and II= Heiberg
_Habitat._--Water.
_Natural vectors._--Cockroaches, India (Pasricha et al., 1938): The
vibrios were found in 16 or 17 percent of 94 cockroaches and resembled
_Vibrio comma_ in their morphology and their main biochemical reactions;
however, serum-agglutination reactions differed.
=Vibrio= sp.
_Habitat._--_Blatta orientalis_, U.S.A. (Leidy, 1853): From intestine.
Family RHIZOBIACEAE
=Chromobacterium violaceum= (Schroeter) Bergonzini
_Synonymy._--_B. violaceus._
_Habitat._--Water.
_Experimental vectors._--Cockroach, U.S.A. (Longfellow, 1913): Recovered
from outer part of body and intestinal tract.
Family MICROCOCCACEAE
* =Micrococcus aurantiacus= (Schroeter) Cohn
_Natural vectors._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952).
* =Micrococcus citreus= Migula
_Natural vectors._--Cockroaches, U.S.A. (Longfellow, 1913).
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
* =Micrococcus epidermidis= (Winslow and Winslow) Hucker
_Natural vectors._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952).
[cross] =Micrococcus nigrofaciens= Northrup
_Source._--Diseased June beetle larvae.
_Experimental infection._--_Periplaneta americana_, U.S.A. (Northrup,
1914): Three of four adults were infected by feeding them bread
saturated with a broth culture of the _Micrococcus_. After 11 days the
tarsi of the cockroaches became infected, and the hind legs split and
broke off. Antennae and setae also were affected and micrococci were
recovered from the feces.
* =Micrococcus pyogenes= var. =albus= (Rosenbach) Schroeter
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al.,
1949).
_Blatta orientalis_, U.S.A. (Tauber, 1940; Tauber and Griffiths, 1942).
_Blattella germanica_, U.S.A. (Herms and Nelson, 1913; Herms, 1939;
Janssen and Wedberg, 1952).
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898); U.S.A.
(Tauber and Griffiths, 1942).
_Micrococcus pyogenes_ var. _albus_ (=_Staphylococcus albus_) and an
unidentified short rod form were found by Tauber (1940) in the hemolymph
of _B. orientalis_. These microorganisms were never found together in
the same insect and caused loss of appetite, sluggishness, irregular
respiratory movements, and paralysis in the cockroach; in the final
stages of the disease the legs were folded under the body, the head was
tucked beneath the forelegs, the whole insect became arched and
maintained this position until death. In some cockroaches infected with
the rod pathogen, conjunctival folds, particularly those between the
dorsal abdominal sclerites, and the joints of the metathoracic legs
ruptured liberating thick white hemolymph filled with bacteria. Tauber
suggested that the infection might be spread by contact, especially to
newly molted individuals or by actual ingestion of the bacteria by the
cockroaches feeding on dead or dying individuals. All the roaches died
after successful inoculation with the _Micrococcus_. The bacterial
infection was associated with high total hemocyte counts and high
percentages of mitotically dividing hemolymph cells (Tauber, 1940);
these responses of the insect were interpreted as a mechanism whereby
the number of hemocytes increases resulting in an increase in the number
of phagocytes for combating the bacteria (Tauber and Griffiths, 1942).
* =Micrococcus pyogenes= var. =aureus= (Rosenbach) Zopf
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949).
_Blatta orientalis_, Italy (Cao, 1906).
_Blattella germanica_, U.S.A. (Herms, 1939).
Cockroaches, U.S.A. (Longfellow, 1913).
=Micrococcus ureae= Cohn
_Habitat._--Stale urine and soil containing urine.
_Natural vectors._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952): From intestinal tract and feces.
* =Micrococcus= spp.
These organisms were obtained from pus or were designated as
staphylococci [i.e., pathogenic micrococci (Blair _in_ Dubos, 1948)].
_Natural vectors._--_Blatta orientalis_, Italy (Spinelli and Reitano,
1932); Germany (Jettmar, 1935).
_Blattella germanica_, Germany (Jettmar, 1935).
_Experimental vectors._--_Blattella germanica_, on shipboard (Morrell,
1911); Germany (Vollbrechtshausen, 1953).
=Micrococcus= sp.
_Natural vectors._--_Periplaneta americana?_ ("_Blatella americana_"),
England (Shrewsbury and Barson, 1948): From intestinal tract.
[cross] =Sarcina alba= Zimmermann
_Habitat._--Water.
_Natural and experimental vectors._--_Blatta orientalis_, Italy (Cao
1898, 1906): From intestinal contents.
=Sarcina aurantiaca= Flügge
_Habitat._--Air and water.
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1906):
Intestinal contents.
=Sarcina lutea= Schroeter
_Habitat._--Air, soil, water, skin surfaces.
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1906): From
intestinal contents.
[cross] =Sarcina symbiotica= Pribram
_Habitat._--Oöthecae of _Blatta orientalis_ and/or _Blattella
germanica_, Germany (Gropengiesser, 1925): It was described as "eine
gelbe _Sarcina_"; Pribram (1933) named the organism.
=Sarcina ventriculi= Goodsir
_Habitat._--Garden soil, dust, mud; isolated from a diseased stomach.
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946):
From intestinal tract.
=Sarcina= sp.
_Natural and experimental vectors._--_Periplaneta americana_, U.S.A.
(Gier, 1947): Organism not pathogenic to the cockroach when injected.
[cross] =Sarcina= sp.
_Synonymy._--_Sarcina_ "blanche" of Sartory and Clerc.
_Natural vectors._--_Blatta orientalis_, France (Sartory and Clerc,
1908): Isolated from intestinal tract.
*[cross] =Sarcina= sp.
_Synonymy._--_Sarcina alba_ "patogena" of Cao.
_Natural vectors._--_Blatta orientalis_, Italy (Cao, 1898, 1906).
*[cross] =Sarcina= sp.
_Synonymy._--_Sarcina_ "bianca" and "gialla" of Cao.
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
Family NEISSERIACEAE
* =Neisseria meningitidis= (Albrecht and Ghon) Holland
_Experimental vectors._--Cockroaches, U.S.A. (Longfellow, 1913).
* =Veillonella parvula= (Veillon and Zuber) Prévot
_Natural vectors._--_Periplaneta americana_, U.S.A. (Hatcher, 1939).
Family LACTOBACTERIACEAE
* =Diplococcus pneumoniae= Weichselbaum
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
Cockroaches, U.S.A. (Longfellow, 1913).
[cross] =Enterococcus= sp.
_Natural vectors._--_Blatta orientalis_, Italy (Spinelli and Reitano,
1932): From intestinal tract.
=Lactobacillus fermenti= Beijerinck
_Habitat._--Fermenting plant or animal products.
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946):
From intestinal canal.
[cross] =Pneumococcus Type I, No. 1231=
_Experimental vectors._--_Blattella germanica_, Germany
(Vollbrechtshausen, 1953).
* =Streptococcus faecalis= Andrewes and Horder
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946).
_Blattella germanica_, U.S.A. (Steinhaus, 1941).
_Periplaneta americana?_ ("_Blatella americana_"), England (Shrewsbury
and Barson, 1948).
Cockroaches [presumably any or all of the above three species], Egypt
(El-Kholy and Gohar, 1945).
* =Streptococcus liquefaciens= Sternberg emend. Orla-Jensen
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946).
*[cross] =Streptococcus microapoika= Cooper, Keller, and Johnson
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946).
*[cross] =Streptococcus non-hemolyticus II= Holman
_Natural vectors._--_Periplaneta americana?_ ("_Blatella americana_"),
England (Shrewsbury and Barson, 1948).
* =Streptococcus pyogenes= Rosenbach
_Natural vectors._--_Blatta orientalis_, Italy (Cao, 1906).
_Experimental vectors._--Cockroaches, U.S.A. (Longfellow, 1913).
* =Streptococcus= sp. (pyogenic group)
_Experimental vectors._--_Blatta orientalis_, Germany (Jettmar, 1935).
* =Streptococcus= sp. (_viridans_ group)
_Experimental vectors._--_Blatta orientalis_, Germany (Jettmar, 1935).
* =Streptococcus= spp.
_Natural vectors._--_Blatta orientalis_, Germany (Jettmar, 1935).
_Blattella germanica_, Germany (Jettmar, 1935); U.S.A. (Janssen and
Wedberg, 1952).
Cockroaches, U.S.A. (Longfellow, 1913).
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
Family CORYNEBACTERIACEAE
* =Corynebacterium diphtheriae= (Flügge) Lehmann and Neumann
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
Cockroaches, U.S.A. (Longfellow, 1913).
Family ACHROMOBACTERIACEAE
[cross] =Achromobacter hyalinum= (Jordan) Bergey et al.
_Natural vectors._--_Periplaneta americana_, U.S.A. (Hatcher, 1939):
Isolated from feces.
=Achromobacter= sp.
_Natural vectors._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952).
=Alcaligenes faecalis= Castellani and Chalmers
_Synonymy._--_Bacillus faecalis alkaligenes_; _Bacillus alcaligenes
faecalis_; _B. alcaligenes faecalis_.
_Habitat._--Intestinal canal of man. Has been isolated from feces,
abscesses related to intestinal canal, and occasionally in the
bloodstream. However, it is generally considered nonpathogenic.
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949):
Isolated from feces.
_Blatta orientalis_, Poland (Nicewicz et al., 1946): Isolated from
intestinal tract.
_Blattella germanica_, U.S.A. (Janssen and Wedberg, 1952): From
intestinal tract and feces.
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949): Isolated
from intestinal tract.
Cockroaches [presumably _Blatta orientalis_, _Blattella germanica_
and/or _Periplaneta americana_], Egypt (El-Kholy and Gohar, 1945): From
suspensions of macerated whole insects.
=Alcaligenes recti= (Ford) Bergey et al.
_Synonymy._--_B. alcaligenes recti._
_Habitat._--Intestinal canal.
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946):
Isolated from intestinal tract.
=Alcaligenes viscosus= (Weldin and Levine) Weldin
_Habitat._--Water, dairy utensils; produces ropiness in milk.
_Natural vectors._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952): Isolated from intestine and feces.
Family ENTEROBACTERIACEAE
=Aerobacter aerogenes= (Kruse) Beijerinck
_Synonymy._--_Bacillus lactis aerogenes._
_Habitat._--Grains, plants, intestinal tract of man and other animals.
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949):
Isolated from feces.
_Blatta orientalis_, Poland (Nicewicz et al., 1946): Isolated from
intestinal tract.
_Blattella germanica_, on shipboard (Morrell, 1911): Isolated from
feces.
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949): Isolated
from intestinal tract.
Cockroaches [presumably _Blatta orientalis_, _Blattella germanica_
and/or _Periplaneta americana_], Egypt (El-Kholy and Gohar, 1945):
Isolated from outer surface of body, intestinal tract, and suspensions
of macerated whole insects.
=Aerobacter cloacae= (Jordan) Bergey et al.
_Synonymy._--_Bacillus cloacae._
_Habitat._--Sewage, soil, water, human and other animal feces.
_Natural vectors._--_Blattella germanica_, on shipboard (Morrell, 1911):
From feces. U.S.A. (Janssen and Wedberg, 1952; Steinhaus, 1941): From
intestinal tract, feces, and oötheca.
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949): From
intestinal tract.
=Aerobacter= sp.
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949): Isolated from intestines.
[cross] =Eberthella oedematiens= Assis
_Habitat._--Intestinal canal.
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949): Intestinal tract.
* =Escherichia coli= (Migula) Castellani and Chalmers
_Natural vectors._--_Blatta orientalis_, Italy (Cao 1898, 1906; Spinelli
and Reitano, 1932); France (Sartory and Clerc, 1908); Europe (Jettmar,
1935); Poland (Nicewicz et al., 1946).
_Blattella germanica_, U.S.A. (Steinhaus, 1941).
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949, 1949a).
Cockroaches [presumably one or all of the above three species], Egypt
(El-Kholy and Gohar, 1945).
Cockroaches, U.S.A. (Longfellow, 1913).
_Experimental vectors._--_Blattella germanica_, Germany
(Vollbrechtshausen, 1953): When injected anally or orally the bacteria
invaded the intestinal cells and in heavy infections killed the
cockroaches.
=Escherichia coli= var. =acidilactici= (Topley and Wilson) Yale
_Synonymy._--_Bacillus acidi lactici._
_Source._--Diseased nun moth larvae.
_Experimental vectors._--_Blatta orientalis_, Europe (Filatoff, 1904):
Organism pathogenic to cockroach when injected but not when fed.
=Escherichia coli= var. =communior= (Topley and Wilson) Yale
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949):
From feces.
=Escherichia freundii= (Braak) Yale
_Habitat._--Soil, water, intestinal canal of man and other animals.
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949):
From feces.
_Blattella germanica_, U.S.A. (Janssen and Wedberg, 1952): From
intestinal canal and feces.
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949): From
intestinal tract.
=Escherichia intermedium= (Werkman and Gillen) Vaughn and Levine
_Habitat._--Soil, water, intestinal canal of man and other animals.
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949): From intestinal canal.
* =Klebsiella pneumoniae= (Schroeter) Trevisan
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
* =Paracolobactrum aerogenoides= Borman, Stuart and Wheeler
_Natural vectors._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952).
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949).
* =Paracolobactrum coliforme= Borman, Stuart and Wheeler
_Natural vectors._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952).
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949).
* =Paracolobactrum= spp.
_Natural vectors._--_Blatta orientalis_, U.S.A. (Olson and Rueger,
1950).
_Blattella germanica_, U.S.A. (Olson and Rueger, 1950).
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949; Olson and
Rueger, 1950).
* =Paracolon bacilli=
_Natural vectors._--Cockroaches [presumably _Blatta orientalis_,
_Blattella germanica_ and/or _Periplaneta americana_], Egypt (El-Kholy
and Gohar, 1945).
* =Proteus mirabilis= Hauser
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949, 1949a).
* =Proteus morganii= (Winslow et al.) Rauss
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949, 1949a).
Cockroaches [presumably _Blatta orientalis_, _Blattella germanica_,
and/or _Periplaneta americana_], Egypt (El-Kholy and Gohar, 1945).
* =Proteus rettgeri= (Hadley et al.) Rustigian and Stuart
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949).
* =Proteus vulgaris= Hauser
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949).
_Blatta orientalis_, Italy (Spinelli and Reitano, 1932).
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949, 1949a).
Cockroaches, U.S.A. (Longfellow, 1913).
* =Proteus= spp.
_Natural vectors._--_Blatta orientalis_, U.S.A. (Olson and Rueger,
1950).
_Periplaneta americana_, U.S.A. (Bitter and Williams, 1949; Olson and
Rueger, 1950).
* =Salmonella anatis= (Rettger and Scoville) Bergey et al.
_Natural vectors._--_Periplaneta americana_, U.S.A. (Eads et al., 1954).
* =Salmonella choleraesuis= (Smith) Weldin
_Experimental vectors._--_Polyphaga saussurei_, U.S.S.R. (Zmeev _in_
Pavlovskii, 1948).
* =Salmonella enteritidis= (Gaertner) Castellani and Chalmers
_Experimental vectors._--_Blatta orientalis_, U.S.S.R. (Rozengolts and
Ȋȗdina _in_ Pavlovskii, 1948).
_Blattella germanica_, U.S.A. (Olson and Rueger, 1950); U.S.S.R.
(Rozengolts and Ȋȗdina _in_ Pavlovskii, 1948).
_Polyphaga saussurei_, U.S.S.R. (Zmeev _in_ Pavlovskii, 1948).
* =Salmonella morbificans= (Migula) Haupt
_Natural vectors._--_Periplaneta americana_, Australia (Mackerras and
Mackerras, 1948, 1949).
_Experimental vectors._--_Nauphoeta cinerea_, _Periplaneta
australasiae_, _Periplaneta ignota_, and _Supella supellectilium_,
Australia (Mackerras and Pope, 1948).
* =Salmonella paratyphi= (Kayser) Castellani and Chalmers
_Experimental vectors._--_Polyphaga saussurei_, U.S.S.R. (Zmeev _in_
Pavlovskii, 1948).
* =Salmonella schottmuelleri= (Winslow et al.) Bergey et al.
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949, 1949a).
_Experimental vectors._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922).
_Polyphaga saussurei_, U.S.S.R. (Zmeev _in_ Pavlovskii, 1948).
* =Salmonella= sp. (=Type Adelaidae=)
_Experimental vectors._--_Nauphoeta cinerea_, _Periplaneta
australasiae_, _Periplaneta ignota_, and _Supella supellectilium_,
Australia (Mackerras and Pope, 1948).
* =Salmonella= sp. (=Type Bareilly=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Eads et al., 1954).
* =Salmonella= sp. (=Type Bredeny=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949, 1949a).
* =Salmonella= sp. (=Type Derby=)
_Experimental vectors._--_Nauphoeta cinerea_, _Periplaneta
australasiae_, and _Supella supellectilium_, Australia (Mackerras and
Pope, 1948).
* =Salmonella= sp. (=Type Kentucky=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Eads et al., 1954).
* =Salmonella= sp. (=Type Kottbus=)
_Experimental vectors._--_Periplaneta australasiae_, Australia
(Mackerras and Pope, 1948).
* =Salmonella= sp. (=Type Meleagris=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Eads et al., 1954).
* =Salmonella= sp. (=Type Montevideo=)
_Experimental vectors._--_Periplaneta americana_, U.S.A. (Jung and
Shaffer, 1952).
* =Salmonella= sp. (=Type Newport=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Eads et al., 1954).
* =Salmonella= sp. (=Type Oranienburg=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949, 1949a; Eads et al., 1954).
_Experimental vectors._--_Blatta orientalis_, _Blattella germanica_, and
_Periplaneta americana_, U.S.A. (Olson and Rueger, 1950).
* =Salmonella= sp. (=Type Panama=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Eads et al., 1954).
* =Salmonella= sp. (=Type Rubislaw=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Eads et al., 1954).
* =Salmonella= sp. (=Type Tennessee=)
_Natural vectors._--_Periplaneta americana_, U.S.A. (Eads et al., 1954).
* =Salmonella typhimurium= (Loeffler) Castellani and Chalmers
_Natural vectors._--_Blattella germanica_, Belgium (Graffar and Mertens,
1950).
_Nauphoeta cinerea_, Australia (Mackerras and Mackerras, 1948).
_Experimental vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al.,
1949).
_Blatta orientalis_, U.S.S.R. (Rozengolts and Ȋȗdina _in_ Pavlovskii,
1948).
_Blattella germanica_, Belgium (Graffar and Mertens, 1950); U.S.A.
(Olson and Rueger, 1950; Janssen and Wedberg, 1952; Beck and Coffee,
1943); U.S.S.R. (Rozengolts and Ȋȗdina _in_ Pavlovskii, 1948).
_Nauphoeta cinerea_, Australia (Mackerras and Pope, 1948).
_Periplaneta americana_, U.S.A. (Beck and Coffee, 1943; Jung and
Shaffer, 1952).
_Periplaneta australasiae_ and _Supella supellectilium_, Australia
(Mackerras and Pope, 1948).
_Polyphaga saussurei_, U.S.S.R. (Zmeev _in_ Pavlovskii, 1948).
* =Salmonella typhosa= (Zopf) White
_Natural vectors._--_Blatta orientalis_, Italy (Antonelli, 1930, 1943).
_Experimental vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al.,
1949).
_Blatta orientalis_, Italy (Spinelli and Reitano, 1932); U.S.A.
(McBurney and Davis, 1930); U.S.S.R. (Rozengolts and Ȋȗdina _in_
Pavlovskii, 1948).
_Blattella germanica_, U.S.A. (Janssen and Wedberg, 1952); Germany
(Jettmar, 1927); U.S.S.R. (Rozengolts and Ȋȗdina _in_ Pavlovskii, 1948).
_Periplaneta americana_, Gold Coast Colony (Macfie, 1922); Netherlands
(Akkerman, 1933); Formosa (Morischita and Tsuchimochi, 1926); U.S.A.
(Olson _in_ Roth and Willis, 1957a).
_Periplaneta australasiae_, Formosa (Morischita and Tsuchimochi, 1926).
_Polyphaga saussurei_, U.S.S.R. (Zmeev _in_ Pavlovskii, 1948).
Cockroaches [presumably _Blatta orientalis_, _Blattella germanica_,
and/or _Periplaneta americana_], Egypt (El-Kholy and Gohar, 1945).
=Serratia marcescens= Bizio
_Synonymy._--_Bacillus prodigiosus_, _Bacterium prodigiosum_.
_Habitat._--Water, soil, milk, foods, and various insects.
_Natural hosts._--_Blatta orientalis_, Poland (Nicewicz et al., 1946):
From intestinal tract. Italy (Spinelli and Reitano, 1932).
_Blattella germanica_, Canada (Heimpel and West, 1959).
_Diploptera punctata_, _Nauphoeta cinerea_, _Neostylopyga rhombifolia_,
_Panchlora nivea_, _Pycnoscelus surinamensis_, and _Supella
supellectilium_, U.S.A. (Roth and Willis, unpublished data, 1958): The
organism was isolated and identified by Dr. Hillel Levinson,
Quartermaster bacteriologist, from dead specimens found in our
laboratory colonies which showed the red coloration characteristic of
insects that have died with infections of _S. marcescens_ (pl. 16, A,
B).
_Leucophaea maderae_, U.S.A. (Levinson, personal communication, 1958):
The organism was isolated from the hemolymph of living insects while
attempting to determine the cause of unexplained mortality in our
laboratory colony of this insect.
_Leucophaea maderae_ or _Periplaneta americana_, Philippine Islands
(Barber, 1912): From hemolymph.
_Periplaneta americana_, U.S.A. (Gier, 1947; Steinhaus, 1959).
_Periplaneta australasiae_ and _Periplaneta brunnea_ (Roth and Willis,
unpublished data [1954]): In laboratory colonies. Isolated from
suspensions of ground insects. In 1954 we received a culture of
_Periplaneta brunnea_ from the Department of Public Health, University
of Minnesota. These insects began to die off rapidly and the normally
lightly pigmented parts of the body became red. Dr. Hillel Levinson,
Quartermaster bacteriologist, cultured _Serratia marcescens_ from
several moribund individuals. The Department of Public Health of
Minnesota had at times in the past cultured _S. marcescens_ but had
discarded the cultures and was unaware that it might be surviving in the
cockroach colonies (Richards, personal communication, 1954).
_Periplaneta_ sp., U.S.A. (Olson _in_ Roth and Willis, 1957a): Isolated
from an undetermined species of _Periplaneta_, received in a shipment
from the South, a strain of _S. marcescens_ which was toxic to mice when
administered intraperitoneally.
_Experimental hosts._--_Blaberus craniifer_, U.S.A. (Wedberg et al.,
1949): When fed in small numbers, _S. marcescens_ increased to such an
extent that the insect's extremities and upper halves of their bodies
turned deep red. The insects died after this color appeared and
practically pure cultures of _Serratia_ were recovered from the reddened
areas.
_Blatta orientalis_, Italy (Cao, 1898): Isolated from intestinal
contents. Passed unchanged through the gut.
_Blattella germanica_, Canada (Heimpel and West, 1959): Not normally
pathogenic per os; LD_{50} by injection, is approximately 38,000
bacteria per insect.
_Periplaneta americana_, U.S.A. (Gier, 1947): Organism toxic to the
cockroach when injected.
Cockroaches, U.S.A. (Longfellow, 1913): Isolated from legs and viscera
after feeding experiments.
* =Shigella alkalescens= (Andrewes) Weldin
_Natural vectors._--_Periplaneta americana_, U.S.A. (Bitter and
Williams, 1949, 1949a).
* =Shigella dysenteriae= (Shiga) Castellani and Chalmers
_Experimental vectors._--_Blatta orientalis_, Italy (Spinelli and
Reitano, 1932).
_Periplaneta americana_, Formosa (Morischita and Tsuchimochi, 1926).
_Polyphaga saussurei_, U.S.S.R. (Zmeev _in_ Pavlovskii, 1948).
* =Shigella paradysenteriae= (Collins) Weldin
_Natural vectors._--_Blatta lateralis_, Tadzhikistan (Zmeev, 1940).
_Experimental vectors._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922).
_Polyphaga saussurei_, U.S.S.R. (Zmeev _in_ Pavlovskii, 1948).
Cockroaches, Venezuela (Tejera, 1926).
Family PARVOBACTERIACEAE
=Bacteroides uncatus= Eggerth and Gagnon
_Habitat._--Probably intestinal canal of mammals; from human feces.
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946):
From intestinal canal.
* =Brucella abortus= (Schmidt and Weis) Meyer and Shaw
_Experimental vectors._--_Periplaneta americana_, U.S.A. (Ruhland and
Huddleson, 1941).
=Fusiformis lophomonadis= Grassé
_Habitat._--Surface of a flagellate (_Lophomonas striata_) which lives
in the intestine of cockroaches (Breed et al., 1948; Grassé 1926,
1926a).
* =Malleomyces mallei= (Zopf) Pribram
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
* =Pasteurella multocida= (Lehmann and Neumann) Rosenbusch and Merchant
_Experimental vectors._--_Blatta orientalis_, Germany (Küster, 1902,
1903); Italy (Cao, 1906).
* =Pasteurella pestis= (Lehmann and Neumann) Holland
_Natural vectors._--_Blatta orientalis_, Hongkong (Hunter, 1906).
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898); Germany
(Küster, 1903).
_Blattella germanica_, Germany (Jettmar, 1927).
_Leucophaea maderae_ and _Periplaneta americana_, Philippine Islands
(Barber, 1912).
Family BACTERIACEAE
[cross] =Bacterium alkaligenes= Nyberg
_Natural vectors._--_Periplaneta americana?_ ("_Blatella americana_"),
England (Shrewsbury and Barson, 1948): From intestinal tract.
[cross] =Bacterium delendae-muscae= Roubaud and Descazeaux
_Source._--Diseased fly larvae.
_Experimental infection._--Cockroach, France (Roubaud and Descazeaux,
1923): Organism pathogenic to cockroach when injected.
=Bacterium haemophosphoreum= Pfeiffer and Stammer
_Habitat._--Diseased larvae of _Mamestra oleracea_.
_Experimental infection._--_Blatta orientalis_ and _Blattella
germanica_, Germany (Pfeiffer and Stammer, 1931): Organism pathogenic,
when injected, to eight _B. orientalis_ and two _B. germanica_.
=Coccobacillus cajae= Picard and Blanc
_Experimental host._--_Blatta orientalis_, France (Picard and Blanc,
1913): The organism was pathogenic to _B. orientalis_ when injected.
Family BACILLACEAE
* =Bacillus anthracis= Cohen emend. Koch
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898, 1906);
Germany (Küster, 1903).
[cross] =Bacillus bütschlii= Schaudinn
_Habitat._--_Blatta orientalis_, Germany (Schaudinn, 1902): Isolated
from intestinal tract. Three percent of the cockroaches from Berlin
bakeries were infected.
=Bacillus cereus= Frankland and Frankland
_Synonymy._--_Bacillus albolactis._
_Habitat._--Soil, dust, milk, plants.
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949):
From feces.
_Periplaneta americana_, U.S.A. (Hatcher, 1939): In feces.
_Experimental host._--_Periplaneta americana_, U.S.A. (Babers, 1938):
The cockroaches died within 96 hours after being injected with 10^{-3}
ml. of a 24-hour broth culture.
=Bacillus circulans= Jordan
_Habitat._--Soil, water, dust.
_Natural vectors._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952): From intestine and feces.
[cross] =Bacillus flacheriae= (Hoffman)
_Source._--Diseased nun moth larvae.
_Experimental infection._--_Blatta orientalis_, Europe (Filatoff, 1904):
The organism was not pathogenic when fed to the cockroach, but killed
the insects when injected into the body cavity; after the insects died
Filatoff reisolated this pathogen together with another bacillus from
the cadavers. He succeeded in culturing the new microorganism and found
it to be pathogenic when injected into, but not when fed to, the
cockroaches. The diseased insects became sluggish, failed to eat or
drink, turned over on their backs, their extremities became totally
paralyzed, and they finally died.
=Bacillus megaterium= De Bary
_Habitat._--Soil, water, decomposing materials.
_Natural vectors._--_Periplaneta americana?_ ("_Blatella americana_"),
England (Shrewsbury and Barson, 1948): From intestinal tract.
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898):
Organism recovered, apparently unchanged, from intestinal contents.
[cross] =Bacillus monachae= (von Tubeuf) Eckstein
_Synonymy._--_Bacterium monache._
_Source._--Diseased larvae of nun moth, _Lymantria monacha_.
_Experimental infection._--_Blatta orientalis_, Europe (Filatoff, 1904):
Organism pathogenic to the cockroach when injected but not when fed.
[cross] =Bacillus periplanetae= Tichomiroff
_Habitat._--_Blatta orientalis_, U.S.S.R.? (Tichomiroff, 1870[?], _in_
Filatoff, 1904): The infected insects suffered from a diarrhea and the
liquid feces were yellow-brown.
[cross] =Bacillus stellatus= Hollande
_Natural infection._--_Blatta orientalis_, France (Hollande, 1934):
Organism observed regularly in the intestine (especially rectum).
Extensive description given.
[cross] =Bacillus radiciformis=
_Experimental vectors._--_Blatta orientalis_, Italy (Cao 1898): Organism
recovered, apparently unchanged, from intestinal contents.
* =Bacillus subtilis= Cohn emend. Prazmowski
_Natural vectors._--_Blaberus craniifer_, U.S.A. (Wedberg et al.,
1949).
_Blatta orientalis_, Italy (Cao, 1898, 1906; Spinelli and Reitano,
1932); France (Sartory and Clerc, 1908); Poland (Nicewicz et al., 1946).
_Cryptocercus punctulatus_, U.S.A. (Hatcher, 1939).
_Periplaneta americana?_ ("_Blatella americana_"), England (Shrewsbury
and Barson, 1948).
Cockroaches, U.S.A. (Longfellow, 1913).
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898, 1906).
"=Bacillus subtilis= group"
_Natural infection._--_Blatta orientalis_, Italy (Ronzoni, 1949):
Isolated from oöthecae.
[cross] =Bacillus tritus= Batchelor
_Habitat._--Isolated from feces (man?).
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946):
Isolated from intestinal tract.
* =Clostridium feseri= Trevisan
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
=Clostridium lentoputrescens= Hartsell and Rettger
_Habitat._--Soil, intestinal tract of man.
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946):
Isolated from intestinal tract.
* =Clostridium novyi= (Migula) Bergey et al. or
* =Clostridium sporogenes= (Metchnikoff) Bergey et al.
_Natural and experimental vectors._--_Blatta orientalis_, Italy (Cao,
1898).
* =Clostridium perfringens= (Veillon and Zuber) Holland
_Natural vectors._--Cockroaches [presumably _Blatta orientalis_,
_Blattella germanica_, and/or _Periplaneta americana_], Egypt (El-Kholy
and Gohar, 1945).
* =Clostridium tetani= (Flügge) Holland
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
* =Clostridium= spp.
_Natural vectors._--_Periplaneta americana?_ ("_Blatella americana_"),
England (Shrewsbury and Barson, 1948).
Order ACTINOMYCETALES
Family MYCOBACTERIACEAE
* =Mycobacterium avium= Chester
_Experimental vectors._--_Blatta orientalis_, U.S.S.R. (Ekzempliarskaia
_in_ Pavlovskii, 1948).
=Mycobacterium friedmannii= Holland
_Habitat._--Parasitic in turtles and possibly sparingly distributed in
soils.
_Natural vectors._--_Periplaneta americana_, U.S.A., Texas (Micks, in
Roth and Willis, 1957a): Organism isolated from batches of intestinal
tracts of cockroaches collected at random.
* =Mycobacterium lacticola= Lehmann and Neumann?
_Natural vectors._--_Periplaneta americana_, U.S.A. (Leibovitz, 1951).
* =Mycobacterium leprae= (Armauer-Hansen) Lehmann and Neumann
_Natural vectors._--_Blattella germanica_, Southern Rhodesia and Kenya
(Moiser, 1945, 1946, 1946a; Anonymous, 1946).
_Periplaneta americana_ and _Periplaneta australasiae_, Formosa
(Arizumi, 1934, 1934a).
Cockroaches, Venezuela (Tejera, 1926); Belgian Congo (Radna, 1939).
_Experimental vectors._--_Blatta orientalis_, Europe (Paldrock _in_
Klingmüller, 1930); Nyasaland (Lamborn, 1940).
_Blattella germanica_, Europe (Paldrock _in_ Klingmüller, 1930);
Southern Rhodesia and Kenya (Moiser, 1945, 1946, 1946a, 1947; Anonymous,
1946).
_Nauphoeta cinerea_, Nyasaland (Lamborn, 1940).
_Periplaneta americana_, Gold Coast Colony (Macfie, 1922); Formosa
(Arizumi, 1934, 1934a).
_Periplaneta australasiae_, Formosa (Arizumi, 1934, 1934a).
Cockroaches, Belgian Congo (Radna, 1939); Venezuela (Tejera, 1926).
* =Mycobacterium lepraemurium= Marchoux and Sorel
_Experimental vectors._--Cockroaches, Belgian Congo (Radna, 1939).
* =Mycobacterium phlei= Lehmann and Neumann
_Natural vectors._--_Periplaneta americana_, U.S.A. (Leibovitz, 1951;
Micks _in_ Roth and Willis, 1957a).
* =Mycobacterium piscium= Bergey et al.
_Natural vectors._--_Periplaneta americana_, U.S.A. (Leibovitz, 1951).
_Experimental vectors._--_Blatta orientalis_, U.S.S.R. (Ekzempliarskaia
_in_ Pavlovskii, 1948).
* =Mycobacterium tuberculosis= (Schroeter) Lehmann and Neumann
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898); Germany
(Küster, 1903); U.S.S.R. (Ekzempliarskaia _in_ Pavlovskii, 1948).
_Blattella germanica_, on shipboard (Morrell, 1911).
_Periplaneta americana_, Gold Coast Colony (Macfie, 1922).
Cockroaches, Venezuela (Tejera, 1926); U.S.A. (Read, 1933).
* =Mycobacterium= spp.
_Natural vectors._--_Periplaneta americana_, U.S.A. (Leibovitz, 1951;
Micks _in_ Roth and Willis, 1957a).
Family ACTINOMYCETACEAE
* =Nocardia= sp.?
_Natural vectors._--_Periplaneta americana_, U.S.A. (Leibovitz, 1951).
Family STREPTOMYCETACEAE
=Streptomyces leidynematis= Hoffman
_Habitat._--Surface of the nematodes _Hammerschmidtiella diesingi_ and
_Leidynema appendiculata_ in _Periplaneta americana_, U.S.A. (Hoffman,
1952, 1953): Eighteen percent of 192 nematodes found in 52 adult
cockroaches were infected with the bacterium.
Order CARYOPHANALES
Family ARTHROMITACEAE
=Arthromitus intestinalis= (Valentin) Peshkoff
_Synonymy._--_Hygrocrocis intestinalis._
_Habitat._--_Blatta orientalis_, Europe (Valentin, 1836; Robin, 1847,
1853; Peshkoff, 1940): Isolated from intestinal tract. The organism
appears as fragments in fecal masses or as fibers adhering to the mucous
membrane of the large intestine (Robin, 1853).
Cockroach, France? (Chatton and Pérard, 1913).
Order SPIROCHAETALES
Family SPIROCHAETACEAE
[cross] =Spirochaeta blattae= Tejera
_Habitat._--_Blaberus atropos_, Venezuela (Tejera, 1926): Isolated from
intestinal tract.
* [cross] =Spirochaeta periplanetae= Laveran and Franchini
_Habitat._--_Blatta orientalis_, France (Laveran and Franchini, 1920a).
Cockroaches, Venezuela (Tejera, 1926): Tejera reported finding
"_Spirochaeta blatarum_ Laveran et Franchini" which may have been a
lapsus.
Family TREPONEMATACEAE
[cross] =Treponema parvum= Dobell
_Habitat._--_Blatta orientalis_, England (Dobell, 1912); U.S.S.R.?
Zasukhin (1930): From intestinal tract.
[cross] =Treponema stylopygae= Dobell
_Synonymy._--_Spirochaeta stylopygae_ Zuelzer.
_Habitat._--_Blatta orientalis_, England (Dobell, 1912); U.S.S.R.?
Zasukhin (1930): From intestinal tract.
=Unidentified spirochaetes=
_Habitat._--_Blatta orientalis_, U.S.S.R. (Yakimov and Miller, 1922):
Spirochaetes and spirilla were found in the intestines of 70 percent of
124 specimens collected in Petrograd.
_Periplaneta americana_, Gold Coast Colony (Macfie, 1922).
ADDITIONAL BACTERIA WHOSE TAXONOMIC POSITION IS UNKNOWN
* "=B. aerobio del pseudoedema maligno=" of Cao
_Natural vectors._--_Blatta orientalis_, Italy (Cao, 1906).
=B. alcaligenes beckeri=
_Natural vectors._--_Blatta orientalis_, Poland (Nicewicz et al., 1946):
Isolated from intestinal tract.
* "=B. del pseudoedema maligno=" of Cao
_Natural vectors._--_Blatta orientalis_, Italy (Cao, 1906).
* "=Bacillo proteisimile=" of Cao
_Natural and experimental vectors._--_Blatta orientalis_, Italy (Cao,
1898, 1906).
* "=Bacillo del barbone dei bufali=" of Cao
_Experimental vectors._--_Blatta orientalis_, Italy (Cao, 1898).
* "=Bacillo similcarbonchio=" of Cao
_Natural and experimental vectors._--_Blatta orientalis_, Italy (Cao,
1898, 1906).
* "=Bacillo similtifo=" or "=Bacillo tifosimile=" of Cao
_Natural and experimental vectors._--_Blatta orientalis_, Italy (Cao,
1898, 1906).
"=Bacillus="
_Natural infection._--_Blatta orientalis_, Germany (Heinecke, 1956):
Disease organism found in the hemolymph of infected cockroaches. It can
be spread by mouth and through wound infection. The animals died with
symptoms of paralysis in 85-90 days. The organism has been isolated and
is in the culture collection of the Institute for Microbiology and
Experimental Therapy, Jena, under the numbers SG 896, Strain A; SG 897,
Strain B; SG 898, Strain C.
_Experimental infection._--_Blattella germanica_, Germany (Heinecke,
1956): Infected animals died in 26-30 days.
_Periplaneta americana_ was unaffected even by heavy inoculations of the
pathogen.
"=Bacterium="
_Source._--(I) Diseased silkworm larvae. (II) Diseased _Ocneria dispar_
larvae and blood of _Blatta orientalis_.
_Experimental infection._--(I)(II) _Blatta orientalis_, Europe
(Filatoff, 1904): Organism pathogenic when injected, nonpathogenic when
ingested.
(I) Cockroach, U.S.A. (Glaser, 1925): Organism pathogenic to cockroach
when injected.
"=Coccobacillus="
_Natural infection._--_Blatta orientalis_, France (Hollande, 1934):
Organism described morphologically.
"=Colon bacilli="
_Natural vectors._--Cockroaches [presumably _Blatta orientalis_,
_Blattella germanica_, and/or _Periplaneta americana_], Egypt (El-Kholy
and Gohar, 1945): From the outer surface, intestinal tract, and
suspensions of macerated insects.
"=Diplococci="
_Natural vectors._--_Blatta orientalis_, Germany (Jettmar, 1935): From
intestinal tract.
_Blattella germanica_, Germany (Jettmar, 1935): From outer surface of
body.
"=Diphtheroid I and II="
_Source._--_Periplaneta americana._
_Natural and experimental infections._--_Periplaneta americana_, U.S.A.
(Gier, 1947): Pathogenicity to the cockroach variable when organism
injected.
"=Gram positive rods="
_Source._--Feces of _Blattella germanica_.
_Experimental vector._--_Blattella germanica_, Germany
(Vollbrechtshausen, 1953): Nonpathogenic to the insect when injected
into the mouth or anus.
"=Silkworm disease bacillus="
Cockroaches that were inoculated with living cultures succumbed in a few
days (Glaser, 1925).
[cross] =Spirillochaeta blattae= Hollande
_Habitat._--_Blatta orientalis_, France (Hollande, 1934; Hollande and
Hollande, 1946): Organism found in hind intestine. This spirillum was
stated to be related in external morphology to _Spirillum
periplaneticum_ Kunstler and Gineste, but it was believed that _S.
blattae_ should be in the Spirochaetaceae rather than the Spirillaceae.
"=Spirochaetoid bacteria="
_Habitat._--_Blatta orientalis_, France (Hollande, 1934): Two kinds
described but not named.
[cross] =Tetragenous= sp.
_Natural and experimental infections._--_Periplaneta americana_, U.S.A.
(Gier, 1947): Pathogenicity to the cockroach variable when organism
injected.
VIII. FUNGI AND YEASTS
By far the greatest number of fungi known to be associated with
cockroaches belong to the Laboulbeniaceae, genus _Herpomyces_, the
species of which are restricted to parasitizing cockroaches (Thaxter,
1908). Most species are hyaline, small and inconspicuous (Thaxter, 1931)
and are usually, but not exclusively, found on the insects' antennae.
Species of _Herpomyces_ are highly, but not completely, host specific
(Richards and Smith, 1954). While attached to the host, these fungi
appear like minute dark-colored, yellow, or white (e.g., _H. arietinus_)
bristles or bushy hairs (pl. 27, A).
[Illustration: FIG. 1.--Diagram illustrating the relationship between a
mature plant of _Herpomyces stylopygae_ and the integument of _Blatta
orientalis_. (Reproduced from Richards and Smith [1956], through the
courtesy of Dr. A. G. Richards.)]
Richards and Smith (1955, 1955a) have studied the life history of
_Herpomyces stylopygae_ on the oriental cockroach. The plants grow only
on living cockroaches, and the infection is disseminated by contact. The
mature plants are found mostly on the antennae (pl. 27, B), either on
setae or on hard or soft cuticle. Spores are ejected from perithecia
singly or in groups of 2 to 4 spores, although groups as large as 12
spores have been found. The presence of single, paired, or multiple
spore groups on the surface of the host was correlated with the presence
of single, paired, or multiple plants on infected cockroaches.
Development from spore to mature perithecia takes about two weeks. The
plant obtains nutriment from the host by means of a tubular haustorium
that extends through the cockroach's cuticle and expands into a large
bulb in the underlying epidermal cells (fig. 1). Infections on nymphs
are lost when the nymph moults, but infections on adults persist
throughout life. However, nymphs which have lost the fungus upon
moulting are readily reinfected. Collart's (1947) statement that nymphs
are never infected with _Herpomyces_ is not true.
Richards and Smith (1956) concluded that there is no evidence of
pathogenicity in _Herpomyces_ infections because heavily infected
cockroaches appear fully active in laboratory colonies; they can run at
the same speed as uninfected cockroaches; they reproduce normally and do
not appear to die prematurely. These workers stated that the infections
cause a dermatitis which is neither pathogenic nor debilitant. So far as
we know there are no comparative data on longevity and reproductive
performance of fungus-infected versus normal cockroaches. However, Gunn
and Cosway (1938) have shown that the presence of these fungi
(identified as _Stigmatomyces_ sp.; see p. 138) on the antennae seemed
to interfere with the humidity reactions of _Blatta orientalis_.
Although Richards and Smith (1956) admit that humidity receptors and
other sense organs on the antennae may be destroyed by the fungus, they
state that "insects possess such a large number of sensilla that the
result may well be more distressing to the sensory physiologist than to
the insect." Yet it seems to us that the loss of sense organs from
fungal infection and concomitant shortening of the antennae (pl. 27, A)
might be considerably more of a handicap to free-living cockroaches than
those in laboratory colonies.
Bode (1936) studied the flora of _Periplaneta americana_ and cultured
Aspergillaceae and Mucorinae from the insect's body surface and
intestinal contents; he also found nonsporulating yeasts in _P.
americana_. To prevent fungal growth on oöthecae of _P. americana_,
Griffiths and Tauber (1942a) autoclaved their rearing containers and
dipped the oöthecae in 70-percent alcohol for 10 seconds.
Mercier (1906) isolated and cultured a pathogenic yeastlike parasite
which had invaded the fat body and blood of _Blatta orientalis_. The
abdomens of the infected insects became swollen, distended, and soft.
McShan (unpublished MS., 1953) consistently isolated Saccharomycetes
from the feces of _Periplaneta americana_.
FUNGI ASSOCIATED WITH COCKROACHES
The use of the asterisk (*) is explained in footnote 3, page 4.
Phylum THALLOPHYTA
Class FUNGI IMPERFECTI
Order MONILIALES
Family PSEUDOSACCHAROMYCETACEAE
=Candida zeylanoides= (Castellani) Langeron and Guerra
_Natural host._--Oötheca of _Blatta orientalis_, Italy (Ronzoni, 1949).
=Torulopsis= sp.
_Natural host._--Oötheca of _Blatta orientalis_, Italy (Ronzoni, 1949).
Family MONILIACEAE
=Spicaria prasina= (Maublanc) Sawada
_Natural host._--_Ischnoptera rufa rufa_, Puerto Rico (Wolcott, 1950): A
dead specimen of this cockroach was found stuck to a leaf and covered
with this fungus.
=Aspergillus flavus= Link
_Natural hosts._--Oöthecae of _Blattella germanica_ and _Eurycotis
floridana_, U.S.A., Pennsylvania (Roth and Willis, unpublished data,
1952): On outer surface. Determination by Miss Mary Downing.
Oöthecae of _Periplaneta americana_, U.S.A., Pennsylvania (Roth and
Willis, unpublished data, 1952): Inside oöthecae. Determination by Miss
Mary Downing.
* =Aspergillus fumigatus= Fresenius
_Natural vector._--_Blatta orientalis_, France (Sartory and Clerc,
1908): From intestine.
* =Aspergillus niger= van Tieghem
_Natural vector._--_Periplaneta americana_, U.S.A., Texas (McShan in
Roth and Willis, 1957a): From feces.
_Experimental vector._--_Blatta orientalis_, Italy (Cao, 1898): Organism
passed unchanged through the gut of the insects.
=Aspergillus sydowi= (Bainier and Sartory) Thom and Church
_Natural host._--Oötheca of _Eurycotis floridana_, U.S.A., Pennsylvania
(Roth and Willis, unpublished data, 1952): On outer surface.
Determination by Miss Mary Downing.
=Aspergillus tamarii= Kita
_Natural host_.--Oöthecae of _Blattella germanica_, U.S.A., Pennsylvania
(Roth and Willis, unpublished data, 1952): On exterior surface.
Determination by Miss Mary Downing.
=Aspergillus= sp.?
_Natural and experimental vector._--_Blattella germanica_, on shipboard
(Morrell, 1911): Isolated from feces. Experimentally Morrell also showed
that the spores of the fungus could be recovered from feces of
cockroaches that had fed on them.
=Aspergillus= sp.
_Natural vector._--_Periplaneta americana_, England (Bunting, 1956): The
fungus was isolated mostly from imperfectly excreted feces.
=Beauveria bassiana= (Balsamo) Vuillemin
_Experimental host._--_Blattella germanica_ and _Periplaneta americana_,
U.S.A. (Dresner, 1949, 1950): The nymphs of American cockroaches became
infected when they (1) were injected with a 1-percent suspension of
spores, (2) ate rat pellets sprayed with the spore suspension, or (3)
were dusted with the fungus spores. The symptoms of the fungus infection
were paralysis followed by death; some of the infected insects
liquefied, others dried up after the appearance of a subcuticular
blackening.
=Cephalosporium= sp.
_Natural vector._--_Periplaneta americana_, U.S.A., Texas (McShan,
unpublished MS., 1953): From feces of cockroaches collected in the
basement of a grain elevator at the docks in Galveston.
* =Geotrichum candidum= Link
_Experimental vector._--_Blatta orientalis_, Italy (Cao, 1898): Organism
retained its pathogenicity after passing through the insect's gut.
=Penicillium= sp.
_Natural vector._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949):
From feces.
_Periplaneta americana_, England (Bunting, 1956): Mostly from
imperfectly excreted feces.
=Metarrhizium anisopliae= (Metschnikoff) Sorokin
_Natural hosts._--_Blattidae_, Seymour (1929); Charles (1941).
_Panesthia australis_, U.S.A., Massachusetts (Roth and Willis,
unpublished data, 1957): Growing on adult specimens that were found dead
in a laboratory colony. Determination by Miss Dorothy Fennell.
_Periplaneta americana_, England (Bunting, 1956): Growing on genitalia
of females where it prevented oöthecal formation.
Cockroach, Puerto Rico (Johnston, 1915): From a "small roach" in the
pathological collection at Rio Piedras (no data).
Family DEMATIACEAE
=Memnoniella echinata= (Rivolta) Galloway
_Natural host._--Oötheca of _Blattella germanica_, U.S.A., Pennsylvania
(Roth and Willis, unpublished data, 1952): On material that had oozed
from a damaged oötheca. Determination by Miss Mary Downing.
=Torula acidophila= Owen and Mobley
_Natural host._--_Periplaneta americana_, U.S.A. (Owen and Mobley,
1948): The digestive tract of this cockroach is the normal habitat of
this yeast which was transmitted to sirup by the insects. The yeast
superimposed a foreign taste, suggestive of malic acid, upon the
original flavor of the sirup.
=Torula gropengiesseri= Lodder
_Natural host._--_Blatta orientalis_, Germany (Gropengiesser, 1925;
Lodder, 1934): Isolated from fat body and oöthecae. Gier (1947) is of
the opinion that the so-called yeasts that supposedly may displace the
bacteroids in the fat body (Mercier, 1907b; Gropengiesser, 1925) may
actually represent poorly fixed and insufficiently stained bacteroids.
=Torula rosea= Preuss
_Experimental host._--_Blaberus craniifer_, U.S.A. (Wedberg et al.,
1949): Upon repeated feeding of massive doses of this yeast to the
cockroach, these workers were able to isolate the organism from the
feces up to six days thereafter. There was no evidence that _T. rosea_
was pathogenic for _B. craniifer_.
Class PHYCOMYCETES
Order MUCORALES
Family MUCORACEAE
=Mucor guilliermondii= Nadson and Filippov
_Natural host._--_Periplaneta americana_, U.S.S.R. (Nadson and Filippov,
1925; Filippov, 1926): Isolated and cultured from intestine.
=Mucor= sp.
_Natural host._--Oötheca of _Periplaneta americana_, U.S.A.,
Pennsylvania (Roth and Willis, unpublished data, 1952): Inside oötheca.
Determination by Miss Mary Downing.
_Pycnoscelus surinamensis_, Germany (Bode, 1936): Isolated from fat body
which it had stained red.
=Rhizopus nigricans= Ehrenberg
_Natural vector._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949):
From feces.
=Rhizopus= sp.
_Natural vector._--_Periplaneta americana_, U.S.A., Texas (McShan,
unpublished MS., 1953): From feces.
=Syncephalastrum= sp.
_Natural vector._--_Periplaneta americana_, U.S.A., Texas (McShan,
unpublished MS., 1953): From feces.
Order ENTOMOPHTHORALES
Family BLASTOCYSTIDACEAE
=Blastocystis hominis= Brumpt
_Natural vector._--_Blatta orientalis_, U.S.S.R. (Zasukhin, 1930): In
hind gut in 40 percent of over 3,000 cockroaches.
=Blastocystis= sp.
_Natural vectors._--_Blatta orientalis_, U.S.S.R. (Yakimov and Miller,
1922): Found in the intestinal contents of 29 percent of 124 _B.
orientalis_.
Cockroaches, Venezuela (Tejera, 1926).
* * * * *
The placement of the following fungus is problematic.
=Coccidioides periplanetae= Avrech
_Natural host._--_Blatta orientalis_, Germany (Avrech, 1931): Found in
cells lining the lumen of midgut and caeca. The whole upper part of the
epithelium was filled with sporangia and spores.
Class ASCOMYCETES
Order ENDOMYCETALES
Family SACCHAROMYCETACEAE
=Saccharomyces cerevisiae= Hansen
_Natural vector._--_Blaberus craniifer_, U.S.A. (Wedberg et al., 1949):
In feces.
=Saccharomyces= sp.
_Natural vector._--_Blattella germanica_, U.S.A. (Janssen and Wedberg,
1952): Found consistently in alimentary tract of _B. germanica_ fed
sucrose solutions.
Order HYPOCREALES
Family HYPOCREACEAE
=Cordyceps amazonica= Hennings
_Natural host._--Cockroaches, British Honduras (Mains, 1940).
=Cordyceps blattae= Petch
_Natural host._--_Blattella germanica_, Ceylon (Petch, 1924): Collected
at Hakgala twice. A slight covering of brown mycelium overran the insect
and fastened it to the underside of a living leaf.
Order LABOULBENIALES
Family LABOULBENIACEAE
=Herpomyces amazonicus= Thaxter
_Natural host._--_Nyctibora obscura_, Brazil, Natal (Thaxter, 1931): On
antennae.
=Herpomyces anaplectae= Thaxter
_Natural hosts._--_Anaplecta_ sp., Venezuela, Caracas (Thaxter, 1905,
1908); Trinidad (Thaxter, 1931): On antennae.
Cockroach, Sumatra (Thaxter, 1931).
=Herpomyces appendiculatus= Thaxter
_Natural host._--_Platyzosteria scabra_, Australia, N.S.W. (Thaxter,
1931): On antennae.
=Herpomyces arietinus= Thaxter
_Natural hosts._--_Ischnoptera_ sp., U.S.A., Georgia (Thaxter, 1908).
_Parcoblatta uhleriana_, U.S.A., Massachusetts (Roth, unpublished data,
1957): The nymphs were in a culture of _Parcoblatta virginica_ which was
infected with this fungus; it is possible that these _P. uhleriana_
became infected by contact with _P. virginica_. Fungus identified by Dr.
R. K. Benjamin.
_Parcoblatta virginica_, U.S.A., Massachusetts (Roth, unpublished data,
1957): Fungus determined by Dr. R. K. Benjamin. Fungus found on
antennae, palpi, legs, body surface (pl. 27, A).
_Parcoblatta_ sp., U.S.A., Kentucky, Massachusetts (Thaxter, 1902,
1908): On antennae.
It is likely that Thaxter's host records (certainly those assigned to
_Temnopteryx_ and possibly those assigned to _Ischnoptera_) were species
of _Parcoblatta_. Hebard (1917) has shown that all the species referred
to _Ischnoptera_ in the United States, except _I. deropeltiformis_, now
belong in the genus _Parcoblatta_. All species originally referred to
the genus _Temnopteryx_ in the United States are now synonymized with
species of _Parcoblatta_.
=Herpomyces chaetophilus= Thaxter
_Natural hosts._--_Periplaneta americana_, Brazil (Thaxter, 1931).
_Periplaneta_ sp., Zanzibar and Mauritius (Thaxter, 1902, 1908): On
spines of legs, antennae, and cerci.
=Herpomyces chilensis= Thaxter
_Natural host._--Cockroach, Chile (Thaxter, 1918): On antennae.
=Herpomyces diplopterae= Thaxter
_Natural hosts._--_Diploptera punctata_, Ascension Island (Thaxter,
1902, 1908): On antennae. This species also was infected experimentally
(Richards and Smith, 1954).
Cockroach, Fiji (Thaxter, 1931).
=Herpomyces ectobiae= Thaxter
_Natural hosts._--_Blattella germanica_, U.S.A., Massachusetts (Thaxter,
1902, 1908); Burma, Tenasserim (Spegazzini, 1915); Argentina, Buenos
Aires (Spegazzini, 1917): On antennae. U.S.A., Minnesota (Richards and
Smith, 1955): Scattered over entire body, wings. France? (Picard, 1913):
On tibial spines. Chile and Philippine Islands (Thaxter, 1931).
"_Ectobia_" spp., Zanzibar and Saint Kitts, B.W.I. (Thaxter, 1902,
1908): Possibly on species that are now in the genus _Blattella_ rather
than in the genus _Ectobius_ as it is known today, because Thaxter also
used the synonym _Ectobia germanica_ for the German cockroach,
_Blattella germanica_.
_Experimental hosts._--_Blattella germanica_ and _Blattella vaga_,
U.S.A. (Richards and Smith, 1954).
=Herpomyces forficularis= Thaxter
_Natural hosts._--Cockroaches, Mauritius? and Fiji (Thaxter, 1902, 1908,
1931): On antennae.
=Herpomyces gracilis= Thaxter
_Natural host._--_Blattella humbertiana_, Philippine Islands, Luzon
(Thaxter, 1931): On antennae.
=Herpomyces grenadinus= Thaxter
_Natural host._--Cockroach, Grenada, B.W.I. (Thaxter, 1931): On antennae
of a "brown wingless blattid."
=Herpomyces leurolestis= Thaxter
_Natural host._--_Leurolestes pallidus_, British Guiana and Trinidad
(Thaxter, 1931): On antennae.
=Herpomyces lobopterae= Thaxter
_Natural host._--_Loboptera_ sp., Argentina (Thaxter, 1931): On
antennae.
=Herpomyces macropus= Spegazzini
_Natural host._--_Blaberus_ sp.?, Argentina (Spegazzini, 1917).
Cockroaches, Peru, Puerto Rico, Ecuador, and Haiti (Spegazzini, 1915,
1917): Material previously assigned by Spegazzini (1915) to _H.
paranensis_ was also placed by him in this new species. However, Thaxter
(1931) believed that _H. macropus_ may be synonymous with _H.
paranensis_, but he provisionally retained _H. macropus_ because he had
not seen Spegazzini's material.
=Herpomyces nyctoborae= Thaxter
_Natural hosts._--_Nyctibora tomentosa_, U.S.A., Texas (Thaxter, 1905,
1908): On antennae. This cockroach is not established in Texas, and the
specimen may have been misidentified (Gurney, personal communication,
1958).
_Nyctibora_ sp., Argentina (Spegazzini, 1917): On antennae.
=Herpomyces panchlorae= Thaxter
_Natural hosts._--_Panchlora nivea_, Trinidad (Thaxter, 1931): On
antennae.
=Herpomyces panesthiae= Thaxter
_Natural host._--_Panesthia lobipennis_, Ceylon (Thaxter, 1915): On
antennae.
=Herpomyces paranensis= Thaxter
_Natural hosts._--_Blaberus_ sp.? Brazil (Thaxter, 1902, 1908): On
antennae.
_Blaberus_ sp., Brazil and Argentina (Spegazzini, 1917): On antennae.
Cockroaches, Trinidad and Argentina (Thaxter, 1931).
=Herpomyces periplanetae= Thaxter
_Natural hosts._--_Blaberus_ sp.?, Argentina (Spegazzini, 1917).
_Blatta orientalis_, U.S.A., Massachusetts (Thaxter, 1902, 1908);
Locality? (Spegazzini, 1915); France? (Picard, 1913).
_Periplaneta americana_, Bermuda and U.S.A., Massachusetts (Thaxter,
1902, 1908); Plains of Biajar, Italian Somaliland, and Argentina
(Spegazzini, 1915, 1917).
_Periplaneta australasiae_, Bermuda (Thaxter, 1902, 1908).
_Periplaneta brunnea_, Brazil (Thaxter, 1931).
_Periplaneta_ sp., Mexico, West Indies, Panama, Brazil, Africa, South
Seas, and China (Thaxter, 1902, 1908).
Cockroaches, Belgium (Collart, 1947).
Additional locality records: Grenada, Trinidad, B.W.I., and Tangier
(Thaxter, 1931).
The fungus was found growing on spines, tegmina, integument, and
antennae, at times abundantly.
_Experimental hosts._--All the following data are from Richards and
Smith (1954):
_Blatta orientalis_: A few plants matured.
_Neostylopyga rhombifolia_: Some development but no mature plants.
_Periplaneta americana_: Fungus developed prolifically with a density
equal to that on original host.
_Periplaneta australasiae_: Some development but no mature plants.
_Periplaneta brunnea_: Fungus developed prolifically with a density
equal to that on original host.
=Herpomyces phyllodromiae= Thaxter
_Natural host._--"_Phyllodromia_" sp., Abyssinia (Thaxter, 1905, 1908):
On antennae.
=Herpomyces platyzosteriae= Thaxter
_Natural host._--"_Eurycotis floridana_," Mexico (Thaxter, 1905, 1908):
On antennal setae.
Since this cockroach is not found in Mexico (J. A. G. Rehn, personal
communication, 1957), _E. floridana_ is undoubtedly not the host for
this fungus. W. B. Brown (personal communication, 1957) searched the
cockroach collection at the Museum of Comparative Zoology but was unable
to find Thaxter's insect for reidentification.
=Herpomyces stylopygae= Spegazzini
_Natural hosts._--_Blatta orientalis_, Argentina (Spegazzini, 1917);
U.S.A. (Richards and Smith, 1955a).
_Experimental hosts._--_Neostylopyga rhombifolia_ (Richards and Smith,
1954): A few plants matured.
_Pycnoscelus surinamensis_ (Richards and Smith, 1954): Some development
but no mature plants.
The fungus (fig. 1) is found on antennae (pl. 27, B, C), palpi, cerci,
and femurs. Thaxter (1931) believed _H. stylopygae_ to be synonymous
with _H. periplanetae_. However, Richards and Smith (1954) concluded
that _H. stylopygae_ would not grow on _P. americana_ under their
laboratory conditions although _H. periplanetae_ would grow on _B.
orientalis_. This indicated a strain or species difference between the
two fungi. Gunn and Cosway (1938) reported a species of _Stigmatomyces_
on the antennae of _B. orientalis_; this fungus was probably _H.
stylopygae_ (Richards and Smith, 1956).
=Herpomyces supellae= (Thaxter)
_Natural host._--_Supella supellectilium_, Trinidad (Thaxter, 1931): On
antennal spines.
=Herpomyces tricuspidatus= Thaxter
_Natural hosts._--_Blaberus craniifer_, U.S.A., Key West (Richards and
Smith, 1955).
_Blaberus_ sp. and _Epilampra?_ sp., Panama (Thaxter, 1902, 1908).
_Epilampra_ sp., Saint Kitts, B.W.I., and Haiti (Thaxter, 1902, 1908).
_Leucophaea maderae_, Fernando Po (Spegazzini, 1915).
_Nauphoeta cinerea_, Brazil (Thaxter, 1931).
Cockroaches, China? (Thaxter, 1902); Philippine Islands, Mindanao
(Thaxter, 1931).
_Experimental hosts._--_Blaberus craniifer_, U.S.A. (Richards and Smith,
1955).
Infections on the antennae. Richards and Smith (1954) were unable to
secure experimental infections in _L. maderae_ with _H. tricuspidatus_.
Experiments with _N. cinerea_ showed some development but no mature
plants although identification of the growing fungus was uncertain
because of simultaneous exposure to _H. ectobiae_, _H. stylopygae_, and
_H. tricuspidatus_.
=Herpomyces zanzibarinus= Thaxter
_Natural hosts._--_Eurycotis manni_, Brazil (Thaxter, 1931): On
antennae.
_Gyna_ sp.?, Isle of Nias (Spegazzini, 1915): On antennae.
Cockroach, Zanzibar (Thaxter, 1902): On antennae.
=INCERTAE SEDIS=
According to Dr. R. K. Benjamin (personal communication, 1957) and Dr.
E. G. Simmons (personal communication, 1957), the phylogenetic position
of the following genus is uncertain.
=Amphoromorpha blattina= Thaxter
_Natural hosts._--Cockroaches, Grenada, B.W.I. (Thaxter, 1920): On the
axis of the antennae of a dark wingless and a pale winged blattid.
=Amphoromorpha= sp.
_Natural host._--Cockroach, Grenada, B.W.I. (Thaxter, 1920): On antennal
setae.
IX. HIGHER PLANTS
The significance of many observed associations between cockroaches and
the higher plants is still obscure. Undoubtedly many associations are
ecological, but lack of adequate supporting evidence makes this
conclusion somewhat tentative. The ecological aspects are covered in
Section III (p. 14). Other associations may be accidental (e.g., certain
unique observations that have never again been confirmed). In the
absence of contrary evidence, most associations are presumed to be
benign; exceptions to this conclusion are found among the cockroaches
that feed on living plants (p. 162) and those allegedly captured as prey
by the carnivorous pitcher plants (_Sarracenia_ and _Nepenthes_). In all
the records cited below the cockroaches were stated to have been on, in,
or feeding on the plant.
The plants are listed below by family according to the taxonomic
arrangement of Lawrence (1951). Botanical nomenclature follows Bailey
(1925), Fernald (1950), or Dr. R. A. Howard (personal communications,
1958, 1959). We take full responsibility for referring to appropriate
taxa certain plants that were reported by common name only in the cited
literature.
Division PTERIDOPHYTA
Family CYATHEACEAE
=Alsophila= sp.
_Associate._--_Pycnoscelus surinamensis_, Louisiana (Anonymous, 1893):
Feeding on heart of tree fern.
Family POLYPODIACEAE
=Asplenium nidus= Linnaeus
_Associate._--_Comptolampra liturata_, Malaya (Karny, 1924): Often found
between dry foliage of the beakers of this fern.
Division EMBRYOPHYTA SIPHONOGAMA
Family PINACEAE
=Pinus australis= Michaux
_Associates._--_Aglaopteryx gemma_ and _Parcoblatta lata_, Alabama
(Hebard, 1917): The former species was common under signs on longleaf
pines, and _P. lata_ was occasional.
_Parcoblatta divisa_, Georgia (Rehn and Hebard, 1916): Under signs.
=Pinus caribaea= Morelet
_Associates._--_Eurycotis floridana_, _Latiblattella rehni_, and
_Parcoblatta fulvescens_, Florida (Hebard, 1917): Many records under
signs on the tree trunks.
=Pinus clausa= Vasey
_Associate._--_Latiblattella rehni_, Florida (Hebard, 1917): Under sign
on tree.
=Pinus echinata= Mill.
_Associates._--_Parcoblatta divisa_, Virginia (Rehn and Hebard, 1916):
Under signs on shortleaf pine.
_Parcoblatta zebra_, Mississippi (Hebard, 1917): Under sign.
=Pinus sylvestris= Linnaeus
_Associate._--_Ectobius pallidus_, England (Milton, 1899; Burr, 1899b):
On Scotch fir.
=Pinus= spp.
_Associates._--_Plectoptera lacerna_ and _Plectoptera vermiculata_, Cuba
(Rehn and Hebard, 1927).
_Latiblattella rehni_, Florida (Rehn and Hebard, 1905): Under signs.
Cuba (Rehn and Hebard, 1927).
Family TAXODIACEAE
=Cryptomeria= sp.
_Associate._--_Diploptera punctata_, Hawaii (Pemberton and Williams,
1938; Zimmerman, 1948).
Family CUPRESSACEAE
=Cupressus macrocarpa= Hartweg
_Associate._--_Diploptera punctata_, Hawaii (Hebard, 1922): "The species
is common and injurious in the territory infesting particularly the
Monterey cypress trees ... and doing particular damage by gnawing away
the bark." Similar injury has been cited by Pemberton (1934), Fullaway
and Krauss (1945), and Zimmerman (1948).
=Juniperus= sp.
_Associate._--_Phyllodromica tartara nigrescens_, Southern Uzbekistan
(Bei-Bienko, 1950): Under bark.
Family PANDANACEAE
=Freycinetia= sp.
_Associate._--_Graptoblatta notulata_ and _Kuchinga remota_, Tahiti
(Hebard, 1933).
=Pandanus= sp.
_Associate._--_Hololeptoblatta_ sp., Seychelles (Scott, 1910, 1912).
Family GRAMINEAE
=Aristida pennata= Trin.
_Associate._--_Phyllodromica pygmaea_, U.S.S.R. (Bei-Bienko, 1950):
Found in the dense turf.
=Bamboo=
_Associate._--_Comptolampra liturata_, Malaya (Karny, 1925).
=Chloris gayana= Kunth
_Associate._--_Blattella vaga_, Texas (Riherd, 1953): This field
cockroach was rather abundant in clumps of Rhodes grass.
=Panicum purpurascens= Raddi
_Synonymy._--_Panicum barbinode_ [Hitchcock, 1936].
_Associate._--_Epilampra abdomen-nigrum_, Puerto Rico (Seín, 1923;
Wolcott, 1936): Abundant in "malojillo" meadow.
=Saccharum officinarum= Linnaeus
_Associates._--_Balta quadricaudata_, _Balta scripta_, _Balta
torresiana_, _Balta verticalis_, _Ellipsidion simulans_, and _Megamareta
verticalis_, Australia, Queensland (Hebard, 1943): All collected by J.
F. Illingworth on sugarcane.
_Blattella humbertiana_, _Ischnoptera schenklingi_, and _Pycnoscelus
surinamensis_, Formosa (Box, 1953).
_Cariblatta stenophrys_, Puerto Rico (Seín, 1923; Wolcott, 1936):
Between the leaves and under the leaf sheaths.
_Panchlora nivea_, Cuba (Rehn and Hebard, 1927): On the leaves.
_Pelmatosilpha coriacea_, Puerto Rico (Wolcott, 1936).
_Phoraspis_ spp., Brazil and Guiana (Doumerc in Blanchard, 1837).
_Plectoptera dorsalis_, _Plectoptora infulata_, and _Plectoptera
rhabdota_, Puerto Rico (Wolcott, 1950): Under the leaf sheaths.
_Symploce ruficollis_, Puerto Rico (Wolcott, 1950): Often found living
under the leaf sheaths.
Cockroaches, Philippine Islands (Uichanco _in_ Williams et al., 1931):
Between cane leaf sheaths.
=Setaria verticillata= (Linnaeus) Beauv.
_Synonymy._--_Chaetochloa verticillata_ (Linnaeus) [Howard, personal
communication, 1958].
_Associate._--_Diploptera punctata_, Hawaii (Severin, 1911): The
cockroach was caught on the barbed awns of this grass.
=Wild oats=
_Associate._--_Ischnoptera deropeltiformis_, Missouri (Rau, 1937).
=Zea mays= Linnaeus
_Associates._--_Cariblatta stenophrys_, Puerto Rico (Seín, 1923;
Wolcott, 1936).
_Ellipsidion bicolor_, Australia, Queensland (Hebard, 1943).
_Lophoblatta arawaka_, Trinidad (Princis and Kevan, 1955).
_Phoraspis_ sp., Brazil and Guiana (Doumerc _in_ Blanchard, 1837).
_Supella supellectilium_, New Caledonia (Cohic, 1956).
Family CYPERACEAE
=Cyperus= sp.
_Associate._--_Maretina uahuka_, Marquesas Islands, Uahuka (Hebard,
1933a).
Family PALMAE
=Acrocomia aculeata= (Jacq.) Lodd.
_Associate._--_Pycnoscelus surinamensis_, Trinidad (Princis and Kevan,
1955): On "gru-gru" fruits.
=Cocos nucifera= Linnaeus
_Associates._--_Aglaopteryx gemma_, Florida (Rehn and Hebard, 1912).
_Cariblatta lutea minima_, Florida, and _Cariblatta delicatula_, San
Domingo (Hebard, 1916a).
_Eurycotis floridana_, Florida (Rehn and Hebard, 1912; Hebard, 1917).
_Periplaneta australasiae_, Jamaica (Rehn and Hebard, 1927).
_Pycnoscelus surinamensis_, Florida (Rehn and Hebard, 1912; Hebard,
1917). Jamaica (Rehn and Hebard, 1927).
=Phoenix dactylifera= Linnaeus
_Associate._--_Blattella germanica_, California (Herms, 1926): On date
palms.
=Pritchardia= sp.
_Associate._--_Periplaneta australasiae_, Nihoa Island (Bryan, 1926).
Hawaii (Zimmerman, 1948).
=Roystonea regia= O. F. Cook
_Associate._--_Cariblatta punctulata_, San Domingo (Hebard, 1916a).
=Sabal palmetto= Lodd.
_Associate._--_Eurycotis floridana_, Florida (Scudder, 1879).
_Periplaneta australasiae_, Florida (Hebard, 1917).
=Undetermined palms=
_Associates._--_Euthlastoblatta abortiva_, Texas (Hebard, 1917).
_Hormetica laevigata_, Brazil (Hancock, 1926).
_Panchlora antillarum_, Dominican Republic (Rehn and Hebard, 1927).
_Periplaneta americana_, Texas (Zimmern _in_ Gould and Deay, 1940).
Family ARACEAE
=Arum= sp.
_Associate._--_Latiblattella vitrea_, Mexico (Hebard, 1921b): In flower
shaft.
=Caladium= sp.
_Associate._--_Plectoptera dorsalis_, Puerto Rico (Rehn and Hebard,
1927).
Family BROMELIACEAE
=Aechmaea porteoides= Britton
_Associate._--_Dryadoblatta scotti_, Trinidad (Princis and Kevan, 1955).
=Ananas comosus= Merr.
_Associates._--_Pycnoscelus surinamensis_, Hawaii (Illingworth, 1927,
1929): Feeding on roots of pineapple.
_Blattella humbertiana_, Formosa (Takahashi, 1940): Imago and grown
nymphs occasionally lie concealed in the leaves.
=Catopsis fulgens= Griseb.
_Associates._--Cockroaches, Costa Rica (Calvert and Calvert, 1917).
=Glomeropitcairnia erectiflora= Mez
_Associate._--_Dryadoblatta scotti_, Trinidad (Princis and Kevan, 1955).
=Grevisia= sp.
_Associate._--_Notolampra antillarum_, Trinidad (Princis and Kevan,
1955): One male only.
=Tillandsia fasciculata= Swartz
_Associate._--_Eurycotis floridana_, Florida (Rehn and Hebard, 1914;
Hebard, 1917).
=Tillandsia usneoides= Linnaeus
_Associates._--_Parcoblatta_ sp., Louisiana (Rainwater, 1941).
_Latiblattella rehni_, Florida (Blatchley, 1920): By beating.
Cockroaches, Louisiana (Rosenfeld, 1911, 1912): One mature and 39
immature blattids were collected from 8 of 12 samples of Spanish moss.
=Tillandsia uttriculata= Linnaeus
_Associate._--_Epilampra mona_, Mona Island, West Indies (Rehn and
Hebard, 1927): The type and one paratypic female of _E. mona_ were
collected in this bromeliad.
_Eurycotis floridana_, Florida (Blatchley, 1920).
=Tillandsia= sp.
_Associates._--_Aglaopteryx gemma_, Texas (Hebard, 1917).
_Dryadoblatta scotti_, Trinidad (Scott, 1912): Found in the leaf bases.
=Undetermined bromeliads=
_Associates._--_Aglaopteryx diaphana_, Jamaica (Hebard, 1917; Rehn and
Hebard, 1927).
_Anaplecta azteca_ and _Anaplecta_ sp., Costa Rica (Picado, 1913).
_Anaplecta mexicana_, Costa Rica (Calvert and Calvert, 1917).
_Audreia bromeliadarum_, Panama (Caudell, 1914).
_Audreia jamaicana_, Jamaica (Rehn and Hebard, 1927).
_Blattella_ sp., Costa Rica (Picado, 1913).
_Buboblatta armata_, Panama (Caudell, 1914): "Probably not a typical
bromeliadicolous species."
_Cariblatta insularis_, Jamaica (Hebard, 1916a, 1917; Rehn and Hebard,
1927).
_Cariblatta nebulicola_, Jamaica (Rehn and Hebard, 1927); One immature
male.
_Dryadoblatta scotti_, Trinidad (Princis and Kevan, 1955).
_Epilampra conspersa_, Dominica (Scott, 1912).
_Epilampra maya_, Panama (Hebard, 1920).
_Epilampra sodalis_, Panama (Caudell, 1914).
_Epilampra_ sp. and _Hormetica laevigata_, Brazil (Hancock, 1926).
_Eurycotis biolleyi_, Costa Rica (Picado, 1913).
_Ischnoptera rufa occidentalis_, Mexico (Caudell, 1914).
_Latiblattella chichimeca_, Costa Rica (Picado, 1913).
_Litopeltis biolleyi_, Costa Rica (Rehn, 1928).
_Litopeltis bispinosa_, Panama (Caudell, 1914).
_Neoblattella brunneriana_, Costa Rica (Calvert and Calvert, 1917).
_Neoblattella dryas_, _Neoblattella eurydice_, _Neoblattella
grossbecki_, and _Neoblattella proserpina_, Jamaica (Rehn and Hebard,
1927).
_Neoblattella fratercula_, Mexico (Hebard, 1921b).
_Neoblattella nahua_, Mexico (Caudell, 1914).
_Nesomylacris relica_, Jamaica (Rehn and Hebard, 1927).
_Nyctibora brunnea_(?), Panama (Caudell, 1914): According to Hebard
(1920) Caudell's specimen was almost certainly incorrectly identified.
It may have been _Nyctibora noctivaga_ or a smaller species of the
genus. Brazil (Hancock, 1926).
_Nyctibora laevigata_, Jamaica (Hebard, 1917; Rehn and Hebard, 1927).
_Nyctibora lutzi_, Puerto Rico (Rehn and Hebard, 1927): "in epiphytes
with pencil-like leaves."
_Pelmatosilpha rotundata_, Panama (Caudell, 1914).
_Pseudomops laticornis_, Costa Rica (Picado, 1913).
_Pycnoscelus surinamensis_, Costa Rica (Picado, 1913). Mexico (Caudell,
1914). Jamaica (Rehn and Hebard, 1927).
"_Rhicnoda_" sp., Costa Rica (Picado, 1913). This genus is now
recognized as not being in the New World fauna. Probably the specimen
was a species of _Epilampra_ or _Hyporhicnoda_ as suggested by Gurney
(personal communication, 1959) and confirmed by Rehn (p.c., 1959).
Cockroaches, Costa Rica (Calvert, 1910): Cockroaches were said to be
common in bromeliads on the moist Atlantic slope.
Family LILIACEAE
=Yucca elata= Engelman
_Associate._--_Latiblattella lucifrons_, Arizona (Ball et al., 1942).
=Easter lilies=
_Associate._--_Pycnoscelus surinamensis_, Connecticut (Zappe, 1918).
Family MUSACEAE
=Bananas=
Cockroaches have been captured in bunches of bananas, in bracts of
banana flowers, under banana leaves, and burrowing in rotten banana
stalks. Although many of the species associated with bananas are
indigenous to the banana-growing areas of the American Tropics, most of
the specimens cited below were captured elsewhere as adventitious
insects that had been imported with the fruit. It is obvious that many
of these insects must have been closely associated with bananas on the
plantations, where, undoubtedly, the growing plants provided attractive
ecological niches. Bunting (1956) deduced, from the presence of healthy
cockroaches on bananas allegedly sprayed with copper arsenate, that the
insects did not feed on stems or fruit but hid among the bananas and
foraged elsewhere; however, certain reports are of cockroaches actually
feeding on bananas. Some of the records cited by Hebard (1917) were
compiled from earlier reports not all of which we have seen. Numbers in
parentheses following certain citations indicate the number of times the
association had been observed. Known or suspected adventive material is
so indicated.
_Aglaopteryx diaphana_, Jamaica (Rehn and Hebard, 1927): Found in bracts
of banana blossoms. England (Bunting, 1955): Adventive, on bananas from
Dominica.
_Aglaopteryx vegeta_, Finland (Princis, 1947): Adventive, in banana box.
_Amazonina emarginata_, Trinidad (Princis and Kevan, 1955): In banana
bunch.
_Archimandrita marmorata_, Denmark (Henriksen, 1939): Adventive (2), in
bananas from Jamaica(?). As Princis (1947) and Gurney (personal
communication, 1959) point out, this is a Central American species, so
Jamaica may be an error.
_Archimandrita tessellate_, Sweden (Princis, 1947): Adventive, from
Honduras.
_Blaberus atropos_(?), Denmark (Henriksen, 1939): Adventive, from
Jamaica. Princis (1947) pointed out that this species was more likely to
have been _Blaberus craniifer_ or _Blaberus discoidalis_, which are West
Indian species, than _B. atropos_ which is a South American species.
_Blaberus boliviensis_, Ecuador (Princis, 1952): In a shipment of
bananas from near Puna.
_Blaberus discoidalis_, Puerto Rico (Rehn and Hebard, 1927): From banana
ripening room. Great Britain (Pearce, 1929): Adventive. England
(Bunting, 1955, 1956): Adventive, from Dominica.
_Capucinella delicatula_, California (Caudell, 1931): Adventive.
_Cariblatta delicatula_, Cuba (Rehn and Hebard, 1927).
_Cariblatta hylaea_, Honduras (Rehn, 1945a): Shaken from hanging dead
banana leaves.
_Cariblatta insularis_, Finland (Frey, 1948): Adventive.
_Cariblatta landalei_, Jamaica (Rehn and Hebard, 1927): All specimens
taken from under drying bracts of banana blossoms.
_Cariblatta punctipennis_ and _Chorisoneura barbadensis_, England
(Bunting, 1956): Adventive, from Dominica.
_Epilampra abdomen-nigrum_ and _Epilampra_ sp., England (Bunting, 1955):
Adventive, from Dominica.
_Epilampra maya_, Massachusetts (Hebard, 1917): Adventive.
_Epilampra mexicana_(?), Denmark (Henriksen, 1939): Adventive (2), from
Danish West Indies. Princis (1947) suggested that this should be
_Epilampra_ sp., because _E. mexicana_ is not a West Indian species.
_Eudromiella calcarata_ and _Eurycotis bananae_, U.S.S.R., Leningrad
(Bei-Bienko, 1947): Adventive, from Colombia.
_Euphyllodromia angustata_, Sweden (Princis, 1947): Adventive.
_Eurycotis caraibea_, New York (Hebard, 1917): Adventive.
_Eurycotis dimidiata_, Washington, D. C. (Caudell, 1931): Adventive.
_Eurycotis lixa_, New York (Rehn, 1930): Adventive, on banana ship from
Jamaica.
_Graptoblatta notulata_, Marquesas Islands, Uahuka (Hebard, 1933a): In
banana leaves.
_Holocompsa nitidula_, Trinidad (Princis and Kevan, 1955): Eating banana
pulp.
_Hormetica laevigata_, Wales (Sandemann, 1934): Adventive, in pile of
banana sacks.
_Hormetica ventralis_, Sweden (Princis, 1947): Adventive, in local
warehouse of banana company.
_Hormetica_ spp., Europe and North America (Bei-Bienko, 1950):
Adventive, introduced with bananas and other tropical fruits.
_Ischnoptera rufa rufa_, Puerto Rico (Wolcott, 1950): Brought into
houses on bunches of bananas.
_Kuchinga remota_, Society Islands, Moorea (Hebard, 1933a): In dead
banana leaves.
_Lamproblatta albipalpus_, Panama Canal Zone (Hebard, 1920): Several
under decayed banana stem.
_Latiblattella_ sp., Finland (Frey, 1948): Adventive.
_Leucophaea maderae_, New York (Hebard, 1917): Adventive. Dominica (Rehn
and Hebard, 1927): Under banana sheaths. England (Palmer, 1928):
Adventive, captured at railroad station after bananas had been unloaded.
England (Bunting, 1955): Adventive, from Dominica. Trinidad (Princis and
Kevan, 1955): Nymph, eating bananas in cupboard. Puerto Rico (Seín,
1923): Seín stated that bananas are the favorite food of _L. maderae_.
_Litopeltis bispinosa_, Panama Canal Zone (Hebard, 1920): From rotting
banana stalks at bases of leaves.
_Litopeltis musarum_, Costa Rica (Rehn, 1928): Shaken from dead banana
leaves.
_Nauclidas nigra_, England (Bunting, 1955, 1956): Adventive, from
Dominica.
_Nauphoeta flexivitta_, Denmark (Vestergaard, 1958): Adventive.
_Neoblattella carcinus_, _Neoblattella celeripes_, and _Neoblattella
laodamia_, England (Bunting, 1956): Adventive, from Dominica. Bunting
(1955) first reported these as _Neoblattella_ spp. and stated that they
were common.
_Neoblattella detersa_, Jamaica (Rehn and Hebard, 1927): From under the
bracts of banana blossoms. Sweden (Princis, 1947): Adventive.
_Neoblattella detersa_ and _Neoblattella tridens_, Finland (Frey, 1948):
Adventive.
_Neoblattella fratercula_, Nebraska (Hebard, 1916b): Adventive.
_Neoblattella semota_, Jamaica (Rehn and Hebard, 1927): From under
drying bracts of banana blossoms.
_Neoblattella vatia_, Cuba (Rehn and Hebard, 1927).
_Neoblattella_ sp., Finland (Princis, 1947): Adventive, from Jamaica.
_Nyctibora azteca_, England (Bunting, 1955): Adventive, from Dominica.
Bunting reported this species as _Nocticola azteca_. Dr. A. B. Gurney
called our attention to the fact that _Nocticola_ is an Old World genus,
presumably combined in error with the New World species _azteca_. The
true identity of the specimen was confirmed by Dr. D. Ragge (personal
communication, 1958), who examined it at the British Museum (Natural
History).
_Nyctibora holoserica_, Canada (Walker, 1912): Adventive.
_Nyctibora laevigata_, Canada, Maine, Massachusetts, Pennsylvania (2)
(Hebard, 1917): Adventive. Taken from banana boat _Annetta_ at
Philadelphia (Rehn and Hebard, 1927). England (Bunting, 1956):
Adventive, from Dominica. Sweden, Denmark (Princis, 1947): Adventive.
_Nyctibora mexicana_(?), Denmark (Henriksen, 1939): Adventive (5), from
Jamaica and West Indies. Princis (1947) suggested that these specimens
were probably the West Indian _Nyctibora noctivaga_, because _N.
mexicana_ is not a West Indian insect.
_Nyctibora noctivaga_, Canada, Idaho, Illinois, Massachusetts, Nebraska
(4), Virginia (Hebard, 1917): Adventive. Nebraska (Hauke, 1949):
Adventive (2). Panama Canal Zone (Hebard, 1920): From banana stalks.
England (Blair _in_ Turner, 1930): Adventive, from Costa Rica.
Washington (Hatch, 1938): Adventive. Sweden (Princis, 1947): Adventive
(2). Finland (Princis, 1947): Adventive, from Jamaica.
_Nyctibora obscura_, Trinidad (Princis and Kevan, 1955): In banana
bunch.
_Nyctibora sericea_, Canada (Stevenson, 1905; Walker, 1912): Adventive;
Hebard (1917) synonymized Walker's specimen under _N. laevigata_. Isle
of Wight (Meade-Waldo, 1910): Adventive, from Jamaica. England (Tulloch,
1939): Adventive, in banana crates from Brazil.
_Nyctibora_ sp., England (Welch, 1935): Adventive, in railway truck that
had carried bananas. England (Tulloch, 1939): Adventive, from Brazil.
_Oxyhaloa deusta_, U.S.S.R., Leningrad (Bei-Bienko, 1947): Adventive,
from Colombia.
_Panchlora antillarum_, England (Bunting, 1955): Adventive, from
Dominica.
_Panchlora exoleta_, Scotland (Distant, 1902): Adventive. Great Britain
(Shaw, 1902): Adventive. England (Coney, 1918): Adventive. Sweden,
Norway (Princis, 1947): Adventive, Norwegian specimen from Brazil.
Germany (Zacher, 1917): Adventive, from Jamaica.
_Panchlora nivea_, Colorado, Nebraska, New Jersey, New York (2), Utah
(Hebard, 1917): Adventive. Nebraska (Hauke, 1949): Adventive. Washington
(Hatch, 1938): Adventive. Massachusetts (Roth and Willis, 1958):
Adventive. England (Bunting, 1955): Adventive, from Dominica. U.S.S.R.
(Bei-Bienko, 1947): Adventive, from Colombia. Sweden (13), Norway (3),
Finland (3) (Princis, 1947): Adventive, mostly females; origin (where
known) Jamaica.
_Panchlora fraterna_(?) and _Panchlora peruana_(?), Denmark (Henriksen,
1939): Adventive; origin (where known) Danish West Indies and Jamaica;
Princis (1947) suggested that both species were probably _Panchlora
nivea_.
_Panchlora sagax_, Puerto Rico (Wolcott, 1936).
_Panchlora virescens_, Canada (Walker, 1912): Adventive; this was
probably _P. nivea_ as we now know it (Gurney, personal communication,
1959).
_Panchlora_ sp., Canada (Walker, 1912): Adventive. England (Tulloch,
1939): Adventive, from Brazil.
_Pelmatosilpha coriacea_, Puerto Rico (Wolcott, 1936).
_Pelmatosilpha marginalis_ and _Pelmatosilpha purpurascens_, England
(Bunting, 1955, 1956): Adventive, from Dominica; both species common.
_Pelmatosilpha vagabunda_, New Zealand (Princis, 1954): Adventive,
probably from South America.
_Periplaneta americana_, Belgium (Schepdael, 1931): Adventive, on
bananas from the American Tropics.
_Periplaneta americana_ and _Periplaneta brunnea_, England (Bunting,
1955, 1956): Adventive, from Dominica.
_Periplaneta americana_ and _Periplaneta australasiae_, England (Watson,
1907): Adventive; they ate ripening bananas in the tropical plant house
of the Royal Botanic Gardens, Kew, where they hid in "the sheathing
bases of palm, banana and pandanus leaves." Sweden (Princis, 1947):
Adventive.
_Periplaneta australasiae_, Canada (Walker, 1912): Adventive. Denmark
(Henriksen, 1939): Adventive (9); origin mostly Jamaica. England
(Tulloch, 1939): Adventive, from Brazil. England (Bunting, 1955, 1956):
Adventive, from Dominica; common.
_Platyzosteria bifida_, Nebraska (Hebard, 1917): Adventive.
_Plectoptera dorsalis_, Puerto Rico (Rehn and Hebard, 1927): Captured by
beating banana plants.
_Pycnoscelus surinamensis_, Canada (Walker, 1912; Hebard, 1917):
Adventive: Marquesas Islands, Nukuhiva (Hebard, 1933a): In banana
leaves. England (Goodliffe, 1958): Adventive, doing considerable damage
to banana plants growing in a conservatory.
_Sibylloblatta panesthoides_, Massachusetts (Rehn, 1937a): Adventive,
from Jamaica.
Family ZINGIBERACEAE
=Renealmia= sp.
_Associate._--_Cariblatta orestera_, Jamaica (Rehn and Hebard, 1927):
The male was taken in a head of wild ginger.
Family CANNACEAE
=Canna= sp.
_Associate._--_Periplaneta americana_, Hawaii (Zimmerman, 1948).
Family ORCHIDACEAE
=Cattleya= sp.
_Associates._--_Periplaneta americana_, U.S.A. (Rau, 1940a).
_Periplaneta australasiae_, England (Lucas, 1918).
=Vanda= sp.
_Associates._--_Periplaneta americana_, U.S.A. (Rau, 1940a).
_Periplaneta australasiae_, England (Lucas, 1918).
=Undetermined orchids=
_Associates._--_Blaberus discoidalis_, _Blatta orientalis_, _Periplaneta
americana_, Hawaii (Swezey, 1945).
_Blatta orientalis_, _americana_, _cinerea_, _maderae_, unidentified
cockroaches, England, in bulb from Ecuador (Westwood, 1876).
_Graptoblatta notulata_, Hawaii (Swezey, 1945): On orchid from India.
_Homalopteryx laminata_ and _Hormetica apolinari_, New York (Hebard,
1912c): In orchids shipped from Colombia.
_Pelmatosilpha coriacea_, Puerto Rico (Wolcott, 1936).
_Periplaneta americana_, Germany (Tashenberg, 1884).
_Periplaneta australasiae_, England (Wainwright, 1898). Pennsylvania
(Skinner, 1905). Massachusetts (Morse, 1920).
_Pycnoscelus surinamensis_, England (Westwood, 1869). Germany (Zacher,
1920). Massachusetts (Morse, 1920). Hawaii (Swezey, 1945).
Family CASUARINACEAE
=Casuarina= sp.
_Associate._--_Diploptera punctata_, Hawaii (Zimmerman, 1948).
Family SALICACEAE
=Populus euphratica= Oliv.
_Synonymy._--_Populus diversifolia_ Schrenk. [Howard, personal
communication, 1959].
_Associate._--_Ectobius semenovi_, Kazakhstan (Bei-Bienko, 1950).
=Populus= sp.
_Associate._--_Ectobius lapponicus_, U.S.S.R. (Stark _in_ Bei-Bienko,
1950): On aspen.
=Salix= sp.
_Associate._--_Ectobius semenovi_, Kazakhstan (Bei-Bienko, 1950): On
willow.
Family MYRICACEAE
=Myrica cerifera= Linnaeus
_Associate._--_Chorisoneura texensis_, Florida (Rehn and Hebard, 1916):
On bayberry. Florida (Blatchley, 1920): Beaten from foliage.
Family FAGACEAE
=Quercus alba= Linnaeus
_Associate._--_Parcoblatta pensylvanica_, Virginia (Rehn and Hebard,
1916): Under signs on white oaks.
=Quercus rubra= Linnaeus
_Associates._--_Parcoblatta divisa_, Virginia, and _Parcoblatta
pensylvanica_, North Carolina (Rehn and Hebard, 1916): Under signs on
red oak.
_Parcoblatta lata_, North Carolina (Hebard, 1917): Under sign.
=Quercus virginiana= Mill.
_Associate._--_Eurycotis floridana_, Georgia (Rehn and Hebard, 1916):
Under dead bark on live-oak tree. Georgia (Hebard, 1917): In cavity in
tree.
=Quercus= spp.
_Associates._--_Aglaopteryx gemma_, Alabama, Georgia, Florida,
Louisiana, Texas (Hebard, 1917): Under signs on oaks.
_Blatta orientalis_, England (Donisthorpe, 1918): Under bark.
_Cariblatta lutea lutea_, Mississippi (Hebard, 1916a): By beating low
oaks on hills.
_Chorisoneura texensis_, Mississippi (Hebard, 1917). Florida (Blatchley,
1920): By beating.
_Ectobius pallidus_, England (Milton, 1899; Burr, 1899b). Massachusetts
(Flint, 1951): Under loose lichens and bark.
_Parcoblatta divisa_, Georgia, Louisiana, and _Parcoblatta
pensylvanica_, Georgia (Hebard, 1917): Under signs.
_Periplaneta australasiae_, Florida (Rehn and Hebard, 1905): Ten
specimens taken from under a tin sign.
_Periplaneta brunnea_, Georgia (Rehn and Hebard, 1916): Under signs.
_Phyllodromica megerlei_, U.S.S.R. (Bei-Bienko, 1950): By shaking oak
branches.
_Plectoptera lacerna_, Cuba (Rehn and Hebard, 1927).
Family MORACEAE
=Cecropia= sp.
_Associate._--_Cariblatta hylaea_, Honduras (Rehn, 1945a).
Family CHENOPODIACEAE
=Beta maritima= Linnaeus
_Associate._--_Ectobius panzeri_, England (Lucas, 1920a).
=Beta vulgaris= var. =cicla= Linnaeus
_Associate._--_Ectobius pallidus_, Massachusetts (Flint, 1951): Many
specimens collected in the bases of Swiss chard plants.
Family LAURACEAE
=Nectandra coriacea= (Sw.) Griseb.
_Synonymy._--_Ocotea catesbyana_ Sarg. [Howard, personal communication,
1959].
_Associate._--_Chorisoneura texensis_, Florida (Rehn and Hebard, 1912).
Family SARRACENIACEAE
Only a few records have been found of cockroaches being trapped in the
pitchers of carnivorous plants of this and the following family. The
insects drown in the fluid within the pitcher where they are apparently
digested by proteinases secreted by the plant (Meyer and Anderson, 1939;
Lloyd, 1942).
=Sarracenia flava= Linnaeus
_Natural prey_--_Cariblatta lutea lutea_, _Ischnoptera deropeltiformis_,
_Parcoblatta lata_, and nymphs of _Parcoblatta_ sp., North Carolina
(Wray and Brimley, 1943): Most of the cockroaches seemed to have been
trapped accidentally with the possible exception of _C. lutea lutea_, 11
of which were found in _Sarracenia_ pitchers.
=Sarracenia purpurea= Linnaeus
_Natural prey._--_Cariblatta lutea lutea_, North Carolina (Wray and
Brimley, 1943).
=Sarracenia minor= Walter
_Synonymy._--_Sarracenia variolaris_ Michx. [Howard, personal
communication, 1958].
_Natural and experimental prey._--_Periplaneta australasiae_, Florida
(Treat, 1876): After the insect imbibed some of the fluid in the pitcher
it became docile; others became highly active and rushed wildly about
before becoming quiescent. See also Treat _in_ Scudder (1877).
Cockroaches, U.S.A. (Riley, 1875).
Family NEPENTHACEAE
=Nepenthes ampularia= Jack
_Natural prey._--Cockroaches, Singapore (Dover, 1928).
=Nepenthes gracilis= Korth.
_Natural prey._--Cockroaches, Singapore (Dover, 1928).
=Nepenthes= sp.
_Natural prey._--Cockroach, Old World Tropics? (Hooker, 1874): The
insect was apparently attracted into the pitcher, where it drowned, by a
piece of cartilage placed there by Hooker.
Family CUNONIACEAE
=Weinmannia= sp.
_Associates._--_Aneurina viridis_, Marquesas Islands, Nukuhiva and
Fatuhiva (Hebard, 1933a).
_Maretina uahuka_, Marquesas Islands, Uahuka (Hebard, 1933a).
Family HAMAMELIDACEAE
=Liquidambar styraciflua= Linnaeus
_Associate._--_Parcoblatta divisa_, Georgia (Rehn and Hebard, 1916):
Under sign on sweet gum.
_Parcoblatta zebra_, Louisiana (Hebard, 1917): In decay cavity.
Family ROSACEAE
=Crataegus= sp.?
=Associates.=--_Plectoptera dorsalis_, _Plectoptera infulata_,
_Plectoptera rhabdota_, Puerto Rico (Wolcott, 1950): In the dry flower
clusters of "espino rubial."
=Rosa= sp.
_Associate._--_Pycnoscelus surinamensis_, Connecticut (Zappe, 1918);
Rhode Island and Pennsylvania (Caudell, 1925); Pennsylvania (Doucette
and Smith, 1926): Feeding on canes in greenhouses.
=Rubus= spp.?
_Associate._--_Hololampra chavesi_, Azores (Chopard, 1932): This species
is exclusively dendricolous and was found only by beating the bushes on
which it abounds. It was very common in hedges, particularly on brambles
(ronces).
Family LEGUMINOSAE
=Acacia farnesiana= Willd.
_Associate._--_Diploptera punctata_, Hawaii (Bridwell and Swezey, 1915;
Zimmerman, 1948): Feeding on pods.
=Acacia= sp.
_Associates._--_Ellipsidion australe_, Australia, New South Wales
(Hebard, 1943).
_Methana curvigera_, Australia, Queensland (Pope, 1953a).
=Ceratonia siliqua= Linnaeus
_Associate._--_Diploptera punctata_, Hawaii (Pemberton and Williams,
1938): Damaging algarroba.
=Erythrina glauca= Willd.
_Associates._--_Aglaopteryx absimilis_, Puerto Rico (Wolcott, 1950): In
abandoned cocoon.
_Aglaopteryx facies_, Puerto Rico (Wolcott, 1936): In empty cocoons.
=Inga laurina= Willd.
_Associate._--_Aglaopteryx facies_, Puerto Rico (Wolcott, 1936): On
trunk.
=Inga vera= Willd.
_Associates._--_Aglaopteryx facies_, Puerto Rico (Wolcott, 1936): In
larval tents.
_Cariblatta stenophrys_, Puerto Rico (Wolcott, 1936): On leaves.
_Plectoptera dorsalis_, _Plectoptera infulata_, and _Plectoptera
rhabdota_, Puerto Rico (Wolcott, 1950): In "butterfly nests" in leaves.
=Mesquite=
_Associate._--_Nyctibora stygia_, Haiti (Rehn and Hebard, 1927).
=Samanea saman= Merr.
_Associate._--_Aglaopteryx absimilis_, Puerto Rico (Wolcott, 1950).
=Tamarindus indica= Linnaeus
_Associate._--_Hemiblabera brunneri_, Puerto Rico (Rehn and Hebard,
1927).
Family GERANIACEAE
=Geraniums=
_Associate._--_Diploptera punctata_, Hawaii (Zimmerman, 1948).
Family ZYGOPHYLLACEAE
=Tribulus= sp.
_Associates._--_Periplaneta americana_ and _Pycnoscelus surinamensis_,
Johnston Island (Bryan, 1926). Zimmerman (1948) lists _Tribulus_ as a
host plant for these cockroaches.
Family RUTACEAE
=Citrus aurantifolia= Swingle
_Associates._--_Plectoptera dominicae_ and _Plectoptera perscita_,
Dominica (Rehn and Hebard, 1927): Beaten from moss-covered lime trees.
=Citrus maxima= Merr.
_Associates._--_Plectoptera dorsalis_, _Plectoptera infulata_,
_Plectoptera rhabdota_, Puerto Rico (Wolcott, 1950).
_Plectoptera rhabdota_, Puerto Rico (Rehn and Hebard, 1927).
=Citrus sinensis= Osbeck
_Associate._--_Diploptera punctata_, Hawaii (Bridwell and Swezey, 1915;
Zimmerman, 1948): Feeding on oranges on tree.
=Citrus= sp.
_Associates._--_Diploptera punctata_, Hawaii (Zimmerman, 1948).
_Riatia_ [=_Lissoblatta_] _fulgida_, Panama, Rio Trinidad (Hebard,
1920).
_Plectoptera porcellana_, Puerto Rico (Sein, 1923).
=Zanthoxylum caribaeum= Lam.
_Associates._--_Plectoptera dorsalis_, _Plectoptera infulata_,
_Plectoptera rhabdota_, Puerto Rico (Wolcott, 1950): In the dry flower
clusters.
Family BURSERACEAE
=Bursera simaruba= (L.) Sarg.
_Associate._--_Chorisoneura texensis_, Florida (Rehn and Hebard, 1912;
Hebard, 1917): Beaten from the lower branches of gumbo limbo.
Family EUPHORBIACEAE
=Poinsettia= sp.
_Associate._--_Pycnoscelus surinamensis_, Connecticut (Zappe, 1918): Ate
bark of greenhouse plants.
Family ANACARDIACEAE
=Mangifera indica= Linnaeus
_Associate._--_Diploptera punctata_, Hawaii (Bridwell and Swezey, 1915;
Zimmerman, 1948): Feeding on mangoes on the tree.
=Spondias mombin= Linnaeus
_Associates._--_Plectoptera dorsalis_, _Plectoptera infulata_,
_Plectoptera rhabdota_, Puerto Rico (Wolcott, 1950): Living on leaves of
"jobo."
=Spondias purpurea= Linnaeus
_Associate._--_Eurycotis biolleyi_, Costa Rica (Rehn, 1918): In the
crown of dry jocoto.
Family AQUIFOLIACEAE
=Ilex cassine= Linnaeus
_Associate._--_Plectoptera poeyi_, Florida (Rehn and Hebard, 1912, 1914;
Hebard, 1917).
=Ilex coriacea= (Pursh) Chapm.
_Synonymy._--_Ilex lucida_ [Fernald, 1950].
_Associate._--_Cariblatta lutea lutea_, Florida (Hebard, 1916a).
Family SAPINDACEAE
=Exothea paniculata= (Juss.) Radlk.
_Associate._--_Aglaopteryx gemma_, Florida (Hebard, 1917).
Family MALVACEAE
=Gossypium= spp.
_Associates._--_Graptoblatta notulata_, Marquesas Islands, Tahuata
(Hebard, 1933a).
_Periplaneta australasiae_, St. Kitts, B.W.I. (Ballou, 1916).
_Periplaneta fuliginosa_ and _Plectoptera poeyi_, Florida (Rainwater,
1941).
_Plectoptera dorsalis_, _Plectoptera infulata_, _Plectoptera rhabdota_,
Puerto Rico (Wolcott, 1950).
=Hibiscus rosa-sinensis= Linnaeus
_Associate._--_Riatia orientis_, Trinidad (Princis and Kevan, 1955).
=Hibiscus= sp.
_Associates._--_Amazonina emarginata_, _Cariblatta antiguensis_,
_Eurycotis kevani_, and _Rhytidometopum dissimile_, Trinidad (Princis
and Kevan, 1955).
=Sida= sp.
_Associate._--_Periplaneta australasiae_, Nihoa Island (Bryan, 1926).
Hawaii (Zimmerman, 1948).
Family STERCULIACEAE
=Theobroma cacao= Linnaeus
_Associate._--_Ceratinoptera picta_, Trinidad (Princis and Kevan, 1955).
Family BIXACEAE
=Bixa= sp.
_Associate._--_Notolampra antillarum_, Trinidad (Princis and Kevan,
1955): Nymphs in dry fruits on "annato" tree.
Family FLACOURTIACEAE
=Xylosma suaveolens= Forst.
_Associate._--_Graptoblatta notulata_, Marquesas Islands, Uahuka
(Hebard, 1933a).
Family PASSIFLORACEAE
=Passiflora= sp.
_Associate._--_Aristiger_ [=_Plumiger_] _histrio_, Malaya (Karny, 1924).
Family CARICACEAE
=Carica papaya= Linnaeus
_Associate._--_Diploptera punctata_, Hawaii (Bridwell and Swezey, 1915;
Zimmerman, 1948): Feeding on papaya fruit on tree.
Family RHIZOPHORACEAE
=Rhizophora mangle= Linnaeus
_Associate._--_Aglaopteryx gemma_, Florida (Hebard, 1917).
Family COMBRETACEAE
=Conocarpus erectus= Linnaeus
_Associate._--_Plectoptera poeyi_, Florida (Rehn and Hebard, 1914):
Running on leaves.
Family MYRTACEAE
=Eucalyptus= sp.
_Associate._--_Ellipsidion australe_, Australia, New South Wales
(Hebard, 1943).
=Eugenia aromatica= Baill.
_Synonymy._--_Syzygium aromaticum_ [Bailey, 1925].
_Associate._--_Plectoptera dorsalis_, Puerto Rico (Wolcott, 1936): On
flowers of "pomarrosa."
=Metrosideros collina= Gray
_Associates._--_Aneurina viridis_, Marquesas Islands: Nukuhiva,
Fatuhiva, and Tahuata (Hebard, 1933a)
_Aneurina tahuata_, Marquesas Islands, Tahuata (Hebard, 1933a).
_Graptoblatta notulata_, Marquesas Islands, Nukuhiva (Hebard, 1933a).
=Psidium guajava= Linnaeus
_Associate._--_Plectoptera rhabdota_, Puerto Rico (Rehn and Hebard,
1927).
Family ONAGRACEAE
=Jussiaea natans= Humb. and Bonpl.
_Associate._--_Epilampra abdomen-nigrum_, Panama (Crowell, 1946): In an
aquarium the cockroach fed on leaves of this aquatic plant which had
been collected in the lagoon where the insect was captured.
Family ERICACEAE
=Calluna vulgaris= Salisb.
_Associates._--_Ectobius lapponicus_, England (Lucas, 1925): "Nymphs of
varying size were beaten out of heather ... on 9 February and later."
_Ectobius panzeri_, England (Lucas, 1927): "numerous imagines of both
sexes were swept from heather."
=Vaccinium meridionale= Sw.
_Associates._--_Chorisoneura formosella_, _Neoblattella dryas_,
_Neoblattella proserpina_, Jamaica (Rehn and Hebard, 1927).
Family SAPOTACEAE
=Sideroxylon foetidissimum= Jacq.
_Associate._--_Pelmatosilpha coriacea_, Puerto Rico (Wolcott, 1941):
Under bark.
Family APOCYNACEAE
=Vinca minor= Linnaeus
_Associate._--_Ectobius pallidus_, Massachusetts (Willis, unpublished
observation, 1958).
Family CONVOLVULACEAE
=Ipomoea tiliasea= Choisy
_Associate._--_Plectoptera dorsalis_, Puerto Rico (Rehn and Hebard,
1927).
Family BORAGINACEAE
=Cordia dentata= Poiret
_Synonymy._--_Calyptracordia alba_ [Howard, personal communication,
1958].
_Associates._--_Cariblatta antiguensis_, _Ischnoptera rufa rufa_,
_Supella supellectilium_, _Symploce ruficollis_ and _Symploce hospes_,
St. Croix, Virgin Islands (Beatty, 1944): On fruits of _C. dentata_
except _S. supellectilium_ which was found at night on the flowers.
Family VERBENACEAE
=Citharexylum villosum= Jacq.
_Associate._--_Chorisoneura texensis_, Florida (Rehn and Hebard, 1912).
Family SOLANACEAE
=Nicotiana= sp.
_Associate._--_Pycnoscelus surinamensis_, Sumatra (Roeser, 1940).
=Solanum tuberosum= Linnaeus
_Associate._--_Pycnoscelus surinamensis_, Haiti (Hoffman, 1927): Feeding
on tubers in field.
Family GESNERIACEAE
=Cyrtandra= sp.
_Associate._--_Aneurina viridis_, Marquesas Islands, Nukuhiva (Hebard,
1933a).
Family RUBIACEAE
=Canthium barbatum= (Forst.) Seem.
_Associate._--_Graptoblatta notulata_, Marquesas Islands, Uahuka
(Hebard, 1933a).
=Cinchona pubescens= Vahl.
_Associate._--_Periplaneta americana_, Puerto Rico (Plank and Winters,
1949): In greenhouse.
=Coffea= sp.
_Associate._--_Plectoptera porcellana_, Puerto Rico (Seín, 1923).
Family COMPOSITAE
=Goldenrod=
_Associate._--_Eurycotis floridana_, Florida (Hebard, 1917): "Climbing
about on top of goldenrod at night."
=Helianthus= sp.
_Associate._--_Pseudomops septentrionalis_, Texas (Hebard, 1917).
=Scorzonera acanthoclada= Franch.
_Associate._--_Phyllodromica tartara nigrescens_, Southern Uzbekistan
(Bei-Bienko, 1950): On the flowers.
DAMAGE TO PLANTS BY COCKROACHES
Cockroaches characteristically feed on dead plant and animal material.
Damage to living plants occurs principally in the Tropics or under
subtropical conditions in greenhouses in temperate regions. Among the
depredations attributed to cockroaches in text books, damage to plants
is seldom emphasized. This is surprising in view of the many records
cited below.
Capt. William Bligh (1792), while collecting breadfruit trees in Tahiti
to take to the West Indies, wrote in his log during January 1789: "This
morning, I ordered all the chests to be taken on shore, and the inside
of the ship to be washed with boiling water, to kill the cockroaches. We
were constantly obliged to be at great pains to keep the ship clear of
vermin, on account of the plants."
Westwood (1869) stated that _Pycnoscelus surinamensis_ was very
destructive in orchid houses feeding on buds and young shoots. Later
Westwood (1876) exhibited the bulb of an orchid from Ecuador which
contained six species of cockroaches: _Blatta orientalis_,
[_Periplaneta_?] _americana_, [_Nauphoeta_?] _cinerea_, [_Leucophaea_?]
_maderae_, and two others unknown to him. Fullaway (1938) stated that
cockroaches damage root tips, buds, and flowers of orchids. _Periplaneta
americana_ has been said to eat the root tips and blossoms of orchids
(Taschenberg, 1884) and to devour the open flower petals of _Cattleya_
orchids as well as the aerial roots and flower spikes of _Vanda_ orchids
(Rau, 1940a). Wainwright (1898) stated that _Periplaneta australasiae_
had been observed in an orchid house in Perthshire where over a period
of three years it had caused a good deal of damage. Skinner (1905)
reported that _P. australasiae_ in greenhouses in Pennsylvania showed no
preference for any one plant but ate both plants and flowers of orchids,
roses, and carnations. Lucas (1918) received specimens of _P.
australasiae_ which had played havoc with orchids especially _Cattleya_
and _Vanda_. Morse (1920) reported that both _P. australasiae_ and
_Pycnoscelus surinamensis_ were obnoxious in a conservatory in
Massachusetts where they gnawed the tips of the aerial roots of orchids.
Swezey (1945) in Hawaii stated that the following cockroaches have been
reported as occasional minor pests on orchids: _Blatta orientalis_,
_Blaberus discoidalis_, _P. americana_, and _P. surinamensis_; he
further stated that _Graptoblatta notulata_ had been intercepted at
Honolulu on orchids from India.
Watson (1907) stated that _Blatta orientalis_, _Periplaneta americana_,
and _Periplaneta australasiae_ were injurious in the tropical plant
houses at Kew: "at night they come out and run or fly about among the
plants, devouring flowers and leaves like rabbits. Such plants as
_Eucharis_, _Crinum_ and _Alpinia_, when in flower, have little chance
in the palm house, where the cockroaches are most abundant; they also
find out the ripening bananas and soon devour them." Raffill (1910)
stated that in plant houses in England _B. orientalis_, _P. americana_,
and _P. australasiae_ commonly, and _Nauphoeta cinerea_, _Nauphoeta
flexivitta_, and _Pycnoscelus surinamensis_ more rarely, are extremely
destructive to plants. Flowers having a strong perfume, such as orchids,
_Eucharis_, _Crinum_, and _Hedychium_, were often attacked while other
flowers nearby were left uninjured.
Plank and Winters (1949) reported that in Puerto Rico the species of
Orthoptera most injurious under greenhouse conditions was _Periplaneta
americana_. Large nymphs destroyed 25 to 30 percent of freshly planted
seed of _Cinchona pubescens_. In Hawaii the host plants of _P.
americana_ are blossoms of _Canna_ and _Tribulus_, and the host plants
of _Periplaneta australasiae_ are _Pritchardia_ and _Sida_ (Zimmerman,
1948). On St. Kitts, B.W.I., young cotton plants were severely attacked
by _P. australasiae_; this caused loss of the stand on a considerable
area and necessitated replanting (Ballou, 1916). _P. australasiae_ was
reported damaging the _Polystichum aristatum_ Presl [=_Lastrea aristata
variegata_] in a greenhouse (Thilow and Riley, 1891). Laing (1946;
British Museum [Natural History], 1951) stated that _P. australasiae_
abounds in greenhouses and forcing pits where it may do considerable
damage to the plants. _Periplaneta fuliginosa_ is also troublesome in
greenhouses because of its tendency to feed on seedlings and succulent
plants (Dodge and Rickett, 1943).
_Ectobius lapponicus_ has been observed feeding in galleries in the
thick skin of young aspen in 25 percent of the trees examined (Stark
_in_ Bei-Bienko, 1950). The aquatic cockroach _Epilampra abdomen nigrum_
fed on the leaves of _Jussiaea natans_ in an aquarium (Crowell, 1946).
_Ischnoptera deropeltiformis_ has been taken while it was feeding on a
fleshy fungus (_Agaricus_ sp.) in dense woods in Indiana (Blatchley,
1920).
_Diploptera punctata_, the cypress roach or beetle roach, has been found
in Hawaii feeding on ripening mangoes and papayas, oranges on the tree,
and the outer covering of the pods of _Acacia farnesiana_ (Bridwell and
Swezey, 1915). Pemberton (1934) stated that _D. punctata_ "disfigures
our cypress trees by eating the bark from the young branches, often
giving them a dead appearance over much of their leaf area." Fullaway
and Krauss (1945) added, "This injury [to cypress] is so severe that
sometimes areas of leaves die and turn brown. The Japanese cedar,
ironwood, citrus and algaroba (kiawe) trees are attacked in a similar
manner." Similar injury to cypress was described by Hebard (1922). In
addition to girdling _Cupressus_, _D. punctata_ injures _Cryptomeria_ in
the same fashion and also attacks algaroba, lime, and other plants
(Pemberton and Williams, 1938). Zimmerman (1948) cited the following
host plants for _D. punctata_ in Hawaii: "_Cupressus macrocarpa_,
_Casuarina_, _Cryptomeria_, _Citrus_, geraniums, _Acacia farnesiana_
pods, mango fruits, orange fruits, papaya fruits."
In the reports of damage to plants by cockroaches, _Pycnoscelus
surinamensis_ has been implicated most often. This species is
undoubtedly one of the economically most important cockroaches, being
the vector of the chicken eyeworm as well as feeding on plants. In
addition to the few reports of damage caused by this species that have
already been mentioned, _P. surinamensis_ has been reported to be very
destructive in New Orleans to palms and ferns, attacking large
alsophilas avidly, eating out the hearts (Anonymous, 1893). Zappe (1918)
in Connecticut reported damage in a greenhouse to roses valued, at that
time, at several hundred dollars; _P. surinamensis_ had girdled the rose
bushes, done much damage to Easter lilies, and in another greenhouse had
eaten the bark from the stems of poinsettias. In Germany this species
bit off the tips of the aerial roots and ate the petals of orchids
(Zacher, 1920). Lucas (1923) reported damage to cucumber plants in a
greenhouse in Surrey. Damage by _P. surinamensis_ to the stems of rose
bushes has been reported in Rhode Island and Pennsylvania; the canes
were attacked both under and above ground (Caudell, 1925). Doucette and
Smith (1926) reported a heavy infestation of _P. surinamensis_ in a
range of greenhouses in Philadelphia: "The roaches were present
literally by the millions.... Although the roaches had been observed in
cabinets and trash barrels for several months, it was not until the
manager had occasion to go through the house one evening that he
discovered that roaches were the cause of the troubles previously
attributed to soil condition, watering, fungus, and other agencies....
About 30,000 to 35,000 rose plants from a total of 200,000 in the three
more heavily infested houses were so badly injured by the gnawing off of
the bark, young buds, and shoots of the main stems, that they were not
in condition to be kept in the beds for another season."
In Haiti _Pycnoscelus surinamensis_ damaged the tubers of growing
potatoes (Hoffman, 1927). Illingworth (1927, 1929) reported that in
Hawaii _P. surinamensis_ was a minor pest of pineapples, feeding on the
roots. This species was very plentiful in a propagating pit in England
where it did much damage to various seeds and seedlings (Lucas, 1930).
Roeser (1940) summarized some of the above-mentioned damage caused by
_P. surinamensis_ and added damage to chrysanthemums in Hawaii and
tobacco in Sumatra where this cockroach destroyed 300,000 plants in a
few days. Roeser was of the opinion that living plants were eaten only
as a substitute when the earth became poor in food material. Zimmerman
(1948) listed as host plants of _P. surinamensis_ in Hawaii: "blossoms
of _Tribulus_; reported feeding at roots of pineapples, and unconfirmed
reports of damage to underground parts of some other plants." Goodliffe
(1958) reported damage by this species to banana plants in a
conservatory in northern England. Cohic (1956) implied that in New
Caledonia "Racines de légumes" were attacked by _P. surinamensis_ and
that _Zea mays_ Linnaeus was attacked by _Supella supellectilium_.
Wolcott (1924a) reported that _P. surinamensis_ damaged transplanted
tobacco plants in Puerto Rico by eating the interior of the stalks.
Tobacco planters in Cuba consider _P. surinamensis_ injurious to the
roots of tobacco plants (Bruner and Scaramuzza, 1936); this belief was
confirmed in the laboratory, where adults and nymphs destroyed the roots
and stems of tobacco plants two inches high and ate into the edges of
the leaves. Dammerman (1929) reported that in Malaya this species often
appeared in large numbers in gardens where it gnawed at the underground
parts of vegetables and ornamental plants. Lever (1947) listed it as a
pest on the leaves of pineapple.
_Blattella vaga_ may occasionally damage seedlings in the laboratory
(Flock, 1941a), but no damage has been reported in the field (Ball et
al., 1942). Heer (1864) reported receiving a shipment of cycads from
Cuba with all stages of _Periplaneta americana_ living in holes in the
branches, apparently subsisting on the starchy tissues. Goldenberg
(1877) stated that sago trees provide cockroaches with their favorite
nourishment. Scudder (1879) found _Eurycotis floridana_ living in the
tops of the cabbage palmetto, on which he presumed it fed. _Parcoblatta
americana_ has been observed feeding on an apple 6 feet above ground
(Fulton, 1930).
X. PROTOZOA ASSOCIATED WITH COCKROACHES
The classification of the Protozoa follows that of Kudo (1954). The use
of the asterisk (*) is explained in footnote 3, page 4.
=Phylum PROTOZOA=
Class MASTIGOPHORA
Order EUGLENOIDINA
Family EUGLENIDAE
=Euglena= sp.
_Experimental host._--_Periplaneta americana_, U.S.A. (Hegner, 1929):
When fed to the insects in concentrated culture, _Euglena_ could
withstand conditions in the crop up to 5 hours and were passed into the
stomach in a viable state up to 6 hours. However, the majority were
killed in the crop within 2 hours and very few reached the stomach
alive.
Order PROTOMONADINA
Family OIKOMONADIDAE
=Oikomonas blattarum Tejera=
_Natural host._--Cockroach, Venezuela (Tejera, 1926).
=Oikomonas= sp.
_Natural host._--_Blatta orientalis_, U.S.S.R. (Yakimov and Miller,
1922): _Oikomonas_ sp. and _Monas_ sp. were found in the intestines of
83 percent of 124 cockroaches.
Cockroach, Venezuela (Tejera, 1926).
Family TRYPANOSOMATIDAE
=Leptomonas blaberae= Tejera
_Natural host._--_Blaberus_ sp., Venezuela (Tejera, 1926).
=Leptomonas= sp.
_Natural hosts._--_Parcoblatta lata_, _Parcoblatta pensylvanica_,
_Parcoblatta virginica_, U.S.A., Ohio (Semans, 1939, 1941): Hind
intestine. Of 70 specimens examined, 86 percent harbored _Leptomonas_
sp.
* =Herpetomonas periplanetae= Laveran and Franchini
_Natural host._--_Blatta orientalis_, Italy, France (Laveran and
Franchini, 1920, 1920a).
Family MONADIDAE
=Monas= sp.
_Natural host._--_Blatta orientalis_, U.S.S.R. (Yakimov and Miller,
1922): _Monas_ sp. and _Oikomonas_ sp. were found in the intestines of
83 percent of 124 cockroaches examined.
Cockroach, Venezuela (Tejera, 1926).
Family BODONIDAE
=Bodo blattae=
_Natural host._--_Blatta orientalis_, England (Lankester, 1865).
=Bodo= sp.
_Natural host._--_Blattella germanica_ and/or _Periplaneta americana_,
South Africa (Porter, 1930).
=Retortamonas blattae= (Bishop)
_Synonymy._--_Embadomonas blattae_ Bishop [Wenrich, 1932].
_Natural host._--_Blatta orientalis_, England (Bishop, 1931): Hind
intestine. The organism occurred in about 40 percent of the cockroaches
examined. L. G. Feo (_in_ Wenrich, 1932) successfully cultured this
protozoan (fig. 2, F).
=Retortamonas= sp.?
_Natural host._--_Leucophaea maderae_, Philippine Islands (Hegner and
Chu, 1930).
[Illustration: FIG. 2.--Representative Protozoa associated with
cockroaches. A, _Monocercomonoides melolonthae_, × 3094 (after Grassé).
B, _Coelosporidium periplanetae_, × 1310 (after Sprague); trophozoite
with spores and chromatoid bodies. C, _Endamoeba blattae_, × 273 (after
Kudo); trophozoite. D, _Lophomonas striata_, × 330 (after Kudo). E,
_Lophomonas blattarum_, × 660 (after Kudo). F, _Retortamonas blattae_,
× 3094 (after Wenrich). G, _Nyctotherus ovalis_, × 175 (after Kudo). H,
_Gregarina rhyparobiae_, c. × 52: mature trophozoite attached to
intestinal wall of _Leucophaea maderae_. (Redrawn from J. M. Watson
[1945].) I, _Diplocystis schneideri_, c. × 14.4 (after Kunstler). J,
_Gregarina blattarum_, c. × 57 (after Kudo). K, _Protomagalhaesia
serpentula_, × 36 (after Pinto). L, _Gamocystis tenax_, magnification
not known (after Schneider). (All figures except H redrawn from Kudo
[1954] after sources indicated.)]
Order POLYMASTIGINA
Family CHILOMASTIGIDAE
=Chilomastix mesnili= (Wenyon)
_Experimental vectors._--_Blatta orientalis_ and _Periplaneta
americana_, South Africa (Porter, 1918): The cockroaches were fed human
excrement that contained cysts of _C. mesnili_. The cysts passed
unharmed through the insects' digestive tract. Rats became infected with
this protozoan on eating food that had been contaminated with feces from
these cockroaches.
Family POLYMASTIGIDAE
=Eutrichomastix= sp.
_Synonymy._--_Trichomastix_ [Kudo, 1954].
_Natural host._--_Blattella germanica_ and/or _Periplaneta americana_,
South Africa (Porter, 1930).
=Monocercomonoides globus= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934): Organism occurs in
practically all hosts.
=Monocercomonoides melolonthae= (Grassi)
(Fig. 2, A)
_Natural host._--_Platyzosteria novae seelandiae_, New Zealand (Laird,
1956): Found in the intestinal tracts of the adult cockroaches, and of
other species of insects.
=Monocercomonoides orthopterorum= (Parisi)
_Synonymy._--_Trichomonas_ (_Trichomastix_) _orthopterorum_ Parisi;
_Monocercomonas orthopterorum_ [Bělǎr, 1916]; _Trichomastic
orthopterum?_ [Zasukhin, 1930]; _Monocercomonoides orthopterorum_
[Travis, 1932; Cleveland et al., 1934]; _Retortamonas orthopterorum_
[Semans, 1943].
_Natural hosts._--_Blatta orientalis_, Italy (Parisi, 1910); U.S.S.R.
(Zasukhin, 1930).
_Ectobius lapponicus_, Italy (Parisi, 1910).
_Periplaneta americana_, Philippine Islands (Hegner and Chu, 1930).
"Küchenschaben," Austria (Bělǎr, 1916).
The protozoan is found in the hind gut. Zasukhin (1930) found the
organism in 85 percent of over 3,000 _B. orientalis_. Parisi (1910)
found the flagellate present in very large numbers.
=Monocercomonoides panesthiae= Kidder
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Kidder,
1937): In hind gut.
=Tetratrichomastix blattidarum= Young
_Natural hosts._--_Blatta orientalis_, _Blattella germanica_,
_Periplaneta americana_, U.S.A. (Young, 1935): The organism when present
occurs in large numbers in the posterior part of the intestine near the
anus. The protozoan was successfully cultivated in a hemoglobin-saline
medium.
Family OXYMONADIDAE
=Oxymonas doroaxostylus= (Cleveland et al.)
_Synonymy._--_Saccinobaculus doroaxostylus_ Cleveland et al. [Cleveland,
1950].
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934).
=Oxymonas nana= Cleveland
_Synonymy._--_Saccinobaculus minor_ Cleveland et al. [Cleveland, 1950].
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934).
Family DINENYMPHIDAE
=Saccinobaculus ambloaxostylus= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934).
=Saccinobaculus lata= Cleveland
_Natural host._--_Cryptocercus punctulatus_, U.S.A. (Cleveland, 1950b):
There are at least two other species of _Saccinobaculus_ in _C.
punctulatus_ that have not been described.
=Notila proteus= Cleveland
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland, 1950b).
Family TRICHOMONADIDAE
* =Trichomonas hominis= (Davaine)
_Experimental vectors._--_Blatta orientalis_, South Africa (Porter,
1918); Italy (Mariani and Besta, 1936).
_Periplaneta americana_, South Africa (Porter, 1918); U.S.A. (Hegner,
1928).
=Trichomonas= sp.
_Natural vector._--Cockroach, Venezuela (Tejera, 1926): Organism found
in digestive tract of the cockroach.
Family HEXAMITIDAE
=Hexamita cryptocerci= Cleveland et al.
_Natural hosts._--_Cryptocercus punctulatus_, U.S.A. (Cleveland et al.,
1934).
_Panesthia angustipennis_, Philippine Islands (Kidder, 1937).
=Hexamita periplanetae= (Bĕlăr)
_Synonymy._--_Octomitus periplanetae_ Bĕlăr [Kudo, 1954].
_Natural hosts._--_Blatta orientalis_, U.S.S.R. (Zasukhin, 1930):
Organism is found in the hind gut. Eighty-five percent of over 3,000 _B.
orientalis_ contained this organism.
_Periplaneta americana_, Philippine Islands (Hegner and Chu, 1930).
"Küchenschaben," Austria (Bĕlăr, 1916).
=Hexamita= sp.?
_Natural host._--_Leucophaea maderae_, Philippine Islands (Hegner and
Chu, 1930): The flagellates were present in large numbers.
* =Giardia intestinalis= (Lambl)
_Experimental vectors._--_Blatta orientalis_, South Africa (Porter,
1918).
_Blattella germanica_, Brazil (Pessôa and Corrêa, 1927).
_Eurycotis floridana_, U.S.A. (Young, 1937).
_Leucophaea maderae_, Brazil (Pessôa and Corrêa, 1927).
_Periplaneta americana_, South Africa (Porter, 1918); Gold Coast Colony
(Macfie, 1922); Brazil (Pessôa and Corrêa, 1927); U.S.A. (Young, 1937).
_Periplaneta brunnea_, U.S.A. (Young, 1937).
Cockroaches, Venezuela (Tejera, 1926); Argentina (Bacigalupo, in Tejera,
1926).
* =Giardia= sp.
_Natural vectors._--Cockroaches, Venezuela (Tejera, 1926).
Order HYPERMASTIGINA
Family HOLOMASTIGOTIDAE
=Leptospironympha eupora= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian area
(Cleveland et al., 1934).
=Leptospironympha rudis= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian area
(Cleveland et al., 1934).
=Leptospironympha wachula= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian area
(Cleveland et al., 1934).
=Macrospironympha xylopletha= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian area
(Cleveland et al., 1934).
Family LOPHOMONADIDAE
=Lophomonas blattarum= Stein
_Natural hosts._--_Blatta orientalis_, Czechoslovakia (Stein, 1860);
Germany (Bütschli, 1878; Schubotz, 1905; Chen, 1933); U.S.A. (Leidy,
1879a; Kudo, 1922, 1925, 1926, 1926b; McAdow, 1931); Europe (Janicki,
1908); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1930); Poland
(Lorenc, 1939).
_Blattella germanica_, U.S.A., Ohio (McAdow, 1931).
_Blattella germanica_ and/or _Periplaneta americana_, Egypt (DeCoursey
and Otto, 1956, 1957).
_Periplaneta americana_, England (Schuster, 1898); Europe (Janicki,
1910); U.S.A. (Kudo, 1926b; McAdow, 1931; Hatcher, 1939; Armer, 1944);
Philippine Islands (Hegner and Chu, 1930).
_Periplaneta_ sp., Goa (Mello and Lima Ribeiro, 1924, 1925).
"Küchenschaben," Austria (Bělǎr, 1916).
The protozoan (fig. 2, E) is found in the host's colon, particularly
anterior portion; encysted stages of organism are found throughout hind
gut. Of 1,400 _B. orientalis_ studied, 32 percent harbored this organism
(Kudo, 1925, 1926). Yakimov and Miller (1922) found 7 percent of 124
_B. orientalis_ infested. Zasukhin (1930) found 10 percent of over 3,000
_B. orientalis_ infested. The flagellate does not harm the host and is
never present in the host tissue; it should be considered a commensal
(Kudo, 1926).
=Lophomonas striata= Bütschli
_Synonymy._--_Lophomonas sulcata_ Schuster is most probably identical
with _L. striata_ (Kudo, 1926b).
_Natural hosts._--_Blatta orientalis_, Germany (Bütschli, 1878;
Schubotz, 1905); Europe (Janicki, 1908, 1910); U.S.A. (Kudo, 1922, 1926,
1926b; McAdow, 1931); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin,
1930); Poland (Lorenc, 1939).
_Blattella germanica_, U.S.A., Ohio (McAdow, 1931).
_Blattella germanica_ and/or _Periplaneta americana_, South Africa
(Porter, 1930).
_Periplaneta americana_, Indochina (Weill, 1929); Philippine Islands
(Hegner and Chu, 1930); U.S.A. (Kudo, 1926b; McAdow, 1931; Armer, 1944).
Cockroach, Venezuela (Tejera, 1926); England or U.S.A.? (Lucas, 1928).
"Küchenschaben," Austria (Bělǎr, 1916).
Found in the host's colon, particularly the anterior portion. _L.
striata_ (fig. 2, D) was found in 29 percent of 1,400 _B. orientalis_
and in 2 of 30 _P. americana_ (Kudo, 1926, 1926b). Yakimov and Miller
(1922) found the organism in 9.6 percent of 124 specimens of _B.
orientalis_. Zasukhin (1930) found 8.6 percent of over 3,000 _B.
orientalis_ infested.
Grassé (1926, 1926a) identified corrugations on the surface of _L.
striata_ as a bacterial parasite which he named _Fusiformis
lophomonadis_.
=Prolophomonas tocopola= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, California, Oregon,
Virginia, West Virginia (Cleveland et al., 1934): Not abundant.
Family HOPLONYMPHIDAE
=Barbulanympha estaboga= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934).
_Barbulanympha coahoma_ (Cleveland et al., 1934) represents the diploid
form of _B. estaboga_ (Cleveland, 1953).
=Barbulanympha laurabuda= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934): This species, _B.
ufalula_, and _Rhynchonympha tarda_ occur in all parts of the colon,
especially in the enlarged, flexed part near the ileum.
=Barbulanympha ufalula= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934).
=Barbulanympha wenyoni= Cleveland
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Pacific coast area
(Cleveland, 1953).
=Rhynchonympha tarda= Cleveland et al.
(Fig. 3, D)
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Pacific coast area
(Cleveland et al., 1934): Fairly abundant in every specimen examined
from Pacific coast.
=Urinympha talea= Cleveland et al.
(Fig. 3, C)
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934): Present in fairly great
numbers in every cockroach examined.
Family STAUROJOENINIDAE
=Idionympha perissa= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian area
(Cleveland et al., 1934): Present in only a few specimens.
Family TRICHONYMPHIDAE
=Trichonympha acuta= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934).
=Trichonympha algoa= Cleveland et al.
(Fig. 3, E)
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934): Fairly abundant and
present in most specimens.
[Illustration: FIG. 3.--Protozoa from the gut of the wood-feeding
cockroach _Cryptocercus punctulatus_. A, _Eucomonympha imla_, female
above, male below, c. × 375. (From Cleveland [1950c].) B,
_Barbulanympha_ sp. (From Cleveland [1953].) C, _Urinympha talea_,
c. × 712. (From Cleveland [1951a].) D, _Rhynchonympha tarda_,
c. × 450. (From Cleveland [1952].) E, _Trichonympha okolona_ or
_T. algoa_, c. × 390.
(From Cleveland [1949].) (All drawings reproduced through the courtesy
of Dr. L. R. Cleveland.)]
=Trichonympha chula= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934).
=Trichonympha grandis= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Pacific coast areas
(Cleveland et al., 1934): Fairly abundant in all specimens from Pacific
area.
=Trichonympha lata= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934).
=Trichonympha okolona= Cleveland et al.
(Fig. 3, E)
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934): Found in only a few
specimens, never abundant.
=Trichonympha parva= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934): This organism is smaller
than any known species of _Trichonympha_; it is more resistant to warm
weather than the other hypermastigotes.
Family EUCOMONYMPHIDAE
=Eucomonympha imla= Cleveland et al.
_Natural host._--_Cryptocercus punctulatus_, U.S.A., Appalachian and
Pacific coast areas (Cleveland et al., 1934): Organism (fig. 3, A)
sometimes becomes attached to the intestinal wall; attached individuals
were seen in 2 to 3 percent of the cockroaches examined.
=Unidentified flagellate=
_Natural host._--_Pycnoscelus surinamensis_, Hawaii (Schwabe, 1950): A
small flagellate was found in the digestive tract and malpighian
tubules.
Class SARCODINA
Order MYCETOZOA
INCERTAE SEDIS
=Peltomyces periplanetae= (Léger)
_Synonymy._--_Peltomyces blattellae._ Sprague (1940a) synonymizes
_Peltomyces periplanetae_, with _Coelosporidium periplanetae_.
_Natural hosts._--_Blatta orientalis_, France (Debaisieux, 1927).
_Blattella germanica_, France (Léger, 1909; Debaisieux, 1927).
The organism inhabits the malpighian tubules of cockroaches. Léger and
Debaisieux concluded that their organism was a mycetozoan, but they may
have erred in synonymizing _Plistophora periplanetae_ with the organism
they studied. Debaisieux found intracellular stages of _Peltomyces
periplanetae_ that have not been found in _Plistophora periplanetae_ or
_Coelosporidium periplanetae_.
Order AMOEBINA
Family AMOEBIDAE
=Hartmannella blattae= Ivanić
_Natural host._--_Blatta orientalis_, Yugoslavia (Ivanić, 1937): Found
in the hind gut.
Family ENDAMOEBIDAE
In the following classification we have accepted the conclusions of
Kirby (1945), Kudo (1954), and others that species of _Endamoeba_ are
generically different from species of _Entamoeba_ and that the latter
genus is not a homonym of _Endamoeba._
=Dobellina= sp.
_Natural vectors._--_Blattella germanica_ and/or _Periplaneta
americana_, Egypt (DeCoursey and Otto, 1956, 1957): Thirty out of 261
cockroaches examined contained this protozoan.
=Endamoeba blattae= (Bütschli)
_Synonymy._--_Amoeba blattae_, _Entamoeba blattae_, _Entamoeba
blattarum_.
_Natural hosts._--_Blatta orientalis_, Germany (Bütschli, 1878;
Schubotz, 1905; Chen, 1933); U.S.A. (Leidy, 1879a, 1880; Kudo, 1922,
1925a, 1926a; Kirby, 1927; McAdow, 1931; Meglitsch, 1938, 1940); France
(Mercier, 1907a, 1908, 1909, 1910); Europe? (Janicki, 1908, 1909);
U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1929, 1930); England
(Thomson and Lucas, 1926; Lucas, 1927, 1927a, 1928); Yugoslavia (Ivanić,
1926a).
_Blattella germanica_ and/or _Periplaneta americana_, South Africa
(Porter, 1930); Egypt (DeCoursey and Otto, 1956, 1957): Seven out of 217
cockroaches examined harbored the protozoan.
_Periplaneta americana_, Philippine Islands (Hegner and Chu, 1930);
U.S.A. (Morris, 1936; Armer, 1944); Gold Coast Colony (Macfie, 1922).
_Periplaneta australasiae_, U.S.A. (Morris, 1936).
Cockroaches, Paraguay? (Elmassian, 1909); Austria (Bělǎr, 1916); U.S.A.
(Morris, 1935, 1936; Balch, 1932); Venezuela (Tejera, 1926).
The habitat of _E. blattae_ (fig. 2, C) is the hind intestine and rectum
of the cockroach. The incidence of infection varies: Kudo (1925a) found
in 1,255 oriental cockroaches infections in 5 percent in March and 50
percent in the summer; Schubotz (1905) found 5 to 20 percent of the
examined cockroaches to be infested; Yakimov and Miller (1922) found 4
percent of 124 oriental cockroaches infested; Zasukhin (1930) found up
to 50 percent of over 3,000 _B. orientalis_ infested; Meglitsch (1938,
1940) found almost 100 percent infection in _B. orientalis_ kept in a
crowded culture for several weeks. Chen (1933) developed two synthetic
media in which _E. blattae_ could be grown for 45 to 50 days.
Mercier (1907a) observed a fungus, _Nucleophaga_ sp., hyperparasitic in
the nucleus of _Endamoeba blattae_.
=Endamoeba javanica= Kidder
_Natural hosts._--_Panesthia angustipennis_, Philippine Islands, and
_Panesthia spadica_, Japan (Kidder, 1937): Occurred in 50 percent of _P.
angustipennis_ examined and in one of four _P. spadica_. The endoplasm
of this amoeba contains large amounts of wood and cellulose fibers.
=Endamoeba philippinensis= Kidder
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Kidder,
1937): Occurred in about 10 percent of the _Panesthia_ examined. The
food vacuoles contained bacteria, no wood.
=Entamoeba coli= (Grassi)
_Synonymy._--_Endamoeba coli_, _Amoeba coli_ [Kirby, 1945].
_Natural vectors._--_Blaberus atropos_, Venezuela (Tejera, 1926): In a
lot of 60 cockroaches captured in latrines, two were found that carried
apparently live cysts similar to cysts of _E. coli_.
_Blattella germanica_ or _Periplaneta americana_, Egypt (DeCoursey and
Otto, 1956, 1957): One out of 44 cockroaches collected in a village
harbored _E. coli_.
_Experimental vector._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922): In nine experiments cysts of _E. coli_ were fed to the
cockroaches. In seven of the experiments cysts of _E. coli_ were found
in the feces. Cysts were observed in the feces for only one to three
days, and eventually disappeared completely. The cysts appeared to be
unharmed. No amoebae were found.
* =Entamoeba histolytica= Schaudinn
_Natural vectors._--_Blatta orientalis_, _Blattella germanica_,
_Periplaneta americana_, _Periplaneta australasiae_, and/or _Supella
supellectilium_, Peru (Schneider and Shields, 1947).
_Blattella germanica_ and/or _Periplaneta americana_, Egypt (DeCoursey
and Otto, 1956, 1957).
Cockroaches, Venezuela (Tejera, 1926).
_Experimental vectors._--_Blatta orientalis_, Italian Somaliland
(Mariani and Besta, 1936).
_Periplaneta americana_, Gold Coast Colony (Macfie, 1922); U.S.A. (Frye
and Meleney, 1936).
Cockroaches, Venezuela (Tejera, 1926).
=Entamoeba pitheci= Prowazek?
_Experimental vector._--_Periplaneta americana_, Formosa (Morischita and
Tsuchimochi, 1926): Eleven of 15 cockroaches fed feces of a monkey
[_Macaca cyclopis_ (Swinhoe)] containing cysts of the amoeba voided live
cysts in their own feces.
=Entamoeba thomsoni= Lucas
_Synonymy._--_Endamoeba thomsoni_ [Kudo, personal communication, 1957].
_Natural hosts._--_Blatta orientalis_, England (Lucas, 1927a, 1928);
U.S.A. (Taliaferro, 1928; McAdow, 1931); U.S.S.R. (Zasukhin, 1930);
Germany (Chen, 1933).
_Blattella germanica_, U.S.A. (McAdow, 1931).
_Periplaneta americana_, England (Lucas, 1927a); U.S.A. (Smith and
Barret, 1928; McAdow, 1931); Philippine Islands (Hegner and Chu, 1930).
The organism is found in the hind intestine and rectum of the cockroach.
Smith and Barret (1928) developed a synthetic medium in which cultures
of _E. thomsoni_ were carried through successive transfers for 24
months.
=Entamoeba= sp.
_Natural vector._--_Periplaneta americana_, Gold Coast Colony (Macfie,
1922): Under the heading "_Entamoeba histolytica_ and _E. coli_" Macfie
(p. 445) stated, "The cockroaches used in these experiments had
previously been carefully examined for amoebic infections a precaution
which was doubly necessary, because some of these insects at Accra had
been found naturally infected."
_Experimental vectors._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922): _Entamoeba_, resembling _E. coli_, from feces of the
monkey [_Erythrocebus patas patas_ (Schreber)] were fed to cockroaches,
and on the second to fourth days thereafter apparently healthy cysts
were recovered in the cockroach feces.
=Endolimax blattae= Lucas
_Natural hosts._--_Blatta orientalis_, England (Lucas, 1927, 1927a);
U.S.S.R. (Zasukhin, 1930); Germany (Chen, 1933).
_Periplaneta americana_, England (Lucas, 1927, 1927a); Indochina (Weill,
1929); U.S.A. (Armer, 1944).
_Periplaneta australasiae_, U.S.A. (Steinhaus, 1946).
Organism is found in the hind gut of the cockroach. Zasukhin (1930)
found 3-percent infestation in over 3,000 _B. orientalis_ examined.
=Endolimax nana= (Wenyon and O'Connor)?
_Synonymy._--_Entamoeba nana._
_Natural host._--_Blaberus atropos_, Venezuela (Tejera, 1926): A small
amoeba greatly resembling _E. nana_ was found in the intestinal contents
of the cockroach. In inoculations this amoeba was not pathogenic.
=Endolimax= sp.
_Natural hosts._--_Blatta orientalis_, U.S.S.R. (Zasukhin, 1930): This
organism was found in the hind gut of 0.3 percent of over 3,000
cockroaches examined.
_Blattella germanica_ and/or _Periplaneta americana_, Egypt (DeCoursey
and Otto, 1956, 1957): Seventy-four out of 261 cockroaches examined
harbored this protozoan.
=Iodamoeba= sp.
_Natural vectors._--_Blattella garmanica_ and/or _Periplaneta
americana_, Egypt (DeCoursey and Otto, 1956, 1957): Fifty-nine of 261
cockroaches examined contained this protozoan. _Iodamoeba_ sp. was
common in human feces in villages in which the cockroaches were
collected.
=Undetermined species of Amoeba=
_Natural host._--_Panesthia angustipennis_, Philippine Islands. (Kidder,
1937): Found in only one specimen.
Class SPOROZOA
Order GREGARINIDA
Family DIPLOCYSTIDAE
=Diplocystis schneideri= Kunstler
(Fig. 2, I)
_Natural hosts._--_Blatta orientalis_, England (Woodcock, 1904; Jameson,
1920).
=Periplaneta americana=, France (Kunstler, 1884, 1887); England
(Jameson, 1920); Germany (Foerster, 1939).
In body cavity of host. Cysts containing spores are ingested during
cannibalistic feeding on infected cockroaches. Sporozoites penetrate the
gut wall which later ruptures, freeing the gregarines into the coelom.
There is no apparent pathogenic effect. Jameson (1920) found 81 percent
of _P. americana_ infested with _D. schneideri_.
=Diplocystis= sp.
_Natural host._--_Periplaneta americana_, U.S.A. (Hertig, 1921): Heavy
infections in body cavity.
Cockroach, India (Ray and Dasgupta, 1955): A large number of
cockroaches, both adults and nymphs, collected in Calcutta were all
infected.
=Diplocystis= sp.?
_Natural host._--_Blaberus craniifer_, U.S.A. (Nutting, 1953): From 1 to
12 or more paired trophozoites or cysts may be found in the hemocoele
and occasionally in the thorax.
Family STENOPHORIDAE
=Stenophora= sp.
_Natural host._--_Blatta orientalis_, India (Bal and Rai, 1955): Organism
found in the midgut of the cockroach.
Family GREGARINIDAE
=Gregarina blattarum= von Siebold
_Synonymy._--_Gregarina blattae orientalis_; _Clepsidrina blattarum_.
_Natural hosts._--_Blatta orientalis_, Germany (Siebold, 1837, 1839;
Stein, 1848; Bütschli, 1881; Wolters, 1891; Marshall, 1892; Wellmer,
1910, 1911; Foerster, 1938; Schubotz, 1905); U.S.A. (Leidy, 1853a;
Crawley, 1903; Watson, 1917; Kudo, 1922; McAdow, 1931; Sprague, 1940,
1941); England (Lankester, 1863); France (Schneider, 1875; Cuénot, 1901;
Laveran and Franchini, 1920a); Brazil (Magalhães, 1900; Pinto, 1919);
U.S.S.R. (Zasukhin, 1929, 1930).
_Blattella germanica_, U.S.A. (Crawley, 1903); South Africa (Fantham,
1929; Porter, 1930: these appear to be the same record).
_Blattella germanica_ and/or _Periplaneta americana_, Egypt (DeCoursey
and Otto, 1956, 1957).
_Periplaneta americana_, Brazil (Magalhães, 1900); U.S.A. (Crawley,
1903; 1907; McAdow, 1931); South Africa (Fantham, 1929; Porter, 1930:
these appear to be the same record); Gold Coast Colony (Macfie, 1922).
_Parcoblatta pensylvanica_, U.S.A., Michigan (Ellis, 1913a).
Cockroaches, Germany (Schiffmann, 1919: probably used the oriental
cockroach); Venezuela (Tejera, 1926).
Organism usually found in the intestinal tract of cockroaches where it
is attached to the gut cells. Cysts are passed in the feces.
Occasionally, _G. blattarum_ (fig. 2, J) is found in the body cavity
(Leidy, 1853a; Hall, 1907). Though this is considered to be one of the
commonest of the Sporozoa encountered in cockroaches, DeCoursey and Otto
(1956) found only 10 of 217 _P. americana_ and _B. germanica_, collected
in restaurants in Egypt, infested with this species. Watson (1917) found
a dozen or more in one specimen of _Blatta orientalis_. Zasukhin (1929,
1930) found 2.6 percent of 3,000 oriental cockroaches infected with this
parasite.
=Gregarina fastidiosa= Harrison
_Natural host._--_Aptera fusca_, South Africa (Harrison, 1955): All
mature females were heavily infected; in all specimens there were over
100 parasites in the gut. All nymphs were infected, the earlier instars
more lightly than the later instars. Gregarines were found in all parts
of the gut except the crop and gizzard.
=Gregarina gibbsi= Harrison
_Natural host._--_Temnopteryx phalerata_, South Africa (Harrison, 1955):
Although the cockroaches were found in groups, only 32 percent were
infected and only 10 percent heavily. The gregarines were found in the
anterior mesenteron but none in the hepatic caeca. All cysts were found
in the hind gut or rectum.
=Gregarina illinensis= M. E. Watson
_Natural host._--_Parcoblatta pensylvanica_, U.S.A., Illinois (Watson,
1915, 1916): The intestine of one cockroach was found to contain 25 of
these gregarines.
=Gregarina impetuosa= Harrison
_Natural host._--_Melanosilpha capensis_, South Africa (Harrison, 1955):
All specimens of this gregarine were found in the anterior mesenteron of
the host.
=Gregarina légeri= Pinto
_Natural host._--_Periplaneta americana_, Brazil (Pinto, 1918, 1918a,
1919): Intestinal canal.
=Gregarina neo-brasiliensis= Al. Cunha
_Natural host._--_Periplaneta americana_, Brazil (R. de Almeida Cunha
_in_ Pinto, 1919; Cunha, 1919).
=Gregarina ohioensis= Semans
_Natural host._--_Parcoblatta virginica_, U.S.A., Ohio (Semans, 1939):
The protozoan was present in large numbers in the insect's midgut.
=Gregarina panchlorae= Frenzel
_Natural host._--_Panchlora exoleta_, Argentina (Frenzel, 1892): Midgut.
=Gregarina parcoblattae= Semans
_Natural hosts._--_Parcoblatta pensylvanica_ and _Parcoblatta
uhleriana_, U.S.A., Ohio (Semans, 1939): Midgut.
=Gregarina rhyparobiae= J. M. Watson
_Natural host._--_Leucophaea maderae_, Uganda (Watson, 1945): Midgut.
Trophozoites could be seen in sections attached to cells of the
intestinal wall (fig. 2, H).
=Gregarina sandoni= Harrison
_Natural host._--_Melanosilpha capensis_, South Africa (Harrison, 1955):
This gregarine was found in the anterior and middle parts of the
mesenteron and in the hepatic caeca.
=Gregarina thomasi= Semans
_Natural host._--_Parcoblatta pensylvanica_, U.S.A., Ohio (Semans,
1939): Enteric caeca and midgut.
=Protomagalhaesia serpentula= (de Magalhães)
_Synonymy._--_Gregarina serpentula_ [Pinto, 1918a, 1919; Semans, 1943].
_Natural host._--_Periplaneta americana_, Brazil (Magalhães, 1900): In
the coelom and alimentary canal. The host of this parasite (fig. 2, K)
was incorrectly cited as _Blatta orientalis_ by Watson (1916).
=Gamocystis tenax= Schneider
(Fig. 2, L)
_Natural hosts._--_Ectobius lapponicus_, France (Schneider, 1875);
Germany (Wellmer, 1910, 1911; Foerster, 1938).
_Ectobius pallidus_, Germany (Foerster, 1938): In intestine.
Family ACTINOCEPHALIDAE
=Pileocephalus blaberae= (Frenzel)
_Synonymy._--_Gregarina blaberae_ [Watson, 1916].
_Natural hosts._--_Blaptica dubia_ and related forms, Argentina
(Frenzel, 1892): In midgut.
=Unidentified Gregarinida=
_Natural hosts._--_Blaberus craniifer_, U.S.A. (Roth and Willis,
unpublished data, 1953): Possibly _Diplocystis_ sp. (pl. 28, A, B).
_Cryptocercus punctulatus_, U.S.A. (Cleveland et al., 1934).
_Leucophaea maderae_, Philippine Islands (Hegner and Chu, 1930): In
intestines of host. U.S.A. (Roth and Willis, unpublished data, 1958):
Cysts in feces (pl. 28, C).
_Gromphadorhina portentosa_, U.S.A., in laboratory colony (Roth and
Willis, unpublished data, 1958): In intestine of adult female.
_Pycnoscelus surinamensis_, Hawaii (Schwabe, 1950): A cephaline
gregarine was found in the cockroach's digestive tract; it was also
claimed to be present in new-born nymphs.
Order COCCIDIA
Family ADELEIDAE
=Adelina cryptocerci= Yarwood
_Natural host._--_Cryptocercus punctulatus_, U.S.A. (Yarwood, 1937):
This intracellular parasite was found in the fat body in light
infestations. In heavy infections the coccidia were found in the head,
antennae, mouthparts, muscles, legs, salivary glands, nerve cord, as
well as fat body. Infection in freshly collected specimens was about 3
percent; when large numbers of cockroaches were kept together in
culture, the rate of infection increased because the insects ate their
dead companions.
Cleveland et al. (1934) mentioned a coccidium which was sometimes
generally distributed through the body (head, legs, antennae, etc.) of
_C. punctulatus_; this parasite was probably the species described by
Yarwood.
Order HAPLOSPORIDIA
=Haplosporidium periplanetae= Georgévitch
_Natural host._--_Blatta orientalis_, Yugoslavia (Georgévitch, 1953):
This organism was described from the malpighian tubules of the cockroach
where it apparently occurred in a mixed infection with the
microsporidian _Plistophora periplanetae_. See synonymy under
_Plistophora periplanetae_.
=Coelosporidium periplanetae= (Lutz and Splendore)
_Synonymy._--_Nosema periplanetae_, _Coelosporidium blattellae_,
_Bertramia blatellae_ [after Semans, 1943]. Some of the observations
cited under _Plistophora periplanetae_ may pertain to _C. periplanetae_
(see Sprague, 1940). See also _Haplosporidium periplanetae_.
_Natural hosts._--_Blatta orientalis_, U.S.S.R. (Epshtein, 1911); U.S.A.
(Kudo, 1922; Sprague, 1940); Yugoslavia (Ivanić, 1926).
_Blattella germanica_, U.S.A. (Crawley, 1905); Germany (Wellmer, 1910,
1911).
_Periplaneta americana_, Brazil (Lutz and Splendore, 1903).
This organism (fig. 2, B) passes its life cycle living free in the
lumina of the malpighian tubules of cockroaches. The elongate
trophozoite is firmly attached to the wall of the tubule as are clusters
of immature spores. Mature spores are freed into the lumina of the
tubules from whence they pass to the exterior. Sprague (1940) examined
about 200 wild-caught _B. orientalis_ and found them to be practically
100 percent infected.
Order MICROSPORIDIA
Family NOSEMATIDAE
=Plistophora kudoi= Sprague and Ramsey
_Natural host._--_Blatta orientalis_, U.S.A., Illinois, West Virginia,
Kentucky (Sprague and Ramsey, 1941, 1942): Found in the epithelial
cells of caeca and midgut. Considerable damage is done to these cells.
Seventy-five percent of 52 _B. orientalis_ harbored the parasite.
=Plistophora periplanetae= (Lutz and Splendore)
_Synonymy._--_Nosema periplanetae_, _Pleistophora periplanetae_ [after
Semans, 1943]. Georgévitch (1953) has pointed out that one may find in
the malpighian tubules of cockroaches a mixed infection of
Microsporidia, Haplosporidia, and Mycetozoa, and that some of the
discrepancies in the earlier literature may be attributed to attempts to
combine in one organism disparate stages belonging to different orders.
See also comments under _Coelosporidium periplanetae_, _Haplosporidium
periplanetae_, and _Peltomyces periplanetae_.
_Natural hosts._--_Blatta orientalis_, France (Mercier, 1906a;
Debaisieux, 1927); England (Perrin, 1906, 1906b); U.S.S.R. (Zhivago,
1909); Yugoslavia (Georgévitch, 1925, 1926, 1926a, 1927); Germany
(Wellmer, 1910, 1911).
_Blattella germanica_, France (Léger, 1909; Debaisieux, 1927); U.S.S.R.
(Zhivago, 1909).
_Periplaneta americana_, Brazil (Lutz and Splendore, 1903).
This organism lives in the lumen of the malpighian tubules of
cockroaches. The cited authors appear to have been convinced that this
organism was a microsporidian. Georgévitch (1927, 1953) described the
polar capsule and filament characteristic of this order.
=Plistophora= sp.
_Natural host._--_Blatta orientalis_, France (Mercier, 1908a): The
organism parasitized the fat body of the cockroach. Mitoses, often
abnormal, were induced in the fat cells. Infected cockroaches were
easily recognizable by their distended abdomens. The fat body became
chalky white and showed through the intersegmental membranes.
Porter (1930) reported finding an unidentified microsporidian in the fat
bodies of _Blattella germanica_ and _Periplaneta americana_ collected in
South Africa. It may or may not have been a species of _Plistophora_.
Class CILIATA
Order HOLOTRICHA
Family PARAMECIIDAE
=Paramecium= sp.
_Natural associate._--Cockroaches, U.S.A., Maryland (Cleveland, 1927):
Three of 30 cockroaches collected in the basement of a department store
had paramecia in their stomachs but none in the rectum.
_Experimental associate._--_Periplaneta americana_, U.S.A. (Hegner,
1929): Paramecia fed to the cockroaches were recovered from the crop at
intervals from one-half to six and one-half hours. In no case were the
protozoa recovered from the stomach alive.
Cockroaches, U.S.A., Maryland (Cleveland, 1927). About 200 starved
cockroaches were fed a culture of _Paramecium_. Few, if any, of the
protozoa were killed in the stomach during the first two hours, but all
were killed within 5 to 6 hours after ingestion.
Family ISOTRICHIDAE
=Isotricha caulleryi= Weill
_Natural host._--_Periplaneta americana_, Indochina (Weill, 1929):
Alimentary canal.
Order SPIROTRICHA
Family BURSARIIDAE
=Balantidium blattarum= Ghosh
_Natural host._--_Periplaneta americana_, India (Ghosh, 1922; Bhatia and
Gulati, 1927); Gold Coast Colony (Macfie, 1922): Intestinal tract.
* =Balantidium coli= (Malmsten)
_Experimental vector._--Cockroach, Venezuela (Tejera, 1926).
=Balantidium ovatum= Ghosh
_Natural host._--_Periplaneta americana_, India (Ghosh, 1922a; Bhatia
and Gulati, 1927); Indochina, Saigon (Weill, 1929): Intestinal tract.
=Balantidium praenucleatum= Kudo and Meglitsch
_Natural host._--_Blatta orientalis_, U.S.A., Illinois (Kudo and
Meglitsch, 1938; Meglitsch, 1940): This protozoan is found in the lumen
of the anterior region of the colon in association with several other
species of protozoa. Only 7.6 percent of 500 cockroaches examined
contained _B. praenucleatum_. The largest number encountered in a single
host was 59, but as a rule each host harbored a smaller number.
=Balantidium= sp.?
_Natural host._--_Periplaneta americana_, Brazil (Magalhães, 1900):
These organisms were numerous in the intestine.
Family SPIROSTOMIDAE
=Nyctotherus buissoni= Pinto
_Natural host._--"Barata sylvestre," Brazil (Pinto, 1926): Organism
found in the cockroach's intestine.
=Nyctotherus ovalis= Leidy
_Synonymy._--_Bursaria blattarum_; _Plagiotoma blattarum_.
_Natural hosts._--_Blatta orientalis_, U.S.A. (Leidy, 1850, 1853, 1853b,
1879a; Kudo, 1922, 1926, 1936; McAdow, 1931; Kudo and Meglitsch, 1938;
Meglitsch, 1940); Germany (Stein, 1860; Schubotz, 1905; Chen, 1933);
England (Lankester, 1865; Schuster, 1898; Lucas, 1927a, 1928); Spain
(Zulueta, 1916); U.S.S.R. (Yakimov and Miller, 1922; Zasukhin, 1928,
1930; Ostroumov, 1929); Portugal (Lima Ribiero, 1924); Brazil (Pinto,
1926); Venezuela (Tejera, 1926).
_Blattella germanica_, South Africa (Porter, 1930); U.S.A. (Balch, 1932;
McAdow, 1931).
_Parcoblatta pensylvanica_, U.S.A. (Semans, 1939, 1941).
_Periplaneta americana_, India (Bhatia and Gulati, 1927); Indochina
(Weill, 1929); Philippine Islands (Hegner and Chu, 1930); South Africa
(Porter, 1930); U.S.A. (McAdow, 1931; Hatcher, 1939; Meglitsch, 1940;
Armer, 1944); Goa (Mello et al., 1934); China (Pai and Wang, 1947);
Czechoslovakia (Low, 1956).
"Barata sylvestre," Brazil (Pinto, 1926).
"Küchenschaben," Austria (Bělǎr, 1916).
_Nyctotherus ovalis_ (fig. 2, G) inhabits the hind gut of cockroaches,
where it occurs almost always in the anterior half of the colon in
association with other species of Protozoa (Kudo, 1936). Ninety percent
of 500 _B. orientalis_ contained _N. ovalis_ (Kudo and Meglitsch, 1938).
Yakimov and Miller (1922) found _N. ovalis_ in 68 percent of 124 _B.
orientalis_. Zasukhin (1930) found this organism in 63 percent of over
3,000 _B. orientalis_. Zasukhin (1928, 1934) found a fungus and possibly
a bacterium hyperparasitic in the cytoplasm of _N. ovalis_. _N. ovalis_
has been cultured outside the cockroach in several media (Lucas, 1928;
Balch, 1932; Chen, 1933; Low, 1956).
=Nyctotherus uichancoi= Kidder
_Natural hosts._--_Panesthia angustipennis_, Philippine Islands, and
_Panesthia spadica_, Japan (Kidder, 1937): About 90 percent of all _P.
angustipennis_ harbored this ciliate in their hindguts.
=Nyctotherus viannai= Pinto
_Natural host._--"Barata sylvestre," Brazil (Pinto, 1926): In the
intestine of the cockroach.
Family CLEVELANDELLIDAE
Most of the Clevelandellidae are parasitized by rod-shaped or spherical
bacteria-like organisms usually in clusters (Kidder, 1937).
_Synonymy._--Clevelandiidae (Kidder, 1938).
Genus CLEVELANDELLA
_Synonymy._--The generic name _Clevelandia_ Kidder (1937) is
preoccupied; it was therefore changed to _Clevelandella_ by Kidder in
1938. All of the following species of _Clevelandella_ were originally
described as _Clevelandia_.
=Clevelandella constricta= (Kidder)
_Natural hosts._--_Panesthia angustipennis_, Philippine Islands, and
_Panesthia spadica_, Japan (Kidder, 1937): In the posterior end of
hindgut.
=Clevelandella contorta= (Kidder)
_Natural hosts._--_Panesthia angustipennis_, Philippine Islands, and
_Panesthia spadica_, Japan (Kidder, 1937).
=Clevelandella elongata= (Kidder)
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Kidder,
1937).
=Clevelandella hastula= (Kidder)
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Kidder,
1937): Common in hindgut.
=Clevelandella nipponensis= (Kidder)
_Natural host._--_Panesthia spadica_, Japan (Kidder, 1937).
=Clevelandella panesthiae= (Kidder)
_Natural hosts._--_Panesthia angustipennis_, Philippine Islands, and
_Panesthia spadica_, Japan (Kidder, 1937): In the hindgut. This
protozoan is commonly parasitized by the microorganism _Sphaerita_.
=Clevelandella parapanesthiae= (Kidder)
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Kidder,
1937).
=Paraclevelandia brevis= Kidder
_Natural hosts._--_Panesthia angustipennis_, Philippine Islands, and
_Panesthia spadica_, Japan (Kidder, 1937): Present in 100 percent of _P.
angustipennis_ and in nearly all _P. spadica_.
=Paraclevelandia simplex= Kidder
_Natural hosts._--_Panesthia angustipennis_, Philippine Islands (Kidder,
1937, 1938): Incidence of infection about 50 percent.
_Panesthia spadica_, Japan (Kidder, 1937).
=Unidentified ciliate=
_Natural host._--_Pycnoscelus surinamensis_, Hawaii (Schwabe, 1950): A
large ciliate was found in the digestive tract and malpighian tubules.
NEGATIVE FINDINGS
In a recent experimental study Schmidtke (1955) failed to demonstrate a
host-parasite relationship between _Periplaneta americana_ and the
haemosporidian _Toxoplasma gondii_ Nicolle and Manceaux. This protozoan
is a blood parasite in a rodent in North Africa (Kudo, 1954).
XI. HELMINTHS
Intestinal nematodes of the family Thelastomatidae have no apparent
pathological effect on their cockroach hosts. Galeb (1878) has shown
experimentally that oxyurids eat the same food as the host insect and
that if one starves them, by withholding food from the host, the
oxyurids die and disappear. In other words, these worms are not
parasites, in the sense that we use the term in this paper, but
commensals. Dobrovolny and Ackert (1934) stated that "all observations
seemed to indicate that the health, fertility and activity of the
heavily infested cockroaches were comparable with those of the
non-parasitised specimens."
Very few papers have dealt with the ecology of the oxyurid parasites of
cockroaches. According to Galeb (1878), usually one species of nematode
is found in a single cockroach, but sometimes two species live together
in the same host (e.g., in _Blatta orientalis_ and _Polyphaga
aegyptiaca_) where they compete for food. Galeb claimed that
_Hammerschmidtiella diesingi_ would replace _Leidynema appendiculata_;
he observed that _H. diesingi_ surpassed _L. appendiculata_ in numbers
and the latter became uncommon in the intestines of the cockroaches. On
the other hand, Sobolev (1937) found that 48 percent of his oriental
cockroaches were infected with both of the above species of nematodes.
The average number of both species was 7.5, and the maximum number was
97; the mean number of _H. diesingi_ was 5.1 and the maximum 64; the
mean number of _L. appendiculata_ was 2.4 and the maximum 33. More than
40 nematodes were found in each cockroach of 1.3 percent of those
examined. These results apparently contradict Galeb's conclusions
inasmuch as the number of each species in mixed infections was
essentially the same as the number found in cockroaches infected by only
one species (see pp. 195 and 197). Dobrovolny and Ackert (1934) found
that 29 percent of 222 _Periplaneta americana_ contained both of the
above species of nematodes; whereas 40 percent contained _L.
appendiculata_ only, and 21 percent contained _H. diesingi_ only. The
infestation ranged from 1 to 36 worms per cockroach with averages of 3.8
per male, 5.1 per female, and 2.7 per nymph.
The eggs of some helminths pass unharmed through the guts of cockroaches
that serve as vectors of these ova and have no effect on the insect.
However, helminths that are secondary parasites in cockroaches damage
the insect to varying degrees depending upon the extent of the
infection. Thus the larvae of _Moniliformis moniliformis_ pass through
the gut wall and some may become embedded in the fat tissue (Moore,
1946). First stage larvae of _Oxyspirura mansoni_ also burrow through
the midgut wall into the fat body; Sanders (1929) believed that
_Pycnoscelus surinamensis_ could be killed if at one time a sufficient
number of migrating larvae of _O. mansoni_ penetrated the cockroach's
intestinal wall. _Gongylonema neoplasticum_ migrates through the
digestive tract and encysts in the muscles of the thorax and legs of the
host (Fibiger and Ditlevsen, 1914). Infective larvae of _Protospirura
muricola_, after hatching from ingested eggs, pass through the
cockroach's gut wall and encyst mainly in the thorax, around the crop,
and at the bases of the large muscles of the prothoracic legs (Foster
and Johnson, 1939). It is probably generally true that nematodes which
are secondary parasites in cockroaches do some damage to the host's
intestinal tract at least, and they probably also damage other organs in
which they may encyst.
Cockroach tissues may react defensively to infections by parasitic
nematodes. For example, encysted third-stage larvae of _Physaloptera
turgida_ have been found enclosed in a thin, brown, chitinous substance
that was undoubtedly deposited by the tissue of the cockroach (Alicata,
1937). Cysts of similar appearance have been found in cockroaches
infected with _Physaloptera rara_, _P. maxillaris_, _P. hispida_ (Petri,
1950; Hobmaier, 1941; Schell, 1952), and _Gongylonema pulchrum_
(Schell, 1952a); in the latter species the deposit eventually completely
surrounded the nematode larva which was killed and "chitinized."
Apparently these pigmented cysts surround unhealthy or dead larvae and
are secreted as a defensive mechanism by the host (Schell, 1952a).
Oswald (1958) has reported finding similar pigmented cysts in _Blatta
orientalis_ and _Periplaneta americana_ that were experimentally
infected with _Rictularia coloradensis_.
Our classification of the helminths follows Hyman (1951, 1951a).
HELMINTHS FOR WHICH COCKROACHES SERVE AS PRIMARY HOSTS
Phylum ASCHELMINTHES
Class NEMATODA
Order MERMITHOIDEA
Family MERMITHIDAE
=Undetermined mermithids=
_Natural Hosts._--_Ectobius pallidus_, U.S.A., Plymouth, Massachusetts
(Roth and Willis, 1957): This mermithid lies coiled in the body cavity
of the host and one end may extend into the thorax. Apparently, the host
is eventually killed and the worms may leave the cockroach ventrally
between the thorax and abdomen (pl. 29, A) or thorax and head.
_Periplaneta americana_, Germany (Bode, 1936): Attacked by "_Mermis_" or
"_Gordius_." It has been suggested that the name _Mermis_ is often
applied without critical identification to immature Nematoda found in
insects (Buxton, 1955).
Order RHABDITOIDEA
Family DIPLOGASTERIDAE
=Diplogaster= sp.
_Synonymy._--_Lycolaimus_ [Goodey, 1951].
_Experimental Host._--_Blattella germanica_, U.S.A. (Christie and
Crossman, 1933).
Family STEINERNEMATIDAE
=Neoaplectana= sp.
_Experimental hosts._--_Blattella germanica_, _Nauphoeta cinerea_, and
_Periplaneta americana_. U.S.A. (Dutky and Hough, 1955): This nematode,
found in codling moth larvae, is close to _Neoaplectana chresima_
Steiner but apparently is a new species. _Nauphoeta cinerea_ was very
susceptible to infection; _B. germanica_ and _P. americana_ were less
susceptible.
Order OXYUROIDEA
Family THELASTOMATIDAE
These nematodes are found in the intestinal tract of cockroaches.
=Aorurus philippinensis= Chitwood and Chitwood, 1934
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Chitwood
and Chitwood, 1934).
=Binema mirzaia= (Basir, 1940) Basir, 1956
_Synonymy._--_Periplaneticola mirzaia_ Basir, 1940.
_Natural host._--_Periplaneta americana_, India, Aligarh (Basir, 1940).
=Blattelicola blattelicola= Basir, 1940
_Natural host._--_Blattella germanica_, India, Aligarh (Basir, 1940).
=Blatticola blattae= (Graeffe, 1860) Chitwood, 1932
_Synonymy._--_Oxyuris blattae_ Graeffe, 1860; _Oxyuris blatticola_
Galeb, 1878; _Blatticola blatticola_ (Galeb, 1877) Schwenck, 1926
[Chitwood, 1930, 1932].
_Natural hosts._--_Blattella germanica_, Brazil (Pessôa and Corrêa,
1926; Schwenck, 1926); U.S.A. (Chitwood, 1930; Bozeman, 1942); Egypt?
(Galeb, 1877, 1878); U.S.S.R. (Sobolev, 1937; Sondak, 1935);
Czechoslovakia (Groschaft, 1956).
_Ectobius lapponicus_, _Ectobius pallidus_, Egypt? (Galeb, 1877, 1878).
_Polyphaga aegyptiaca_, France (Graeffe, 1860).
The life cycle has been studied by Bozeman (1942): He found never more
than four worms in the large intestine of each cockroach. Embryos
developed to "resting" stage in vitro. The resting stage was infective
while the active stage was not. The worms seemed to have no effect on
the vital activities of the host. Alicata (1934b) found that the embryo
undergoes a molt before hatching.
Chitwood (1930) found 75 percent of the German cockroaches examined from
houses in Washington infected. As a rule, one adult female, one or two
males, and possibly two larval females are found in a single individual,
apparently only in the rectum.
Sobolev (1937) found 72 percent of _Blattella germanica_ collected in
Gorkov (U.S.S.R.) infected with _Blatticola blattae_. The mean number of
worms per host was 1.8, the maximum 5. Sondak (1935) found about 30
percent of 788 _B. germanica_ collected in Leningrad to be infected with
_B. blattae_. Groschaft (1956) regularly found only single worms in _B.
germanica_, collected in a laboratory in Prague, except for two females
that contained 2 and 3 worms each.
=Blattophila sphaerolaima= Cobb, 1920
_Synonymy._--_Aorurus sphaerolaima_ (Cobb, 1920) Travassos, 1929.
Although Chitwood (1932) indicated that the taxonomic position of this
nematode is questionable, Chitwood and Chitwood, 1934, apparently
accepted it as a valid species in describing the variety _javanica_.
_Natural host._--_Panesthia laevicollis_ [Cobb recorded the host as
_Panesthia brevicollis_, but no such cockroach exists. Van Zwaluwenburg
(1928) and Caudell (in Chitwood, 1932) believed that Cobb meant
_Panesthia laevicollis_. According to Gurney (personal communication,
1957) Caudell's notes show that in 1933 he wrote to Dr. Chitwood and
explained that he had compared Cobb's figure of the cockroach with
_laevicollis_ Saussure (figures and description) and had found them the
same.] Australia, New South Wales (Cobb, 1920).
=Blattophila sphaerolaima var. javanica= Chitwood and Chitwood, 1934
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Chitwood
and Chitwood, 1934).
=Blattophila supellaima= Basir, 1941
_Natural host._--_Supella supellectilium_, India, Aligarh (Basir, 1941).
=Cephalobellus brevicaudatum= (Leidy, 1851) Christie, 1933
_Synonymy._--_Thelastoma brevicaudatum_ Leidy, 1851 [Christie, 1933].
_Thelastoma indiana_ Basir, 1940 [Basir, 1949].
_Natural host._--_Leucophaea_ sp., India, Aligarh (Basir, 1940, 1949).
=Cephalobellus magalhāesi= (Schwenck, 1926) Basir, 1956
_Synonymy._--_Bulhỡesi magalhāesi_ Schwenck, 1926; _Thelastoma
magalhāesi_ (Schwenck, 1926) Travassos, 1929 [Basir, 1956].
_Natural host._--"Barata selvagem," Brazil, São Paulo (Schwenck, 1926).
=Euryconema paradisa= Chitwood, 1932
_Natural host._--_Eurycotis floridana_, U.S.A., Florida (Chitwood,
1932).
=Galebia aegyptiaca= (Galeb, 1878) Chitwood, 1932
_Synonymy._--_Oxyuris aegyptiaca_ Galeb, 1878; _Blatticola aegyptiaca_
(Galeb, 1878) Schwenck, 1926 [Chitwood, 1932].
_Natural hosts._--_Blattella germanica_, Brazil (Schwenck, 1926).
_Polyphaga aegyptiaca_, Egypt? (Galeb, 1878).
=Hammerschmidtiella diesingi= (Hammerschmidt, 1838) Chitwood, 1932
_Synonymy._--_Anguillula macrura_ Diesing, 1851; _Aorurus diesingi_
(Hammerschmidt, 1838) Travassos, 1929; _Streptostomum gracile_ Leidy,
1850; _Oxyuris diesingi_ Hammerschmidt, 1838; _Oxyuris blattae
orientalis_ Hammerschmidt, 1838 [Chitwood, 1932]. _Oxyuris macrura_ of
Lankester (1865).
_Natural hosts._--_Blatta orientalis_, Europe (Hammerschmidt 1838, 1847;
Bütschli, 1871); Egypt? (Galeb, 1878); England (Lankester, 1865; Lee,
1958); U.S.A. (Leidy, 1850a); U.S.S.R. (Yakimov and Miller, 1922;
Sobolev, 1937; Sondak, 1935); Brazil (Travassos, 1929); China (Chitwood,
1932); Czechoslovakia (Groschaft, 1956).
_Leucophaea maderae_, Brazil (Pessôa and Corrêa, 1926).
_Periplaneta americana_, Brazil (Magalhães, 1900; Pessôa and Corrêa,
1926). U.S.A.: Texas (Todd, 1943); Kansas (Dobrovolny, 1933; Dobrovolny
and Ackert, 1934); North Carolina (Hatcher, 1939); Iowa, North Dakota,
Michigan (Hoffman, 1953). China (Chitwood, 1932). India (Basir, 1940).
Czechoslovakia (Groschaft, 1956). England (Lee, 1958).
_Periplaneta australasiae_, Brazil (Pessôa and Corrêa, 1926).
_Polyphaga aegyptiaca_ (Linstow, 1878).
Cockroaches (_Blatta orientalis_, _Blattella germanica_, and/or
_Periplaneta americana_), U.S.A. (McAdow, 1931).
Cockroach, Venezuela (Tejera, 1926).
According to Hammerschmidt (1847) this worm may be found throughout the
intestinal canal but especially in the small intestine. It is frequently
found in adults and seldom in the nymphs. There were seldom more than 5
to 10 worms in one cockroach and female worms were found more frequently
than males; the male worms were found only in winter and spring while
the females were present at all times of the year. Bütschli (1871)
stated that all stages from those just hatching to mature males and
females are found.
Yakimov and Miller (1922) found _H. diesingi_ in 50.8 percent of 124 _B.
orientalis_ collected in Petrograd. Sobolev (1937) found 96 percent of
_B. orientalis_ infected with _H. diesingi_ with a mean number of 5.6
and maximum number of 22 in one cockroach. Groschaft (1956) found 18 in
one specimen of _B. orientalis_. Dobrovolny and Ackert (1934) found
about 50 percent of 222 _P. americana_ infected with _H. diesingi_.
Sondak (1935) found about 36 percent of 412 _B. orientalis_ infected
with either or both _H. diesingi_ and _Leidynema appendiculata_.
Two molts occur during development of the eggs; the first takes place
outside the host resulting in a resting or infective stage. After the
egg in the infective stage is eaten by the host, the second molt occurs
before the egg hatches. Completion of the second molt and hatching
perhaps are connected with ammonia present in the digestive tract; the
ammonia seems to arise from the bacteria present in the gut. There
appears to be a relationship between the intestinal bacteria of the
cockroach and development and hatching of nematode eggs (Todd, 1944).
At the time of oviposition the nematode eggs are in the very earliest
stages of cleavage. In 36 hours a motile, tadpole-like stage is reached
and in a few days the embryo becomes quiescent and nonmotile. This
nonmotile stage is infective whereas the motile embryonic stage is not.
Feeding experiments proved that transmission of the nematode is direct.
The worm reaches sexual maturity in 20 or 30 days after being ingested
by the cockroach (Dobrovolny, 1933).
The bacterium _Streptomyces leidynematis_ Hoffman grows on the cuticle
of _H. diesingi_ (Hoffman, 1953). The bacterium apparently is only
anchored to the nematode and probably obtains its food from the
intestinal contents of the cockroach. See notes under _Leidynema
appendiculata_.
=Hammerschmidtiella neyrai= Serrano Sánchez, 1945
_Synonymy._--_Hammerschmidtiella neyrae_ Serrano Sánchez, 1947.
[According to M. B. Chitwood, personal communication, 1957, Serrano
Sánchez's emendation is apparently an error.]
_Natural host._--_Blatta orientalis_, Spain, Grenada (Serrano Sánchez,
1947): Of 2,943 specimens examined, 1,143 were parasitized by oxyurids
and of these 45 percent contained _H. neyrai_.
=Leidynema appendiculata= (Leidy, 1850) Chitwood, 1932
_Synonymy._--_Oxyuris blattae orientalis_ Hammerschmidt, 1847, of
Bütschli, 1871, and _Oxyuris blattae-orientalis_ of Magalhães, 1900;
_Oxyuris blattae_ Hammerschmidt, 1847, of Galeb, 1878; _Aorurus_
(_Thelastoma_) _appendiculatus_ Leidy, 1850. [Chitwood, 1932.] Serrano
Sánchez (1947) has divided this species into three geographical
varieties as follows: _L. appendiculata_ (Leidy, 1852) (Dobrovolny and
Ackert, 1934) var. _indiana_; _L. appendiculata_ (Leidy, 1852)
(Chitwood, 1932) var. _americana_; _L. appendiculata_ (Serrano Sánchez,
1947) var. _hispana_. However, Basir (1956) does not recognize these
varieties. The Russians recognize _hispana_ (M. B. Chitwood, personal
communication, 1957).
_Natural hosts._--_Blaberus atropos_, South America (Chitwood, 1932).
_Blatta orientalis_, Egypt? (Galeb, 1878); Europe (Bütschli, 1871);
U.S.S.R. (Sobolev, 1937; Sondak, 1935); U.S.A., Nebraska (Todd, 1944);
Spain (Serrano Sánchez, 1947): Recorded as var. _hispana_.
Czechoslovakia (Groschaft, 1956). England (Lee, 1958a).
_Blatta orientalis_ or _Periplaneta americana_, Brazil (Magalhães,
1900).
_Periplaneta americana_, U.S.A.: Texas (Todd, 1943); Nebraska (Todd,
1944); Kansas (Dobrovolny, 1933; Dobrovolny and Ackert, 1934); North
Carolina (Hatcher, 1939); Iowa, North Dakota, Michigan (Hoffman, 1953).
Czechoslovakia (Groschaft, 1956). England (Lee, 1958a).
Cockroach, Venezuela (Tejera, 1926).
Cockroaches (_Blatta orientalis_, _Blattella germanica_, and/or
_Periplaneta americana_), U.S.A. (McAdow, 1931).
Chitwood (1932) also listed China for distribution of the worm, but we
could not tell which host was involved.
The worms are found in the colon and rectum of the host. Galeb (1878)
found as many as 20 individuals in a single _B. orientalis_. Sobolev
(1937) found 52 percent of _B. orientalis_ infected with _L.
appendiculata_; the mean number of worms per roach was 1.5 and the
maximum 2. Dobrovolny and Ackert (1934) found 69 percent of 222 _P.
americana_ infected with this species.
Two molts occur within the egg during development of the larva. The
first molt occurs outside the host resulting in the formation of an
infective resting stage. The second molt occurs inside the cockroach
(Todd, 1941, 1944).
Transmission of the nematode is direct, eggs in the resting embryonated
stage being infective (Dobrovolny and Ackert, 1934).
Hoffman (1953) described a filamentous bacterium, _Streptomyces
leidynematis_ Hoffman, which grows on the cuticle of _L. appendiculata_
in _P. americana_. Leidy (1853) noted the presence of simple,
inarticulate, amorphous filaments, growing from nematodes infecting _B.
orientalis_. Bütschli (1871) and Magalhães (1900) described similar
filaments adhering to the surface of oxyurids from cockroaches.
=Leidynema appendiculata= (Leidy, 1850) Chitwood, 1932?
_Natural host._--_Eurycotis floridana_, U.S.A., Massachusetts (Roth and
Willis, unpublished data, 1955): Determined by Dr. G. Steiner who wrote
us, "In _Eurycotis floridana_ there were ten specimens of the nematode
_Leidynema appendiculata_ (Leidy, 1850). This cockroach is obviously a
new host for this nematode. I am not sure that the nematode exactly
agrees with the description as given in the literature."
=Leidynema cranifera= Chitwood, 1932
_Natural hosts._--_Blaberus craniifer_, U.S.A., Florida (Chitwood,
1932); Massachusetts (Roth and Willis, unpublished data, 1955).
Determined by Dr. G. Steiner.
_Blaberus atropos_?, U.S.A., Florida (Chitwood, 1932): _B. craniifer_
has generally been recorded as _B. atropos_ of Stoll which is a closely
related but distinct South American species (Rehn and Hebard, 1927).
=Leidynema delatorrei= Chitwood, 1932
_Natural host._--_Leucophaea maderae_, Cuba, Havana (Chitwood, 1932).
=Leidynema nocalum= Chitwood and Chitwood, 1934
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Chitwood
and Chitwood, 1934).
=Leidynemella fusiformis= Cobb, 1934
_Natural hosts._--_Panesthia laevicollis_?, Philippine Islands (Cobb
_in_ Chitwood and Chitwood, 1934).
_Panesthia angustipennis_, Philippine Islands (Chitwood and Chitwood,
1934).
=Leidynemella panesthiae= (Galeb, 1878) Chitwood and Chitwood, 1934
_Synonymy._--_Oxyuris panesthiae_ Galeb, 1878, in part; _Thelastoma
panesthiae_ (Galeb, 1878) Travassos, 1929. [Chitwood, 1932; Chitwood and
Chitwood, 1934.]
_Natural host._--_Panesthia_ sp., New Guinea (Galeb, 1878): About 40
nematodes may be found in one insect.
=Leidynemella paracranifera= Chitwood and Chitwood, 1934
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Chitwood
and Chitwood, 1934).
=Oxyuris= (?) =heterogamiae= Galeb, 1878
_Synonymy._--_Thelastoma heterogamiae_ (Galeb, 1878) Travassos, 1929.
The taxonomic position of this species is questionable; it might
possibly belong in _Blatticola_ or _Protrellina_ (Chitwood, 1932). Basir
(1956) placed it in an appendix to the family Thelastomatidae.
_Natural host._--_Polyphaga aegyptiaca_, Egypt? (Galeb, 1878).
=Protrelleta floridana= Chitwood, 1932
_Natural host._--_Blaberus craniifer_, U.S.A., Florida (Chitwood, 1932).
=Protrellus aureus= Cobb, 1920
_Synonymy._--The taxonomic position of this nematode is questionable
(Chitwood, 1932).
_Natural host._--_Polyzosteria melanaria_?, Australia, New South Wales
(Cobb, 1920). [Caudell (_in_ Chitwood, 1932) stated that this host was
probably _Platyzosteria analis_.]
=Protrellus aurifluus= (Chitwood, 1932) Chitwood, 1933
_Synonymy._--_Protrellina aurifluus_ Chitwood, 1932.
_Natural hosts._--_Parcoblatta lata_, U.S.A., North Carolina, Maryland
(Chitwood, 1932).
_Parcoblatta uhleriana_, North Carolina (Hatcher, 1939).
=Protrellus australasiae= (Pessôa and Corrêa, 1926) Travassos, 1929
_Synonymy._--_Oxyuris australasiae_ Pessôa and Corrêa, 1926;
_Protrellina australasiae_ (Pessôa and Corrêa, 1926) Chitwood, 1932
[Chitwood, 1933].
_Natural host._--_Periplaneta australasiae_, Brazil (Pessôa and Corrêa,
1926, 1927).
=Protrellus galebi= Schwenck, 1926
_Synonymy._--_Protrellina galebi_ (Schwenck, 1926) Chitwood, 1932
[Chitwood, 1933].
_Natural host._--"Barata selvagem," Brazil (Schwenck, 1926).
=Protrellus künckeli= (Galeb, 1878) Travassos, 1929
_Synonymy._--_Oxyuris künckeli_ Galeb, 1878; _Protrellina künckeli_
(Galeb, 1878) Chitwood, 1932 [Chitwood, 1933].
_Natural hosts._--_Periplaneta americana_, Egypt? (Galeb, 1877, 1878).
[Chitwood (1932) questioned the determination of this host because he
failed to find this nematode in a large number of specimens from U.S.A.
and China.] Brazil (Pessôa and Corrêa, 1926).
_Periplaneta australasiae_, Brazil (Pessôa and Corrêa, 1926).
=Protrellus manni= (Chitwood, 1932) Chitwood, 1933
_Synonymy._--_Protrellina manni_ Chitwood, 1932.
_Natural host._--_Aglaopteryx diaphana_, Cuba (Chitwood, 1932).
=Protrellus phyllodromi= (Basir, 1942) Basir, 1956
_Synonymy._--_Protrellina phyllodromi_ Basir, 1942.
_Natural host._--_Blattella humbertiana_, India, Aligarh (Basir, 1942):
Found in the rectum.
=Protrelloides paradoxa= Chitwood, 1932
_Natural host._--_Eurycotis floridana_, U.S.A., Florida (Chitwood,
1932).
=Schwenkiella icemi= (Schwenck, 1926) Basir, 1956
_Synonymy._--_Bulhõesia icemi_ Schwenck, 1926; _Thelastoma icemi_
(Schwenck, 1926) Travassos, 1929; _Thelastoma aligarhica_ Basir, 1940.
[Basir, 1956.]
_Natural hosts._--"Barata selvagem," Brazil, São Paulo (Schwenck, 1926).
_Periplaneta americana_, India, Aligarh (Basir, 1940); U.S.A., Nebraska
(Todd, 1943).
_Periplaneta brunnea_, U.S.A., Louisiana (Todd, 1943).
=Severianoia magna= Pereira, 1935
_Natural host._--"Blattidae sylvestres," Brazil (Pereira, 1935).
=Severianoia severianoi= (Schwenck, 1926) Travassos, 1929
_Synonymy._--_Bulhõesia severianoi_ Schwenck, 1926 [Travassos, 1929].
_Natural hosts._--"Baratas de pau podre," Brazil (Schwenck, 1926).
_Pycnoscelus surinamensis_, U.S.A., Florida (Chitwood, 1932).
=Suifunema caudelli= Chitwood 1932
_Natural host._--_Steleopyga? sinensis_, Asia: Suifu, Szchuen, China
(Chitwood, 1932).
=Thelastoma pachyjuli= (Parona, 1896) Travassos, 1929
_Synonymy._--_Oxyuris bulhõesi_ de Magalhães, 1900; _Bulhõesia bulhõesi_
(Magalhães, 1900) Schwenck, 1926 [Travassos, 1929; Chitwood, 1932];
_Thelastoma bulhõesi_ (Magalhães, 1900) Travassos, 1929; although this
last combination (from Chitwood, 1932) is not given by Basir (1956), it
is implied by the synonymy that he does cite under _T. pachyjuli_.
_Natural hosts._--_Blatta orientalis_, Czechoslovakia (Groschaft, 1956).
_Periplaneta americana_, Brazil (Magalhães, 1900); North America
(Chitwood, 1932); U.S.A., North Carolina (Hatcher, 1939).
=Thelastoma palmettum= Chitwood and Chitwood, 1934
_Natural host._--_Panesthia angustipennis_, Philippine Islands (Chitwood
and Chitwood, 1934).
=Thelastoma riveroi= Chitwood, 1932
_Natural host._--_Periplaneta_ sp., Cuba (Chitwood, 1932).
=Undetermined nematodes=
_Natural host._--_Cutilia_ sp. near _sedilloti_, U.S.A. (hosts imported
from New Zealand) (Roth, unpublished data, 1957).
Class NEMATOMORPHA
Order GORDIOIDEA
The immature stages of the following gordian worms have been found in
the body cavity of cockroaches.
Family CHORDODIDAE
=Chordodes morgani= Montgomery, 1898
_Synonymy._--_Chordotes_ [sic] _puerilis_ Montgomery, 1898 [Ward, 1918].
_Natural host._--Cockroach, U.S.A. (Montgomery, 1898); Pennsylvania,
Maryland, Michigan, Ohio, Florida, Iowa, Nebraska (Ward, 1918).
Family GORDIIDAE
=Gordius aquaticus= Linnaeus, 1758
_Natural host._--_Blatta_ sp., U.S.A. (Stiles and Hassall, 1894).
Leidy (1879) identified a 9-inch-long nematode which came from a
cockroach (_Blatta orientalis?_) as probably being _Gordius aquaticus_.
Ransom (_in_ Pierce, 1921) states that _G. aquaticus_ may be an
accidental parasite of man. Faust (1955) summarizes the few reported
cases of human parasitism. Dorier (1930) reported that the regurgitated
liquid of _Blatta orientalis_ had no effect on cysts of _G. aquaticus_
after one hour.
=Gordius blattae orientalis= Diesing, 1851
_Synonymy._--_Gordius orientalis_ of Lankester (1865).
_Natural host._--_Blatta orientalis_, Germany (Siebold, 1842; Linstow,
1878): Found in abdomen. Von Siebold called this "Filarien" but did not
otherwise name or describe the worm.
=Gordius pilosus= (Möbius, 1855) Diesing, 1861
_Synonymy._--_Chordodes pilosus_ Möbius, 1855 [Diesing, 1861.]
_Natural host._--_Blaberus giganteus_, Venezuela (Möbius, 1855): From
the insect's abdomen.
=Gordius= sp.
_Natural hosts._--_Periplaneta americana_, South Africa (Porter, 1930);
Germany (Bode, 1936): Bode's record may have referred to a _Mermis_ or
other nematode.
Cockroaches, Venezuela (Miall and Denny, 1886; Burr, 1899a; Tejera,
1926).
=Parachordodes raphaelis= (Camerano, 1893) Camerano, 1897
_Synonymy._--_Gordius raphaelis_ Camerano, 1893 [Camerano, 1897].
_Natural hosts._--_Symploce parenthesis_ and _Kuchinga hemerobina_,
French Equatorial Africa (Camerano, 1893, 1897).
=Undetermined gordian worms=
_Natural hosts._--_Eurycotis floridana_, Florida (T. Eisner, personal
communication, 1958): See plate 29, B.
_Parahormetica bilobata_, Brazil (Pessôa and Corrêa, 1929): Worm
referred to as "gordiaceo."
Cockroaches, Australia (E. F. Riek, personal communication, 1953): Three
undescribed gordian worms were found in undetermined cockroaches of the
subfamily Blattinae.
HELMINTHS[4] FOR WHICH COCKROACHES SERVE AS INTERMEDIATE HOSTS
The use of the asterisk (*) is explained in footnote 3, page 4.
Phylum ACANTHOCEPHALA
Order ARCHIACANTHOCEPHALA
Family OLIGACANTHORHYNCHIDAE
* =Prosthenorchis elegans= (Diesing, 1851) Travassos, 1915
_Natural host._--_Blattella germanica_, France (Brumpt and Urbain, 1938,
1938a; Brumpt et al., 1939).
_Experimental hosts._--_Blaberus atropos_ and _Leucophaea maderae_,
France (Brumpt and Desportes, 1938).
* =Prosthenorchis spirula= (Olfers in Rudolphi, 1819) Travassos, 1917
_Natural host._--_Blattella germanica_, France (Brumpt and Urbain, 1938,
1938a; Brumpt et al., 1939); Netherlands (Thiel and Wiegand Bruss,
1946).
_Experimental hosts._--_Blattella germanica_, Netherlands (Thiel and
Wiegand Bruss, 1946).
_Blaberus atropos_ and _Leucophaea maderae_, France (Brumpt and
Desportes, 1938).
Family MONILIFORMIDAE
* =Moniliformis dubius= Meyer, 1932
_Natural hosts._--_Periplaneta americana_, Brazil (Magalhães, 1898;
Travassos, 1917); Gold Coast (Southwell, 1922); India (Pujatti, 1950);
U.S.A. (Burlingame and Chandler, 1941; Moore, 1946).
_Periplaneta australasiae_, India (Pujatti, 1950).
_Experimental hosts._--_Blattella germanica_, Japan (Yamaguti and
Miyata, 1942).
_Periplaneta americana_, U.S.A. (Chandler, 1941; Moore, 1946); Japan
(Yamaguti and Miyata, 1942).
* =Moniliformis kalahariensis= Meyer, 1931
_Natural host._--_Blattella germanica_, India (Meyer, 1931, 1932).
* =Moniliformis moniliformis= (Bremser _in_ Rudolphi, 1819) Travassos,
1915
_Natural hosts._--_Periplaneta americana_, Argentina (Bacigalupo, 1927,
1927a, 1928); Brazil (Pessôa and Corrêa, 1929); Algeria (Seurat, 1912);
Burma (Subramanian, 1927); South Africa (Porter, 1930); Madras (Sita,
1949).
_Periplaneta_ spp., New Caledonia (Rageau, 1956).
Cockroaches, Venezuela (Tejera, 1926).
_Experimental hosts._--_Blaberus atropos_, _Blatta orientalis_,
_Blattella germanica_, _Leucophaea maderae_, France (Brumpt and Urbain,
1938a).
_Periplaneta americana_, Japan (Yamaguti and Miyata, 1942); France
(Brumpt, 1949); Madras (Sita, 1949).
Phylum ASCHELMINTHES
Class NEMATODA
Order OXYUROIDEA
Family SUBULURIDAE
* =Subulura jacchi= (Diesing, 1861) Railliet and Henry, 1914
_Synonymy._--_Subulura jacchi_ (Marcel, 1857) [Dr. J. T. Lucker,
personal communication, 1957].
_Experimental host._--_Blaberus atropos_, France (Chabaud and Larivière,
1955).
Order SPIRUROIDEA
Family THELAZIIDAE
* =Oxyspirura mansoni= (Cobbold, 1879) Ransom, 1904
_Natural hosts._--_Pycnoscelus surinamensis_, Australia (Fielding, 1926,
1927, 1928, 1928a); U.S.A. (Sanders, 1927, 1928, 1929; Shealy, 1927);
Formosa (Kobayashi, 1927); Antigua (Hutson, 1938, 1943); Hawaii
(Illingworth, 1931; Schwabe, 1950, 1950a, 1950b, 1951); New Caledonia
(Rageau, 1956).
We have recently found (Roth and Willis, 1960) that two strains of
_Pycnoscelus surinamensis_ exist; a parthenogenetic strain (from
Florida), and a bisexual strain (from Hawaii) which does not reproduce
parthenogenetically. The parthenogenetic strain is undoubtedly the form
that has been shown to be the host of _O. mansoni_ in the United States
and Antigua, because only this form is found in the New World. Probably
the parthenogenetic strain was the form involved in most Pacific areas.
However, from internal evidence in his papers, we concluded that
Schwabe, in Hawaii, may well have been working with the bisexual strain
and possibly also with the parthenogenetic strain; if this is true, then
both parthenogenetic and bisexual strains of _Pycnoscelus surinamensis_
may serve as intermediate hosts of the eyeworm.
_Experimental hosts._--_Periplaneta americana_, Antigua (Hutson, 1943).
_Pycnoscelus surinamensis_, U.S.A. (Sanders, 1929); Australia (Fielding,
1927, 1928a); Hawaii (Schwabe, 1951).
=Rictularia coloradensis= Hall, 1916
_Natural hosts._--_Parcoblatta pensylvanica_ and _Parcoblatta
virginica_, U.S.A., Ohio (Oswald, 1958): Of 49 wood roaches collected,
one of each species contained a single larva each.
_Experimental hosts._--_Blatta orientalis_, _Blattella germanica_,
_Parcoblatta pensylvanica_, _Parcoblatta virginica_, _Periplaneta
americana_, and _Supella supellectilium_, U.S.A. (Oswald, 1958): The
larvae underwent normal development in all species of cockroaches except
_B. orientalis_ and _P. americana_ in which cysts developed that
contained a reddish-brown pigment; larvae in such cysts were dead or
dying. Eggs of _R. coloradensis_ hatched in the midgut of _B. germanica_
and first-stage larvae entered the hindgut epithelium within 24 hours.
The larvae underwent two molts within a cyst formed by tissues of the
host's gut, the second molt occurring during the twelfth or thirteenth
day. In _Parcoblatta_, cysts were found free in the body cavity as well
as attached to the hindgut. In _B. germanica_ and _S. supellectilium_
the cysts remained attached to the hindgut. Usually over 20 cysts
developed in each infected _Parcoblatta_; fewer than 10 per insect
developed in the other species. Larvae became infective to the
definitive host, the white-footed mouse [_Peromyscus leucopus
noveboracensis_ (Fischer)], as early as the tenth day.
Family SPIRURIDAE
* =Agamospirura parahormeticae= Pessôa and Corrêa, 1929
_Natural host._--_Parahormetica bilobata_, Brazil (Pessôa and Corrêa,
1929).
* =Gongylonema ingluvicola= Ransom, 1904
_Experimental host._--_Blattella germanica_, U.S.A. (Cram, 1935).
* =Gongylonema neoplasticum= (Fibiger and Ditlevsen, 1914) Ransom and
Hall, 1916
_Natural hosts._--_Blatta orientalis_, Netherlands (Baylis, 1925).
_Blattella germanica_, U.S.A. (Hitchcock and Bell, 1952).
_Periplaneta americana_, Denmark and St. Croix (Fibiger, 1913, 1913a;
Fibiger and Ditlevsen, 1914); Netherlands (Baylis, 1925); Argentina
(Bacigalupo, 1930); England (Leiper, 1926); South Africa (Porter, 1930);
U.S.A. (Hitchcock and Bell, 1952); Formosa (Yokagawa, 1924, 1925,
1925a).
_Periplaneta australasiae_, Formosa (Yokagawa, 1924, 1925, 1925a).
_Experimental hosts._--_Blattella germanica_, Denmark (Fibiger and
Ditlevsen, 1914); U.S.A. (Hitchcock and Bell, 1952); France (Brumpt,
1949).
_Blatta orientalis_, Denmark (Fibiger and Ditlevsen, 1914).
_Periplaneta americana_, Denmark and St. Croix (Fibiger, 1913; Fibiger
and Ditlevsen, 1914); U.S.A. (Hitchcock and Bell, 1952).
* =Gongylonema pulchrum= Molin, 1857
_Experimental hosts._--_Blattella germanica_, U.S.A. (Ransom and Hall,
1915, 1916, 1917; Stiles and Baker, 1927; Schwartz and Lucker, 1931;
Lucker, 1932; Alicata, 1934a, 1935); Europe (Baylis et al., 1925, 1926,
1926a; Sambon, 1926).
_Parcoblatta_ sp., Alicata (1934, 1935).
* =Gongylonema= sp.
_Natural host._--_Periplaneta americana_, Brazil (Magalhães, 1900);
Algeria (Seurat, 1916); England? (Leiper, 1926).
* =Microtetrameres helix= Cram, 1927
_Experimental host._--_Blattella germanica_, U.S.A. (Cram, 1934).
* =Protospirura bonnei= Ortlepp, 1924
_Natural host._--_Leucophaea maderae_, Venezuela (Brumpt, 1931).
_Experimental hosts._--_Blatta orientalis_, _Blattella germanica_,
_Leucophaea maderae_, France (Brumpt, 1931).
* =Protospirura columbiana= Cram, 1926
_Experimental host._--_Blattella germanica_, U.S.A. (Cram, 1926).
* =Protospirura muricola= Geodoelst, 1916
_Natural host._--_Leucophaea maderae_, Panama (Foster and Johnson, 1938,
1939).
* =Seurocyrnea colini= (Cram, 1927) Cram, 1931
_Experimental host._--_Blattella germanica_, U.S.A. (Cram, 1931, 1931a,
1933a).
* =Spirura gastrophila= (Müller, 1894) Seurat, 1913
_Natural hosts._--_Blatta orientalis_, Europe? (Deslongchamps, 1824,
_in_ Seurat, 1911); Italy (Grassi, 1888); Algeria (Seurat, 1916).
_Periplaneta americana_, Brazil (Pessôa and Corrêa, 1929).
Cockroach, Venezuela (Tejera, 1926).
_Experimental hosts._--_Blatta orientalis_, France (Galeb, 1878a);
"Cafards," Algeria (Roger, 1906, 1907).
* =Tetrameres americana= Cram, 1927
_Natural host._--_Blattella germanica_, U.S.A. (Cram, 1931b, personal
communication, 1956); Hawaii (Alicata, 1938, 1947).
_Experimental host._--_Blattella germanica_, U.S.A. (Cram, 1931b).
* =Tetrameres pattersoni= Cram, 1933
_Experimental host._--_Blattella germanica_, U.S.A. (Cram, 1933).
Family PHYSALOPTERIDAE
* =Physaloptera hispida= Schell, 1950
_Experimental host._--_Blattella germanica_, U.S.A. (Schell, 1952,
1952a).
* =Physaloptera maxillaris= Molin, 1860
_Experimental host._--_Blattella germanica_, U.S.A. (Hobmaier, 1941).
* =Physaloptera praeputialis= von Linstow, 1889
_Experimental host._--_Blattella germanica_, U.S.A. (Petri and Ameel,
1950).
* =Physaloptera rara= Hall and Wigdor, 1918
_Experimental host._--_Blattella germanica_, U.S.A. (Petri and Ameel,
1950; Petri, 1950).
* =Physaloptera turgida= Rudolphi, 1819
_Experimental host._--_Blattella germanica_, U.S.A. (Alicata, 1937;
Schell, 1952).
HELMINTHS WHOSE EGGS HAVE BEEN CARRIED BY COCKROACHES
The use of the asterisk (*) is explained in footnote 3, page 4.
Phylum PLATYHELMINTHES
Class TREMATODA
Order DIGENEA
Family SCHISTOSOMATIDAE
* =Schistosoma haematobium= (Bilharz, 1852) Weinland, 1858
_Experimental vector._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922).
Class CESTODA
Order TAENIOIDEA
Family HYMENOLEPIDIDAE
* =Hymenolepis= sp.
_Natural vectors._--_Periplaneta americana_, Formosa (Morischita and
Tsuchimochi, 1926).
_Polyphaga saussurei_, Tadzhikistan (Zmeev, 1936).
Family TAENIIDAE
* =Taenia saginata= Goeze, 1782
_Experimental vector._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922).
=Echinococcus granulosis= (Batsch, 1786) Rudolphi, 1805
_Synonymy._--_Taenia echinococcus_ (Zeder, 1803) [Faust, 1939].
_Experimental vector._--_Periplaneta americana_, Uruguay (Pérez Fontana,
1955): Eggs were recovered from the feces of artificially infested
cockroaches under "natural" conditions.
Family Unknown
* =Undetermined tapeworm ova=
_Natural vector._--_Polyphaga saussurei_, Tadzhikistan (Zmeev, 1936).
Phylum ASCHELMINTHES
Class NEMATODA
Order OXYUROIDEA
Family OXYURIDAE
* =Enterobius vermicularis= (Linnaeus, 1758) Leach in Baird, 1853
_Natural vectors._--_Blatta orientalis_ and _Blattella germanica_,
U.S.S.R. (Sondak, 1935).
Order ASCAROIDEA
Family ASCARIDAE
* =Ascaris lumbricoides= Linnaeus, 1758
_Natural vector._--_Periplaneta americana_, South Africa (Porter, 1930):
The eggs may have been those of _A. suum_ Goeze, 1782.
_Experimental vectors._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922); India (Chandler, 1926).
_Periplaneta americana_, _Periplaneta australasiae_, _Neostylopyga
rhombifolia_, Formosa (Morischita and Tsuchimochi, 1926).
* =Ascaris= sp.
_Natural vector._--_Blatta orientalis_, Italian Somaliland (Mariani and
Besta, 1936).
_Experimental vector._--_Periplaneta americana_, Uruguay (Pérez Fontana,
1955): Eggs recovered from the insects' feces.
Order STRONGYLOIDEA
Family ANCYLOSTOMIDAE
* =Ancylostoma caninum= (Ercolani, 1859) Hall, 1913
_Experimental vector._--_Periplaneta americana_, Netherlands (Akkerman,
1933).
* =Ancylostoma ceylanicum= (Looss, 1911) Leiper, 1915
_Experimental vector._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922); Netherlands (Akkerman, 1933).
* =Ancylostoma duodenale= (Dubini, 1843) Creplin, 1845
_Natural vector._--_Periplaneta americana_, South Africa (Porter, 1929,
1930).
_Experimental vector._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922).
* =Necator americanus= (Stiles, 1902) Stiles, 1906
_Natural vector._--_Periplaneta americana_, India (Chandler, 1926).
_Experimental vector._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922).
* =Hookworm ova=
_Experimental vectors._--_Periplaneta americana_, _Periplaneta
australasiae_, _Neostylopyga rhombifolia_, Formosa (Morischita and
Tsuchimochi, 1926).
Family TRICHOSTRONGYLIDAE
* =Trichostrongylus= sp.
_Natural vector._--_Blatta orientalis_, Italian Somaliland (Mariani and
Besta, 1936).
Order TRICHUROIDEA
Family TRICHURIDAE
* =Capillaria hepatica= (Bancroft, 1893) Travassos, 1915
_Experimental vector._--_Blatta orientalis_, Italy? (Giordano, 1950).
* =Trichuris trichiura= (Linnaeus, 1771) Stiles, 1901
_Natural vectors._--_Blatta orientalis_, Italian Somaliland (Mariani and
Besta, 1936); U.S.S.R. (Sondak, 1935).
_Blattella germanica_, U.S.S.R. (Sondak, 1935).
_Periplaneta americana_, Gold Coast Colony (Macfie, 1922); Formosa
(Morischita and Tsuchimochi, 1926).
_Experimental vectors._--_Periplaneta americana_, Gold Coast Colony
(Macfie, 1922); India (Chandler, 1926); Uruguay (Pérez Fontana, 1955).
_Periplaneta americana_, _Periplaneta australasiae_, and _Neostylopyga
rhombifolia_, Formosa (Morischita and Tsuchimochi, 1926).
XII. ARTHROPODA
The classification follows Brues et al. (1954) with the following
exceptions. The Acarina are arranged according to Dr. J. H. Camin
(personal communication, 1955). Family Eupelmidae of the Hymenoptera
follows the classification of Peck (1951).
Class ARACHNIDA
In this class, representatives of at least four orders have utilized
cockroaches as food: the whip scorpions, scorpions, spiders, and mites.
Apparently none of these feed exclusively on cockroaches, but the
Philippine forest scorpion _Heterometrus_ (=_Palamnaeus_) _longemanus_
seems to prefer blattids to other insects (Schultze, 1927).
Order PEDIPALPIDA
Family THELYPHONIDAE
=Mastigoproctus giganteus= (Lucas)
_Synonymy._--_Thelyphonus giganteus_ Lucas [Dr. R. E. Crabill, personal
communication, 1958].
_Experimental prey._--Cockroaches, U.S.A. (Marx, 1892, 1894): Immature
whip scorpion captured and fed on one or two cockroaches a week. It
lived on this diet for about two years.
=Mastigoproctus= sp.
_Common name._--Whip scorpion.
_Experimental prey._--_Blattella germanica_, U.S.A., Florida [Dr. B. J.
Kaston, personal communication, 1953].
Order SCORPIONIDA
Pocock (1893) noticed that a scorpion whose pectines had come in contact
with a cockroach immediately turned back and ate the insect. He
concluded that the scorpion detected the cockroach by means of the
pectines. However, Cloudsley-Thompson (1955) has demonstrated that the
main function of the pectines is probably the detection of ground
vibrations. He accounted for Pocock's observation by the presence of
sensory spines (presumably tactile) which project from beneath the
pectines. In a house in Arizona, Lyon (1951) observed over 60 scorpions
living in a kitchen cabinet that enclosed a sink. They were apparently
thriving on a heavy infestation of cockroaches. Stahnke (1953) stated
that he used _Periplaneta americana_ as the principal food for scorpions
at the Poisonous Animals Research Laboratory of Arizona State College.
Cloudsley-Thompson (1955a) cited cockroaches as one of the arthropods
that scorpions feed upon.
Family BUTHIDAE
=Buthus australis= (Linnaeus)
_Synonymy._--_Androctonus australis_ [Crabill, personal communication,
1957].
_Experimental prey._--Cockroaches, England (Cloudsley-Thompson, 1955a):
This African species ate at least one cockroach per week during the
summer months. It can, however, survive four months' starvation and is
particularly adapted to a dry climate (Cloudsley-Thompson, personal
communication, 1956).
=Centruroides gracilis= (Latreille)
_Experimental prey._--_Periplaneta americana_, U.S.A. (Roth, unpublished
data, 1953): Scorpion collected in Florida by Roth and identified by Dr.
M. H. Muma.
=Centruroides hentzi= (Banks)
_Experimental prey._--_Periplaneta australasiae_ and _Pycnoscelus
surinamensis_, U.S.A. (Muma, personal communication, 1953): This
scorpion occurs in large numbers in the Florida citrus groves, together
with _P. australasiae_ which is probably an important natural prey.
=Centruroides vitattus= (Say)
_Natural prey._--_Parcoblatta pensylvanica_ (?), U.S.A., Florida (Muma,
personal communication, 1953). This may have been another species of
this genus, possibly _P. divisa_, as _P. pensylvanica_ is not known from
Florida (Rehn, personal communication, 1958).
_Experimental prey._--_Blatta orientalis_, _Blattella germanica_,
_Periplaneta americana_, and _Pycnoscelus surinamensis_, U.S.A., Florida
(Muma, personal communication, 1953).
=Parabuthus capensis= (Hemprich and Ehrenberg)
_Experimental prey._--"Common house-cockroach" (Pocock, 1893): The
scorpions were collected in Cape Town, South Africa.
Family CHACTIDAE
=Euscorpius germanus= (Koch)
_Synonymy._--_Euscorpius carpathicus_ [Cloudsley-Thompson, 1955a].
_Experimental prey._--_Blattella germanica_, England? (Pocock, 1893).
_Periplaneta americana_, nymphs, England? (Cloudsley-Thompson, personal
communication, 1956): This scorpion is found naturally in southern
Europe and North Africa.
=Euscorpius italicus italicus= (Herbst)
_Experimental prey._--Cockroaches including nymphs of _Periplaneta_,
England? (Cloudsley-Thompson, 1951): The cockroaches had to be disabled
before the scorpion would feed on them. Prey is apparently detected by
tactile and auditory senses, sight being poorly developed and not used.
The scorpion is found in southern Europe and North Africa.
Family VEJOVIDAE
=Hadrurus arizonensis= Ewing
_Experimental prey._--_Periplaneta americana_, U.S.A. (Stahnke, 1949):
This record is a photograph showing the scorpion eating the cockroach.
Family ISCHNURIDAE
=Hormurus caudicula= (Koch)
_Experimental prey._--Cockroach, Australia (McKeown, 1952): This record
is a photograph showing the scorpion feeding on a cockroach.
Family SCORPIONIDAE
=Heterometrus longimanus= (Herbst)
_Synonymy._--_Palamnaeus longimanus_ [Cloudsley-Thompson, personal
communication, 1956].
_Natural prey._--"Large wood cockroach," Philippine Islands (Schultze,
1927): On several occasions Schultze found fragments of wings and legs
of the large wood cockroach in a scorpion cavity, under a rotten log.
_Experimental prey._--_Leucophaea maderae_, _Periplaneta americana_, and
other species of Blattidae, Philippine Islands (Schultze, 1927):
Blattids seemed to be the favored food. This scorpion is usually found
in humid, damp places in forest and jungle. Schultze describes in detail
feeding behavior of the scorpion and method of capturing its prey.
=Urodacus novaehollandiae= Peters
_Experimental prey._--_Periplaneta americana_, Australia (Glauert,
1946): An injured cockroach was accepted at once by the scorpion, which
held the insect in its claws and tore it with the alternately moving
chelicerae. The scorpion ate all the soft parts and most of the
sclerotized exoskeleton.
Order ARANEIDA
Many observations of spiders feeding on cockroaches are quite general,
and many observers have failed to identify either the spider or its
prey. Belt (1874) stated that "the cockroaches that infest houses in the
tropics ... have numerous enemies--birds, rats, scorpions, and spiders."
When Belt tried to drive a cockroach toward a large cockroach-eating
spider, the insect rushed away from him until it came within a foot of
the spider when it would double back, never advancing nearer.
Beebe (1925) watched a giant "wood roach," which was in the grasp of a
2-inch ctenid spider, fly through the window of his British Guiana
laboratory. While the spider ate the cockroach, the insect gave birth to
51 nymphs. Sonan (1924) reported that large gray spiders devour nymphs
and adults of _Periplaneta americana_ and _P. australasiae_ in Formosa;
this spider also occurs on Hiyakejima Island and Okinawa. Passmore
(1936), who has produced some excellent photographs of tarantulas,
stated that they destroy cockroaches. Rau (1940) stated that American
and oriental cockroaches were the principal item of diet of a friend's
pet tarantula for several years. Kaston (personal communication, 1953)
successfully fed a tarantula with _Periplaneta americana_.
Bristowe (1941) found that the British species of _Ectobius_ are readily
accepted by _Xysticus_, _Clubiona_, _Drassodes_, _Zelotes_, _Tarantula_,
and the web-builders _Ciniflo_ and _Aranea_. The British domestic
cockroaches were accepted by _Tegenaria_ and _Ciniflo_, spiders large
enough to overpower them, and were useful as food for tropical
avicularids, ctenids, and sparassids in captivity.
Family THERAPHOSIDAE
=Avicularia avicularia= (Linnaeus) and =Avicularia= sp.?
_Common name._--Bird-eating spider.
_Natural prey._--_Periplaneta americana_, Trinidad (Main, 1924, 1930):
The remains of the host were compressed into globular form by the spider
after it had extracted the nutritive parts.
=Phormictopus cancerides= (Latreille)
_Experimental prey._--Cockroach, West Indies (Wolcott, 1953).
Family SPARASSIDAE
=Heteropoda venatoria= (Linnaeus)
_Synonymy._--_Heteropoda regia_ Fabricius.
_Common names._--Banana spider (Comstock, 1912); huntsman spider
(Gertsch, 1949); big brown house spider (Bryan, 1915).
_Natural and experimental prey._--Cockroaches, Bermuda (Verrill, 1902);
Puerto Rico (Sein, 1923; Wolcott, 1924a; Petrunkevitch, 1930a); Hawaii
(Bryan, 1915, Williams et al., 1931); British Guiana (Moore _in_
Williams et al., 1931); Panama (Gertsch, 1949); New Zealand (adventive)
(Parrott, 1952); England (Cloudsley-Thompson, 1953); Comstock (1912);
Hawaii (Pemberton, 1917).
This (pl. 30, A) is a tropical species frequently imported into northern
localities with bunches of bananas (Comstock, 1912; Cloudsley-Thompson,
1953). Adults measure 3 to 4 inches across with bodies over an inch
long. They seldom leave their resting places during the day, but are
active at night and search for food. The female does not spin a web
(Bryan, 1915; Gertsch, 1949). The spider turns the cockroach over onto
its back at the instant of seizure and holds it firmly against the
substrate. The cockroach dies in 10 minutes and is gradually rolled up
by the spider as it sucks out the nutriment (Moore _in_ Williams et al.,
1931). The spider does not attempt to bite when captured, but if it
does, its bite is said to be painful but not dangerous
(Cloudsley-Thompson, 1953). Zimmerman (1948) found scores of
_Periplaneta australasiae_ breeding in rock piles in Hawaii; also
present were large numbers of these spiders and centipedes which
presumably preyed upon the cockroaches.
Family THERIDIIDAE
=Latrodectus indistinctus= Pickard-Cambridge
_Common name._--Button spider.
_Natural prey._--_Karnyia discoidalis_, South Africa, Western Cape
Province (Hesse, 1942): The nest is constructed on the ground among
grass stems or other vegetation. Preferred sites are slight hollows,
hoof imprints, etc. Nests are roughly tubular. The remains of insects
are entangled in the walls of the nest where they form dense
accumulations. Predatory activities of the spider are limited to an area
close to the tubular entrance to the nest and do not extend beyond the
trapping strands near the entrance. Capture is dependent upon accidental
contact of the insect with sticky threads surrounding the entrance. This
spider apparently attacks any insect or arachnid that becomes entangled
in the nest. In an examination of 40 nests, remains of 6 _K.
discoidalis_ were found.
=Latrodectus mactans= (Fabricius)
_Common names._--Black widow, hourglass, or shoe-button spider.
_Natural prey._--Cockroaches, Puerto Rico (Petrunkevitch, 1930). U.S.A.,
Florida, on shipboard (Anonymous, 1939): This is a presumptive host
record, as the spiders were not reported as having been seen eating
cockroaches; however, heavy infestations of both were found together.
Family LYCOSIDAE
=Lycosa helluo= Walckenaer
_Experimental prey._--Young nymphs of _Diploptera punctata_, U.S.A.
(Eisner, 1958): Larger nymphs and adults repelled the spider by ejecting
a repellent secretion, which has been identified as a mixture of
_p_-benzoquinone and its derivatives by Roth and Stay (1958).
=Lycosa= sp.
_Experimental prey._--_Supella supellectilium_, U.S.A. (Roth and Willis,
unpublished data, 1953): The lycosid (pl. 30, B-E) was probably _L.
avida_ Walckenaer (tentatively identified by Dr. B. J. Kaston from a
photograph).
Order ACARINA
Family PHYTOSEIIDAE
=Blattisocius tineivorus= (Oudemans)
_Synonymy._--_Blattisocius triodons_ Keegan [Baker and Wharton, 1952].
_Natural host._--_Blattella germanica_, U.S.A. (Keegan, 1944): Three
mites found on 238 cockroaches examined; others taken in debris from
floor of cockroach cage (Keegan, 1944). Members of this family are
predaceous (Baker and Wharton, 1952).
Family LAELAPTIDAE
=Blattilaelaps nauphoetae= Womersley
_Natural host._--_Nauphoeta cinerea_, Australia, Brisbane (Womersley,
1956).
=Coleolaelaps= (?) sp.
_Natural host._--_Gromphadorhina portentosa_, the hosts were imported
into U.S.A. from Madagascar via Europe (Roth and Willis, unpublished
data, 1958): The mites (pl. 12, C) were tentatively determined by Dr. E.
W. Baker.
=Hypoaspis= sp.
_Natural host._--_Panesthia australis_, imported into U.S.A. from
Australia (Roth and Willis, unpublished data, 1955): Cockroach
determined by J. A. G. Rehn. Generic determination of mite made by Dr.
R. W. Strandtmann (Camin, personal communication, 1955).
Family UROPODIDAE
=Uropoda= sp.
_Natural host._--_Blattella humbertiana_, Formosa (Takahashi, 1940).
Nymphs of the cockroach may be destroyed (Takahashi, 1940). Uropodids
frequently attach themselves to insects, especially in nymphal stages
but probably are harmless (Baker and Wharton, 1952).
Family DIPLOGYNIIDAE
=Undetermined diplogyniid=
_Natural host._--_Panesthia australis_, imported into U.S.A. from
Australia (Roth and Willis, unpublished data, 1955): Cockroach
determined by J. A. G. Rehn. According to Dr. J. H. Camin (personal
communication, 1955) this is a new genus and new species in the
subfamily Diplogyniinae, and is most closely related to the genus
_Lobogynioides_. Mites of this family live as ectoparasites and
commensals on beetles and possibly other insects (Baker and Wharton,
1952).
Family ANOETIDAE
=Histiostoma feroniarum= (Dufour)
_Natural host._--_Pycnoscelus surinamensis_, Germany (Roeser, 1940):
Though not parasitic, the mites at times became so numerous that the
insects were hindered in their movement, were unable to feed, and died.
The mites were introduced with soil and leaves and had originally been
attached to millipedes, waterfleas, and sowbugs.
The deutonymphs, hypopial forms, or travelers are found on insects; the
other stages are found in decaying organic matter (Baker and Wharton,
1952).
Family ACARIDAE
=Caloglyphus spinitarsus= (Hermann)
_Natural host._--_Pycnoscelus surinamensis_, Germany (Roeser, 1940): See
notes following _Histiostoma feroniarum_ above.
=Caloglyphus= sp.
_Natural hosts._--_Blattella germanica_ and _Periplaneta americana_,
U.S.A. (Piquett and Fales, 1952): Mite feeds on organic matter but can
reduce the vigor of a cockroach colony.
=Tyrophagus lintneri= (Osborne)
_Common name._--Mushroom mite.
_Associate._--_Pycnoscelus surinamensis_, U.S.A. (Roth and Willis,
unpublished data, 1953): Mite determined by Dr. E. W. Baker (personal
communication, 1953). Although this mite was found on the cockroach, it
is a known pest in stored foods (Baker and Wharton, 1952) and probably
was brought into the culture with food. Rau (1924) reported that the
food of _Blatta orientalis_ often became infested with this species, but
it did not affect the health or mortality of the cockroaches in his
culture.
=Tyrophagus noxius= A. Z.
_Natural host._--_Periplaneta americana_, U.S.A. (Roth and Willis,
unpublished data, 1953): Mite determined by Dr. E. W. Baker (personal
communication, 1953). Mites were found in the oöthecal cavity of a
female cockroach that had been isolated for her entire adult life. The
mites were in a closely packed mass behind a plug of what appeared to be
feces, disintegrated eggs, and dried blood; none of the mites were
visible until this plug was removed. Baker (personal communication,
1953) stated that the mite is probably not parasitic and that species of
the genus feed on organic matter.
=Rhizoglyphus tarsalus= Banks
_Natural host._--_Periplaneta americana_, U.S.A. (Rau, 1940a): Not
normally parasitic on cockroaches, but the mites became so numerous at
times they would attack living as well as dead and dying cockroaches.
Family GLYCIPHAGIDAE
=Chaetodactylus= sp.
_Synonymy._--_Trichotarsus_ sp. [Baker and Wharton, 1952].
_Natural host._--_Leucophaea maderae_, Puerto Rico (Seín, 1923): Mites
found on cockroach's thorax and particularly among the folds of the
wings (Seín, 1923). Mites of this genus are found infesting organic
matter (Baker and Wharton, 1952).
Family PODAPOLIPODIDAE
=Locustacarus= sp.
_Natural hosts._--_Diploptera punctata_ and _Nauphoeta cinerea_, U.S.A.
(Roth and Willis, unpublished data, 1954): Mite genus determined by Dr.
E. W. Baker (personal communication, 1954). The mites cluster thickly on
intersegmental membranes, particularly around the coxae and neck.
Despite a heavy infestation, the colony of _Nauphoeta_ thrived for
several years. This mite was found first on _N. cinerea_ and possibly
transferred to _D. punctata_ when the latter was brought into the
laboratory from Hawaii.
Family IOLINIDAE
=Iolina nana= Pritchard
_Natural hosts._--_Blaberus craniifer_ (originally from a culture at
Harvard University) and _Diploptera punctata_ (originally from Hawaii),
U.S.A., Pennsylvania (Roth and Willis, unpublished data, 1953;
Pritchard, 1956): The mites usually attached near the wing bases of the
insects. Morphologically, the species is intermediate between certain
predaceous and phytophagous mites (Pritchard, 1956).
Family PTERYGOSOMIDAE
=Pimeliaphilus podapolipophagus= Trägårdh
_Common name._--Cockroach mite.
_Natural hosts._--_Parcoblatta_ sp., U.S.A. (Edmunds, 1953a).
_Periplaneta americana_, U.S.A. (Piquett and Fales, 1952).
Cockroaches. U.S.A. (Baker and Wharton, 1952).
_Experimental hosts._--_Blatta orientalis_, _Blattella germanica_, and
_Periplaneta americana_, U.S.A. (Cunliffe, 1952).
Eggs of this mite (fig. 4) are usually laid indiscriminately in the
rearing cages, rarely on the host. Eggs are coated with a sticky
secretion which enables those laid on the host to adhere. Hatching
occurs in 6-11 days at 90-95° F., and in 9-11 days at 80° F. The newly
hatched larva starts to feed immediately on the cockroach. Larval stage
lasts 4-6 days, rests 2-3 days, and molts. During the single nymphal
instar, the mite feeds on the host and moves about for 6-7 days. The
mite then rests 3-4 days before molting. Entire life cycle covers a
period of 28-32 days. Adult mite lives 2-3 weeks, during which time it
can produce 2-3 batches of from 1 to 20 eggs; the usual batch is about
12 eggs. The mites are unable to live on cockroach feces, cast skins, or
dead cockroaches. Mites died within 4-5 days unless live cockroaches
were supplied. Parasitism was proved by detecting radioactivity in mites
that had fed on cockroaches which had been previously fed radioactive
NaCl (Cunliffe, 1952).
The mites can destroy laboratory cultures of cockroaches (Piquett and
Fales, 1952; Edmunds, 1953a). A cockroach attacked by 25 mites succumbed
after about an hour, falling on its back; it died after 5 hours
(Cunliffe, 1952).
When found in homes and offices, these mites are an indication of the
presence of cockroaches; the mite has been twice accused of biting
people (Baker et al., 1956).
RECORDS OF UNIDENTIFIED MITES
_Natural hosts._--_Aglaopteryx facies_, Puerto Rico (Seín, 1923): Four
red "tick" nymphs found under wings of female.
_Blaberus craniifer_, U.S.A., Florida (Hebard, 1917): "A number of lice
[mites] are present on many of these specimens [28[F][F]]."
_Blaberus discoidalis_, adventive from West Indies, taken in Scotland
(Stewart, 1925): A considerable number of mites were all over the body
and hind wings.
_Blatta orientalis_, Germany (Cornelius, 1853): Ex sexual organs of
male.
_Blattella germanica_, U.S.A., in laboratory (Parker, 1939): Under
conditions of high humidity, the cockroaches became heavily infested
with mites. In cages where the infestation was heavy, an abnormally
large number of females dropped their oöthecae, and the percentage of
eggs hatching was low.
_Parcoblatta uhleriana_, U.S.A., North Carolina (Hatcher, 1939): Hypopi
of mites were found deeply embedded in the fat body of two individuals.
Mites in the hypopial stage attach to insects by which they are
dispersed. Hypopi have been found in the gill chambers of a mollusk and
in the gonads of a millipede (Baker and Wharton, 1952).
_Periplaneta americana_, U.S.A., in laboratory (Fisk, 1951): The insects
were sluggish and molted with difficulty. Gold Coast Colony (Macfie,
1922): Larvae of a tarsonemid mite were found in the feces.
_Pycnoscelus surinamensis_, Hawaii (Illingworth, 1915): During the
summer the soil was literally swarming with young of various stages.
Early in September most of the adults were dead and all were covered
with mites. U.S.A., Connecticut, in laboratory (Zappe, 1918a). Hawaii,
in laboratory (Schwabe, 1950): Some of the cockroaches apparently died
from mite infestations.
[Illustration: FIG. 4.--The cockroach mite, _Pimeliaphilus
podapolipophagus_. (From Baker et al. [1956]; reproduced through the
courtesy of Dr. E. W. Baker and the National Pest Control
Association.)]
Cockroach, England? (Ealand, 1915): Cockroaches may carry the hypopial
stage of the cheese mite.
CONTROL OF MITES IN COCKROACH COLONIES
Fisk (1951) eliminated the mites [possibly _Pimeliaphilus
podapolipophagus_ (Baker et al., 1956)] in his cockroach colony by using
a 5-percent spray and a 5-percent dust of p-chlorophenyl,
p-chlorobenzene sulfonate. The exterior of the cockroach containers were
sprayed with the solution and the interior, including the insects, were
dusted. Within a month the mites had disappeared and the vigor of the
cockroach colony improved. Piquett and Fales (1952) used flowers of
sulfur and general sanitary procedures for eliminating the mites in
laboratory colonies of _Blatta orientalis_; they cleaned the dishes
every few days and applied grease around the edges of the containers to
prevent new mite invasions. Qadri (1938) employed similar control
measures.
Class CHILOPODA
Large centipedes which entered houses in India probably sought out
cockroaches (Maxwell-Lefroy, 1909). In Puerto Rico, centipedes entered
homes to which they were attracted by cockroaches (Seín, 1923). In
Hawaii, centipedes preyed on insects generally but especially on
cockroaches (Bryan, 1915). Sonan (1924) reported that in Formosa and
Okinawa Islands a species of centipede 5 to 6 inches long comes into the
houses and devours both adults and nymphs of _Periplaneta americana_ or
_P. australasiae_. Zimmerman (1948) found _P. australasiae_ breeding by
scores in rock piles in Hawaii accompanied by large numbers of
_Scolopendra_ and large spiders that probably preyed upon the
cockroaches.
Order SCUTIGEROMORPHA
Family SCUTIGERIDAE
=Scutigera coleoptrata= (Linnaeus)
_Synonymy._--_Scutigera forceps_ Rafinesque [Crabill, 1952].
_Common name._--House centipede.
_Natural prey._--Cockroaches, U.S.A. (Felt, 1909; Back, 1947; Auerbach,
1951; Crabill, 1952; and others): This predator-prey relationship seems
to be based on good circumstantial evidence (Crabill, personal
communication, 1953).
_Experimental prey._--_Blattella germanica_, newly hatched nymphs and
adult female, U.S.A. (Snodgrass, 1930; Roth and Willis, unpublished
data, 1953).
_Periplaneta americana_, U.S.A. (Roth and Willis, unpublished data,
1953).
_Supella supellectilium_, U.S.A. (Roth and Willis, unpublished data,
1953): See plate 31.
Our specimen caught a small American cockroach nymph that we placed in
its jar. Before it had finished its meal, it caught and held two other
nymphs with its legs while it continued to feed on the first. The body
of this centipede reaches a maximum length of 27 mm. and it is usually
found in basements, dark corners, or in spaces in the walls (Auerbach,
1951). Introduced from Europe, this species is now widespread in the
United States (Crabill, 1952).
=Allothereua maculata= (Newport)
_Synonymy._--_Scutigera maculata_ [Crabill, personal communication,
1957].
_Natural prey._--Cockroaches, Malay peninsula, Batu caves (Ridley in
Annandale et al., 1913): This is a presumptive host record.
Order SCOLOPENDROMORPHA
Family SCOLOPENDRIDAE
=Scolopendra cingulata= Latreille
_Experimental prey._--Cockroaches, England (Cloudsley-Thompson, 1955):
After capture in France, this specimen was kept for four weeks without
food. She was then fed medium-sized nymphal cockroaches of which she ate
an average of about one per week throughout the summer. Adult
cockroaches were attacked only after they had been disabled.
=Scolopendra morsitans= Linnaeus
_Natural prey._--Cockroaches, Guadeloupe (Lherminier, 1837).
_Experimental prey._--Cockroaches, India, Nagpur (Jangi, 1955): As soon
as the centipede became aware of its prey, it rapidly embraced the
cockroach within its legs and with its fangs gripped the insect's
thorax. The predator continued to hold the prey with its fangs while its
mouth parts prodded the victim's body. After feeding on an adult
cockroach, the centipede is not inclined to kill another for 2-3 days.
=Scolopendra subspinipes= Leach
_Natural prey._--Cockroaches, Hawaii (Williams et al., 1931): This is a
common species with a body length of 6 or more inches. It is reported to
be a great enemy of cockroaches.
=Scolopendra= sp.
_Natural prey._--_Ectobius panzeri_, England (Lucas, 1911, 1920): When
captured, the centipede was holding a live cockroach which it had
apparently just caught. The insect was held beneath its captor's body,
ventral surface upward, by several of the anterior legs while the
centipede fed.
Class INSECTA
We have found representatives of only 10 orders that have preyed on or
parasitized cockroaches: Beetles, flies, bugs, ants, wasps, stylops, and
cockroaches occurred in nature; the others resulted from feeding
cockroaches to captive insects or were laboratory observations.
Order ODONATA
Family AESHNIDAE
=Anax strenuus= Hagen
_Common name._--Giant Hawaiian dragonfly.
_Experimental prey._--Cockroaches, Hawaii (Williams, 1936): The
dragonfly nymph was fed with medium large cockroaches and other insects.
Order BLATTARIA
In this chapter the relations of other arthropods to cockroaches are
either as parasites or as predators. Certain cockroaches have turned the
tables on their adversaries and become predators themselves. This aspect
of cockroach behavior is discussed in chapter XVI. Other associations of
cockroaches, as commensals with other insects and as associates of other
cockroaches, are discussed in chapters XV and XVII.
Order ORTHOPTERA
Family MANTIDAE
=Hierodula tenuidentata= (Saussure) (?) (Serville)
(Pl. 32)
_Experimental prey._--_Blatta orientalis_, _Diploptera punctata_,
_Eurycotis floridana_, _Leucophaea maderae_, _Nauphoeta cinerea_,
_Neostylopgya rhombifolia_, and _Periplaneta americana_, U.S.A.
(Rilling, personal communication, 1957): Mrs. Rilling wrote us that with
the exception of _N. rhombifolia_, all the above cockroaches were
readily eaten. All the mantids initially rejected _N. rhombifolia_ after
grasping and making a brief attempt to chew the cockroaches. However, if
specimens of _N. rhombifolia_ were left in the jars with the mantids,
the cockroaches were usually eaten within the next 24 hours. _N.
rhombifolia_ ejects an odorous substance when seized and the mantids
probably ate these insects after most of this secretion had been
depleted. It is highly probable that the secretion of _N. rhombifolia_
may deter the mantid's attack, but it should be pointed out that, with
the possible exception of _N. cinerea_, all the other species fed to
these mantids give off odorous substances when seized or disturbed.
Apparently, certain naturally repellent compounds will deter this
mantid, whereas others that are presumed to be repellent will not;
however, the nutritional state of the mantid is undoubtedly a factor
which may limit the effectiveness of certain repellent secretions
against this predator.
_Byrsotria fumigata_, teneral males, and _Periplaneta australasiae_,
nymphs, U.S.A. (Roth and Willis, unpublished data, 1958).
_Diploptera punctata_, U.S.A. (Eisner, 1958).
=Mantis religiosa= Linnaeus
_Common name._--European mantis.
_Experimental prey._--_Nauphoeta cinerea_, and _Periplaneta americana_,
U.S.A. (Rilling, personal communication, 1957).
=Metallyticus semiaeneus= Westwood
_Experimental prey._--Cockroaches, Borneo (Shelford, 1916).
=Sphodromantis viridus= (Forskål)
_Synonymy._--_Sphodromantis bioculata_ Burmeister [Gurney, personal
communication, 1958].
_Experimental prey._--_Blatta orientalis_, Egypt (Adair, 1923): This
species of cockroach was apparently used regularly as food for the
mantid in the laboratory.
=Stagmomantis carolina= (Johansson)
_Common name._--Carolina mantis.
_Experimental prey._--_Blattella germanica_ and _Periplaneta americana_,
U.S.A. (Breland, 1941): The mantids were fed 1-2 German cockroaches
daily. One female mantid consumed 10 adult German cockroaches plus one
oötheca and part of another in 2.5 hours. An adult German cockroach was
consumed in an average of 8.5 minutes (range 5.5-15 minutes).
_Blatta orientalis_, nymphs, and _Diploptera punctata_, U.S.A. (Roth and
Willis, unpublished data, 1953).
=Tarachodes maurus= (Stal)
_Experimental prey._--Cockroaches, South Africa (Faure, 1940).
=Tenodera aridifolia sinensis= Saussure
_Common name._--Chinese mantis.
_Experimental prey._--_Nauphoeta cinerea_ and _Periplaneta americana_,
U.S.A. (Rilling, personal communication, 1957).
Family GRYLLACRIDIDAE
=Diestrammena apicalis= Br. v. Wattenwyl
and
=Diestrammena japanica= Blatchley
_Natural prey._--Cockroach eggs, Japan (Asano, 1937): These are
questionable records. Asano found _D. apicalis_ and _D. japanica_
beneath his house near several empty cockroach oöthecae which appeared
to have been eaten into. He assumed from the condition of the oöthecae
and the proximity of the stone crickets that the insects had devoured
the cockroach eggs.
_Experimental prey._--Eggs of _Blattella germanica_ and _Periplaneta
japanica_, Japan (Asano, 1937): Seven eggs of _B. germanica_ (obtained
from an oötheca being carried by a female) and eggs of _P. japanica_
(presumably in oöthecae) were fed to both species of stone crickets in
the evening. The eggs were devoured by the next morning.
Order DERMAPTERA
Family FORFICULIDAE
=Undetermined earwigs=
_Experimental prey._--Cockroaches, France (Chopard, 1938): According to
Chopard, Brisout de Barneville in 1848 indicated that earwigs in
captivity can be fed small cockroaches.
Order HEMIPTERA
Family LYGAEIDAE
=Clerada apicicornis= Signoret
_Natural prey._--Cockroach, Hawaii (Illingworth, 1917): This predaceous
bug is commonly found about buildings. Illingworth says that Kirkaldy
suspected that it fed on small blattids and that Dr. Perkins saw it
feeding on a dead cockroach.
Family REDUVIIDAE
=Spiniger domesticus= Pinto
_Natural prey._--_Periplaneta americana_, Brazil, Matto Grosso (Pinto,
1927, 1927a): This bug preys principally on cockroaches and was observed
infesting the walls of dwellings where it preyed on _P. americana_.
=Triatoma arthurneivai= Lent and Martins
_Natural prey._--_Monastria_ sp., Brazil, Minas Gerais (Martins, 1941):
This bug probably feeds on cockroaches of this genus, as well as on
rodents.
=Undetermined reduviids=
_Natural prey._--_Arenivaga roseni_ and _Polyphaga saussurei_, Turkmen
S.S.R. (Vlasov and Miram, 1937): These desert cockroaches are found in
burrows of rodents and desert turtles around Ashkhabad. Reduviids are
their main enemies. Vlasov (1933) found nymphs of _Reduvius christophi_
Jak. and _R. fedtschenkianus_ Osch. in similar burrows in this same
area, although he did not specifically cite them as enemies of the
desert cockroaches.
Family NEPIDAE
=Ranatra= sp.
_Experimental prey._--Cockroaches, U.S.A. (Hoffman, 1924).
Order NEUROPTERA
Family ASCALAPHIDAE
=Undetermined larva=
_Experimental prey._--_Blattella germanica_, Kenya Colony (Someren,
1924).
Order DIPTERA
From the few observations that have come to our attention, it seems that
flies are comparatively rare parasites in cockroaches.
Family PHORIDAE
=Megaselia= sp.
_Host._--Eggs of _Parcoblatta_ sp., Ohio (Edmunds, 1952a).
Family CONOPIDAE
=Stylogaster stylata= (Fabricius)
_Hosts._--Cockroaches, Brazil (Souza Lopes, 1937): L. Travassos was
quoted as having observed this species pursue cockroaches that were
escaping columns of the army ant _Eciton_ sp. Souza Lopes (1937) stated
that the female deposits eggs on the cuticle of the host near the end of
the body; the egg is barely inserted and two recurrent hooks prevent it
from falling off. Souza Lopes (1937) also observed other species of
_Stylogaster_ pursue Orthoptera, but he was unable to devote proper
attention to the behavior of the flies.
=Stylogaster= spp.
_Hosts._--_Chorisoneura_ sp., Brazil (Souza Lopes, 1937): An adult
specimen was found in a museum collection with an egg of _Stylogaster_
attached to the posterior end of its abdomen.
Cockroaches, Panama (C.W. Rettenmeyer, personal communication, 1959):
"Seven species were collected hovering over army ant swarms and a few
flies were seen apparently attacking cockroaches that had been flushed
by the ants."
Family LARVAEVORIDAE
=Calodexia= (?) =venteris= Curran
_Hosts._--_Periplaneta americana_, Brazil (Souza Lopes, 1937): Obtained
complete evolution of the parasite in this host. This may have been an
experimental host.
=Calodexia= spp.
_Hosts._--Cockroaches, Panama (Rettenmeyer, personal communication,
1959): Swarms of army ants are accompanied by about 20 species of
_Calodexia_. These flies larviposit on the cockroaches, crickets, and
possibly other arthropods that are flushed from cover by the ants.
Larvae were found in one(?) cockroach. Larvae from an adult of
_Calodexia_ were introduced experimentally into a cockroach and
successfully reared.
=Undetermined tachinids=
_Hosts._--_Eurycotis floridana_, from Florida (Roth, unpublished data,
1953): Three larvae (det. by W.W. Wirth) were found in a living adult
male.
_Panesthia australis_, from Australia (Roth, unpublished data, 1957):
Reared from a wild-caught cockroach that was maintained in a laboratory
colony.
Cockroaches, Australia (E. F. Riek, personal communication, 1955):
Reared from some of the larger species.
Family MUSCIDAE
=Coenosia basalis= Stein
_Host._--Eggs of _Parcoblatta_ sp., Ohio (Edmunds, 1952a).
Family SARCOPHAGIDAE
=Sarcophaga omani= Hall
_Host._--_Arenivaga bolliana_, Texas (Wirth, personal communication,
1953): Specimens in U.S. National Museum.
=Sarcophaga lambens= Wied.
_Synonymy._--_Sarcophaga sternodontis_ (Towns.).
Hoffman (1927) claimed that approximately 40 percent of some specimens
of _Pycnoscelus surinamensis_ collected in southern Haiti were
parasitized by _S. lambens_. However, according to entomologists at the
University of Puerto Rico Agricultural Experiment Station, Hoffman was
incorrect in his observations: _S. lambens_ was never reared from a
living insect and had been recovered only from dead cockroaches and
other dead insects and was considered saprophytic rather than parasitic
(Schwabe, 1950b).
=Sarcophaga= spp.
Sanjean (1957) reared various species of sarcophagid larvae on
_Periplaneta americana_ which were freshly killed or chopped up; first
instar larvae were also introduced into the body cavity of cockroaches
which had their heads and legs removed. Adult sarcophagids were
collected and freshly killed American cockroaches used as bait.
Order COLEOPTERA
Family CARABIDAE
=Harpalus pennsylvanicus= De Geer
_Experimental prey._--_Cryptocercus punctulatus_, U.S.A. (Cleveland et
al., 1934): This beetle is often found in the galleries of _C.
punctulatus_ in nature. In the laboratory it killed and devoured
cockroaches as large as itself.
Family DYTISCIDAE
=Rhantus pacificus= Boisduval
_Experimental prey._--Cockroaches, disabled, Hawaii (Williams, 1936):
This beetle, which is common in mountain streams, located wounded
cockroaches in an aquarium by sense of smell or taste rather than sight.
Family LAMPYRIDAE
=Undetermined larva=
_Experimental prey._--_Parcoblatta virginica_, adult female (pl. 33, C),
U.S.A. (Roth and Willis, unpublished data, 1953).
Family RIPIPHORIDAE[5]
=Neonephrites partiniger= Riek
_Natural host._--Cockroach (undescribed genus belonging to the
Pseudomopinae), Australia Capital Territory (Riek, 1955).
=Neorhipidius neoxenus= Riek
_Natural host._--_Robshelfordia longiuscula_ or _Robshelfordia
circumducta_, Australia Capital Territory (Riek, 1955).
=Paranephrites xenus= Riek
_Natural host._--_Oniscosoma granicollis_, Australia Capital Territory
(Riek, 1955).
=Rhipidioides ableptus= Riek
_Natural host._--_Balta patula_, Australia, Victoria (Riek, 1955): Pupal
stage lasted only 3 days.
=Rhipidioides adynatus= Riek
_Natural host._--_Escala_ sp. or an undescribed genus of Pseudomopinae,
Australia, Victoria (Riek, 1955).
=Rhipidioides fuscatus= Riek
_Natural host._--_Ellipsidion affine_, Australia, New South Wales (Riek,
1955).
=Rhipidioides helenae= Riek
_Natural host._--_Robshelfordia longiuscula_ or _Robshelfordia
circumducta_, Australia Capital Territory (Riek, 1955).
=Rhipidioides mollis= Riek
_Natural host._--_Robshelfordia longiuscula_ or _Robshelfordia
circumducta_, Australia Capital Territory (Riek, 1955).
=Rhipidioides rubricatus= Riek
_Natural host._--_Choristima_ sp. and _Choristimodes_ sp., Australia
Capital Territory (Riek, 1955).
=Riekella australis= (Riek)
_Synonymy._--_Nephrites australis_ Riek [Selander, 1957].
_Natural host._--_Cutilia_ sp., Australia Capital Territory (Riek,
1955): Two females emerged from one host.
=Riekella nitidioides= Selander
_Synonymy._--_Nephrites nitidus_ of Riek not Shuckard [Selander, 1957].
_Natural host._--_Platyzosteria_ sp., Tasmania (Riek, 1955).
=Riekella= sp.
_Synonymy._--_Nephrites_ sp. [Selander, 1957].
_Natural host._--_Platyzosteria castanea_, Australia Capital Territory
(Riek, 1955).
_Biology of Australian Ripidiini._--The Australian species of Ripidiini
are parasites of apparently endemic, ground-dwelling species of
cockroaches. There is some correlation between host subfamily and
parasite genus: _Riekella_ spp. [= _Nephrites_] have only been bred from
Blattinae. _Rhipidioides_ spp. occur only in the closely related
Ectobiinae and Pseudomopinae. _Neonephrites_ and _Neorhipidius_ also
occur in the Pseudomopinae. _Paranephrites_ occurs in the Panchlorinae.
There is some evidence that the parasitized cockroaches migrate onto
trees when the larval parasite is mature, as pupae have only been found
on the trunks of eucalyptus trees. In all species the larva leaves the
host dorsally through an intersegmental membrane. The host continues to
live for a few days after the parasite emerges. The larva attaches
itself to bark on the tree trunk by a few strands of silk before
pupating. The larviform, wingless female remains near the pupal skin and
is sought out by the winged male. The eggs are laid in a mass around the
pupal skin (Riek, 1955).
=Ripidius[5] boissyi= Abeille
Balduf (1935) lists _Ripidius boissyi_ as parasitic on nymphs of
_Ectobius pallidus_ giving Abeille de Perrin (1909) as a source for
this information. However, Abeille de Perrin simply presumed that _R.
boissyi_ parasitized _E. pallidus_ because he collected this cockroach
in the same habitat as the beetle. Abeille de Perrin suggested that the
species of the genus _Ripidius_ lived in the bodies of cockroaches, but
there are no rearing records, as far as we know, of _R. boissyi_ from
cockroach hosts.
=Ripidius denisi= Chobaut
Chobaut (1919), in France, collected both _R. denisi_ and _Ectobius
pallidus_ when beating an oak tree. Because of the known association of
other species of _Ripidius_ with cockroaches, he presumed that this
beetle was parasitic on _E. pallidus_, a cockroach common in this
beetle's habitat.
=Ripidius pectinicornis= Thunberg
_Synonymy._--_Symbius blattarum_ Sundevall [Leng, 1920].
_Natural hosts._--_Blattella germanica_, on shipboard (Sundevall, 1831);
Germany (Aclogue and Fowler, _in_ Burr, 1899a); on steamship "Samui"
(Stamm, 1936); on cruiser "Duguay-Trouin" (Barbier, 1947); Hawaii
(Williams, 1946a): This last record was based on a specimen dissected
from an adult German cockroach collected on an airplane from the South
Pacific. The parasite was reported as _Ripidius_ sp. by Williams, but
Weber (1948) made the specific identification.
_Ectobius pallidus_? Abeille de Perrin (1909) stated that _R.
pectinicornis_ was first described by Sunders [sic] as _blattarum_
because it had been captured in the body of _Ectobia livida_. We presume
that Abeille de Perrin was referring to Sundevall's work in which the
host was given as _Blattella germanica_.
_Periplaneta americana_, on shipboard (Sundevall, 1831): One nymph only.
With the exception of the single nymph of _P. americana_, _R.
pectinicornis_ apparently attacks only adult females and nymphs of _B.
germanica_. Barbier (1947) found only _B. germanica_ parasitized,
although both _Blatta orientalis_ and _Supella supellectilium_ were
prevalent on board the ship. Primary larvae of the parasite failed to
parasitize _Supella_.
_Adult behavior._--The winged male is relatively active compared to the
apterous female; it runs around, flies well, and jumps on the female
when in her vicinity. The female remains stationary and lays eggs around
her by bending her long ovipositor (Sundevall, 1831). The eggs (50-100)
are laid among a network of silk fibers secreted by the female. The
female dies after completing oviposition (Barbier, 1947).
_Development._--The eggs hatch after 14 days, and the primary
(triungulin) larvae ascend the host's legs to its body; the larvae then
cut the intersegmental membrane between the metasternum and first
abdominal segment of the cockroach, in order to enter the host's abdomen
(Barbier, 1947). Chobaut (1892) first suggested this method of attack by
the ripiphorid larva. As the parasites develop, the abdomen of the host
becomes swollen. Developing larvae apparently eat the host's fat body,
leaving the vital organs until the last. Parasitized female hosts were
sterile and the eggs, when formed, never hatched. Development of the
oötheca was also inhibited. There were usually two larval parasites per
host, but three or four were found several times (Barbier, 1947).
Sundevall (1831) found only one larva per cockroach except one host
which, when crushed, yielded five. Stamm (1936) found three hosts
infested with five larvae each. In a little over 100 cockroaches, Stamm
found 10 that were parasitized.
The day before the parasite leaves the host, the cockroach shows an
abrupt uneasiness and runs about, finally falling over on its back. The
parasite larva emerges from the host through an opening it makes in the
membrane between penultimate and last tergite. The host dies a few hours
after the larva has left. The larva seeks a sheltered area and pupates
within 48 hours. Adults emerge in 9 days (females) and 13 days (males)
(Barbier, 1947).
_Distribution._--Adult males have been collected in light traps in
Hawaii (Van Zwaluenburg, 1946), and the first female was reported by
Weber (1948); the parasite is now established in the islands around
Pearl Harbor (Dr. F. X. Williams, personal communication, 1953). The U.
S. National Museum has specimens of _R. pectinicornis_ from England,
Guatemala, Hawaii, Panama, and from Florida and Georgia in the U. S.
(Dr. E. A. Chapin, personal communication, 1953). Kono (_in_ Asano,
1937) reported two species in Japan. It is noteworthy that all these
records are from localities adjacent to oceans and on ships; none are
from interiors of continents. The only biological data were obtained
from parasites found on board ships. Sundevall (1831) believed that the
parasites boarded his ship with their hosts during loading in Calcutta,
since before that not any were seen on board. Barbier (1947) suggested
that the parasite must be spread very easily in ports between
neighboring ships by parasitized cockroaches in baskets or sacks of
provisions.
=Ripidius scutellaris= Heller
_Natural hosts._--Blattidae, Philippine Islands (Schultze, 1925).
Family DERMESTIDAE
=Dermestes ater= De Geer
_Common name._--Black larder beetle.
_Natural prey._--_Blatta orientalis_, U.S.A. (Roth and Willis,
unpublished data, 1953): _Dermestes ater_ is generally a scavenger, but
we have seen adult beetles, which had developed in our cockroach colony,
clinging to and feeding on living oriental cockroaches, eventually
killing them; the beetles probably attack only the weakened or injured
cockroaches in a culture. This was a natural infestation of a laboratory
culture by a predator.
_Experimental prey._--_Blattella germanica_, oöthecae, U.S.A. (Roth and
Willis, 1950): The beetle larvae can penetrate unhatched oöthecae of the
German but not those of the American or oriental cockroaches.
=Dermestes= sp.
_Natural prey._--_Blatta orientalis_, oöthecae, U.S.A., Missouri: Rau
(1924) stated that _Dermestes_ larvae often infest the egg cases of this
cockroach; it is probable that Rau was referring to cockroaches in
laboratory cultures.
Order STREPSIPTERA
Pierce (1909) predicted that the Blattoidea and the Grylloidea would be
the only groups of the Orthoptera which would be parasitized by
Strepsiptera. Essig (1926) made the statement that certain cockroaches
are among the hosts of Strepsiptera. E. F. Riek (personal communication,
1952) found a strepsipteron in a late nymph of _Cutilia_ sp. from
Waroona, Western Australia; he wrote us, "The female parasite is
extruded between a pair of sternites towards the base of the abdomen and
appears to belong to the family Halictophagidae." This is the only
record that we have been able to find of a strepsipteron parasitizing
cockroaches.
Order HYMENOPTERA
PREDATORS AND PARASITES OF COCKROACH EGGS
Wasps from at least six families of Hymenoptera have been recorded as
developing on cockroach eggs. All the Evaniidae are presumed to be
parasitic in the egg capsules of cockroaches (Clausen, 1940; Townes,
1951), although hosts for many of the described species have yet to be
discovered. The presence of evaniids in dwellings indicates the presence
of cockroaches (Gross, 1950). At times these wasps may become a
nuisance; a family in Worthington, Ohio, complained of the evaniid wasps
that they found on the windows and in other areas of their home, but
they were apparently not annoyed by the oriental cockroaches in the
basement (Edmunds, 1953).
The known parasites of cockroach eggs are listed below with summaries of
their biology.
Family EVANIIDAE
=Acanthinevania princeps= (Westwood)
_Synonymy._--_Evania princeps_ [Dr. H. Townes, personal communication,
1956].
_Natural host._--Cockroach eggs, Australia (Froggatt, 1906).
=Brachygaster minutus= (Olivier)
_Synonymy._--_Evania minuta_ Olivier [Kieffer, 1920].
_Natural hosts._--_Blattella germanica_, Europe? (Schletterer, 1889;
Kiefer, 1912; Crosskey, 1951.)
_Ectobius lapponicus_, Europe? (Schletterer, 1889; Kieffer, 1912;
Crosskey, 1951.)
_Ectobius panzeri_ var. _nigripes_? Great Britain (Blair, 1952): This is
a presumptive record. The wasp was collected at Niton and Headon Hill,
Isle of Wight, an area in which this variety of _E. panzeri_ was the
only species of cockroach known to occur.
_Ectobius_ sp., England (Cameron, 1955, 1957): Natural History Museum
records.
Adult wasps have been collected on _Asparagus officinalis_ Linnaeus
(Schmiedeknecht _in_ Schletterer, 1889; Crosskey, 1951). Thompson's
(1951) citation of records of _B. minutus_ and _Evania appendigaster_
from _Blatta orientalis_ and _Blattella germanica_, and Cameron's (1957)
citation of these records and one from _Ectobius lapponicus_, all
attributed to Kadocsa (1921), are almost certainly in error. Kadocsa
(1921, p. 33) listed these wasps as egg parasites of cockroaches but not
necessarily in Hungary and did not name specific cockroach hosts.
The present writers have found no information, other than host reports,
on the biology of _Brachygaster minutus_. The records of this wasp
parasitizing _B. germanica_ may trace back to Schletterer, but his
listing may not have been an original observation. Since the female of
_B. germanica_ carries its oötheca attached to the abdomen until or just
before the eggs hatch, it would seem that the female of _B. minutus_ (if
the host records are valid) must oviposit into the oötheca of this
species while it is still being carried by the female; this would not
necessarily be true for the other hosts which drop the egg case long
before the eggs hatch.
_Distribution._--Europe: Sweden, Russia, England, France, Germany,
Austria, Hungary, Switzerland, Italy (Kieffer, 1920).
=Evania appendigaster= (Linnaeus)
_Synonymy._--_Evania desjardinsii_ Bordage, _Evania laevigata_ Latreille
[Dalla Torre, 1901-1902].
_Natural hosts._--_Blatta_, "exotic species" (Westwood, 1854, 1954a).
_Blatta orientalis_, Europe? (Schletterer, 1886; Howard, 1888, Kieffer,
1912); Egypt? (Alfieri, 1914; Adair, 1923). [Girault (1907, 1914)
erroneously attributed another record to Marlatt (1902);[see footnote 6].
See also notes under _Brachygaster minutus_ with respect to Kadocsa.]
_Blattella germanica_? (Girault 1907, 1914). [This record is obviously
an error. Girault attributed the record to Marlatt (1902); see footnote
6.]
_Cutilia soror_, Hawaii (Swezey, 1929; Zimmerman, 1948).
_Leucophaea maderae_ (Schletterer, 1889; Bordage, 1896; Kieffer, 1912):
These records are probably erroneous inasmuch as this cockroach
incubates its eggs internally (Roth and Willis, 1954). Later, after
finding that _L. maderae_ is ovoviviparous, Bordage (1913) admitted
having misidentified a parasitized oötheca from some other species; he
concluded that the developing eggs of this species are protected against
egg parasites because they are carried within the female. Clausen
(1940), in classifying the placement of parasitic wasp eggs in relation
to the host, erected the category: Egg placed in the embryo while the
latter is still within the parent. He stated that although this behavior
was not definitely known to occur, it probably could occur. However, the
records cited above do not indicate that the alleged parasitization
followed this pattern.
_Neostylopyga rhombifolia_, Hawaii (Swezey, 1929).
_Periplaneta americana_, Europe (Schletterer, 1889; Bordage, 1896;
Kieffer, 1912); Réunion Island (Bordage, 1913); Puerto Rico (Seín,
1923); Jamaica (Gowdey, 1925); Hawaii (Swezey, 1929); Palestine
(Bodenheimer, 1930); U.S.A., Florida (Ashmead, 1900); Maryland (Piquett
and Fales, 1952); Saudi Arabia, Jedda (Cameron, 1957); Canton Island and
Samoa (Dumbleton, 1957).
_Periplaneta americana_ or _P. australasiae_, Formosa (Sonan, 1924).
_Periplaneta australasiae_, U.S.A., Florida (Ashmead, 1900); Hawaii
(Swezey, 1929; Zimmerman, 1948). [Girault (1914) erroneously attributed
another record to Marlatt (1902); see footnote 6, above.]
_Experimental host._--_Blatta orientalis_, U.S.A. (Haber, 1920).
Relatively little detailed information was known about this wasp (fig.
5), one of the earliest parasites of cockroach eggs to be discovered,
until Cameron (1957) studied its biology. Arnold (Kirby and Spence,
1826) discovered that the genus _Evania_ parasitized _Blatta_, but did
not know whether the wasp developed on the cockroach eggs or in the
nymphs. MacLeay (Westwood, 1843) determined that _Evania_ developed
within the oöthecae of cockroaches. Westwood (1854a) found the larvae,
pupae, and adults of _E. appendigaster_ in egg cases of an unidentified
species of cockroach found on orchids received from Calcutta.
_Adult behavior._--Adult wasps visited flowers of parsley, _Petroselium
crispum_, and fennel, _Foeniculum vulgare_ (Margretti _in_ Schletterer,
1886; Crosskey, 1951). In Hawaii the adult wasps have been seen resting
on leaves coated with honey dew (Williams et al., 1931); _Evania_ sp.
were attracted to the honey dew secreted by a diaspine scale insect
(Williams, 1931). Adults lived two to three weeks in captivity with
ample food and water (Cameron, 1957).
_Oviposition._--Shelford (1912, 1916) erroneously supposed that
_Evania_, by means of her cleaverlike abdomen, opened the oötheca at the
crista and then deposited her egg or eggs on the eggs of the cockroach.
Haber (1920) observed and described oviposition. The female wasp
crawled over the surface of the oötheca, actively vibrating her
antennae, and settled with the axis of her body parallel to the axis of
the egg case as it lay upon its right side. Lying on her right side, the
wasp extended her ovipositor and punctured the oötheca in the fifth cell
on the left side; she remained in this position for about 15 minutes.
Cameron (1957) described similar oviposition behavior that lasted about
half an hour. Kieffer (1912) and Crosskey (1951) stated that the female
deposits her eggs before the walls of the oötheca harden.
[Illustration: FIG. 5.--_Evania appendigaster._ Left, dorsal view, × 8.
Right, side view, × 5. (Reproduced with permission. British Museum
[Natural History], 1951, figs. 1A and 1B.)]
_Development._--Kieffer (1912) stated that the larvae in this family eat
the cockroach eggs and pupate in the oötheca without forming a cocoon.
Smith (1945) stated that the larva feeds on one cockroach egg after
another until all are destroyed; by that time it is full grown and it
pupates within the oötheca. Cameron (1957) found that there are five
larval instars and that in material from Saudi Arabia there are three or
possibly four generations a year.
_Distribution._--Tropical and subtropical parts of the world as far
north as New York City, and all of Europe except the northern part
(Kieffer, 1920; Townes, 1949). The wide distribution of _Evania_ has
been attributed to the abundance of host cockroaches on ships between
the Tropics (Haldeman, 1847). Kieffer (1903) appears to have shown some
correlation between the numbers of species of cockroaches found in
various geographical regions and the numbers of species of evaniids
found in similar regions. However, the number of blattids he listed is
small.
=Evania dimidiata= Fabricius
_Synonymy._--_Evania abyssinica_ Westwood [Schletterer, 1889].
_Natural host._--_Blatta orientalis_, Egypt? (Alfieri, 1914).
=Evania subspinosa= Kieffer
_Natural host._--_Periplaneta_ sp., Fiji (Lever, 1946): Although Lever
(1946) listed this species as a cockroach-egg parasite, he did not state
that he actually reared it from _Periplaneta_ oöthecae.
=Hyptia dorsalis= of Ashmead
_Synonymy._--Dr. H. Townes, (personal communication, 1956) believes that
this wasp was probably either _H. reticulata_, _H. harpyoides_, or _H.
thoracica_; it is not possible to tell which without reexamining
Ashmead's specimens; these apparently have been lost.
_Natural host._--_Parcoblatta pensylvanica_, U.S.A., Mississippi
(Ashmead, 1900).
=Hyptia harpyoides= Bradley
_Natural hosts._--_Parcoblatta virginica_, U.S.A., Ohio (Edmunds, 1952a,
1953a, 1954).
_Parcoblatta pensylvanica_, U.S.A. (Muesebeck, 1958).
_Parcoblatta uhleriana_, U.S.A., Natick, Mass.: Oötheca collected by L.
Roth, May 17, 1956; wasp emerged June 12, 1956 (pl. 33, B); determined
by Dr. H. Townes. The keel region of the oötheca of _P. uhleriana_ (pl.
18, B) is different from that of any other species of _Parcoblatta_
(Hebard, 1917; Lawson, 1954) so there can be no doubt as to the species
of cockroach parasitized by this wasp.
_Development._--The last instar larva overwinters inside the cockroach
oötheca (Edmunds, 1954). Five oöthecae yielded one parasite each
(Edmunds, 1953a).
_Distribution._--Canada, Ontario. U.S.A.: New Hampshire and Minnesota to
South Carolina, Mississippi, Texas, and Kansas. Upper and Lower Austral
Zones (Townes, 1951).
=Hyptia reticulata= Say
_Natural host._--_Parcoblatta pensylvanica_, U.S.A., Missouri (Rau,
1940).
Adult wasps have been taken on parsnip, _Pastinaca sativa_ (Robertson,
1928).
_Distribution._--U.S.A.: Pennsylvania to Florida and Louisiana. Mexico.
Upper Austral to Tropical Zones (Townes, 1951).
=Hyptia thoracica= (Blanchard)
_Natural host._--_Parcoblatta pensylvanica_, U.S.A., Ohio (Edmunds,
1952a, 1953a, 1954).
_Adult behavior._--Copulation was rapid, lasting only a few seconds.
Blooms of _Asmorrhiza longistylis_ were placed in a cage with adult
wasps. The insects were attracted to and fed on the flowers (Edmunds,
1954).
_Development._--Entire contents of oötheca are eaten by the single
larva. Last instar larva overwinters inside the oötheca. Emergence in
Ohio was around the middle of June. The emergence hole made by this
genus was about 2 mm. in diameter. The hole was made at the top side of
the oötheca near one end. Adult took about 65 minutes to emerge from the
time its mandibles first broke through the oöthecal wall. (Edmunds,
1954.)
_Distribution._--Canada, Ontario, U.S.A.: Connecticut to Wisconsin,
south to Florida and Texas. Upper Austral to Tropical Zones. (Townes,
1951.)
=Hyptia= sp.
_Natural host._--_Cariblatta delicatula_, Cuba (Hebard, 1916a); Parasite
identified by Ashmead.
=Hyptia= sp. (undescribed)
_Natural host._--_Parcoblatta_ sp., U.S.A., Ohio (Edmunds, 1952a).
=Prosevania punctata= (Brullé)
_Synonymy._--_Evania punctata_ Brullé [Townes, 1949].
_Natural and experimental hosts._--_Blatta orientalis_, Istrian
Peninsula (Fahringer, 1922); Algeria (Cros, 1942); U.S.A., Ohio
(Edmunds, 1954).
_Blattella germanica_? Europe? (Girault 1907, 1914); Europe (Fahringer,
1922). [The records on this host are extremely doubtful. Girault
erroneously cited Marlatt (1902) as the source of this record; see
footnote 6, page 236. Fahringer, however, claimed that he obtained
seven female parasites from oöthecae of _Blattella germanica_. He placed
female parasites with adults of _B. germanica_ in a glass cage. As soon
as oöthecae could be seen between folds of a woolen rag, he removed all
the larger cockroaches and held the oöthecae until the parasites
emerged. Fahringer may have been dealing with a different species of
cockroach, because placing oöthecae in crevices (or between folds of
rag) is a habit foreign to _B. germanica_, the female of which usually
carries her oötheca until hatching or until about a day before. Edmunds
(1953b) could not induce this wasp to parasitize eggs of _B.
germanica_.]
_Periplaneta americana_, Istrian Peninsula (Fahringer, 1922); Palestine
(Bodenheimer, 1930); U.S.A., Ohio (Edmunds, 1952, 1953b, 1954).
_Adult behavior._--The wasps (pl. 33, A) are very active; they walk
about a great deal and fly short distances. They are often found in
abundance in buildings infested with the larger domiciliary cockroaches
where they may reproduce for many generations without leaving the
premises. Specimens have also been collected outdoors. (Edmunds, 1953,
1954.) As the adult walks about, the laterally compressed abdomen moves
up and down like a waving flag; because of this behavior, these insects
are commonly known as ensign-flies. Cros (1942) maintained adults 17
days without food. Edmunds (1954) fed adults on unidentified flowers in
the laboratory. He also maintained them for 20 days after capture on a
5-percent honey solution.
_Oviposition._--A female _P. punctata_ selected oöthecae of _P.
americana_ for oviposition and ignored those of _B. orientalis_ and
_Parcoblatta pensylvanica_ in the same cage. Oviposition was
accomplished as described for _Evania appendigaster_. One oötheca was
turned over onto its right side by the wasp before she oviposited.
(Edmunds, 1952.) Although there seemed to be a "preferred" position for
oviposition, it was not obligatory. The usual position was for the
female to face the keel of the oötheca, but she also oviposited from the
opposite side or, rarely, directly down into the side of the oötheca.
The average time spent by females in 10 ovipositions was 29 minutes
(range 16-62 minutes). The wasp apparently could not determine whether
the eggs had been previously parasitized. The wasp laid her egg between
the cockroach eggs rather than in them and she oviposited into oöthecae
that had just been dropped and those two weeks old or older. On three
occasions nymphal cockroaches emerged within a few hours after the wasp
had oviposited. (Edmunds, 1954.) Apparently, for successful
parasitization the wasp must oviposit before the cockroaches have
reached the final stages of preemergence development. Edmunds (1954)
placed females of _Periplaneta americana_ that were carrying oöthecae,
into cages with _Prosevania_; some of the female wasps showed
considerable interest in the attached oöthecae, but he observed
oviposition only into egg cases that had been dropped by the
cockroaches.
Cros (1942) described an interesting reaction that he called
"instinctive hostility" of the oriental cockroach toward _Prosevania_. A
wasp was placed in a jar in which a cockroach had just deposited its
oötheca. The wasp tried to oviposit into the egg case but was upset and
pursued by the cockroach. The cockroach placed herself over the oötheca,
standing high on her legs, and remained there motionless. The wasp then
approached from the rear, slipped under the cockroach, and, unnoticed by
the cockroach, climbed on the oötheca and oviposited successfully.
_Development._--In _Blatta orientalis_: The developmental period was
completed in 40-57 days in summer and fall (Cros, 1942). Time from
oviposition to emergence of adult varied from 45-177 days; three
parthenotes from an oviposition by an unfertilized female wasp developed
in 45-53 days (Edmunds, 1954). In _Blattella germanica_: Almost 4 weeks
spent in development (Fahringer, 1922). In _Periplaneta americana_:
Three wasps developed in 127 days (Edmunds, 1952). Only one parasite
develops in each oötheca. There were three generations a year in Ohio.
(Edmunds, 1954.) In Algeria there were two to three generations per
year. The adult emerged from the oötheca through a hole 4 mm. in
diameter. (Cros, 1942.) Parthenogenesis exists; the unfertilized eggs
produced only males (Edmunds, 1954).
_Distribution._--Eastern U.S.A., from New York and Ohio south to Georgia
(Townes, 1949). Europe, Syria, Palestine (Kieffer, 1920).
=Szepligetella sericea= (Cameron)
_Synonymy._--_Evania sericea_ Cameron [Townes, 1949, personal
communication, 1956]. _Evania impressa_ Schletterer [Townes, p. c.,
1956].
_Natural hosts._--_Cutilia soror_ and _Neostylopyga rhombifolia_, Hawaii
(Swezey, 1929).
_Periplaneta americana_ and _Periplaneta australasiae_, Hawaii (Swezey,
1929; Zimmerman, 1948).
_Periplaneta_ sp., Fiji (Lever, 1943, 1946).
Adults are sometimes found resting on leaves covered with honey dew
(Williams et al., 1931).
=Zeuxevania splendidula= Costa
_Natural hosts._--_Loboptera decipiens_, France (Lavagne, 1914; Genieys,
1924).
Picard (1913) believed that _Z. splendidula_ parasitized _L. decipiens_
and not its eggs; however, Lavagne (1914) explained the true
relationship by dissecting two specimens of _Z. splendidula_ from
oöthecae of _L. decipiens_.
The following information is taken from Genieys (1924):
_Oviposition._--Wasp egg is introduced into the still-soft oötheca
before the wall hardens. Some oöthecae had four oviposition scars but
never contained more than two parasite eggs. _Development._--Larva
commences development in July or August. Only one larva completes
development, but it eats all the eggs in the oötheca. The wasp passes
the winter as a last instar larva and pupates in the spring; the adult
emerges during the spring or in June. _Hyperparasitism._--About 10
percent of the oöthecae of _Loboptera decipiens_ that were parasitized
by _Z. splendidula_ were also hyperparasitized by an eulophid (see
_Syntomosphyrum ischnopterae_, p. 249).
Family CLEONYMIDAE
=Agamerion metallica= Girault
_Natural hosts._--_Ellipsidion australe_, Australia, Queensland (Dodd,
1917): "the parasite when ready to emerge fully occupies the whole space
of the destroyed eggs."
Cockroach, Australia, New South Wales (Dr. B. D. Girault, 1915a).
Family ENCYRTIDAE
=Blatticida pulchra= Ashmead
_Natural host._--Cockroach eggs on orange leaves, Australia, New South
Wales (Gahan and Peck, 1946). According to Dr. A. B. Gurney the oötheca
associated with the type specimens of the wasps in the United States
National Museum is possibly _Balta_ sp. (Burks, personal communication,
1956).
=Blatticidella ashmeadi= (Girault)
_Synonymy._--_Blatticida ashmeadi._ _Blatticida_ Girault, 1915, is
preoccupied by _Blatticida_ Ashmead, 1904. In 1923 Gahan and Fagan
renamed _Blatticida_ Girault, _Blatticidella_. [Burks, p. c., 1956.]
_Natural host._--Cockroach, Australia, Queensland (Girault, 1915).
=Cheiloneurus viridiscutum= (Girault)
_Synonymy._--_Cristatithorax_ Girault = _Cheiloneurus_ Westwood [Mercet,
1921].
_Natural host._--_Ellipsidion australe_, Australia, Queensland (Dodd,
1917).
=Comperia merceti= (Compere)
_Synonymy._--_Comperia merceti_ var. _falsicornis_ Gomes [Peck, 1951].
_Natural hosts._--_Blattella germanica_, Brazil, Distrito Federal
(Gomes, 1941): In the English summary of his paper, Gomes states that
_C. merceti_ var. _falsicornis_ was reared from _B. germanica_. However,
in the body of the paper, he states that the _supposed_ origin of the
parasite was the oötheca of _B. germanica_. Burks (personal
communication, 1956) does not believe that this wasp parasitizes the
eggs of _B. germanica_. We (unpublished data, 1957) exposed six oöthecae
of _B. germanica_ to _C. merceti_. In order to retard water loss the
oöthecae were removed from the females by cutting the insects in two so
that each oötheca remained attached to the posterior part of the
abdomen. No wasps developed in these oöthecae.
_Supella supellectilium_, U.S.A., Kansas (Lawson, 1954a); Hawaii
(Zimmerman, 1944; Compere, 1946; Keck, 1951).
_Adult behavior._--Males and nonovipositing females showed a flea-like
jumping tendency. Adults were attracted to light and were found near
windows. Both sexes pursued an erratic course in walking and continually
touched the surface with their antennae. (Lawson, 1954a.)
_Oviposition._--The wasp (pl. 34, B) selected a site on an oötheca with
the sheath of her ovipositor; it was uncertain whether there was a
definite preference for oviposition sites. Wasp tended to choose a
nearly horizontal position for oviposition. She preferred to oviposit
into eggs about 2 weeks old, although she would place eggs in oöthecae
less than a week old and in embryos in the green band stage. There were
1-50 oviposition punctures per oötheca. (Lawson, 1954a.)
_Development._--If enough wasp larvae were present, they ate all eggs in
an oötheca. Occasionally wasps developed in one end of an oötheca while
cockroaches developed in the other; when this occurred, the cockroach
nymphs always emerged last. The developmental period was 30-41 days at
room temperature. There were 5-25 parasites per oötheca. The single exit
hole in the oötheca varied from 0.6 to 0.9 mm. in diameter. (Lawson,
1954a.)
_Distribution._--U.S.A.: New Jersey south to Florida, west to Illinois,
Kansas, and Arizona. West Indies; Central and South America; Hawaii.
(Burks, personal communication, 1956.)
=Dicarnosis alfierii= Mercet
_Natural hosts._--"_Phyllodromia_" sp., Egypt (Mercet, 1930): According
to Mercet, Dr. Alfieri claimed that this wasp parasitized one of the
species of "_Phyllodromia_" found in Egypt, namely, _Phyllodromia_
[= _Blattella_] _germanica_, _Phyllodromia_ [= _Supella_] _supellectilium_
and/or _Phyllodromia treitliana_. We do not know to which modern genus
the host of this wasp belonged.
Cockroach, Egypt? (Mercet in Compere, 1938.)
=Eutrichosomella blattophaga= Girault
_Natural host._--Cockroach, Australia, Queensland (Girault, 1915).
Family EUPELMIDAE[7]
=Anastatus blattidifurax= Girault
_Natural host._--Cockroach, Australia, Queensland (Girault, 1915).
=Anastatus floridanus= Roth and Willis
_Natural host._--_Eurycotis floridana_, U.S.A., Florida (Roth and
Willis, 1954a).
_Experimental hosts._--_Blatta orientalis_, _Eurycotis floridana_, and
_Periplaneta americana_, U.S.A. (Roth and Willis, 1954a).
_Adult behavior._--Female wasps are sexually receptive almost
immediately on leaving the oötheca. Mating takes 3-4 seconds. Males mate
repeatedly and may fertilize several females; females may also mate more
than once. At about 80° F. the female wasps lived 2-4 days, males one
day.
_Oviposition._--The female wasp first probes the oötheca with her
sheathed ovipositor until she finds an acceptable spot; she then drills
through the wall of the oötheca with her ovipositor. One female
oviposited for 5 hours, but briefer periods were more usual. We have
seen six or more females ovipositing simultaneously into an oötheca of
_Eurycotis floridana_. One female was seen to feed on material that
oozed from the oviposition puncture. The wasp (pl. 34, A) may oviposit
into the oötheca of _E. floridana_ while it is still being carried by
the female, as well as in oöthecae that have been dropped and which have
hard walls. Eggs 36 days old were successfully parasitized.
_Development._--In _Eurycotis floridana_: In the laboratory, development
was completed in 34-36 days at about 85° F. This time was regulated to
some extent by the number of parasites in the oötheca.
There is evidence that larvae eat unhatched wasp eggs or other larvae.
In 34 oöthecae exposed to many female wasps, the maximum number of
parasites to emerge was 306; yet an average of 601 wasp larvae were
dissected from four oöthecae that had each been exposed to 50 female
wasps one week earlier. The larvae usually eat all the host eggs.
Cockroach eggs that were not eaten by the wasp larvae sometimes
developed but usually failed to hatch. Adult wasps made one to six
emergence holes in the oötheca; the average number in 42 oöthecae was
two holes.
_Number of parasites per oötheca._--In _Blatta orientalis_: One of 111
oöthecae exposed to female wasps yielded 48 parasites. In _Eurycotis
floridana_: One oötheca parasitized in the field yielded 68 parasites; 8
oöthecae exposed to single wasps for their entire lifespan yielded an
average of 50 ± 6 parasites (range 23-81); 34 oöthecae exposed to many
wasps for their entire lifespan yielded an average of 198 ± 8 parasites
(range 93-306). In _Periplaneta americana_: Nine oöthecae of 152 exposed
to the wasps were found to be parasitized when dissected; 11 adults
emerged from one oötheca; no parasites emerged from the other 8
oöthecae.
_Sex ratio._--4 [F][F]:1 [M] from ovipositions by isolated females. In
the one oötheca collected in the field, the ratio was 21.6 [F][F]:1 [M].
Parthenogenesis exists; the unfertilized eggs produced only males.
=Anastatus tenuipes= Bolívar y Pieltain
_Synonymy._--_Anastatus blattidarum_ Ferrière. Dr. C. Ferrière (personal
communication, 1957) is of the opinion that his _A. blattidarum_ is a
synonym of _A. tenuipes_. He stated "I have never been able to see the
unique type of _A. tenuipes_ B. y P., which is in Madrid, but the
description agrees with _A. blattidarum_. I had not yet knowledge of
Bolívar's description, when describing my species. The parasite of
cockroaches eggs [_Supella supellectilium_] should be called _A.
tenuipes_ Bol." Mani (1938) synonymized _Solindenia blattiphagus_ Mani
with _Anastatus blattidarum_.
_Natural hosts._--_Supella supellectilium_, Anglo-Egyptian Sudan
(Ferrière, 1930, 1935); U.S.A., Arizona (Flock, 1941); Egypt (Alfieri,
_in_ Hafez and Afifi, 1956). Ohio (Hull and Davidson, 1958).
_Periplaneta americana_, India (Burks _in_ Roth and Willis, 1954a).
Cockroach, Hawaii (Weber, 1951); India (Mani, 1936).
The following is based on parasites that developed on eggs of _Supella
supellectilium_ (Flock, 1941): _Adult behavior._--Wasp may be seen
running rapidly on walls in buildings infested with the cockroach host.
The wasp rarely flies but hops proficiently; when disturbed it can hop
from several inches to several feet. The female licks up the drop of
fluid that oozes from the oviposition puncture. Females die in a few
days, but if fed honey and water may live two weeks. _Oviposition._--The
female selects an oötheca by feeling with her antennae. Flock stated,
without citing experimental evidence, that the age of the egg case was
apparently the chief factor determining choice. The wasp took 15-45
minutes to oviposit. Three females oviposited simultaneously into a
single oötheca; a single female repeatedly oviposited into one oötheca
at intervals. _Development._--Completed in an average of 32.6 days at a
constant temperature of 82° F.
_Number of parasites per oötheca._--Average about 10.7 (range 4-16)
(Flock, 1941); 15 (Ferrière, 1935).
_Sex ratio._--4 [F][F]:1 [M] (Ferrière, 1935); average of 6 [F][F]:1 [M]
(Flock, 1941). Parthenogenesis occurs; the unfertilized eggs produced
only males (Flock, 1941).
_Distribution._--U.S.A.: Maryland, south to Florida, west to Illinois,
Kansas, and Arizona. Guatemala; Hawaii; India; Egypt; Sudan. (Burks,
personal communication, 1956).
=Eupelmus atriflagellum= Girault
_Natural host._--_Blattella germanica_, Australia, Queensland (Girault,
1924).
=Eupelmus= sp.
_Natural host._--"Tree cockroach," U.S.A., Florida (Howard, 1892).
=Solindenia picticornis= Cameron
_Natural hosts._--_Allacta similis_, Hawaii (Perkins, 1906, 1913;
Timberlake, 1924; Swezey, 1929; Zimmerman, 1948).
Other species of cockroaches, Hawaii (Perkins, 1913).
Family PTEROMALIDAE
=Pteromalus= sp.?
_Natural host._--_Leucophaea maderae_?, Jamaica (Westwood, 1839; Sells,
1842). [This host is undoubtedly an error. Sells stated that the oötheca
which contained 96 unidentified chalcids had 16 dentations at the edge;
the description fits the oötheca of an oviparous cockroach and not that
of _L. maderae_ (see Roth and Willis, 1954). Westwood (1839, footnote p.
423) stated that at the meeting of the Entomological Society in 1838 Mr.
Sells exhibited 94 specimens of a small _Pteromalus_ (apparently
identified by Westwood) obtained from one cockroach oötheca. This same
record of Sells was published posthumously in 1842, although in this
paper he identified the host oötheca as "_Blaberus_" _maderae_. Cameron
(1955) lists a European record of _Pteromalus_ sp. from _Periplaneta
americana_ citing Girault (1914) as the source of the record. Girault's
record was apparently taken from Westwood's footnote mentioned above.]
=Systellogaster ovivora= Gahan
_Natural hosts._--_Blatta orientalis_, U.S.A., Illinois (Gahan, 1917).
_Parcoblatta pensylvanica_, Canada, Ontario (Judd, 1955).
_Parcoblatta_ sp., U.S.A., Ohio (Edmunds, 1952a, 1953a).
"Blattid," U.S.A., Maryland (Gahan, 1917).
One oötheca of _P. pensylvanica_ yielded 14 parasites with a sex ratio
of 2.5 [F][F]: 1[M] (Judd, 1955). The average number of parasites in 11
oöthecae of _Parcoblatta_ sp. collected in 1950-51 was 27 wasps
(Edmunds, 1952a, 1953a). The adults made two to three emergence holes in
the oötheca (Edmunds, 1953a; Judd, 1955).
Family EULOPHIDAE
=Melittobia chalybii= Ashmead
_Natural host._--_Periplaneta americana_, U.S.A., Missouri (Rau, 1940a):
_M. chalybii_ is normally a parasite of Coleoptera and Hymenoptera
(Peck, 1951). This is the only record from cockroach eggs. Burks
(personal communication, 1956) stated that this species will attack any
insect to which it is exposed and can be a serious pest in insect
cultures of practically any insect order. In nature it seems to prefer
the nests of aculeate Hymenoptera; Rau suggested that the parasites were
probably brought into his laboratory with mud nests of _Sceliphron
caementarium_ (Drury).
=Mestocharomyia oophaga= Dodd
_Natural host._--_Ellipsidion australe_, Australia, Queensland (Dodd,
1917).
=Syntomosphyrum blattae= Burks
_Natural hosts._--_Parcoblatta_ sp., U.S.A., Ohio (Burks, 1952; Edmunds,
1952a, 1953a): Ten oöthecae yielded an average of 92 wasps (Edmunds,
1952a). Five oöthecae, collected a year later, yielded an average of 74
wasps; adults sometimes made two to three exit holes in the oötheca
(Edmunds, 1953a).
Cockroach, U.S.A., West Virginia (Burks, 1952).
=Syntomosphyrum ischnopterae= (Girault)
_Synonymy._--_Epomphaloides ischnopterae_ Girault [Peck, 1951].
Parker and Thompson (1928) called their hyperparasite _Tetrastichus_ sp.
However, Dr. B. D. Burks (personal communication, 1955) has examined the
teneral specimens which Parker and Thompson deposited in the U.S.
National Museum; he stated that the species is apparently
_Syntomosphyrum ischnopterae_. In view of the experimental work by
Parker and Thompson (see below), this wasp may prove to be a
hyperparasite on evaniids in cockroach oöthecae rather than a primary
parasite on cockroach eggs. (See _Zeuxevania splendidula_, p. 243.)
_Natural hosts._--_Ischnoptera_ sp. [probably _Parcoblatta_ sp. (Rehn,
personal communication, 1958)]. U.S.A., Maryland (Girault, 1917).
_Zeuxevania splendidula_ Costa (an evaniid in the oöthecae of _Loboptera
decipiens_), France (Parker, 1924; Parker and Thompson, 1928).
The following information is from Parker and Thompson (1928): _Adult
behavior._--Courtship and mating were accomplished as soon as adults
emerged, and in a manner similar to that in other chalcids. The females
oviposited only into oöthecae that were parasitized by _Zeuxevania_,
never into normal, nonparasitized oöthecae. _Oviposition._--Oviposition
occurred two days after mating. The female wasp stroked the oötheca with
her antennae, selected a site, and bored into the oötheca with her
ovipositor. She inserted the ovipositor deeply and oviposited for 10-30
minutes. The eggs were deposited randomly on the evaniid larva, some
upright and others lying down. _Development._--Eggs of the hyperparasite
hatched within 3 days and the larvae commenced feeding on the host
larva. There were 30 and 50 hyper-parasites in two oöthecae. _Sex
ratio._--5 [F][F]:1 [M] (from 3 oöthecae).
_Distribution._--U.S.A., District of Columbia, Maryland (Burks, 1952).
=Tetrastichus australasiae= Gahan
_Natural host._--_Periplaneta australasiae_, Sumatra (Gahan, 1923).
=Tetrastichus hagenowii= (Ratzeburg)
_Synonymy._--_Entedon hagenowii_ Ratzeburg, _Blattotetrastichus
hagenowii_ (Ratzeburg) [Burks, 1943]. _Tetrastichodes asthenogmus_
Waterston. G. J. Kerrich (personal communication, 1957) compared the
type of _Tetrastichodes asthenogmus_ Waterston with authentically
determined material of _Tetrastichus hagenowii_ and concluded that _T.
asthenogmus_ is only a weakly developed specimen of _T. hagenowii_. He
stated, "The longitudinal dorsal grooves of the scutellum, which are
strongly developed in normal _hagenowii_, are only rather faintly
developed in Waterston's type and also the second specimen, which was
dissected and mounted on a series of ten microscope slides. No doubt it
was this faint development that caused Waterston to describe the species
in _Tetrastichodes_, a segregate that has since been recognized by Dr.
Burks (Proc. U. S. Nat. Mus., 1943) as being not truly generically
distinct from _Tetrastichus_."
_Natural hosts._--_Blatta orientalis_, Seychelles (Ratzeburg, 1852);
India (Usman, 1949).
_Blatta_ sp., U.S.A., Louisiana (Gahan, 1914).
_Blattella germanica_ (Burks, 1943; Peck, 1951). [In personal
communications, Burks and Peck cite Howard (1892) and Marlatt (1902, and
the 1908 revision of 1902) as sources for this host record. However, _B.
germanica_ is not mentioned specifically as a host of _T. hagenowii_ in
the sources cited nor in the 1915 revision of Marlatt's 1902 paper cited
by Burks (1943); see footnote 6, p. 236.]
_Neostylopyga rhombifolia_, Hawaii (Pemberton, 1941): This record is
based on one parasitized oötheca. We have exposed, at three different
times, groups of 10 to 20 oöthecae of _N. rhombifolia_ to many newly
emerged _T. hagenowii_, but none of the eggs was parasitized (Roth and
Willis, unpublished data, 1957).
_Parcoblatta_ sp., U.S.A., Ohio (Edmunds, 1953a).
_Periplaneta americana_, Africa (Crawford, 1910; Nash, 1955): Nash's
record was incorrectly attributed to _Syntomosphyrum glossinae_ Wtstn.,
a parasite of tse-tse fly pupae (Jordan, 1956), Formosa (Takahashi,
1924; Sonan, 1924); Palestine (Bodenheimer, 1930); Puerto Rico (Seín,
1923; Plank, 1947, 1950; Wolcott, 1951); St. Croix, Virgin Islands
(Beatty, 1944); Hawaii (Schmidt, 1937); U.S.A.: Missouri (Rau, 1940a);
Ohio (Edmunds, 1955); Florida (parasitized oöthecae were collected near
Orlando by members of the Orlando Laboratory, Entomology Research
Branch, U.S. Department of Agriculture; the parasites were identified by
Burks, personal communication, 1955). Fiji (Lever, 1943); India (Mani,
1936; Usman, 1949); Trinidad and Saudi Arabia (Cameron, 1955). Westwood
(1839) stated that 70 parasites belonging to the genus _Eulophus_
emerged from an oötheca of _P. americana_ collected on shipboard. Burks
(personal communication, 1955) stated that the wasp was probably _T.
hagenowii_.
_Periplaneta australasiae_, Australia (Shaw, 1925); India (Usman, 1949);
Saudi Arabia, Trinidad (Cameron, 1955); Formosa (Sonan, 1924).
_Periplaneta brunnea_, U.S.A., Florida (parasitized oöthecae were
collected near Orlando, by members of the Orlando Laboratory, Entomology
Research Branch, U.S. Department of Agriculture. The parasites were
identified by Burks, p. c., 1955).
Cockroach eggs, Formosa (Maki, 1937); Ceylon (Waterston, 1914): Taken on
an oötheca.
"Domestic cockroaches," U.S.A., Louisiana (Girault, 1917).
"Roach egg cases," Panama Canal Zone (Rau, 1933).
_Evania_ sp., Hawaii (Ashmead, 1901; Perkins, 1913); Guam (Fullaway,
1912); Fiji (Lever, 1946); Europe, Cuba, Florida (Marlatt, 1902, 1915).
_Experimental hosts._--_Blatta orientalis_, _Eurycotis floridana_, and
_Periplaneta americana_, U.S.A. (Roth and Willis, 1954b): We have
maintained _T. hagenowii_ for over two years through more than 30
generations on eggs of both _B. orientalis_ and _P. americana_.
_Periplaneta fuliginosa_, U.S.A., Pennsylvania (Roth and Willis, 1954b);
Massachusetts (Roth and Willis, unpublished data, 1957).
Schmidt (1937) deduced that _T. hagenowii_ was a primary parasite of
eggs of _P. americana_ because the parasitized oötheca was obtained from
a cage covered with screen too fine to permit entry of a larger
parasite, such as an evaniid. As noted above, we have reared _T.
hagenowii_ for more than 30 generations on cockroach eggs, none of which
was ever exposed to parasitization by an evaniid. If _T. hagenowii_ were
ever hyperparasitic on _Evania_, this relationship would be accidental,
the eulophid happening to oviposit into an oötheca already containing an
evaniid, or vice versa.
_Adult behavior._--The male mates soon after becoming adult; he mounts
the female from behind, grasps her antennae with his own antennae, and
vibrates his wings during copulation. Mating is accomplished in from
"several" to 20 seconds (Takahashi, 1924; Edmunds, 1955). The adults are
positively phototactic and are capable of hopping for some distance
(Edmunds, 1955). The females feed on material that oozes through the
oviposition puncture (Roth and Willis, 1954b). Females lived 10 days
(Seín, 1923). Without food, females lived 7.8 days and males 3.4 days,
but when fed dilute honey females lived 12.5 days (Usman, 1949). Females
lived 5-11 days (Roth and Willis, 1954b). Fed water and sugar, the wasps
lived 2-6 weeks at 65°F. (Cameron, 1955). Without food, 9 females lived
an average of 3.5 days and 9 males an average of 1.7 days, but when fed
on raisins, 9 females lived an average of 25 days and 9 males 15 days
(Edmunds, 1955). In Formosa there were six generations from April to
December (Maki, 1937).
In Hawaii, Severin and Severin (1915) caught 571 _T. hagenowii_ in 10
kerosene traps that were set up to sample populations of Mediterranean
fruitfly. Apparently the parasite is attracted by the odor of kerosene.
_Oviposition._--The female wasp explores the surface of the oötheca with
vibrating antennae (Edmunds, 1955). She bends her abdomen ventrad and
repeatedly touches the surface of the oötheca with her valvae; when she
finds an acceptable oviposition site, the wasp unsheathes her ovipositor
and bores through the wall of the oötheca (Roth and Willis, 1954b). The
wasp deposited her eggs in 2-5 minutes (Edmunds, 1955). Wasps oviposited
(pl. 34, C) into young or old eggs of _P. americana_ (Roth and Willis,
1954b). A single wasp parasitized more than one oötheca and more than
one wasp oviposited into the same oötheca (Roth and Willis, 1954b;
Edmunds, 1955). We found freshly laid wasp eggs in 34 empty but
previously parasitized oöthecae from which the wasps had emerged (Roth
and Willis, 1954b).
_Development._--In _Periplaneta americana_: Development is completed in
an average of 36 days (range 29-58 days) (Maki, 1937); 29-40 days
(Lever, 1943); average of 23.6 days (range 22-26 days) at 62°-85° F.
(Usman, 1949); about 3 months at 60°-65° F. (Cameron, 1955); 31-60 days
at 70°-80° F. (Edmunds, 1955). We found that the wasps completed
development in 23-56 days at about 85° F., but the period depended on
the number of wasps in the oötheca; the larger the number of wasps (up
to an average of about 70 wasps per oötheca), the shorter the time
required to complete development. Wasps in oöthecae containing 70 or
more parasites developed in an average of about 32 days (Roth and
Willis, 1954b). Wasp larvae eat the contents of the cockroach egg in
which they start development, then rupture the chorion and attack
adjoining eggs (Cameron, 1955; Edmunds, 1955). All eggs are consumed
when the parasite density is high, but if too few larvae develop per
oötheca, some cockroach eggs survive and the embryos complete
development (Roth and Willis, 1954b). However, a certain number of
cockroach nymphs must complete development to enable the survivors to
force open the crista and emerge from the oötheca; fewer than this
number of surviving nymphs will be trapped and killed as effectively as
if they had been eaten by the parasite. The adult parasites emerge from
one to three holes cut through the wall of the oötheca (Usman, 1949;
Roth and Willis, 1954b).
_Number of offspring per female._--In _Blatta orientalis_: In the
laboratory, 5 oöthecae were left with each of 25 female wasps for their
entire lifespans; of the 125 oöthecae, 32 were parasitized. The average
number of offspring per female was 66 (range 5-164) (Roth and Willis,
1954b). In _Periplaneta americana_: Each of 206 oöthecae was exposed to
a single female wasp for 24 hours; the average number of offspring per
female was 103 (range 50-139). Five oöthecae were left with each of 38
females for their entire lifespans; of the 190 oöthecae, 81 were
parasitized. The average number of offspring per female was 94 (range
45-168 [from original data]) (Roth and Willis, 1954b).
_Number of parasites per oötheca._--In _Eurycotis floridana_: In the
laboratory, 3 oöthecae that had been exposed to 20 female wasps yielded
an average of 648 parasites (range 606 [from original data] to 685)
(Roth and Willis, 1954b). In _Neostylopyga rhombifolia_: One oötheca
yielded 73 parasites (Pemberton, 1941). In _Parcoblatta_ sp.: Two
oöthecae yielded an average of 100 parasites (Edmunds, 1953a). In
_Periplaneta americana_: 100 parasites per oötheca (Seín, 1923); 140
(Schmidt, 1937); 25 (Rau, 1940a); 7-38, average 33 (Usman, 1949); 71
(Wolcott, 1951); 4 oöthecae exposed to 20 female wasps yielded an
average of 204 wasps (range 164 [from original data] to 261) (Roth and
Willis, 1954b); average of 30-40 (Cameron, 1955); 39 oöthecae yielded an
average of 93 parasites (range 12-187) (Edmunds, 1955). In _Periplaneta
australasiae_: Oöthecae yielded an average of 40-50 adult parasites
(Cameron, 1955); about 50 (Shaw, 1925).
_Sex ratio._--3 [F][F]:1 [M] (Usman, 1949); 4 [F][F]:1 [M] (Cameron,
1955); 2-8 [F][F]:1 [M] (Roth and Willis, 1954b); 1.2 [F][F]:1 [M]
(Edmunds, 1955). Parthenogenesis exists; the unfertilized eggs produced
only males (Roth and Willis, 1954b; Edmunds, 1955).
_Distribution._--Probably worldwide. Eastern and southern U.S.A.;
Central and South America; Europe; Arabia; Africa; India; Formosa;
Hawaii.
=Tetrastichus periplanetae= Crawford
_Natural hosts._--_Periplaneta americana_, Mozambique (Crawford, 1910);
Union of South Africa (parasites reared from oöthecae collected in
Durban, Natal, by the City Health Department): The parasites were
identified by Burks (personal communication, 1956). Jamaica (Gowdey,
1925); Réunion Island (Bordage, 1913).
"Domestic cockroach," Puerto Rico (Wolcott, 1951).
=Tetrastichus= sp. I
_Taxonomy._--Burks (personal communication, 1956) stated that this
species (specimens of which are in the U.S. National Museum) is very
close to _T. hagenowii_.
_Natural hosts._--_Periplaneta americana_, Union of South Africa
(parasites reared from oöthecae collected in Durban, Natal, by the City
Health Department [Burks, p. c., 1956]).
_Periplaneta australasiae_, Manila, Philippine Islands (Burks, p. c.,
1956).
=Tetrastichus= sp. II
_Synonymy._--Because of the war, Cros (1942) could not determine this
insect specifically. He designated it provisionally and with reserve
under the name _Eulophus_ sp. However, Burks (p. c., 1956) stated that
the species is most certainly a _Tetrastichus_ from the description
given; but, it is apparently not _T. hagenowii_ because of its brilliant
steel-blue color.
_Natural host._--_Blatta orientalis_, Algeria (Cros, 1942): _Adult
behavior._--Mating began as soon as wasps emerged from an oötheca. Males
mated repeatedly. Adults lived up to 5 days in summer and up to 12 days
in fall. There were up to four generations per year in the laboratory.
_Oviposition._--Wasps oviposited into oöthecae 6, 22, 40, and 43 days
old, and the parasites developed successfully. More than one female
oviposited into the same oötheca. Oviposition was of long duration.
_Development._--From egg to eclosion took an average of 34 days in
summer (range: 30-38 days, 5 oöthecae), and an average of 67 days in
fall (range: 58-73 days, 3 oöthecae). An average of 55 parasites
developed per oötheca (range 21-105, 5 oöthecae); over 130 wasps emerged
from a sixth oötheca. _Sex ratio._--10-20 [F][F]:1 [M].
HOST SELECTION BY EGG PARASITES
The nature of the oviposition stimulus(i) for the wasp parasites of
cockroach eggs is unknown. Edmunds (1954) noted that _Prosevania
punctata_ showed more interest in oöthecae that had been cemented to the
substrate than in clean oöthecae that had simply been dropped. Cros
(1942) experimented with two females of _P. punctata_ to see if the
wasps could find oöthecae that had been buried in sand by the oriental
cockroach. After prospecting the sand with their antennae, the wasps dug
deep excavations with their front legs but always mistook the location
of the oöthecae. Cros suggested that the wasps were misled by the odor
left in the jar by the cockroaches. It is quite possible that odor helps
the wasp find the host oötheca.
The extent of host selection varies among these parasites; some species
will oviposit into the eggs of more than one species of cockroach, but
others show some degree of host specificity. Positive selection of
specific hosts by certain parasites appears in correlative data from
different investigators on pages 235 to 254. There is a small body of
data that shows nonacceptance of certain hosts by some of these wasps.
For example, _Comperia merceti_ would not parasitize eggs of _Blatta
orientalis_ or _Periplaneta americana_ in the laboratory (Lawson,
1954a). We (unpublished data, 1957) exposed a soft oötheca, recently
removed from _Eurycotis floridana_, to _C. merceti_; no wasps developed;
we had similar negative results with _C. merceti_ and oöthecae of _B.
germanica_. We (1954b) could not induce _Tetrastichus hagenowii_ to
parasitize eggs of _Blattella germanica_, _B. vaga_, or _Parcoblatta
virginica_ in the laboratory. In our experiments, _T. hagenowii_
oviposited into eggs of _Supella supellectilium_, but the wasp eggs
either failed to hatch, or if they hatched, the larvae died before
completing development. Neither would _T. hagenowii_ parasitize eggs of
_N. rhombifolia_ (Roth and Willis, unpublished data, 1957). _Anastatus
tenuipes_ would not parasitize the eggs of _Latiblattella lucifrons_
Hebard, _Periplaneta americana_, _B. germanica_, or _B. vaga_ (Flock,
1941). _Anastatus floridanus_ would not oviposit into eggs of _S.
supellectilium_ and only rarely into eggs of _P. americana_ or _B.
orientalis_ (Roth and Willis, 1954a); in the laboratory, this wasp could
not be maintained beyond one generation on the eggs of _P. americana_.
Edmunds (1953b) could not induce _Prosevania punctata_ to parasitize
eggs of _B. germanica_. Cros (1942) induced _P. punctata_ to oviposit
into a mantid oötheca, but neither mantids nor parasite developed.
COCKROACH-HUNTING WASPS
A number of wasps of the families Ampulicidae, Sphecidae, and a very few
species of Pompilidae have been found to provision their nests with
nymphal or adult cockroaches. This habit of preying on cockroaches is
primitive (Leclercq, 1954); Leclercq (personal communication, 1955)
stated that this habit is always associated with the conservation of a
number of structures considered as archaic from a purely morphological
point of view.
The records of wasps of the genus _Astata_ capturing cockroaches (e.g.,
Sickmann, 1893; St. Fargeau _in_ Sharp, 1899) "all trace back to a
questionable record by Lepeletier (1841) which probably was a
misidentification of the predator" (K. V. Krombein, personal
communication, 1956). Marshall (1866) suggested that the braconid
_Paxylomma buccata_ Bréb., which he found frequenting cockroach runs in
Pembrokeshire, was parasitic on _Ectobius nigripes_ Stephens; however,
this wasp is undoubtedly parasitic on ants, probably on ant larvae
(Donisthorpe and Wilkinson, 1930).
The wasps that are known to capture cockroaches, and summaries of their
biology, are listed below.
WASPS THAT PROVISION THEIR NESTS WITH COCKROACHES
Family POMPILIDAE
=Pompilus bracatus= Bingham
_Natural hosts._--Cockroaches, India (Bingham, 1900).
=Pompilus= sp.
_Natural host._--Cockroach, Nyasaland (Lamborn _in_ Poulton, 1926): The
wasp was collected leading a nymph of the cockroach by its antenna. The
cockroach was in a stupefied state, and its antennae were bitten off to
about half their length.
=Salius verticalis= Smith
_Natural hosts._--Cockroaches, India (Bingham, 1900).
Family AMPULICIDAE
The species of _Ampulex_ do not appear to make special nests in which to
lay their eggs but drag their prey to any convenient hole, or crack in
the ground (Arnold, 1928). Although many species of _Ampulex_ have been
described, the prey of only a small number of species have been
discovered, but the known prey are all cockroaches.
=Ampulex amoena= Stål
_Synonymy._--_Ampulex novarae_ Saussure [Krombein, personal
communication, 1957].
_Natural hosts._--_Periplaneta americana_ and _Periplaneta
australasiae_, both as small nymphs, Formosa (Sonan, 1924, 1927): The
wasp stings a nymph about one inch long and carries it to a suitable
place, such as bamboo pipes, folds of newspaper, or books (in houses),
for oviposition.
_Periplaneta picea_, Japan (Kamo, 1957; Kohriba, 1957).
_Experimental hosts._--_Periplaneta picea_, Japan (Kamo, 1957; Kohriba,
1957).
Kamo (1957) observed that in the field both males and females sucked
juices from wounds they made in the stems of _Clerodendron_
_trichotomum_ Thunberg or _Ilex rotunda_ Thunberg. Kohriba (1957), on
the other hand, found both sexes sucking sap of _Abies_ sp. and other
trees from points injured by the rostrum of cicadas. Kamo (1957)
observed that the female wasp grasped the cockroach by a tergum and
stung it several times in the thorax. The wasp always amputated the
antennae of the prey and sucked up the fluid oozing from the cut
antennae. The wasp egg was placed on the mesocoxa of the cockroach. In
the laboratory as many as three cockroaches, each with a wasp egg, were
stored in artificial nests per day. Kohriba (1957) observed similar
behavior in the laboratory and made these additional notes. The
paralyzed cockroach could move its legs and was led to the nest by the
wasp which seized its antennae. The egg hatched in 2 days, and after
sucking up body fluid for 2 days the larva began to devour the prey.
Three days later the larva spun its cocoon, and about one month after
spinning a female wasp emerged.
=Ampulex assimilis= Kohl
_Natural hosts._--_Blatta lateralis_, wingless females, Iraq (Hingston,
1925): Nesting sites are holes in palm trees, galleries of beetles, or
tunnels in ground. The wasp first seizes a cockroach by the edge of its
thorax and stings it in the thoracic region, then seizes the cockroach
by an antenna and pulls and leads it to the nest. The wasp deposits her
egg on the outer surface of the femur of the cockroach's midleg. The
nest is closed with debris; later the cockroach recovers from the sting.
The wasp larva first feeds externally, then bores into the cockroach and
devours the internal organs. Pupation occurs inside the exoskeleton of
the cockroach.
=Ampulex canaliculata= (Say)
_Synonymy._--_Rhinopsis caniculatus._
_Natural hosts._--_Ischnoptera_ sp., U.S.A. (Krombein, 1951).
_Lobopterella dimidiatipes_, Hawaii (Williams, 1928a, 1929).
_Parcoblatta pensylvanica_? MacNay (1954) referred to a rare sphecoid
wasp in eastern Canada which provisioned its nest with nymphs and adults
of _P. pensylvanica_. Dr. W. R. M. Mason (personal communication, 1957)
wrote us that although this wasp was _Ampulex canaliculata_, it was not
reared from the cockroach but was swept from a pine tree. There are no
positive records linking _A. canaliculata_ with _P. pensylvanica_.
_Experimental host._--_Parcoblatta virginica_, females, U.S.A., Missouri
(Williams, 1928a, 1929): figure 6.
Nesting sites are in twigs (Krombein, 1951). The adult behavior is
similar to that of _A. compressa_; the female wasp imbibes blood that
oozes from the amputated antennae of the cockroach; the egg hatches in
2-3 days, and the development of one male was completed in 33 days
(Williams, 1929).
[Illustration:
FIG. 6. _Ampulex canaliculata_ attacking _Parcoblatta virginica_. A,
Female wasp stinging her prey, c. × 4.8. B, Wasp's egg attached to the
coxa of the mesothoracic leg of the cockroach. C, Larva of _A.
canaliculata_ (about three-quarters grown) feeding on the internal
organs of the host from the exterior, c. × 4. (Reproduced from F. X.
Williams [1929], through the courtesy of Dr. F. X. Williams and F. A.
Bianchi, Hawaiian Sugar Planters' Association.)]
_Distribution._--U.S.A.: Connecticut south to Georgia; Ohio, Wisconsin,
Missouri, Kansas; in open woods (Krombein, 1951).
=Ampulex compressa= (Fabricius)
(Pl. 35)
_Synonymy._--Guĕpe ichneumon of Réaumur [Williams, 1929]; _Chlorion_
(_Ampulex_) _compressum_.
_Natural hosts._--_Periplaneta americana_, New Caledonia (Lucas, 1879);
India (Dutt, 1912); Reunion (Bordage, 1912).
_Periplaneta australasiae_, Hawaii (Swezey, 1944).
_Periplaneta_ sp., India (Maxwell-Lefroy, 1909).
Cockroach. Mauritius (Réaumur, 1742); Burma (Bingham, 1897).
_Experimental hosts._--_Neostylopyga rhombifolia_, _Periplaneta
americana_, and _Periplaneta australasiae_, Hawaii (Williams, 1942,
1942a). Zimmerman's (1948) listings probably were taken from Williams.
_Nesting sites._--Holes in walls; holes in banyan and fig trees; in
houses in drawers and cartons. _Behavior._--Similar to that of _A.
assimilis_. Bordage (1912) gives a complete description of capture of
prey. The female wasp cuts off part of the cockroach's antennae, legs,
and wings; she sticks her egg onto the host's mesothoracic coxa. The
wasp frequents houses in search of prey. Five [F][F], supplied with a
cockroach per day, stored an average of 57±14 cockroaches; 8 [F][F]
stored an average of 45±3 cockroaches; these latter wasps were not
supplied with a cockroach per day throughout (mean values computed from
Williams, 1942). This wasp will not attack _Nauphoeta cinerea_
(Williams, 1942a) or _Pycnoscelus surinamensis_ (Schwabe, 1950b). On one
occasion, _A. compressa_ stung _Diploptera punctata_, but did not
oviposit (Williams, 1942a). _Development._--Minimum 34 days, maximum 140
days (Williams, 1942). About 6 weeks (Swezey, 1944). _Longevity of
adults._--13 [F][F] lived an average of 110±11 days (minimum 31, maximum
159); several [M][M] lived 2 months (Williams, 1942).
=Ampulex fasciata= Jurine
_Natural host._--_Ectobius pallidus_, France (Picard, 1911, 1919):
Nesting sites are in brier or bramble stems, or in crevices in fig
trees; the female possibly uses old nests of leaf-cutter bees. The
feeding of the wasp larva is similar to that of other _Ampulex_. Adult
wasp emerges by cutting open a passage through its cocoon and through
the anus of the cockroach.
=Ampulex ruficornis= (Cameron)
_Natural hosts._--Cockroaches, Oriental region (Rothney _in_ Sharp,
1899): Nesting sites are in crevices in bark. The female grasps the
cockroach by an antenna to drag it to her nest.
=Ampulex sibirica= Fabricius
_Synonymy._--Perkins referred to this species as _Ampulex sibirica_.
Williams (1942a), referring to Perkins's observations, mentions the
species as "_A. compressiventris_ Guérin (=_A. siberica_ Sauss.)."
Krombein (personal communication, 1956) has commented upon this synonymy
as follows: _Ampulex siberica_ Sauss. is apparently a misidentification
by Saussure of _sibirica_ Fab. Kohl (1893) in his revision of the genus
_Ampulex_ considered _A. compressiventris_ Guérin to be the correct name
for this common African species and that _sibirica_, described from
Siberia, must be another species. However, Turner (1912) stated that he
had seen Fabricius's type specimen and that it was identical with what
had been called _compressiventris_; he considered the Siberian locality
given by Fabricius as an error. Krombein suggested that Williams's use
of the combination _siberica_ Sauss. was a lapsus and that the valid
name, if Turner is correct, is _sibirica_ Fab.
_Natural hosts._--Cockroaches, West Africa (Perkins _in_ Sharp, 1899):
Nesting sites are keyholes. Enters apartments in search of cockroaches.
Wasp cocoon protrudes from dead body of cockroach.
=Ampulex sonnerati= Kohl
_Synonymy._--"La mouche bleue" of Sonnerat (Kohl, 1893).
_Natural host._--"Kakkerlac," Philippine Islands (Sonnerat, 1776):
Nesting sites are readymade crevices. The wasp seizes the cockroach by
an antenna and stings the host many times in the "abdomen." She drags
the cockroach by an antenna to the nest, and, after depositing her egg,
plugs the opening with moistened earth.
=Dolichurus bicolor= Lepeletier
_Synonymy._--Schulz (1912) considered this to be _Dolichurus
corniculus_. Berland (1925) stated that this is possibly a color variety
of _D. corniculus_. Soyer (1947), from a study of the behavior of the
wasps, believed that both _D. bicolor_ and _D. haemorrhous_ are
varieties of _D. corniculus_. Krombein (personal communication, 1956)
stated that _D. corniculus_ and _D. bicolor_ differ in characters other
than color alone and that _D. bicolor_ is considered a valid species
today.
_Natural host._--Cockroach, France (Benoist, 1927): The wasp was
observed closing the entrance to its burrow. Its egg was attached to the
coxa of the midleg of the cockroach.
Maneval (1932) stated that _D. bicolor_ is found at the edge of dry
woods along with _D. corniculus_ and that the wasp will also accept the
prey of _D. corniculus_ if presented to it.
=Dolichurus corniculus= (Spinola)
_Synonymy._--_Dolichurus haemorrhous_ Costa [Schulz, 1912]. Berland
(1925) listed _D. haemorrhous_ separately but stated that it is perhaps
a color variety of _D. corniculus_.
_Natural hosts._--_Blattella germanica_, France (Benoist, 1927).
_Ectobius lapponicus_, Germany (Sickmann, 1893); Denmark (Nielsen,
1903); Sweden (Adlerz, 1903); Italy (Grandi, 1931, 1954); France
(Benoist, 1927; Maneval, 1928).
_Ectobius pallidus_, France (Maneval, 1932; Soyer, 1947).
_Ectobius panzeri_, France (Soyer, 1947).
_Ectobius_ sp., Italy (Grandi, 1954).
_Hololampra punctata_, Pitten (Handlirsch, 1889).
_Loboptera decipiens_, France (Ferton, 1894).
Cockroach, Netherlands (Bouwman, 1914).
_Nesting sites._--The wasp uses already-made cavities such as rotting
dead branches on ground, fissures in the earth, abandoned ant holes,
chinks in stone, or the empty cocoon of the ichneumon _Ophion luteus_
(Ferton, 1894; Maneval, 1932).
_Behavior._--The prey is immobile while being dragged to the nest but
recovers sufficiently from the sting so that if dug up it will run
around (Ferton, 1894; Bouwman, 1914; Benoist, 1927; Grandi, 1954). The
wasp cuts off about two-thirds of the cockroach's antennae prior to
putting its prey in its nest (Adlerz, 1903; Bouwman, 1914; Soyer, 1947).
One cockroach is placed in the nest and the wasp's egg is attached to
the midcoxa (Ferton, 1894). Oviposition takes 5 to 6 minutes (Maneval,
1939). Wasp fills and seals its nest with bits of earth and stones
(Ferton, 1894; Grandi, 1954). The wasp larva feeds externally and
devours the entire cockroach, including its exoskeleton (Ferton, 1894).
_Development._--Hatching occurs in 3 to 4 days (Ferton, 1894) or longer
during cooler weather (Maneval, 1939). Larval development takes 6 days
(Grandi, 1954), 8 days (Ferton, 1894), or 10 to 25 days depending on
season (Maneval, 1939).
=Dolichurus gilberti= Turner
_Natural hosts._--"Small Blattidae," India (Turner, 1917).
=Dolichurus greenei= Rohwer
_Natural host._--_Parcoblatta_ sp., U.S.A., Virginia (Krombein, 1951,
1955): Nesting sites are under leaf litter. The prey was a paralyzed
third-instar nymph. _Distribution._--Ontario. U.S.A. from Canadian
border south to Florida in coastal States (Krombein, 1951).
=Dolichurus ignitus= Sm.
_Natural hosts._--Cockroaches, Natal and Southern Rhodesia (Arnold,
1928): The wasp is "usually seen running up and down the trunks of
trees searching for small cockroaches in the crevices of the bark."
=Dolichurus stantoni= (Ashmead)
_Natural hosts._--_Allacta similis_, nymphs, Hawaii (Williams et al.,
1931; Zimmerman, 1948).
_Blattella lituricollis_, usually nymphs, Philippine Islands, Hawaii
(Williams, 1919).
_Cutilia soror_, nymphs, Hawaii (Williams et al., 1931; Zimmerman,
1948).
"_Phyllodromia_" sp., Philippine Islands, Hawaii (Williams, 1918;
Bridwell, 1920).
_Experimental hosts._--"Field cockroaches," Philippine Islands
(Williams, 1944).
_Nesting site._--Readymade crevices or holes in ground; porosity in
lava. _Behavior._--The wasp seizes the cockroach by a cercus or leg and
stings it in the thorax. She (fig. 7, A) then drags the cockroach to the
nest by the base of an antenna. Wasp bites off distal part of host's
antennae. She deposits her egg on one of the host's midcoxae. Nest is
plugged with lumps of soil. The larva eats the entire host.
_Development._--Eggs hatched in about a day and a half. Adults emerged
about 3 weeks later. About five generations per year. (Williams, 1918,
1919; Williams et al., 1931.)
=Dolichurus= sp.
_Natural hosts._--Cockroaches, nymphs, South Africa (Bridwell, 1917).
Adult female cockroach carrying an oötheca, France (Deleurance, 1943).
_Nesting site._--Plant stem, or in ground possibly an old abandoned nest
of _Ammophile_. _Behavior._--Bridwell noted that one wasp larva ate two
cockroach nymphs before pupating; the adult emerged about 4 months after
cocoon formation. Deleurance observed the wasp close its nest with small
pebbles, balls of earth, and small dead branches. The wasp egg was
placed on the femur of the midleg. The prey in the nest is alert when
disturbed. Deleurance believed the wasp was a variety of _D.
corniculus_.
=Trirhogma caerulea= Westwood
_Natural hosts._--_Periplaneta americana_ and _Periplaneta
australasiae_, Formosa (Sonan, 1924): The wasp stings a nymph about one
inch long and carries it to a suitable place (bamboo pipe) for
oviposition.
[Illustration:
FIG. 7.--Cockroach-hunting wasps. A, _Dolichurus stantoni_ leading a
nymph of _Blattella lituricollis_ to her nest, c. × 4. (Reproduced
from F. N. Williams [1919].) B, _Podium haematogastrum_ attaching her
egg to an _Epilampra_ sp. while on the side of a termite mound that
contains the wasp's nest, c. × 1.6. C, _Epilampra_ sp. parasitized
by _P. haematogastrum_ showing the wasp's egg attached to the right
fore coxa, c. × 3.2. (B and C reproduced from Williams [1928],
through the courtesy of Dr. F. X. Williams and F. A. Bianchi.)]
=Trirhogma= sp.
_Natural hosts._--Cockroaches, Oriental region (Williams, 1918, 1928):
As far as is known species of this genus of wasps hunt cockroaches.
Family SPHECIDAE
=Tachysphex blatticidus= Williams
_Natural hosts._--_Chorisoneura_ sp., adults, Trinidad, St. Augustine
(Callan, 1942): The wasps nest gregariously in sandy places. The wasp
itself is parasitized by the mutillid _Timulla_ (_Timulla_) _eriphyla_
Mickel.
Cockroaches, Trinidad (Williams, 1941a; Callan, 1950).
=Tachysphex coriaceus= Costa
_Natural hosts._--Cockroaches, Italy (Beaumont, 1954).
=Tachysphex fanuiensis= Cheesman
_Natural hosts._--_Graptoblatta notulata_, Society Islands (Cheesman,
1927, 1928).
Cockroach ("except for its smaller size [it] much resembles
_Graptoblatta notulata_."), New Caledonia (Williams, 1945).
_Nesting sites._--Patches of dry soil (Cheesman, 1928); coarse sand at
base of a bank (Williams, 1945). _Behavior._--The female wasp pounces on
the cockroach and stings it into immobility; she carries her prey in
flight to the nest. Two to 13 cockroaches may be found in one nest; and
one or more wasp eggs may be deposited in one nest. The egg is attached
at one end to the host's thorax behind a forecoxa. Nest is sealed with
dry pellets of soil. The cockroaches apparently do not recover from the
wasp's sting.
=Tachysphex lativalvis= (Thomson)
_Natural hosts._--_Ectobius lapponicus_, adults, Sweden (Adlerz, 1906);
France (Maneval, 1932).
_Ectobius pallidus_, nymphs, France (Ferton, 1894, 1901; Maneval, 1932;
Deleurance, 1946); Italy (Grandi, 1928).
_Ectobius panzeri_, Netherlands (Bouwman, 1914).
_Ectobius_ sp., Denmark (Nielsen, 1933).
Ferton (1914) stated that he had reported in 1912 that this species
hunted Hemiptera, but that this observation was a lapsus. _Nesting
site._--In the ground of sandy woodlot or border of dry woods; the nest
is a hole 5.5 to 8 cm. long ending in a horizontal cell. Grandi (1928)
stated that the entrance to the nest descended obliquely for 5 to 6 cm.
and ended 4 cm. below the surface of the ground. _Behavior._--Two
cockroaches, either sex, adults or nymphs, were stored in the cell
(Adlerz, 1903; Grandi, 1928). The wasp laid her egg on the first prey
brought, attaching it behind the front coxa. The cockroaches were not
excitable and their antennae had not been injured. Grandi (1928) stated
that the claws of the hind tarsi of the victims may be amputated. The
hatched larva may consume one of its victims in four days leaving only
the head, pronotum, tegmina, wings, and the urosternum.
=Podium abdominale= (Perty)
_Synonymy._--_Trigonopsis abdominalis_ Perty [Kohl, 1902].
_Natural hosts._--Cockroaches, nymphs, Ecuador (Williams, 1928): These
wasps are apparently mainly arboreal mud daubers. The female wasp
constructs a mud nest on underside of a palm leaf. Wasp egg is attached
behind one of the forecoxae of the cockroach. Several cockroaches are
stored in each nest. The prey is not immobilized as a result of the
sting, and its antennae are left intact.
=Podium carolina= Rohwer
_Natural host._--_Parcoblatta pensylvanica_, nymphs (Rau, 1937): Nesting
sites are mud nests of _Sceliphron caementarium_ (Drury). One to three
cockroach nymphs are stored per nest; mud partitions are placed in tube;
the nest is plugged with mud which is coated with resin.
_Distribution._--U.S.A., New York to North Carolina (Murray, 1951);
Florida (Krombein and Evans, 1955).
=Podium dubium= Taschenberg
_Natural hosts._--Epilamprine cockroaches, Brazil (Williams, 1928):
Burrows, lenticular in cross section, are found on shaded trails. The
wasp's habits are similar to those of _P. flavipenne_ and _P.
haematogastrum_.
=Podium flavipenne= Lepeletier
_Natural host._--_Epilampra abdomen-nigrum_, British Guiana (Williams,
1928): _Nesting site._--Burrows, about 2 inches deep and lenticular in
cross section, are dug in the ground in well-drained, partly sheltered
areas; also old _Podium_ nests are used. _Behavior._--The wasp stings
the cockroach to helplessness and flies with it back to her nest where
the host may recover from the sting; one or more cockroaches are stored
per nest; the egg is deposited behind the forecoxa while the cockroach
is still outside the burrow. The nest is sealed with mud. The larva
feeds on most of the cockroach and leaves only some heavily sclerotized
portions in the cell. In 153 nests examined, there was an average of 2.2
± 0.08 [standard error computed from cited data] cockroaches per cell;
four nests contained five cockroaches apiece. Of the 331 cockroaches in
the nests, only 6 percent were adults. _Development._--Egg hatches in
about 2 days; larva feeds about 4 days and pupates about 2 weeks later;
adult emerges about 10-12 days later.
=Podium haematogastrum= Spinola
_Natural host._--_Epilampra_ sp., Brazil, Pará (Williams, 1928): The
female wasp (fig. 7, B) burrows into the surface of termite mounds, in
banks, and in level ground. This wasp's behavior is similar to that of
_P. flavipenne_. There was an average of 1.6 cockroaches (fig. 7, C) per
cell in 74 nests examined. Of the 121 cockroaches collected, 28 percent
were adults. Under artificial conditions, the life cycle varied from
about a month to 45 days or more.
=Podium luctuosum= Smith
_Natural host._--_Parcoblatta virginica_, female, U.S.A., New York
(Pate, 1949).
_Distribution._--U.S.A.: New York to Texas (Murray, 1951).
=Podium rufipes= Fabricius
_Natural hosts._--"Wood roaches," British Guiana (Howes, 1917, 1919);
Brazil (Williams, 1928): Nesting sites were clay column nests on houses,
sides of stumps, or forest trees; banks; termite mound. Variable numbers
of cockroaches were placed in the nests with one wasp egg attached
behind forecoxa of the last host. The egg hatches in 2 days, the larva
pupates about 2 weeks later, and the adult emerges 24 days later.
=Podium= sp.
_Natural host._--_Epilampra conferta_, Brazil (Poulton, 1917): The
burrow contained several cockroaches of the same species.
ANTS PREDACEOUS ON COCKROACHES
A large roach endeavored to escape by crossing the main front of
the army. The creature made several powerful jumps, but each time
it touched the ground ... its legs were grasped by the fearless
ants.... In the end it fell ... and was instantly torn to bits and
carried to the rear.... Another ant with the body of a wood roach
was assisted by a worker who held the carrier's abdomen high in the
air out of the way of her burden, all the way to the nest.
HOWES (1919)
Family FORMICIDAE
From the known entomophagous habits of the lower ants (Wheeler, 1928),
we wonder that there are not more records of ants feeding on
cockroaches, because this act must occur frequently. Kirby and Spence
(1822) stated that R. Kittoe had observed in Antigua that ants which
nested in the roofs would seize a cockroach by the legs so it could not
move, kill it, and carry it up to their nest. Hotchkiss (1874) observed
ants kill cockroaches on shipboard. Cockroaches attracted to sugar in
the pantry were killed and carried off by the ants. The destruction of
cockroaches by army ants has been recorded by Bates (1863), Wallace
(1891), Beebe (1917, 1919), Howes (1919), and others. Dead and mutilated
specimens of _Ischnoptera_ sp. [undoubtedly _Parcoblatta americana_
(Gurney, personal communication, 1958)] are common in the nests of
species of _Formica_ in California (Mann, 1911).
=Aphaenogaster picea= Emery
_Natural prey._--_Ectobius pallidus_, U.S.A., Massachusetts (Roth and
Willis, 1957).
=Camponotus pennsylvanicus= (De Geer)
_Common name._--Carpenter ant.
_Natural prey._--_Parcoblatta pensylvanica_, U.S.A. (Rau, 1940): The
ants entered traps set up to capture the cockroach and carried off about
a dozen adults of both sexes.
=Dorylus (Anomma) nigricans= subsp. =sjöstedi Emery=
_Natural prey._--Small cockroach, Belgian Congo (Raignier and van Boven,
1955).
=Dorylus (Anomma) wilverthi= Emery
_Natural prey._--Small cockroaches, Belgian Congo (Raignier and van
Boven, 1955).
=Dorylus= sp.
_Common name._--"Safari ant."
_Natural prey._--Cockroaches, Africa, Lake Victoria (Carpenter, 1920):
When the "Safari ants" were hunting, many species of cockroaches were
driven from hiding among dead leaves in the forest. The cockroaches
rushed about but easily fell prey to the ants which tore them to bits.
=Eciton burchelli= (Westwood)
_Common name._--Army ant.
_Natural prey._--Cockroaches, Panama Canal Zone (Johnson, 1954;
Schneirla, 1956).
=Formica omnivora=
_Synonymy_.--The identity of this form is unknown. There are no species
of _Formica_ on Ceylon. There was another _Formica omnivora_ described
from tropical America, whose identity is also unknown (W. L. Brown,
personal communication, 1956).
_Natural prey._--Cockroaches, Ceylon (Kirby and Spence, 1822).
=Iridomyrmex humilis= Mayr
_Common name._--Argentine ant.
_Natural prey._--Cockroaches, injured individuals only (Ealand, 1915).
=Lasius alienus= (Förster)
_Natural prey._--_Ectobius pallidus_, U.S.A., Massachusetts (Roth and
Willis, 1957).
=Pheidole megacephala= (Fabricius)
_Common name._--Big-headed ant.
_Natural prey._--_Holocompsa fulva_, Hawaii (Illingworth, 1916).
_Nauphoeta cinerea_ and _Pycnoscelus surinamensis_, Hawaii (Illingworth,
1914, 1942): The ants followed and killed _N. cinerea_ and _P.
surinamensis_ as they burrowed in moist soil and attacked and destroyed
_N. cinerea_ in breeding cages.
XIII. VERTEBRATA
Class PISCES
In British Guiana, Beebe (1925a) found undetermined cockroach remains in
the stomachs of four species of fish belonging to three families, as
follows:
Family POTAMOTRYGONTIDAE
=Potamotrygon humboldti= (Duméril)
(= _Potamotrygon hystrix_)
Family PIMELODIDAE
=Rhamdia sebae= Cuvier and Valenciennes
Family CHARACIDAE
=Cyrtocharax magdalenae essequibensis= (Eigenmann)
(= _Cynopotamus essequibensis_)
=Chalceus macrolepidotus= Cuvier and Valenciennes
The only other records of cockroaches being eaten by fish pertain to the
use of cockroaches as bait.[8] Captain William Owen (_in_ Webster, 1834)
stated that the Chinese used cockroaches as bait in their fishing
excursions. At Reelfoot Lake, Tennessee, _Blatta orientalis_ were kept
in large numbers by bait dealers and were sold to fishermen who used
them for catching _Lepomis pallidus_, a sunfish locally known as bream,
blue bream, or bluegill (Rau, 1944). In Indiana, oriental cockroaches
were collected at a city dump by fishermen (Gould, 1941). Peterson
(1956) states that cockroaches are satisfactory bait for bluegills,
crappies, channel cat, blue heads, and large mouth black bass.
Class AMPHIBIA[9]
Order CAUDATA
Family PLETHODONTIDAE
=Plethodon glutinosus= (Green)
_Natural prey._--_Cryptocercus punctulatus_, U.S.A. (Honigberg, 1953):
Protozoa which are normally only found in _C. punctulatus_ were present
in the intestine of the salamander indicating that this cockroach had
been eaten by the amphibian.
Order SALIENTIA
Family BUFONIDAE
=Bufo funereus= Bocage
_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): The stomachs
of 62 out of 72 specimens contained food; this included 3 cockroaches.
=Bufo ictericus= Spix
_Natural prey._--Cockroaches, Brazil (Valente, 1949): Stomach contents
revealed the prothorax, legs, and wings of cockroaches, and fragments of
wood-cockroaches. This toad frequently feeds at night.
=Bufo marinus= (Linnaeus)
_Common name._--Giant toad, marine toad, Surinam toad.
_Natural prey._--_Epilampra abdomen-nigrum_, Trinidad (Weber, 1938):
Found in the stomachs of two toads.
_Diploptera punctata_, Hawaii (Pemberton and Williams, 1938).
_Periplaneta_ sp., Fiji (Lever, 1939): Many householders in Suva have
seen the toad eat considerable numbers of these cockroaches.
_Pycnoscelus surinamensis_, Hawaii (Alicata, 1938; Illingworth, 1941).
Cockroaches, Nicaragua (Noble, 1918): Stomach contents of toads captured
at street lamps in Rio Grande consisted chiefly of large cockroaches.
Puerto Rico (Wolcott, 1937).
=Bufo valliceps= Wiegmann
_Experimental prey._--_Periplaneta americana_, U.S.A. (Moore, 1946):
Cockroaches containing infective acanthellas of _Moniliformis dubius_
were fed to three toads.
Family HYLIDAE
=Hyla cinerea= (Schneider)
_Common name._--Green tree frog.
_Natural prey._--_Ischnoptera deropeltiformis_, _Periplaneta americana_,
and undetermined cockroaches, U.S.A., Georgia (Haber, 1926): Cockroaches
were found in 11 of 100 stomachs.
Family RANIDAE
=Arthroleptis variabilis= Matschie
_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): Of 52
specimens examined, the stomach contents of 17 contained food, including
3 cockroaches.
=Hyperolius picturatus= Peters
_Natural prey._--Cockroach, Belgian Congo (Noble, 1924): The stomachs of
12 of 56 specimens examined contained food, including one cockroach.
=Leptodactylus albilabris= (Günther)
_Natural prey._--Cockroach, Puerto Rico (Schmidt, 1920): One of 25
stomachs contained a medium-sized cockroach.
=Leptodactylus pentadactylus= (Laurenti)
_Common name._--"Smoky jungle frog" or "pepper frog."
_Natural prey._--Cockroaches, Nicaragua (Noble, 1918): Cockroach wings
were found in the stomach of a frog caught around human habitation.
Brazil (Valente, 1949).
=Leptopelis calcaratus= (Boulenger)
_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): The stomachs
of 35 specimens were examined of which 13 contained food, including 2
cockroaches.
=Leptopelis rufus= Reichenow
_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): Forty-five of
83 stomachs examined contained food, including 2 cockroaches.
=Megalixalus fornasinii= (Bianconi)
_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): The stomachs
of 3 of 40 specimens contained food, including 2 cockroaches.
=Rana catesbeiana= Shaw
_Common name._--Bullfrog.
_Natural prey._--Cockroaches, Puerto Rico (Derez, 1949).
=Rana mascareniensis= Duméril and Bibron
_Natural prey._--Cockroaches, Belgian Congo (Noble, 1924): The stomach
contents of 138 specimens were examined, 39 of which contained food,
including 2 cockroaches.
=Rana pipiens= Schreber
_Common name._--Leopard frog.
_Experimental prey._--_Periplaneta americana_, U.S.A. (Moore, 1946):
Cockroaches containing infective acanthellas of _Moniliformis dubius_
were fed to two frogs.
_Neostylopyga rhombifolia_, U.S.A. (Dr. T. Eisner, personal
communication, 1958.)
=Frogs=
_Natural prey._--_Blatta orientalis_, U.S.A. (Rau, 1924): Frogs which
escaped from a tank in the cellar consumed quantities of this cockroach.
_Parcoblatta pensylvanica_, U.S.A. (Frost, 1924): One adult specimen
recovered from alimentary canal of a frog, probably _Rana_ sp.
=Unidentified batrachians=
_Experimental prey._--_Blattella germanica_, Germany, Frankfurt am Main,
Zoological Garden (Lederer, 1952): These insects were preferred by all
the insect eaters in the zoo.
_Periplaneta americana_, Germany, Frankfurt am Main, Zoological Garden
(Lederer, 1952): Newly molted individuals were accepted as food, but
others were usually passed by or consumed unwillingly.
Class REPTILIA[10]
Order CHELONIA
Family EMYDIDAE
=Chrysemys picta= (Schneider)
_Common name._--Painted turtle.
_Natural prey._--_Periplaneta australasiae_, England (Lucas, 1916,
1920): The cockroach, apparently injured, fell into water in the
tortoise house, Zoological Gardens, Regent's Park, and the terrapin ate
it.
Order SAURIA
Family GEKKONIDAE
=Gekko gecko= (Linnaeus)
_Natural prey._--Cockroaches, Philippine Islands, Laguna (Villadolid,
1934): The geckos frequent holes in trees and underside of bark which
are favorable haunts of cockroaches. Stomach contents mostly Blattidae
and "Locustidae."
=Hemidactylus frenatus= Duméril and Bibron
_Common name._--House lizard.
_Natural prey._--Cockroaches, Philippine Islands, Laguna (Villadolid,
1934): Bulk of stomach contents of 22 lizards consisted of Orthoptera,
mostly cockroaches.
=Sphaerodactylus= sp.
_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above
lizard is found in houses.
=Thecadactylus= sp.
_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above
lizard is found in houses.
=Undetermined geckos=
_Natural prey._--Cockroaches, Australia, Flinders River (Froggatt,
1906): The lizard lived in the walls of the hut and hunted cockroaches
upon the roof at night. Arno Atoll (Usinger and La Rivers, 1953).
Family IGUANIDAE
=Anolis carolinensis= Voigt
_Experimental prey._--_Diploptera punctata_, U.S.A. (Eisner, 1958).
=Anolis cristatellus= Duméril and Bibron
_Natural prey._--_Blattella_ sp., _Cariblatta delicatula_, _Epilampra
wheeleri_, _Periplaneta americana_, _Periplaneta australasiae_, and
_Symploce flagellata_, Puerto Rico (Wolcott, 1924): The last-named
cockroach may have been _S. ruficollis_ Rehn and Hebard, the females of
which are hard to distinguish from _flagellata_. Rehn and Hebard (1927)
stated that in all probability _flagellata_ does not occur on the island
of Puerto Rico. Wolcott (1950) stated that _Symploce ruficollis_
[= _bilabiata_] serves as food for the crested lizard.
Cockroaches, Puerto Rico (Schmidt, 1920): Of 100 stomachs examined, 16
contained Orthoptera, including cockroaches. Puerto Rico (Wolcott,
1924): One hundred _A. cristatellus_ had eaten 8 cockroaches, 4.14
percent of the total food, or 25 percent of the food for 8 lizards.
=Anolis pulchellus= Duméril and Bibron
_Natural prey._--Cockroaches, Puerto Rico (Wolcott, 1924): Two small
cockroaches found in 50 lizards examined.
=Anolis equestris= Merrem
_Experimental prey._--_Neostylopyga rhombifolia_, U.S.A. (Eisner,
personal communication, 1958.)
=Anolis grahami= Garman
_Natural prey._--_Periplaneta_ spp. and Blattidae, Bermuda (Simmonds,
1958): Stomachs of 176 lizards yielded 6 cockroaches.
=Anolis leachi= Duméril and Bibron
_Natural prey._--_Periplaneta_ spp. and Blattidae, Bermuda (Simmonds,
1958): Stomachs of 46 lizards yielded 31 cockroaches.
=Anolis sagrei= Cocteau
_Natural and experimental prey._--_Pycnoscelus surinamensis_, Cuba
(Darlington, 1938): This species was eaten both in captivity and in
nature. The lizard ate most readily soft, immature cockroaches.
_Pycnoscelus surinamensis_ is probably a staple food of the lizard in
nature, as Darlington observed wild lizards catch the nymphs.
=Anolis stratulus= Cope
_Natural prey._--_Aglaopteryx facies_, Puerto Rico (Wolcott, 1924): One
cockroach was found in 50 lizards examined.
Cockroach, Puerto Rico (Schmidt, 1920): One of 25 stomachs contained a
cockroach.
=Anolis= sp.
_Natural prey._--"Wood roaches," British Guiana (Beebe, 1925a): The
above lizard is arboreal on foliage in low jungle.
Family SCINCIDAE
=Leiolopisma laterale= Say
_Common name._--Brown skink.
_Natural prey._--Woodroaches, U.S.A., Louisiana (Slater, 1949): Analysis
of stomach contents of 84 adult skinks showed that nymphal and adult
woodroaches comprised the majority of Orthoptera.
=Tropidophorus grayi= Günther
_Common name._--Spiny lizard.
_Natural prey._--Cockroaches, Philippine Islands, Laguna (Villadolid,
1934): Food of this species was mostly Blattidae.
=Unidentified skinks=
_Natural prey._--Cockroaches, Arno Atoll (Usinger and La Rivers, 1953).
Family AGAMIDAE
_Experimental prey?_--Cockroaches, Australia (Lee and Mackerras, 1955):
A general statement was made that in captivity Agamidae were observed
feeding avidly on cockroaches and other insects. Three agamids studied
by these workers were _Amphibolurus barbatus_ (Gray), _Physignathus
lesueurii_ Gray, and _Chlamydosaurus kingii_ Gray.
Family CHAMAELEONTIDAE
=Chamaeleon chamaeleon= (Linnaeus) and =Chamaeleon oustaleti= Mocquard
_Experimental prey._--Cockroaches, Amsterdam (Portielje, 1914): Large
cockroaches were fed to these lizards in the reptile house of Artis.
Family TEIIDAE
=Ameiva exsul= Cope
_Common name._--Iguana, ground lizard.
_Natural prey._--Cockroach (nymph), _Epilampra wheeleri_, and
_Periplaneta americana_, Puerto Rico (Wolcott, 1924): Stomach contents
of 15 lizards were analyzed. _E. wheeleri_ formed 30 percent of the food
of one lizard. The cockroach nymph formed 5 percent of the food of one
lizard. One _P. americana_ formed 20 percent of the food of one lizard;
another formed 50 percent of the food of a second lizard.
_Experimental prey._--Cockroach nymphs, Puerto Rico (Wolcott, 1924).
=Ameiva= sp.
_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above
lizard is terrestrial and found near clearings. The stomach contents of
18 out of 40 reptiles contained cockroach remains.
=Cnemidophorus= sp.
_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): This is a
terrestrial lizard found near clearings. The stomach contents of 4 out
of 40 lizards contained cockroaches.
=Unidentified lizards=
_Natural prey._--Cockroaches, West Indies (H., 1800).
_Experimental prey._--_Blatta orientalis_, U.S.A. (Rau, 1924): Rau
called the predator a common gray lizard.
_Periplaneta americana_, Germany, Frankfurt am Main, Zoological Garden
(Lederer, 1952): Newly molted cockroaches were accepted as food, but
others were usually passed by or consumed unwillingly.
Order SERPENTES
Family COLUBRIDAE
=Heterodon platyrhinos= Latreille
_Synonymy._--_Heterodon contortrix_ [Dr. Doris M. Cochran, personal
communication, 1957].
_Common name._--Hog-nosed snake.
_Experimental prey._--_Periplaneta americana_, U.S.A. (Moore, 1946):
Cockroaches containing infective acanthellas of _Moniliformis dubius_
were fed to one snake.
=Garter Snake=
_Experimental prey._--_Blatta orientalis_, U.S.A. (Rau, 1924).
Class AVES
The cockroach is always wrong when arguing with a chicken.
_Spanish proverb_ (HARTNACK, 1939)
Arboreal cockroaches hidden in and under bark are much more likely to be
encountered by birds than by other predators, and insectivorous birds
undoubtedly consume many more cockroaches than the few records would
indicate. Most of the records we have located identify the birds at
least by common name. Where possible we have given the scientific names
for those birds whose common names are recognizably specific. We have
followed the systematic classification of Wetmore (1940).
Figuier (1869) stated that poultry and owls are very fond of
cockroaches. Perkins (1913) made the general statement that some of the
native birds of Hawaii are partial to the endemic _Allacta similis_.
Asano (1937) stated that in Japan natural enemies of cockroaches may be
found in the Galliformes, Strigiformes, Passeriformes, and Piciformes.
Although Lederer (1952) successfully fed newly molted _Periplaneta
americana_ to insectivorous birds in the Zoological Garden, Frankfurt am
Main, _Blattella germanica_ were preferred by these birds. The following
records are of specific birds feeding on cockroaches.
Order ANSERIFORMES
Family ANATIDAE
=Domestic duck=
_Natural prey._--_Pycnoscelus surinamensis_, Australia (Fielding 1926):
The ducks became infected with Manson's eye worm of which _P.
surinamensis_ is the only known intermediate host.
Cockroach, Bermuda (Jones, 1859): "All kinds of poultry feed greedily
upon the cockroach; tame ducks spending entire moonlight nights in their
capture."
Order GALLIFORMES
Family PHASIANIDAE
=Bambusicola thoracica= Temminck
_Common name._--Kojukei.
_Natural prey._--Cockroaches, Japan (Asano, 1937).
=Gallus= sp.
_Common name._--Jungle fowl.
_Natural prey._--_Periplaneta australasiae_, Hawaii (Schwartz and
Schwartz, 1949).
=Phasianus calchicus karpowi= Buturlin
_Common name._--Korean pheasants.
_Experimental prey._--_Blattella germanica_ and _Periplaneta picea_,
Japan (Asano, 1937): Adults of these cockroaches were devoured at once
when they were fed with the heads cut off.
=Phasianus= sp.
_Common name._--Pheasant.
_Natural prey._--Blattidae, unidentified (below 1 percent of the diet),
_Cutila soror_ (below 1 percent of the diet), _Diploptera punctata_
(above 6 percent of the diet), and _Pycnoscelus surinamensis_ (6 percent
of the diet), Hawaii (Schwartz and Schwartz, 1949).
=Coturnix coturnix japonica= (Temminck and Schlegel)
_Common name._--Japanese quail.
_Natural prey._--Blattidae (unidentified) and _Lobopterella
dimidiatipes_, Hawaii (Schwartz and Schwartz, 1949).
=Domestic chicken=
_Natural and experimental prey._--_Blaberus craniifer_, U.S.A., Key
West, Florida. J.A.G. Rehn in 1912 (personal communication) observed
chickens feeding on nymphs of _B. craniifer_ which had dropped to the
ground from among stacked coffins in an undertaker's shack.
_Blatta orientalis_, U.S.A. (Rau, 1924): The chickens ate cockroaches
that were caught in traps.
_Hebardina concinna_, Japan (Asano, 1937): Experimental feeding to white
Leghorn chickens.
_Periplaneta americana_, Surinam (Stage, 1947): Several cockroaches ran
off the floor of a house, which was being sprayed with DDT, and were
eaten by chickens. Although some chickens had DDT tremors the next day,
all appeared normal two days later.
_Pycnoscelus surinamensis_, Australia (Fielding, 1926); Formosa,
experimental feeding (Kobayashi, 1927); Australia, experimental feeding
(Fielding, 1927, 1928); U.S.A., Florida, experimental feeding (Sanders,
1928); Antigua (Hutson, 1943); Hawaii (Illingworth, 1931; Schwabe, 1949,
1950a, 1950b). This cockroach is the intermediate host of _Oxyspirura
mansoni_, the chicken eye worm.
Cockroaches, Guadeloupe (Dutertre, 1654); Africa (Moiser, 1947):
"Poultry" ate cockroaches which had been killed by DDT and sodium
fluoride. Hawaii (Zimmerman, 1948).
=Partridge=
_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The food of
two small species of leaf-colored partridges that lived on the jungle
floor, consisted chiefly of cockroaches and beetles.
Family MELEAGRIDIDAE
=Meleagris gallopavo= (Linnaeus)
_Common name._--Turkey.
_Natural prey._--_Pycnoscelus surinamensis_, Antigua (Hutson, 1943):
Turkeys were found heavily infected with Manson's eye worm of which _P.
surinamensis_ is the only known intermediate host. These turkeys
therefore were presumed to have fed on this cockroach.
Order COLUMBIFORMES
Family COLUMBIDAE
=Streptopelia chinensis= (Scopoli)
_Common name._--Chinese dove.
_Natural prey._--_Pycnoscelus surinamensis_, Hawaii (Schwabe, 1950b).
=Pigeon=
_Experimental prey._--_Pycnoscelus surinamensis_, Australia (Fielding,
1927); U.S.A., Florida (Sanders, 1928).
Order STRIGIFORMES
Family STRIGIDAE
=Gymnasio nudipes= (Daudin)
_Common name._--Bare-legged owl.
_Natural prey._--_Epilampra_ sp., Puerto Rico (Wetmore, 1916): One
specimen identified in stomach of a wild-caught owl.
Cockroaches, Puerto Rico (Wetmore, 1916): These insects were found in
stomachs of five owls.
Order CORACIFORMES
Family TODIDAE
=Todus mexicanus= Lesson
_Common name._--Porto Rican tody.
_Natural prey._--_Plectoptera poeyi?_, Puerto Rico (Wetmore, 1916): The
stomachs of 89 birds were examined; a single bird had eaten the above
cockroach. According to Wolcott (1950) _P. poeyi_ could be _Plectoptera
dorsalis_, _P. rhabdota_, or _P. infulata_.
Family BUCEROTIDAE
=Tockus birostris= (Scopoli)
_Synonymy._--_Lophocerus birostris_ [Dr. H. Friedmann, personal
communication, 1957].
_Common name._--Common gray hornbill.
_Natural prey._--Cockroaches, India, Central Provinces (D'Abreu, 1920).
Order PICIFORMES
Family PICIDAE
=Dendrocopus mahrattensis= (Latham)
_Synonymy._--_Liopicus mahrattensis_ [Friedmann, p. c. 1957].
_Common name._--Yellow-fronted pied woodpecker.
_Natural prey._--Cockroaches, India, Central Provinces (D'Abreu, 1920).
=Melanerpes portoricensis= (Daudin)
_Common name._--Puerto Rican woodpecker.
_Natural prey._--_Pycnoscelus surinamensis_, Puerto Rico (Wetmore,
1916): One specimen found in 59 bird stomachs examined.
Order PASSERIFORMES
Family FORMICARIIDAE
=Gymnopithys leucaspis= (Sclater)
_Common name._--Bicolored antbird.
_Natural prey._--Cockroaches, Panama Canal Zone (Johnson, 1954): This
bird feeds on small cockroaches, and other arthropods, which are flushed
from their hiding places by swarms of the army ant, _Eciton burchelli_.
Family ORIOLIDAE
=Icterus portoricensis= (Bryant)
_Common name._--Puerto Rican oriole.
_Natural prey._--Cockroaches, Puerto Rico (Wetmore, 1916): Cockroaches
and oöthecae found in the birds' stomachs.
Family CORVIDAE
=Aphelocoma coerulesens= (Bosc)
_Common name._--Florida jay.
_Experimental prey._--_Pycnoscelus surinamensis_, U.S.A., Florida
(Sanders, 1928).
=Cyanocitta cristata= (Linnaeus)
_Common name._--Blue jay.
_Experimental prey._--_Diploptera punctata_, U.S.A. (Eisner, 1958).
_Eurycotis floridana_, _Neostylopyga rhombifolia_, and _Periplaneta
americana_, U.S.A. (Eisner, personal communication, 1958): _E.
floridana_ was only eaten after the odor of 2-hexenal, which was
released by the insect on being attacked by the bird, had dissipated.
Family PARADISEIDAE
=Paradisea papuana= Bechstein
_Experimental prey._--Cockroaches, Malaya and on shipboard (Wallace,
1869): Two adult males fed voraciously on rice, bananas, and
cockroaches. Wallace collected cockroaches every night on board ship to
feed the birds. "At Malta ... I got plenty of cockroaches from a
bakehouse, and when I left, took with me several biscuit-tins full, as
provision for the voyage home."
Family TROGLODYTIDAE
=Troglodytes aedon= Vieillot
_Common name._--House wren.
_Natural prey._--Cockroaches, U.S.A. (Greenewalt and Jones, 1955): The
wren carried three small cockroaches to nestlings; the records probably
represent incidental captures.
=Troglodytes audax= Tschudi
_Common name._--Cucarachero.
_Natural prey._--Cockroach (called Chilicabra by Peruvian Indians), Peru
(Tschudi, 1847): The bird seized the cockroach and bit off its head then
devoured the body discarding the wings.
Family LANIIDAE
=Lanius ludovicianus= Linnaeus
_Common name._--Loggerhead shrike.
_Experimental prey._--_Pycnoscelus surinamensis_, U.S.A., Florida
(Sanders, 1928).
Family STURNIDAE
=Acridotheres tristis= Bonnaterre and Vieillot
_Common name._--Myna, mynah.
_Natural prey._--Cockroaches, Hawaii, Lanai (Illingworth, 1928):
Illingworth reported that he had never seen as many cockroaches anywhere
else in Hawaii. The birds followed tractors that were destroying cactus
and kept close to the chain that turned over the stumps. The following
species were collected: _Allacta similis_, _Blattella germanica_,
_Cutilia soror_, _Diploptera punctata_, _Leucophaea maderae_,
_Periplaneta americana_, _Periplaneta australasiae_, _Pycnoscelus
surinamensis_. Illingworth did not state whether the birds ate all these
species indiscriminately.
_Pycnoscelus surinamensis_, Hawaii (Williams et al., 1931; Schwabe,
1950b): In many places this species forms an important fledgling food
for mynah birds.
Family VIREONIDAE
=Vireo latimeri= Baird
_Common name._--Latimer's vireo.
_Natural prey._--_Periplaneta_ sp., Puerto Rico (Wetmore, 1916):
Cockroaches were found in one of 43 stomachs examined.
Family ICTERIDAE
=Agelaius xanthomus= (Sclater)
_Common name._--Yellow-shouldered blackbird.
_Natural prey._--Cockroaches, Puerto Rico (Wetmore, 1916): Oöthecae and
remains of adult cockroaches found in stomachs.
=Dolichonyx oryzivorus= (Linnaeus)
_Common name._--Bobolink.
_Experimental prey._--_Pycnoscelus surinamensis_, U.S.A., Florida
(Sanders, 1928).
=Holoquiscalus brachypterus= (Cassin)
_Common name._--Puerto Rican blackbird.
_Natural prey._--Cockroaches, Puerto Rico (Wetmore, 1916): A few eggs
(oöthecae) of cockroaches in stomachs.
=Black bird=
_Experimental prey._--_Pycnoscelus surinamensis_, U.S.A., Florida
(Sanders, 1928).
Family FRINGILLIDAE
=Passer domesticus= (Linnaeus)
_Common name._--English sparrow.
_Natural prey._--_Pycnoscelus surinamensis_, Hawaii (Illingworth, 1931;
Schwabe, 1950b): Remains of this cockroach were found in the stomach of
the sparrow.
=Sparrow=
_Natural prey._--_Periplaneta americana_, England (Lucas, 1908, 1920).
Cockroaches, Japan (Asano, 1937).
=Tiaris bicolor omissa= (Jardine)
_Common name._--Carib grassquit.
_Natural prey._--Cockroaches, Puerto Rico (Wetmore, 1916): Animal food
was found in 5 of 72 stomachs examined; one bird had eaten two
cockroaches among other insects.
Class MAMMALIA[11]
Order MARSUPIALIA
Family DIDELPHIDAE
=Monodelphis= sp.
_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above
opossum is nocturnal and arboreal but nests on the ground in grass.
Order INSECTIVORA
Family ERINACEIDAE
=Erinaceus europaeus= Linnaeus
_Experimental prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): Two hedgehogs were fed cockroaches infested with _Prosthenorchis
elegans_ and _P. spirula_.
=Erinaceus= sp.
_Common name._--Hedgehog.
_Natural prey._--Cockroaches, England (Samouelle, 1841; Cowan, 1865).
Order CHIROPTERA
Family MOLOSSIDAE
=Molossus= sp.
_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a): The above
bat is a common house bat of the area.
Order PRIMATES
Family LEMURIDAE
=Lemur coronatus= Gray
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): The monkey apparently became infested naturally with
_Prosthenorchis spirula_ for which _B. germanica_ was the intermediate
host in the monkey house.
=Lemur fulvus= E. Geoffroy
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comment under _Lemur coronatus_.
Family LORISIDAE
=Loris tardigradus= (Linnaeus)
_Synonymy._--_Lemur tardigradus_ [Dr. D. H. Johnson, personal
communication, 1957].
_Natural prey._--Cockroaches, on board ship (Cowan, 1865).
=Perodicticus potto= (P. L. S. Müller)
_Common name._--Potto.
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comment under _Lemur coronatus_.
_Experimental prey._--Blattidae, East Africa (Pitman, 1931): Both sexes
of the potto ate freely of all types of cockroaches.
Family TARSIIDAE
=Tarsius= sp.
_Experimental prey._--Cockroaches, Borneo (Shelford, 1916).
Family CEBIDAE
=Aotes zonalis= Goldman
_Synonymy._--_Aotus_ [Simpson, 1945].
_Common name._--Canal Zone night monkey.
_Natural prey._--_Leucophaea maderae_, Panama (Foster and Johnson,
1939): Captive monkeys became naturally infested with _Protospirura
muricola_ by eating cockroaches that contained infective larvae of the
worm.
=Ateles dariensis= Goldman
_Common name._--Darien black spider monkey.
_Natural prey._--_Leucophaea maderae_, Panama (Foster and Johnson,
1939): See comment under _Aotes zonalis_.
=Cebus apella= (Linnaeus)
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comment under _Lemur coronatus_.
=Cebus capucinus= (Linnaeus)
_Common name._--White-faced monkey.
_Natural prey._--_Leucophaea maderae_, Panama (Foster and Johnson,
1939): Favorite item of food in the laboratory. See comment under _Aotes
zonalis_.
=Saimiri sciurea= Linnaeus
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comment under _Callithrix chrysoleucos_.
Family CALLITHRICIDAE
=Callithrix chrysoleucos= (Natterer)
_Synonymy._--_Callithrix chrysolevea_ [Johnson, personal communication,
1957].
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): The monkey apparently became infested naturally with
_Prosthenorchis elegans_ for which _B. germanica_ was the intermediate
host in the monkey house.
=Callithrix jacchus= (Linnaeus)
_Synonymy._--_Simia jacchus._
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comment under _Lemur coronatus_.
Cockroaches, on board ship (Neill, 1829; also cited by Samouelle, 1841,
and Cowan, 1865): "It was quite amusing to see it at its meal. When he
had got hold of one of the largest cockroaches, he held it in his fore
paws, and then invariably nipped the head off first; he then pulled out
the viscera and cast them aside, and devoured the rest of the body,
rejecting the dry elytra and wings, and also the legs of the insect,
which are covered with short stiff bristles. The smaller cockroaches he
eat[s] without such fastidious nicety."
=Leontocebus oedipus= (Linnaeus)
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comment under _Callithrix chrysoleucos_.
=Leontocebus rosalia= (Linnaeus)
_Synonymy._--_Midas rosalia_ [Simpson, 1945].
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comments under _Lemur coronatus_ and _Callithrix
chrysoleucos_.
=Leontocebus ursulus= (E. Geoffroy)
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comments under _Callithrix chrysoleucos_.
Family CERCOPITHECIDAE
=Cercopithecus= sp.
_Experimental prey._--Cockroaches, East Africa (Carpenter, 1921, 1925):
The monkey rarely tasted and usually ignored cockroaches offered to it.
In one experiment the monkey had to be deprived of food before it would
eat the cockroach.
=Macaca mulatta= (Zimmermann)
_Synonymy._--_Macaca rhesus_ [Johnson, personal communication, 1957].
_Experimental prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938, 1938a): The macaque was fed cockroaches infested with
_Prosthenorchis elegans_ and _P. spirula_.
=Macaca sylvanus= (Linnaeus)
_Synonymy._--_Inuus sylvanus_ [Simpson, 1945].
_Common name._--Macaque.
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): See comment under _Lemur coronatus_.
=Papio papio= (Desmarest)
_Experimental prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938, 1938a): This baboon was fed cockroaches infested with
_Prosthenorchis elegans_ and _P. spirula_.
Family PONGIDAE
=Pan= sp.
_Common name._--Chimpanzee.
_Natural prey._--_Blattella germanica_, Netherlands (Thiel and Wiegand
Bruss, 1946): Indirect evidence for this relationship was shown by these
workers who found two animals heavily infected with _Prosthenorchis
spirula_ in a zoo in Rotterdam; the intermediate host of the worm was
shown to be _B. germanica_.
Family HOMINIDAE
=Homo sapiens= Linnaeus
_Natural prey._--Oöthecae of _Blatta orientalis_ and _Neostylopyga
rhombifolia_, Thailand (Bristowe, 1932).
_Periplaneta americana_, Formosa (Takahashi, 1924).
_Periplaneta americana_ and _Periplaneta australasiae_, Australia,
China, and Japan (Bodenheimer, 1951).
Cockroaches, Annam and French Guinea (Brygoo, 1946).
In addition to the above records of cockroaches being used as food by
man these insects have also been eaten for medicinal purposes (see Roth
and Willis, 1957a).
Order EDENTATA
Family DASYPODIDAE
=Dasypus novemcinctus= Linnaeus
_Synonymy._--_Tatu novemcinctum_ [Johnson, personal communication,
1958].
_Natural prey._--_Ischnoptera deropeltiformis_, Texas (Hebard, 1917): A
specimen of this cockroach in the U. S. National Museum was taken from
the stomach of the armadillo.
Order RODENTIA
Family MURIDAE
=Mus musculus= Linnaeus
_Experimental prey._--_Diploptera punctata_, U.S.A. (Eisner, 1958).
=Rattus norvegicus= (Berkenhout)
_Synonymy._--_Mus decumanus; Epimys norvegicus._
_Natural prey._--_Leucophaea maderae_, Venezuela (Brumpt, 1931): Rats
infested with _Protospirura bonnei_ presumably ate this cockroach which
is the intermediate host of the worm.
_Periplaneta americana_, Brazil (Magalhães, 1898): Remains found in the
stomachs of brown rats. Denmark (Fibiger and Ditlevsen, 1914): This
cockroach was found to be the intermediate host of _Gongylonema
neoplasticum_, a parasite of rats.
=Rattus rattus= (Linnaeus)
_Natural prey._--_Periplaneta americana_, Denmark (Fibiger and
Ditlevsen, 1914): See comment after these authors under _Rattus
norvegicus_.
=Rattus= spp.
_Natural prey._--Cockroaches, India (Maxwell-Lefroy, 1909); Burma
(Subramanian, 1927).
Family CAVIIDAE
=Cavia= sp.
_Experimental prey._--_Blattella germanica_, U.S.A. (Hobmaier, 1941):
Guinea pigs were fed cockroaches infested with _Physaloptera
maxillaris_.
Order CARNIVORA
Family CANIDAE
=Canis familiaris= Linnaeus
_Experimental prey._--_Blattella germanica_, U.S.A. (Hobmaier, 1941):
Dogs were fed cockroaches infested with _Physaloptera maxillaris_.
U.S.A. (Petri and Ameel, 1950): Cockroaches infested with _Physaloptera
rara_ were fed to a dog.
=Canis latrans= Say
_Experimental prey._--_Blattella germanica_, U.S.A. (Petri and Ameel,
1950): Cockroaches infested with _Physaloptera rara_ were fed to a
coyote.
=Vulpes= sp.
_Experimental prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938a): A fox was successfully infected when fed cockroaches infested
with _Prosthenorchis elegans_ and _P. spirula_.
Family PROCYONIDAE
=Bassariscus astutus= (Lichtenstein)
_Common names._--Cacomistle, ring-tailed cat.
_Natural prey._--Cockroaches, U.S.A., Arizona (Dr. H. Stahnke, personal
communication, 1953): The ring-tailed cat enters dwellings located on
the desert and feeds on cockroaches and other arthropods.
=Nasua narica= (Linnaeus)
_Natural prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938): The coati apparently became infested naturally with
_Prosthenorchis spirula_ for which _B. germanica_ was the intermediate
host in the laboratory.
=Nasua nasua= (Linnaeus)
_Natural prey._--_Blattella germanica_, on board ship (Myers, 1931):
This insect was eaten when other insects were absent.
Cockroach, a small outdoor species, Trinidad (Myers, 1931).
=Nasua= sp.
_Natural prey._--Cockroaches, British Guiana (Beebe, 1925a).
Family MUSTELIDAE
=Meles= sp.
_Experimental prey._--_Blattella germanica_, France (Brumpt and Urbain,
1938a): A badger was successfully infected when fed cockroaches infested
with _Prosthenorchis elegans_ and _P. spirula_.
Family VIVERRIDAE
=Herpestes javanicus auropunctatus= Hodgson
_Natural prey._--_Epilampra wheeleri_, _Eurycotis improcera_, _Panchlora
nivea_, _Pycnoscelus surinamensis_, and others unidentified to species,
St. Croix and Puerto Rico (Wolcott, 1953): Based on 37 or more
cockroaches obtained from stomachs of 42 mongooses collected in St.
Croix (by Seaman) and 56 collected in Puerto Rico (by Pimentel).
Pimentel (personal communication, 1958) has given us the following
percentage occurrence of cockroach species in the total number of
mongoose stomachs that he examined in Puerto Rico: _Epilampra wheeleri_
1.8, _Ischnoptera rufa rufa_ 3.6, _Panchlora nivea_ 1.8, _Periplaneta
americana_ 1.8, and _Pycnoscelus surinamensis_ 19.6.
=Herpestes= sp.
_Natural prey._--_Periplaneta americana_ and _Periplaneta australasiae_,
Hawaii (Perkins, 1913): Large numbers of these cockroaches are devoured.
Cockroach, East Africa (Loveridge, 1923): Cockroach remains found in
stomach of mongoose.
Family FELIDAE
=Felis catus= Linnaeus
_Natural prey._--_Periplaneta americana_, Hawaii (Williams et al.,
1931).
Cockroaches, U.S.A., Arizona (Stahnke, personal communication, 1953).
_Experimental prey._--_Blattella germanica_, U.S.A. (Hobmaier, 1941):
Cats were fed cockroaches infested with _Physaloptera maxillaris_.
U.S.A. (Petri and Ameel, 1950): Cockroaches infested with _Physaloptera
rara_ were fed to a kitten. France (Brumpt and Urbain, 1938a): A young
cat was fed cockroaches infested with _Prosthenorchis elegans_ and _P.
spirula_.
=Felis pardalis mearnsi= J. A. Allen
_Natural prey._--Cockroaches, Panama (Dr. H. L. Sweetman, personal
communication, 1958): An ocelot was seen collecting and feeding on
cockroaches, possibly _Blaberus_ sp. "The ocelot was quite efficient and
seemed to relish the roaches."
XIV. CHECKLIST OF COCKROACHES AND SYMBIOTIC ASSOCIATES
Only naturally occurring associations are included in this list.
Commensal cockroaches are listed on page 315. Bacteroids are not listed
because they undoubtedly occur in all species. The higher plants were
excluded because most of the associations may be too casual to
constitute symbiosis; however, many of the plant associations were
included in the chapter on ecology. The cockroaches and the associates
within each category are arranged alphabetically by genus and species.
Page references are to citations in the classified sections where
details of the associations and/or sources of the records are given.
=Aglaopteryx facies=
Mite: Undetermined, p. 220.
Reptile: _Anolis stratulus_, p. 274.
=Aglaopteryx diaphana=
Nematode: _Protrellus manni_, p. 200.
=Allacta similis=
Insects:
_Dolichurus stantoni_, p. 262.
_Solindenia picticornis_, p. 247.
=Anaplecta= sp.
Fungus: _Herpomyces anaplectae_, p. 134.
=Aptera fusca=
Protozoan: _Gregarina fastidiosa_, p. 182.
=Arenivaga bolliana=
Insect: _Sarcophaga omani_, p. 229.
=Arenivaga roseni=
Insects: Undetermined reduviids, p. 227.
=Balta patula=
Insect: _Rhipidioides ableptus_, p. 230.
=Blaberus atropos=
Bacterium: _Spirochaeta blattae_, p. 125.
Protozoa:
_Endolimax nana_, p. 180.
_Entamoeba coli_, p. 178.
Nematodes:
_Leidynema appendiculata_, p. 197.
_Leidynema cranifera_, p. 198.
=Blaberus craniifer=
Bacteria:
_Aerobacter aerogenes_, p. 111.
_Alcaligenes faecalis_, p. 111.
_Bacillus cereus_, p. 120.
_Bacillus subtilis_, p. 121.
_Escherichia coli_ var. _communior_, p. 113.
_Escherichia freundii_, p. 113.
_Micrococcus pyogenes_ var. _albus_, p. 106.
_Micrococcus pyogenes_ var. _aureus_, p. 107.
_Proteus vulgaris_, p. 114.
_Pseudomonas aeruginosa_, p. 104.
Fungi:
_Herpomyces tricuspidatus_, p. 138.
_Penicillium_ sp., p. 131.
_Rhizopus nigricans_, p. 133.
_Saccharomyces cerevisiae_, p. 133.
Protozoan: _Diplocystis_ (?) sp., pp. 181, 184.
Nematodes:
_Leidynema cranifera_, p. 198.
_Protrelleta floridana_, p. 199.
Mite: _Iolina nana_, p. 219.
Undetermined, p. 220.
Bird: Chicken, p. 278.
=Blaberus discoidalis=
Mite: Undetermined, p. 220.
=Blaberus giganteus=
Hair worm: _Gordius pilosus_, p. 202.
=Blaberus= sp.
Fungi:
_Herpomyces macropus_, p. 136.
_Herpomyces paranensis_, p. 136.
_Herpomyces periplanetae_, p. 137.
_Herpomyces tricuspidatus_, p. 138.
Protozoan: _Leptomonas blaberae_, p. 167.
=Blaptica dubia=
Protozoan: _Pileocephalus blaberae_, p. 184.
=Blatta (Shelfordella) lateralis=
Bacterium: _Shigella paradysenteriae_, p. 119.
Insect: _Ampulex assimilis_, p. 257.
=Blatta orientalis=
Bacteria:
_Aerobacter aerogenes_, p. 111.
_Alcaligenes faecalis_, p. 111.
_Alcaligenes recti_, p. 111.
_Arthromitus intestinalis_, p. 124.
B. aerobio del pseudoedema maligno, p. 125.
_B. alcaligenes beckeri_, p. 125.
B. del pseudoedema maligno, p. 125.
Bacillo proteisimile, p. 126.
Bacillo similcarbonchio, p. 126.
Bacillo similtifo (Bacillo tifosimile), p. 126.
_Bacillus bütschlii_, p. 120.
_Bacillus periplanetae_, p. 121.
_Bacillus stellatus_, p. 121.
_Bacillus subtilis_, p. 122.
_Bacillus tritus_, p. 122.
_Bacteroides uncatus_, p. 119.
_Clostridium lentoputrescens_, p. 122.
_Clostridium novyi_ or _Clostridium sporogenes_, p. 122.
_Enterococcus_ sp., p. 109.
_Escherichia coli_, p. 112.
_Lactobacillus fermenti_, p. 109.
_Micrococcus pyogenes_ var. _albus_, p. 107.
_Micrococcus pyogenes_ var. _aureus_, p. 107.
_Micrococcus_ sp., p. 108.
_Paracolobactrum_ sp., p. 113.
_Pasteurella pestis_, p. 119.
_Proteus_ sp., p. 114.
_Proteus vulgaris_, p. 114.
_Pseudomonas aeruginosa_, p. 104.
_Pseudomonas eisenbergii_, p. 105.
_Pseudomonas fluorescens_, p. 105.
_Salmonella typhosa_, p. 117.
_Sarcina alba_, p. 108.
_Sarcina_ sp., pp. 108, 109.
_Sarcina symbiotica_ (host may have been _B. germanica_), p. 108.
_Sarcina ventriculi_, p. 108.
_Serratia marcescens_, p. 117.
_Spirillochaeta blattae_, p. 127.
_Spirillum_ α, β and γ, p. 105.
_Spirillum_ sp., p. 105.
_Spirochaeta periplanetae_, p. 125.
_Streptococcus faecalis_, p. 109.
_Streptococcus liquefaciens_, p. 110.
_Streptococcus microapoika_, p. 110.
_Streptococcus pyogenes_, p. 110.
_Streptococcus_ sp., p. 110.
_Treponema parvum_, p. 125.
_Treponema stylopygae_, p. 125.
_Vibrio_ sp., p. 106.
Fungi:
_Aspergillus fumigatus_, p. 130.
_Blastocystis hominis_, p. 133.
_Blastocystis_ sp., p. 133.
_Candida zeylanoides_, p. 129.
_Coccidioides periplanetae_, p. 133.
_Herpomyces periplanetae_, p. 137.
_Herpomyces stylopygae_, p. 137.
_Torula gropengiesseri_, p. 132.
_Torulopsis_ sp., p. 130.
Protozoa:
_Balantidium praenucleatum_, p. 187.
_Bodo blattae_, p. 167.
_Coelosporidium periplanetae_, p. 185.
_Diplocystis schneideri_, p. 181.
_Endamoeba blattae_, p. 177.
_Entamoeba thomsoni_, p. 179.
_Endolimax blattae_, p. 180.
_Endolimax_ sp., p. 180.
_Gregarina blattarum_, p. 181.
_Haplosporidium periplanetae_, p. 185.
_Hartmannella blattae_, p. 177.
_Herpetomonas periplanetae_, p. 167.
_Hexamita periplanetae_, p. 171.
_Lophomonas blattarum_, p. 172.
_Lophomonas striata_, p. 173.
_Monas_ sp., p. 167.
_Monocercomonoides orthopterorum_, p. 169.
_Nyctotherus ovalis_, p. 188.
_Oikomonas_ sp., p. 166.
_Peltomyces periplanetae_, p. 177.
_Plistophora kudoi_, p. 185.
_Plistophora periplanetae_, p. 186.
_Plistophora_ sp., p. 186.
_Retortamonas blattae_, p. 167.
_Stenophora_ sp., p. 181.
_Tetratrichomastix blattidarum_, p. 170.
Helminths:
_Ascaris_ sp., p. 209.
_Enterobius vermicularis_, p. 209.
_Gongylonema neoplasticum_, p. 206.
_Gordius blattae orientalis_, p. 202.
_Hammerschmidtiella diesingi_, p. 195.
_Hammerschmidtiella neyrai_, p. 196.
_Leidynema appendiculata_, p. 197.
_Spirura gastrophila_, p. 207.
_Thelastoma pachyjuli_, p. 201.
_Trichostrongylus_ sp., p. 210.
_Trichuris trichiura_, p. 210.
Mite: Undetermined, pp. 220, 222.
Insects:
_Dermestes ater_, p. 234.
_Dermestes_ sp., p. 234.
_Evania appendigaster_, p. 236.
_Evania dimidiata_, p. 239.
_Prosevania punctata_, p. 240.
_Systellogaster ovivora_, p. 248.
_Tetrastichus hagenowii_, p. 250.
_Tetrastichus_ sp., p. 254.
Bird: Chicken, p. 278.
Mammal: _Homo sapiens_, p. 286.
=Blatta= sp.
Hair worm: _Gordius aquaticus_, p. 201.
Insects:
_Evania appendigaster_, p. 236.
_Tetrastichus hagenowii_, p. 250.
=Blattella germanica= and/or =Blattella vaga=
Viruses: Unspecified strains of poliomyelitis virus, p. 103.
=Blattella germanica=
Bacteria:
_Achromobacter_ sp., p. 110.
_Aerobacter aerogenes_, p. 111.
_Aerobacter cloacae_, p. 112.
_Alcaligenes faecalis_, p. 111.
_Alcaligenes viscosus_, p. 111.
_Bacillus circulans_, p. 120.
_Escherichia coli_, p. 112.
_Escherichia freundii_, p. 113.
_Micrococcus aurantiacus_, p. 106.
_Micrococcus epidermidis_, p. 106.
_Micrococcus pyogenes_ var. _albus_, p. 107.
_Micrococcus pyogenes_ var. _aureus_, p. 107.
_Micrococcus ureae_, p. 107.
_Micrococcus_ sp., p. 108.
_Mycobacterium leprae_, p. 123.
_Paracolobactrum aerogenoides_, p. 113.
_Paracolobactrum coliforme_, p. 113.
_Paracolobactrum_ sp., p. 113.
_Pseudomonas aeruginosa_, p. 104.
_Sarcina symbiotica_ (host may have been _B. orientalis_), p. 108.
_Salmonella typhimurium_, p. 116.
_Serratia marcescens_, p. 117.
_Streptococcus faecalis_, p. 109.
_Streptococcus_ sp., p. 110.
Fungi:
_Aspergillus flavus_, p. 130.
_Aspergillus tamarii_, p. 130.
_Aspergillus_ sp., p. 130.
_Cordyceps blattae_, p. 134.
_Herpomyces ectobiae_, p. 135.
_Memnoniella echinata_, p. 132.
_Saccharomyces_ sp., p. 133.
Protozoa:
_Bodo_ sp. (host may have been _P. americana_), p. 167.
_Coelosporidium periplanetae_, p. 185.
_Dobellina_ sp. (host may have been _P. americana_), p. 177.
_Endamoeba blattae_ (host may have been _P. americana_), p. 177.
_Endolimax_ sp. (host may have been _P. americana_), p. 180.
_Entamoeba coli_ (host may have been _P. americana_), p. 178.
_Entamoeba histolytica_ (host may have been _P. americana_), p. 179.
_Entamoeba thomsoni_, p. 179.
_Eutrichomastix_ sp. (host may have been _P. americana_), p. 169.
_Gregarina blattarum_, p. 182.
_Iodamoeba_ sp. (host may have been _P. americana_), p. 180.
_Lophomonas blattarum_, p. 172.
_Lophomonas striata_, p. 173.
_Nyctotherus ovalis_, p. 188.
_Peltomyces periplanetae_, p. 177.
_Plistophora periplanetae_, p. 186.
_Tetratrichomastix blattidarum_, p. 170.
Helminths:
_Blattelicola blattelicola_, p. 193.
_Blatticola blattae_, p. 193.
_Enterobius vermicularis_, p. 209.
_Galebia aegyptiaca_, p. 195.
_Gongylonema neoplasticum_, p. 206.
_Moniliformis kalahariensis_, p. 203.
_Prosthenorchis elegans_, p. 203.
_Prosthenorchis spirula_, p. 203.
_Tetrameres americana_, p. 207.
_Trichuris trichiura_, p. 210.
Mites:
_Blattisocius tineivorus_, p. 216.
_Caloglyphus_ sp., p. 218.
Undetermined, p. 220.
Insects:
_Brachygaster minutus_, p. 235.
_Eupelmus atriflagellum_, p. 247.
_Dolichurus corniculus_, p. 261.
_Ripidius pectinicornis_, p. 232.
Mammals:
_Callithrix chrysoleucos_, p. 285.
_Callithrix jacchus_, p. 285.
_Cebus apella_, p. 284.
_Lemur coronatus_, p. 283.
_Lemur fulvus_, p. 283.
_Leontocebus oedipus_, p. 285.
_Leontocebus rosalia_, p. 285.
_Leontocebus ursulus_, p. 285.
_Macaca sylvanus_, p. 286.
_Nasua narica_, p. 288.
_Nasua nasua_, p. 288.
_Pan_ sp., p. 286.
_Perodicticus potto_, p. 284.
_Saimiri sciurea_, p. 285.
=Blattella humbertiana=
Fungus: _Herpomyces gracilis_, p. 135.
Nematode: _Protrellus phyllodromi_, p. 200.
Mite: _Uropoda_ sp., p. 217.
=Blattella lituricollis=
Insect: _Dolichurus stantoni_, p. 262.
=Blattella= sp.
Reptile: _Anolis cristatellus_, p. 273.
=Cariblatta delicatula=
Insect: _Hyptia_ sp., p. 240.
Reptile: _Anolis cristatellus_, p. 273.
=Cariblatta lutea lutea=
Plants:
_Sarracenia flava_, p. 154.
_Sarracenia purpurea_, p. 154.
=Chorisoneura= sp.
Insects:
_Stylogaster_ sp., p. 228.
_Tachysphex blatticidus_, p. 264.
=Choristima= sp.
Insect: _Rhipidioides rubricatus_, p. 231.
=Choristimodes= sp.
Insect: _Rhipidioides rubricatus_, p. 231.
=Cryptocercus punctulatus=
Bacterium: _Bacillus subtilis_, p. 122.
Protozoa:
_Adelina cryptocerci_, p. 184.
_Barbulanympha estaboga_, p. 173.
_Barbulanympha laurabuda_, p. 174.
_Barbulanympha ufalula_, p. 174.
_Barbulanympha wenyoni_, p. 174.
_Eucomonympha imla_, p. 176.
_Hexamita cryptocerci_, p. 171.
_Idionympha perissa_, p. 174.
_Leptospironympha eupora_, p. 172.
_Leptospironympha rudis_, p. 172.
_Leptospironympha wachula_, p. 172.
_Macrospironympha xylopletha_, p. 172.
_Monocercomonoides globus_, p. 169.
_Notila proteus_, p. 170.
_Oxymonas doroaxostylus_, p. 170.
_Oxymonas nana_, p. 170.
_Prolophomonas tocopola_, p. 173.
_Rhynchonympha tarda_, p. 174.
_Saccinobaculus ambloaxostylus_, p. 170.
_Saccinobaculus lata_, p. 170.
_Trichonympha acuta_, p. 174.
_Trichonympha algoa_, p. 174.
_Trichonympha chula_, p. 176.
_Trichonympha grandis_, p. 176.
_Trichonympha lata_, p. 176.
_Trichonympha okolona_, p. 176.
_Trichonympha parva_, p. 176.
_Urinympha talea_, p. 174.
Undetermined gregarine, p. 184.
Amphibian: _Plethodon glutinosus_, p. 269.
=Cutilia soror=
Insects:
_Dolichurus stantoni_, p. 262.
_Evania appendigaster_, p. 236.
_Szepligetella sericea_, p. 242.
Bird: _Phasianus_ sp., p. 277.
=Cutilia= sp.
Nematode: Undetermined, p. 201.
Insects:
_Riekella australis_, p. 231.
Undetermined strepsipteron, p. 234.
=Diploptera punctata=
Bacterium: _Serratia marcescens_, p. 117.
Fungus: _Herpomyces diplopterae_, p. 135.
Mites:
_Iolina nana_, p. 219.
_Locustacarus_ sp., p. 219.
Insect: _Nauphoeta cinerea_, p. 324.
Amphibian: _Bufo marinus_, p. 270.
Bird: _Phasianus_ sp., p. 277.
=Ectobius lapponicus=
Protozoa:
_Gamocystis tenax_, p. 184.
_Monocercomonoides orthopterorum_, p. 169.
Nematode: _Blatticola blattae_, p. 193.
Insects:
_Brachygaster minutus_, p. 235.
_Dolichurus corniculus_, p. 261.
_Tachysphex lativalvis_, p. 264.
=Ectobius pallidus=
Protozoan: _Gamocystis tenax_, p. 184.
Helminths:
_Blatticola blattae_, p. 193.
Undetermined mermithid, p. 192.
Insects:
_Ampulex fasciata_, p. 259.
_Aphaenogaster picea_, p. 267.
_Dolichurus corniculus_, p. 261.
_Lasius alienus_, p. 268.
_Ripidius boissyi_ (presumptive record), p. 231.
_Ripidius denisi_ (presumptive record), p. 232.
_Tachysphex lativalvis_, p. 264.
=Ectobius panzeri=
Centipede: _Scolopendra_ sp., p. 224.
Insects:
_Dolichurus corniculus_, p. 261.
_Tachysphex lativalvis_, p. 264.
=Ectobius= sp.
Insects:
_Brachygaster minutus_, p. 235.
_Dolichurus corniculus_, p. 261.
_Tachysphex lativalvis_, p. 264.
=Ellipsidion affine=
Insect: _Rhipidioides fuscatus_, p. 230.
=Ellipsidion australe=
Insects:
_Agamerion metallica_, p. 243.
_Cheiloneurus viridiscutum_, p. 244.
_Mestocharomyia oophaga_, p. 248.
=Epilampra abdomen-nigrum=
Insect: _Podium flavipenne_, p. 265.
Amphibian: _Bufo marinus_, p. 270.
=Epilampra conferta=
Insect: _Podium_ sp., p. 266.
=Epilampra= sp.
Fungus: _Herpomyces tricuspidatus_, p. 138.
Insect: _Podium haematogastrum_, p. 266.
Bird: _Gymnasio nudipes_, p. 279.
=Epilampra wheeleri=
Reptiles:
_Ameiva exsul_, p. 275.
_Anolis cristatellus_, p. 273.
Mammal: _Herpestes javanicus auropunctatus_, p. 289.
=Escala= (?) sp.
Insect: _Rhipidioides adynatus_, p. 230.
=Eurycotis floridana=
Fungi:
_Aspergillus flavus_, p. 130.
_Aspergillus sydowi_, P. 130.
Helminths:
_Euryconema paradisa_, p. 194.
_Leidynema appendiculata_(?), p. 198.
_Protrelloides paradoxa_, p. 200.
Undetermined gordian worm, p. 202.
Insects:
_Anastatus floridanus_, p. 245.
Undetermined tachinid, p. 228.
=Eurycotis improcera=
Mammal: _Herpestes javanicus auropunctatus_, p. 289.
=Eurycotis manni=
Fungus: _Herpomyces zanzibarinus_, p. 138.
=Graptoblatta notulata=
Insect: _Tachysphex fanuiensis_, p. 264.
=Gromphadorhina portentosa=
Protozoan: Undetermined gregarine, p. 184.
Mite: _Coleolaelaps_ ? sp., p. 216.
=Gyna= sp.(?)
Fungus: _Herpomyces zanzibarinus_, p. 138.
=Holocompsa fulva=
Insect: _Pheidole megacephala_, p. 268.
=Hololampra punctata=
Insect: _Dolichurus corniculus_, p. 261.
=Ischnoptera deropeltiformis=
Plant: _Sarracenia flava_, p. 154
Amphibian: _Hyla cinerea_, p. 270.
Mammal: _Dasypus novemcinctus_, p. 287.
=Ischnoptera rufa rufa=
Fungus: _Spicaria prasina_, p. 130.
Mammal: _Herpestes javanicus auropunctatus_, p. 289.
=Ischnoptera= sp.
Fungus: _Herpomyces arietinus_, p. 134.
Insects:
_Ampulex canaliculata_, p. 257.
_Syntomosphyrum ischnopterae_, p. 249.
=Karnyia discoidalis=
Spider: _Latrodectus indistinctus_, p. 215.
=Kuchinga hemerobina=
Hair worm: _Parachordodes raphaelis_, p. 202.
=Leucophaea maderae=
Bacterium: _Serratia marcescens_, p. 117.
Fungus: _Herpomyces tricuspidatus_, p. 138.
Protozoa:
_Gregarina rhyparobiae_, p. 183.
_Hexamita_ (?) sp., p. 171.
_Retortamonas_ (?) sp., p. 167.
Undetermined gregarine, p. 184.
Nematodes:
_Hammerschmidtiella diesingi_, p. 195.
_Leidynema delatorrei_, p. 198.
_Protospirura bonnei_, p. 206.
_Protospirura muricola_, p. 206.
Mite: _Chaetodactylus_ sp., p. 218.
Mammals:
_Aotes zonalis_, p. 284.
_Ateles dariensis_, p. 284.
_Cebus capucinus_, p. 284.
_Rattus norvegicus_, p. 287.
=Leucophaea= sp.
Nematode: _Cephalobellus brevicaudatum_, p. 194.
=Leurolestes pallidus=
Fungus: _Herpomyces leurolestis_, p. 136.
=Loboptera decipiens=
Insects:
_Dolichurus corniculus_, p. 261.
_Zeuxevania splendidula_, p. 243.
=Loboptera= sp.
Fungus: _Herpomyces lobopterae_, p. 136.
=Lobopterella dimidiatipes=
Insect: _Ampulex canaliculata_, p. 257.
Bird: _Coturnix coturnix japonica_, p. 277.
=Melanosilpha capensis=
Protozoa:
_Gregarina impetuosa_, p. 183.
_Gregarina sandoni_, p. 183.
=Monastria= sp.
Insect: _Triatoma arthurneivai_, p. 227.
=Nauphoeta cinerea=
Bacteria:
_Salmonella typhimurium_, p. 116.
_Serratia marcescens_, p. 117.
Fungus: _Herpomyces tricuspidatus_, p. 138.
Mites:
_Blattilaelaps nauphoetae_, p. 216.
_Locustacarus_ sp., p. 219.
Insect: _Pheidole megacephala_, p. 268.
=Neostylopyga rhombifolia=
Bacterium: _Serratia marcescens_, p. 117.
Insects:
_Evania appendigaster_, p. 237.
_Szepligetella sericea_, p. 242.
_Tetrastichus hagenowii_, p. 250.
Mammal: _Homo sapiens_, p. 286.
=Nyctibora obscura=
Fungus: _Herpomyces amazonicus_, p. 134.
=Nyctibora= sp.
Fungus: _Herpomyces nyctoborae_, p. 136.
=Nyctibora tomantosa=
Fungus: _Herpomyces nyctoborae_, p. 136.
=Oniscosoma granicollis=
Insect: _Paranephrites xenus_, p. 230.
=Panchlora exoleta=
Protozoan: _Gregarina panchlorae_, p. 183.
=Panchlora nivea=
Bacterium: _Serratia marcescens_, p. 117.
Fungus: _Herpomyces panchlorae_, p. 136.
Mammal: _Herpestes javanicus auropunctatus_, p. 289.
=Panesthia angustipennis=
Protozoa:
_Clevelandella constricta_, p. 189.
_Clevelandella contorta_, p. 189.
_Clevelandella elongata_, p. 189.
_Clevelandella hastula_, p. 189.
_Clevelandella panesthiae_, p. 189.
_Clevelandella parapanesthiae_, p. 189.
_Endamoeba javanica_, p. 178.
_Endamoeba philippinensis_, p. 178.
_Hexamita cryptocerci_, p. 171.
_Monocercomonoides panesthiae_, p. 170.
_Nyctotherus uichancoi_, p. 188.
_Paraclevelandia brevis_, p. 190.
_Paraclevelandia simplex_, p. 190.
Undetermined amoeba, p. 181.
Nematodes:
_Aorurus philippinensis_, p. 193.
_Blattophila sphaerolaima_ var. _javanica_, p. 194.
_Leidynema nocalum_, p. 198.
_Leidynemella fusiformis_, p. 198.
_Leidynemella paracranifera_, p. 198.
_Thelastoma palmettum_, p. 201.
=Panesthia australis=
Fungus: _Metarrhizium anisopliae_, p. 131.
Mite:
_Hypoaspis_ sp., p. 217.
Undetermined diplogyniid, p. 217.
Insect: Undetermined tachinid, p. 228.
=Panesthia laevicollis=
Nematode: _Blattophila sphaerolaima_, p. 194.
=Panesthia laevicollis= (?)
Nematode: _Leidynemella fusiformis_, p. 198.
=Panesthia lobipennis=
Fungus: _Herpomyces panesthiae_, p. 136.
=Panesthia spadica=
Protozoa:
_Clevelandella constricta_, p. 189.
_Clevelandella contorta_, p. 189.
_Clevelandella nipponensis_, p. 189.
_Clevelandella panesthiae_, p. 189.
_Endamoeba javanica_, p. 178.
_Nyctotherus uichancoi_, p. 188.
_Paraclevelandia brevis_, p. 190.
_Paraclevelandia simplex_, p. 190.
=Panesthia= sp.
Nematode: _Leidynemella panesthiae_, p. 198.
=Parahormetica bilobata=
Helminths:
_Agamospirura parahormeticae_, p. 205.
Undetermined gordian worm, p. 202.
=Parcoblatta lata=
Plant: _Sarracenia flava_, p. 154.
Protozoan: _Leptomonas_ sp., p. 167.
Nematode: _Protrellus aurifluus_, p. 199.
=Parcoblatta pensylvanica=
Protozoa:
_Gregarina blattarum_, p. 182.
_Gregarina illinensis_, p. 183.
_Gregarina parcoblattae_, p. 183.
_Gregarina thomasi_, p. 184.
_Leptomonas_ sp., p. 167.
_Nyctotherus ovalis_, p. 188.
Nematode: _Rictularia coloradensis_, p. 205.
Scorpion: _Centruroides vittatus_ (?), p. 212.
Insects:
_Camponotus pennsylvanicus_, p. 267.
_Hyptia dorsalis_, p. 239.
_Hyptia harpyoides_, p. 239.
_Hyptia reticulata_, p. 240.
_Hyptia thoracica_, p. 240.
_Podium carolina_, p. 265.
_Systellogaster ovivora_, p. 248.
Amphibian: _Rana_ (?) sp., p. 272.
=Parcoblatta uhleriana=
Fungus: _Herpomyces arietinus_, p. 134.
Protozoan: _Gregarina parcoblattae_, p. 183.
Nematode: _Protrellus aurifluus_, p. 199.
Mite: Undetermined, p. 220.
Insect: _Hyptia harpyoides_, p. 239.
=Parcoblatta virginica=
Fungus: _Herpomyces arietinus_, p. 134.
Protozoa:
_Gregarina ohioensis_, p. 183.
_Leptomonas_ sp., p. 167.
Nematode: _Rictularia coloradensis_, p. 205.
Insects:
_Hyptia harpyoides_, p. 239.
_Podium luctuosum_, p. 266.
=Parcoblatta= sp.
Fungus: _Herpomyces arietinus_, p. 134.
Plant: _Sarracenia flava_, p. 154.
Mite: _Pimeliaphilus podapolipophagus_, p. 219.
Insects:
_Coenosia basalis_, p. 229.
_Dolichurus greenei_, p. 261.
_Hyptia_ sp., p. 240.
_Megaselia_ sp., p. 227.
_Syntomosphyrum blattae_, p. 248.
_Systellogaster ovivora_, p. 248.
_Tetrastichus hagenowii_, p. 250.
=Periplaneta americana=
Viruses: Unspecified strain(s) of poliomyelitis virus
(host may have been _P. brunnea_), p. 103.
Bacteria:
_Achromobacter hyalinum_, p. 110.
_Aerobacter aerogenes_, p. 111.
_Aerobacter cloacae_, p. 112.
_Aerobacter_ sp., p. 112.
_Alcaligenes faecalis_, p. 111.
_Bacillus cereus_, p. 120.
_Bacillus megaterium_, p. 121.
_Bacillus subtilis_, p. 122.
_Bacterium alkaligenes_, p. 119.
_Clostridium_ sp., p. 122.
_Eberthella oedematiens_, p. 112.
_Escherichia coli_, p. 112.
_Escherichia freundii_, p. 113.
_Escherichia intermedium_, p. 113.
_Mycobacterium friedmannii_, p. 123.
_Mycobacterium lacticola_, p. 123.
_Mycobacterium leprae_, p. 123.
_Mycobacterium phlei_, p. 123.
_Mycobacterium piscium_, p. 124.
_Mycobacterium_ sp., p. 124.
_Nocardia_ sp. (?), p. 124.
_Paracolobactrum aerogenoides_, p. 113.
_Paracolobactrum coliforme_, p. 113.
_Paracolobactrum_ sp., p. 113.
_Proteus mirabilis_, p. 114.
_Proteus morganii_, p. 114.
_Proteus rettgeri_, p. 114.
_Proteus vulgaris_, p. 114.
_Proteus_ sp., p. 114.
_Pseudomonas aeruginosa_, p. 104.
_Pseudomonas fluorescens_, p. 105.
_Salmonella anatis_, p. 114.
_Salmonella morbificans_, p. 115.
_Salmonella schottmuelleri_, p. 115.
_Salmonella_ sp. (Type Bareilly), p. 115.
_Salmonella_ sp. (Type Bredeny), p. 115.
_Salmonella_ sp. (Type Kentucky), p. 115.
_Salmonella_ sp. (Type Meleagris), p. 116.
_Salmonella_ sp. (Type Newport), p. 116.
_Salmonella_ sp. (Type Oranienburg), p. 116.
_Salmonella_ sp. (Type Panama), p. 116.
_Salmonella_ sp. (Type Rubislaw), p. 116.
_Salmonella_ sp. (Type Tennessee), p. 116.
_Sarcina_ sp., p. 108.
_Serratia marcescens_, p. 118.
_Shigella alkalescens_, p. 118.
_Spirillum periplaneticum_, p. 105.
_Streptococcus faecalis_, p. 109.
_Streptomyces leidynematis_, pp. 196, 197.
_Tetragenous_ sp., p. 127.
_Veillonella parvula_, p. 109.
Fungi:
_Aspergillus flavus_, p. 130.
_Aspergillus niger_, p. 130.
_Aspergillus_ sp., p. 131.
_Cephalosporium_ sp., p. 131.
_Herpomyces chaetophilus_, p. 135.
_Herpomyces periplanetae_, p. 137.
_Metarrhizium anisopliae_, p. 131.
_Mucor guilliermondii_, p. 132.
_Mucor_ sp., p. 132.
_Penicillium_ sp., p. 131.
_Rhizopus_ sp., p. 133.
_Syncephalastrum_ sp., p. 133.
_Torula acidophila_, p. 132.
Protozoa:
_Balantidium blattarum_, p. 187.
_Balantidium ovatum_, p. 187.
_Balantidium_ sp., p. 187.
_Bodo_ sp. (host may have been _B. germanica_), p. 167.
_Coelosporidium periplanetae_, p. 185.
_Diplocystis schneideri_, p. 181.
_Diplocystis_ sp., p. 181.
_Dobellina_ sp. (host may have been _B. germanica_), p. 177.
_Endamoeba blattae_, p. 177.
_Endolimax blattae_, p. 180.
_Endolimax_ sp. (host may have been _B. germanica_), p. 180.
_Entamoeba coli_ (host may have been _B. germanica_), p. 178.
_Entamoeba histolytica_ (host may have been _B. germanica_), p. 179.
_Entamoeba_ sp., p. 179.
_Entamoeba thomsoni_, p. 179.
_Eutrichomastix_ sp. (host may have been _B. germanica_), p. 169.
_Gregarina blattarum_, p. 182.
_Gregarina légeri_, p. 183.
_Gregarina neo-brasiliensis_, p. 183.
_Hexamita periplanetae_, p. 171.
_Iodamoeba_ sp. (host may have been _B. germanica_) p. 180.
_Isotricha caulleryi_, p. 187.
_Lophomonas blattarum_, p. 172.
_Lophomonas striata_, p. 173.
_Monocercomonoides orthopterorum_, p. 169.
_Nyctotherus ovalis_, p. 188.
_Plistophora periplanetae_, p. 186.
_Protomagalhaesia serpentula_, p. 184.
_Tetratrichomastix blattidarum_, p. 170.
Helminths:
_Ancylostoma duodenale_, p. 209.
_Ascaris lumbricoides_ or _Ascaris suum_, p. 209.
_Binema mirzaia_, p. 193.
_Gongylonema neoplasticum_, p. 206.
_Gongylonema_ sp., p. 206.
_Gordius_ sp., p. 202.
_Hammerschmidtiella diesingi_, p. 195.
_Hymenolepis_ sp., p. 208.
_Leidynema appendiculata_, p. 197.
_Moniliformis dubius_, p. 203.
_Moniliformis moniliformis_, p. 204.
_Necator americanus_, p. 210.
_Protrellus künckeli_, p. 199.
_Schwenkiella icemi_, p. 200.
_Spirura gastrophila_, p. 207.
_Thelastoma pachyjuli_, p. 201.
_Trichuris trichiura_, p. 210.
Mites:
_Caloglyphus_ sp., p. 218.
_Pimeliaphilus podapolipophagus_, p. 219.
_Rhizoglyphus tarsalus_, p. 218.
_Tyrophagus noxius_, p. 218.
Undetermined, p. 220.
Spiders:
_Avicularia avicularia_, p. 214.
_Avicularia_ sp., p. 214.
Centipedes: Undetermined, p. 222.
Insects:
_Ampulex amoena_, p. 256.
_Ampulex compressa_, p. 259.
_Anastatus tenuipes_, p. 246.
_Calodexia_ (?) _venteris_, p. 228.
_Evania appendigaster_, p. 237.
_Melittobia chalybii_, p. 248.
_Prosevania punctata_, p. 241.
_Ripidius pectinicornis_, p. 232.
_Spiniger domesticus_, p. 227.
_Szepligetella sericea_, p. 242.
_Tetrastichus hagenowii_, p. 250.
_Tetrastichus periplanetae_, p. 253.
_Tetrastichus_ sp., p. 254.
_Trirhogma caerulea_, p. 262.
Amphibian:
_Hyla cinerea_, p. 270.
Reptiles:
_Ameiva exsul_, p. 275.
_Anolis cristatellus_, p. 273.
Birds:
Chicken, p. 278.
Sparrow, p. 282.
Mammals:
_Felis catus_, p. 289.
_Herpestea javanicus auropunctatus_, p. 289.
_Herpestes_ sp., p. 289.
_Homo sapiens_, p. 286.
_Rattus norvegicus_, p. 287.
_Rattus rattus_, p. 287.
=Periplaneta australasiae=
Bacteria:
_Mycobacterium leprae_, p. 123.
_Serratia marcescens_, p. 118.
Fungus: _Herpomyces periplanetae_, p. 137.
Plant: _Sarracenia minor_, p. 154.
Protozoa:
_Endamoeba blattae_, p. 177.
_Endolimax blattae_, p. 180.
Nematodes:
_Gongylonema neoplasticum_, p. 206.
_Hammerschmidtiella diesingi_, p. 195.
_Moniliformis dubius_, p. 203.
_Protrellus australasiae_, p. 199.
_Protrellus künckeli_, p. 200.
Centipedes: Undetermined, p. 222.
Insects:
_Ampulex amoena_, p. 256.
_Ampulex compressa_, p. 259.
_Evania appendigaster_, p. 237.
_Szepligetella sericea_, p. 242.
_Tetrastichus australasiae_, p. 249.
_Tetrastichus hagenowii_, p. 250.
_Tetrastichus_, sp., p. 254.
_Trirhogma caerulea_, p. 262.
Reptiles:
_Anolis cristatellus_, p. 273.
_Chrysemys picta_, p. 272.
Bird: _Gallus_ sp., p. 277.
Mammals:
_Herpestes_ sp., p. 289.
_Homo sapiens_, p. 287.
=Periplaneta brunnea=
Viruses: Unspecified strain(s) of poliomyelitis virus
(host may have been _P. americana_), p. 103.
Bacterium: _Serratia marcescens_, p. 118.
Fungus: _Herpomyces periplanetae_, p. 137.
Nematode: _Schwenkiella icemi_, p. 200.
Insect: _Tetrastichus hagenowii_, p. 251.
=Periplaneta fuliginosa=
Insect: _Ampulex amoena_, p. 256.
=Periplaneta= sp.
Bacterium: _Serratia marcescens_, p. 118.
Fungi:
_Herpomyces chaetophilus_, p. 135.
_Herpomyces periplanetae_, p. 137.
Protozoan: _Lophomonas blattarum_, p. 172.
Helminths: _Thelastoma riveroi_, p. 201.
_Moniliformis moniliformis_, p. 204.
Insects:
_Ampulex compressa_, p. 259.
_Evania subspinosa_ (presumptive record?), p. 239.
_Szepligetella sericea_, p. 242.
Amphibian: _Bufo marinus_, p. 270.
Reptiles:
_Anolis grahami_, p. 274.
_Anolis leachi_, p. 274.
Bird: _Vireo latimeri_, p. 281.
"=Phyllodromia=" sp.
Fungus: _Herpomyces phyllodromiae_, p. 137.
Insects:
_Dicarnosis alfierii_, p. 245.
_Dolichurus stantoni_, p. 262.
=Platyzosteria castanea=
Insect: _Riekella_ sp., p. 231.
=Platyzosteria novae seelandiae=
Protozoan: _Monocercomonoides melolonthae_, p. 169.
=Platyzosteria scabra=
Fungus: _Herpomyces appendiculatus_, p. 134.
=Platyzosteria= sp.
Insect: _Riekella nitidioides_, p. 231.
=Plectoptera= sp.
Bird: _Todus mexicanus_, p. 279.
=Polyphaga aegyptiaca=
Nematodes:
_Blatticola blattae_, p. 193.
_Galebia aegyptiaca_, p. 195.
_Hammerschmidtiella diesingi_, p. 195.
_Oxyuris_ (?) _heterogamiae_, p. 199.
=Polyphaga saussurei=
Helminths:
_Hymenolepis_ sp., p. 208.
Undetermined tapeworm ova, p. 208.
Insects: Undetermined reduviids, p. 227.
=Polyzosteria melanaria= (?) or =Platyzosteria analis=
Nematode: _Protrellus aureus_, p. 199.
=Pycnoscelus surinamensis=
Bacterium: _Serratia marcescens_, p. 117.
Fungus: _Mucor_ sp., p. 132.
Protozoa:
Undetermined ciliate, p. 190.
Undetermined flagellate, p. 176.
Undetermined gregarine, p. 184.
Nematodes:
_Oxyspirura mansoni_, p. 204.
_Severianoia severianoi_, p. 200.
Mites:
_Caloglyphus spinitarsus_, p. 217.
_Histiostoma feroniarum_, p. 217.
_Tyrophagus lintneri_, p. 218
Undetermined, p. 220.
Insects:
_Pheidole megacephala_, p. 268.
Undetermined ants, p. 350.
Amphibian: _Bufo marinus_, p. 270.
Reptile: _Anolis sagrei_, p. 274.
Birds:
_Acridotheres tristis_, p. 281.
Ducks, p. 277.
_Melanerpes portoricensis_, p. 280.
_Meleagris gallopavo_, p. 278.
_Passer domesticus_, p. 282.
_Phasianus_ sp., p. 277.
_Streptopelia chinensis_, p. 278.
Mammal: _Herpestes javanicus auropunctatus_, p. 289.
=Robshelfordia circumducta= or =Robshelfordia longiuscula=
Insects:
_Neorhipidius neoxenus_, p. 230.
_Rhipidioides helenae_, p. 230.
_Rhipidioides mollis_, p. 231.
=Steleopyga= (?) =sinensis=
Nematode: _Suifunema caudelli_, p. 200.
=Supella supellectilium=
Viruses: Unspecified strain(s) of poliomyelitis virus, p. 103.
Bacterium: _Serratia marcescens_, p. 117.
Fungus: _Herpomyces supellae_, p. 138.
Nematode: _Blattophila supellaima_, p. 194.
Insects:
_Anastatus tenuipes_, p. 246.
_Comperia merceti_, p. 244.
=Symploce flagellata= or =Symploce ruficollis=
Reptile: _Anolis cristatellus_, p. 273.
=Symploce parenthesis=
Hair worm: _Parachordodes raphaelis_, p. 202.
=Temnopteryx phalerata=
Protozoan: _Gregarina gibbsi_, p. 182.
=Undetermined Cockroaches=
Bacteria:
_Arthromitus intestinalis_, p. 124.
_Bacillus subtilis_, p. 122.
_Clostridium perfringens_, p. 122.
_Escherichia coli_, p. 112.
_Fusiformis lophomonadis_, p. 119.
_Micrococcus citreus_, p. 106.
_Micrococcus pyogenes_ var. _aureus_, p. 107.
_Mycobacterium leprae_, p. 123.
Paracolon bacilli, p. 113.
_Proteus morganii_, p. 114.
_Proteus vulgaris_, p. 114.
_Spirillum_ sp., p. 105.
_Spirochaeta periplanetae_, p. 125.
_Streptococcus_ sp., p. 110.
_Vibrio_ Types I and II Heiberg, p. 106.
Fungi:
_Amphoromorpha blattina_, p. 139.
_Amphoromorpha_ sp., p. 139.
_Blastocystis_ sp., p. 133.
_Cordyceps amazonica_, p. 134.
_Herpomyces anaplectae_, p. 134.
_Herpomyces chilensis_, p. 135.
_Herpomyces diplopterae_, p. 135.
_Herpomyces forficularis_, p. 135.
_Herpomyces grenadinus_, p. 136.
_Herpomyces macropus_, p. 136.
_Herpomyces paranensis_, p. 136.
_Herpomyces periplanetae_, p. 137.
_Herpomyces platyzosteriae_, p. 137.
_Herpomyces tricuspidatus_, p. 138.
_Herpomyces zanzibarinus_, p. 138.
_Metarrhizium anisopliae_, p. 131.
Plants:
_Nepenthes ampularia_, p. 154.
_Nepenthes gracilis_, p. 154.
_Nepenthes_ sp., p. 154.
Protozoa:
_Diplocystis_ sp., p. 181.
_Endamoeba blattae_, p. 177.
_Entamoeba histolytica_, p. 179.
_Giardia_ sp., p. 172.
_Gregarina blattarum_, p. 182.
_Hexamita periplanetae_, p. 171.
_Lophomonas blattarum_, p. 172.
_Lophomonas striata_, p. 173.
_Monas_ sp., p. 167.
_Monocercomonoides orthopterorum_, p. 169.
_Nyctotherus buissoni_, p. 188.
_Nyctotherus ovalis_, p. 188.
_Nyctotherus viannai_, p. 189.
_Oikomonas blattarum_, p. 166.
_Oikomonas_ sp., p. 166.
_Paramecium_ sp., p. 186.
_Trichomonas_ sp., p. 171.
Helminths:
_Cephalobellus magalhãesi_, p. 194.
_Chordodes morgani_, p. 201.
_Gordius_ sp., p. 202.
_Hammerschmidtiella diesingi_, p. 195.
_Leidynema appendiculata_, p. 197.
_Moniliformis moniliformis_, p. 204.
_Protrellus galebi_, p. 199.
_Schwenkiella icemi_, p. 200.
_Severianoia magna_, p. 200.
_Severianoia severianoi_, p. 200.
_Spirura gastrophila_, p. 207.
Undetermined gordian worms, p. 202.
Scorpion: _Heterometrus longimanus_, p. 213.
Spiders:
Ctenid, p. 214.
_Heteropoda venatoria_, p. 215.
_Latrodectus mactans_, p. 216.
Mite: _Pimeliaphilus podapolipophagus_, p. 219.
Centipedes:
_Allothereua maculata_ (circumstantial evidence), p. 223.
_Scolopendra morsitans_, p. 223.
_Scolopendra subspinipes_, p. 224.
_Scutigera coleoptrata_ (circumstantial evidence), p. 222.
Insects:
_Acanthinevania princeps_, p. 235.
_Agamerion metallica_, p. 243.
_Ampulex compressa_, p. 259.
_Ampulex ruficornis_, p. 259.
_Ampulex sibirica_, p. 260.
_Ampulex sonnerati_, p. 260.
_Anastatus blattidifurax_ p. 245.
_Anastatus tenuipes_, p. 246.
_Blatticida pulchra_, p. 243.
_Blatticidella ashmeadi_, p. 243.
_Calodexia_ spp., p. 228.
_Clerada apicicornis_, p. 226.
_Dicarnosis alfierii_, p. 245.
_Diestrammena apicalis_, p. 226.
_Diestrammena japonica_, p. 226.
_Dolichurus bicolor_, p. 260.
_Dolichurus corniculus_, p. 261.
_Dolichurus gilberti_, p. 261.
_Dolichurus ignitus_, p. 261.
_Dolichurus_ sp., p. 262.
_Dorylus nigricans sjöstedi_, p. 267.
_Dorylus_ sp., p. 267.
_Dorylus wilverthii_, p. 267.
_Eciton burchelli_, p. 268.
_Eupelmus_ sp., p. 247.
_Eutrichosomella blattophaga_, p. 245.
"_Formica omnivora_," p. 268.
_Iridomyrmex humilis_, p. 268.
_Neonephrites partiniger_, p. 230.
_Podium abdominale_, p. 265.
_Podium dubium_, p. 265.
_Podium rufipes_, p. 266.
_Pompilus bracatus_, p. 256.
_Pompilus_ sp., p. 256.
_Salius verticalis_, p. 256.
_Ripidius scutellaris_, p. 233.
_Solindenia picticornis_, p. 247.
_Stylogaster_ spp., p. 228.
_Stylogaster stylata_, p. 228.
_Syntomosphyrum blattae_, p. 248.
_Systellogaster ovivora_, p. 248.
_Tachysphex blatticidus_, p. 264.
_Tachysphex coriaceus_, p. 264.
_Tachysphex fanuiensis_, p. 264.
_Tetrastichus hagenowii_, p. 251.
_Tetrastichus periplanetae_, p. 253.
_Trirhogma_ sp., p. 264.
Undetermined tachinid, p. 229.
Fish:
_Chalceus macrolepidotus_, p. 269.
_Cyrtocharax magdalenae essequibensis_, p. 269.
_Potamotrygon humboldti_, p. 268.
_Rhamdia sebae_, p. 268.
Amphibians:
_Arthroleptis variabilis_, p. 270.
_Bufo ictericus_, p. 270.
_Bufo funereus_, p. 270.
_Bufo marinus_, p. 270.
_Hyla cinerea_, p. 270.
_Hyperolius picturatus_, p. 271.
_Leptodactylus albilabris_, p. 271.
_Leptodactylus pentadactylus_, p. 271.
_Leptopelis calcaratus_, p. 271.
_Leptopelis rufus_, p. 271.
_Megalixalus fornasinii_, p. 271.
_Rana catesbeiana_, p. 271.
_Rana mascareniensis_, p. 271.
Tree frogs, p. 351.
Reptiles:
_Ameiva exsul_, p. 275.
_Ameiva_ sp., p. 275.
_Anolis cristatellus_, p. 273.
_Anolis grahami_, p. 274.
_Anolis leachi_, p. 274.
_Anolis pulchellus_, p. 273.
_Anolis_ sp., p. 274.
_Anolis stratulus_, p. 274.
_Cnemidophorus_ sp., p. 275.
Geckos, p. 273.
_Gekko gecko_, p. 272.
_Hemidactylus frenatus_, p. 272.
_Leiolopisma laterale_, p. 274.
Lizards, p. 275.
Skinks, p. 274.
_Sphaerodactylus_ sp., p. 273.
_Thecadactylus_ sp., p. 273.
_Tropidophorus grayi_, p. 274.
Birds:
_Acridotheres tristis_, p. 281.
_Agelaius xanthomus_, p. 282.
_Bambusicola thoracica_, p. 277.
Chickens, p. 278.
_Coturnix coturnix japonica_, p. 277.
_Dendrocopus mahrattensis_, p. 279.
Ducks, p. 277.
_Gymnasio nudipes_, p. 279.
_Gymnopithys leucaspis_, p. 280.
_Holoquiscalus brachypterus_, p. 282.
_Icterus portoricensis_, p. 280.
Owl, p. 351.
Partridge, p. 278.
_Phasianus_ sp., p. 277.
Sparrow, p. 282.
_Tiaris bicolor omissa_, p. 282.
_Tockus birostris_, p. 279.
_Troglodytes aedon_, p. 281.
_Troglodytes audax_, p. 281.
Mammals:
_Bassariscus astutus_, p. 288.
_Callithrix jacchus_, p. 285.
_Erinaceus_ sp., p. 283.
_Felis catus_, p. 289.
_Felis paradalis mearnsi_, p. 290.
_Herpestes javanicus auropunctatus_, p. 289
_Herpestes_ sp., p. 289.
_Homo sapiens_, p. 287.
_Loris tardigradus_, p. 284.
_Molossus_ sp., p. 283.
_Monodelphis_ sp., p. 283.
_Nasua nasua_, p. 288.
_Nasua_ sp., p. 289.
_Rattus_ sp., p. 287.
XV. COCKROACHES AS COMMENSALS
These particular associations may well have been accidental and due
to a predilection for the same type of nesting site. But this fact
in no way detracts from the interest of such records. Chance must
play a very considerable part in first bringing symbiotic or
commensal partners together. Once such a partnership between
species has been firmly established, it is on the whole, fairly
obvious, ... On the other hand, in the early stages before the
relationship has become fixed as a specific habit, individual cases
are generally dismissed as coincidences. It is, however, unwise to
disregard such isolated observations or dismiss them lightly.
ROTHSCHILD AND CLAY (1957)
The following social insects have been found harboring cockroaches in a
state of commensalism in which the cockroaches presumably benefit by
acquiring food from their hosts. Benefits accruing to the hosts are not
apparent. Unfortunately, biological details are not always sufficient to
substantiate the suspected association. However, it seems significant
that the cockroach commensals of the insects listed below have been
found only in association with their hosts and, so far as we know, have
never been found apart from them. Chopard (1938) has pointed out that
the myrmecophilous cockroaches are all small, being only a very few
millimeters long; they are apterous or subapterous; their eyes are
reduced; and they are all of American origin.
HOSTS OF COMMENSAL COCKROACHES
Order ISOPTERA
Family RHINOTERMITIDAE
=Coptotermes ceylonicus= Holmgren
_Commensal._--_Sphecophila ravana_, Ceylon (Fernando, 1957): Six
females, 50 males, and nymphs of both sexes were found among decaying
timber in the ground in association with a colony of this termite. The
antennae of most specimens were mutilated unsymmetrically.
Family TERMITIDAE
=Macrotermes barneyi= Light
_Commensal._--_Nocticola sinensis_, Kowloon (Silvestri, 1947): Among
specimens of termites collected from a nest.
=Macrotermes bellicosus= (Smeathman)
or
=Macrotermes natalensis= (Haviland)
_Synonymy._--_Termes bellicosus_ [Snyder, 1949].
_Commensal._--_Sphecophila termitium_, Kibonoto, East Africa (Shelford,
1910): Two males were collected in a termite mound.
=Macrotermes malaccensis= (Haviland)
_Synonymy._--_Termes malaccensis_ Haviland [Snyder, 1949].
_Commensal._--_Nocticola termitophila_, Tonkin (Silvestri, 1946): The
cockroach was found in the termite nest.
=Odontotermes= sp.
_Commensals._--_Nocticola sinensis_, Kowloon (Silvestri, 1946): In
termite nest.
_Nocticola termitophila_, Penang (Silvestri, 1946): In termite nest.
=Termes= sp.
_Commensal._--_Nocticola sinensis_, Repulse Bay, Australia(?)
(Silvestri, 1946): In a termite gallery.
=Termites=
_Commensal._--_Ergaula capensis_ [= _Dyscologamia wollastoni_] French
Equatorial Africa, Brazzaville (Rehn, 1926; Chopard, 1949).
Order HYMENOPTERA
Family FORMICIDAE
Subfamily FORMICINAE
=Camponotus femoratus= (Fabricius)
_Commensal._--_Phorticolea boliviae_, Bolivia, Cachuela Esperanza
(Caudell, 1923): Three males collected in the joint nests of _C.
femoratus_ and _Crematogaster limata_.
=Camponotus maculatus= (Fabricius)?
_Note._--Dr. W. L. Brown (personal communication, 1957) states that this
ant is an Old World species only. So presumably Mann's record pertains
to a different species.
_Commensal._--_Myrmecoblatta rehni_, Mexico (Mann, 1914): "They were
very abundant, several occurring in almost every nest, where they are no
doubt very efficient scavengers."
=Camponotus rufipes= (Fabricius)
_Commensals._--_Atticola mortoni_, _Nothoblatta wasmanni_, and
_Phorticolea testacea_, Brazil, San Leopoldo (Bolívar, 1905): Found in
the formicaries of _C. rufipes_.
=Formica rufibarbis= Fabricius
and
=Formica subcyanea= Wheeler
_Commensal._--_Myrmecoblatta rehni_, Mexico (Mann, 1914): "They were
very abundant, several occurring in almost every nest."
Subfamily MYRMICINAE
=Acromyrmex lobicornis= Emery
_Commensal._--_Attaphila bergi_, or possibly a variety of this species,
Huasán, Argentina? (Bruch, 1916).
=Acromyrmex lundi= (Guérin)
_Synonymy._--_Atta lundi_ [Brown, personal communication, 1957].
_Commensal._--_Attaphila bergi_, Argentina and Uruguay (Bolívar, 1901):
The cockroach was found in the nests of the ants sitting on the back,
neck, or head of sexual individuals. It remains attached to the ant
during swarming. The antennae seem always to be mutilated. Bruch (1916)
stated that in La Plata _A. bergi_ is encountered by hundreds in every
nest of _A. lundi_.
=Acromyrmex niger= (F. Smith)
_Synonymy._--_Atta nigra_ Schupp [Brown, p.c., 1957].
_Commensal._--_Attaphila schuppi_, Brazil, Porto Alegre (Bolívar, 1905):
Found outside the nest of the ant and mixed in the columns of ants on
the march.
=Acromyrmex octospinosus= (Reich)
_Synonymy._--_Atta octospinosa_ [Brown, p.c., 1957].
_Commensals._--_Attaphila fungicola_, Panama (Wheeler, 1928): Taken in
the fungus gardens of the ant.
_Attaphila aptera_, Esperanza, Dibulla, Colombia (Bolívar, 1905).
=Acromyrmex silvestrii= Emery
_Commensal._--_Attaphila bergi_, or possibly a variety of this species,
San Luis Province, Argentina (Bruch, 1916): According to Bruch, the
behavior of this species of _Attaphila_ is identical with the one
encountered in Huasán in the nests of _Acromyrmex lobicornis_ Emery; it
differed from _A. bergi_ in size and color.
=Atta cephalotes= (Linnaeus)
_Commensal._--_Attaphila fungicola_, British Guiana (Wheeler, 1928):
Taken in the fungus gardens of the ant.
_Attaphila_ sp., British Guiana (Beebe, 1921): 7 of 12 queens in one
nest had cockroaches hanging on them.
=Atta sexdens= (Linnaeus)
_Commensal._--_Attaphila sexdentis_, Brazil, San Leopoldo (Bolívar,
1905): Found in nests of the ant.
=Atta texana= (Buckley)
_Synonymy._--_Atta fervens_ Say [Wheeler, 1910].
_Commensal._--_Attaphila fungicola_, U.S.A., Texas (Wheeler, 1900,
1910): The cockroach does not feed on the fungus in the ants' nest, as
Wheeler (1900) first supposed, but mounts the back of the soldiers and
licks their surfaces. It is tolerated by the ants with no signs of
hostility. The antennae of the cockroach are clipped short. Although
Wheeler (1910) stated that this is probably accidental or unintentional,
it is peculiar that Bolívar (1905) noticed the same invariable
mutilation of the antennae of _Attaphila bergi_. Wheeler (1900) had
originally suggested that the antennae were probably clipped off by the
ants which are continuously trimming the fungus hyphae. Louisiana
(Moser, personal communication, 1959): Numerous specimens were
encountered in some nests of _A. texana_. This cockroach is the most
closely associated inquiline in the nest and maintains very intimate
terms with the ants. It is found living in the fungus cavities and
tunnels.
=Crematogaster limata parabiotica= Forel
_Commensal._--_Phorticolea boliviae_, Bolivia, Cachuela Esperanza
(Caudell, 1923): Collected in joint nests of _C. limata_ and _Componotus
femoratus_.
=Solenopsis geminata= (Fabricius)
_Commensal._--_Myrmecoblatta wheeleri_, Guatemala (Hebard, 1917a):
Collected from a colony of this ant under a stone on the shores of Lake
Atitlan, altitude 11,719 feet.
=Unknown host=
_Cockroach._--_Attaphila flava_, British Honduras (Gurney, 1937):
Because the known hosts of the other five species of _Attaphila_ are
ants, we presume that this species also lives in the nest of some
myrmecine ant.
Subfamily PONERINAE
=Odontomachus affinis= (Guérin)
_Commensal._--_Myrmeblattina longipes_, Brazil, Rio de Janeiro (Chopard,
1924, 1924a; Hancock, 1926): Originally described as _Phileciton
longipes_ by Chopard (1924) from the nest of an ant mistakenly
identified as _Eciton_ sp.
Family VESPIDAE
=Polybia pygmaea= Fabricius
_Commensal._--_Sphecophila polybiarum_, French Guiana (Shelford, 1906a):
Shelford stated that it was probable that the cockroaches living on the
floor of the paper nest fed on small fragments of insects and spiders
that were dropped by the wasp larvae feeding in the cells above.
Family MEGACHILIDAE
=Melipona nigra= Lepeletier
_Commensal._--_Oulopteryx meliponarum_, Brazil (Hebard, 1921): According
to Hebard, this cockroach is the first one to be known to inhabit the
nests of bees. Nothing is known of the relationship between the
cockroach and the bees. [See comment by Sonan (1924) on page 318.]
CHECKLIST OF COMMENSAL COCKROACHES WITH THEIR HOSTS
The cockroaches are arranged alphabetically by genus and species. The
page references are to citations in the classified section above, where
details and/or sources of the records are given.
=Attaphila aptera=
Ant: _Acromyrmex octospinosus_, p. 313.
=Attaphila bergi=
Ants: _Acromyrmex lobicornis_, p. 312.
_Acromyrmex lundi_, p. 312.
_Acromyrmex silvestrii_, p. 313.
=Attaphila flava=
Host unknown, presumably an ant, p. 314.
=Attaphila fungicola=
Ants: _Acromyrmex octospinosus_, p. 313.
_Atta cephalotes_, p. 313.
_Atta texana_, p. 313.
=Attaphila schuppi=
Ant: _Acromyrmex niger_, p. 312.
=Attaphila sexdentis=
Ant: _Atta sexdens_, p. 313.
=Attaphila= sp.
Ant: _Atta cephalotes_, p. 313.
=Atticola mortoni=
Ant: _Camponotus rufipes_, p. 312.
=Ergaula capensis=
Termites, p. 311.
=Myrmeblattina longipes=
Ant: _Odontomachus affinis_, p. 314.
=Myrmecoblatta rehni=
Ants: _Camponotus maculatus_(?), p. 312.
_Formica rufibarbis_, p. 312.
_Formica subcyanea_, p. 312.
=Myrmecoblatta wheeleri=
Ant: _Solenopsis geminata_, p. 314.
=Nocticola sinensis=
Termites: _Macrotermes barneyi_, p. 311.
_Odontotermes_ sp., p. 311.
=Nocticola termitophila=
Termites: _Macrotermes malaccensis_, p. 311.
_Odontotermes_ sp., p. 311.
_Termes_ sp., p. 311.
=Nothoblatta wasmanni=
Ant: _Camponotus rufipes_, p. 312.
=Oulopteryx meliponarum=
Bee: _Melipona nigra_, p. 314.
=Phorticolea boliviae=
Ants: _Camponotus femoratus_, p. 311.
_Crematogaster limata parabiotica_, p. 312.
=Phorticolea testacea=
Ant: _Camponotus rufipes_, p. 312.
=Sphecophila polybiarum=
Wasp: _Polybia pygmaea_, p. 314.
=Sphecophila ravana=
Termite: _Coptotermes ceylonicus_, p. 310.
=Sphecophila termitium=
Termite: _Macrotermes bellicosus_ or _Macrotermes natalensis_, p. 311.
OBSCURE ASSOCIATIONS
Cockroaches that are sometimes found in the nests of, or in association
with, other animals are not necessarily commensals. This is particularly
true of cockroaches that normally are found unassociated with other
animals or that merely occupy the same habitat with the other animals
because of similar microclimatic requirements (see Chopard, 1924c).
McCook (1877) excavated in February a nest of _Formica rufa_ in
Pennsylvania. A hundred or more lively cockroaches occupied a part of
the nest that contained few ants. Near the cockroaches McCook also found
a colony of _Termes flavipes_. _Ischnoptera deropeltiformis_ has been
found in the company of ants, but it is probably not myrmecophilous
(Donisthorpe, 1900). Mann (1911) found an "_Ischnoptera_" sp. (probably
a species of _Parcoblatta_) abundant in the nests of, and tolerated by,
_Camponotus maccooki_ Forel in California. Dead and mutilated specimens
of this cockroach were common in the nests of "Formicas."
"_Ischnoptera_" sp. was also common in the nests of _Veromessor andrei_
(Mayr) [= _Stenamma andrei_]. Hebard (1917) reported that W. M. Wheeler
collected _Eremoblatta subdiaphana_ in Arizona as an ant guest. Rehn
(1906a; Rehn and Hebard, 1927) reported that _Pholadoblatta inusitata_
had also been taken by Wheeler from the galleries of a jumping ant,
_Odontomachus clarus_ Roger [= _O. haematodes insularis_ Guérin var.
_pallens_ Wheeler; Brown (personal communication, 1958)], on Andros
Island, Bahamas; Rehn and Hebard (1927) stated that "This genus and
species is the only blattid, which is presumably a myrmecophile, known
from the West Indies." Rehn (1932a) reported _Dendroblatta sobrina_ as
taken in an ant nest in a tree in the Amazon Basin. _Tivia australica_
was taken in an ant nest in Australia (Princis, 1954). The male of
_Compsodes schwarzi_ was taken in an ant nest in the Santa Rita
Mountains of Arizona (Ball et al., 1942). A male and female of
_Stilpnoblatta minuta_ were taken in a migrating column of the ant
_Myrmicaria natalensis_ Sm. subsp. _eumenoides_ Gerst. in Nyasaland
(Princis, 1949). Princis cautioned that it is premature to derive any
inference from this, possibly accidental, association. Four females of
_Parcoblatta desertae_ were taken about a nest of an ant, _Ischnomyrmex_
sp. (Hebard, 1943a). A nymph of _Parcoblatta virginica_ was found in a
nest of _Formica_ sp. (Hauke, 1949).
_Chorisoneura texensis_ has been found in nests of webworm in Florida
(Rehn and Hebard, 1916). Karny (1924) in Malaya found an oötheca of
_Aristiger histrio_ (sp.?) between leaves (_Costus_ sp.) that had been
stuck together by a thysanopteron, _Anaphothrips_ sp. He pointed out
that the oötheca would not adhere to leaves that were not stuck together
but would fall to the ground.
Seín (_in_ Rehn and Hebard, 1927) in Puerto Rico found _Aglaopteryx
facies_ in abandoned cocoons of _Megalopyge krugii_ (Dewitz) and in
leaves webbed together by caterpillars and in abandoned spiders nests.
Wolcott (1950; and _in_ Rehn and Hebard, 1927) also found _A. facies_ in
the empty cocoons of _M. krugii_ and in the larval tents of _Tetralopha
scabridella_ Ragonot on _Inga vera_ (coffee shade tree); and "Where
there are no butterfly-nests, it lives in abandoned spider-nests on the
leaves of other forest trees." Cotton (_in_ Wolcott, 1950) found the
type of _Aglaopteryx absimilis_ also living in the abandoned cocoon of
_M. krugii_ on bucare trees in Puerto Rico. Wolcott (1950) reported that
_Plectoptera dorsalis_, _Plectoptera infulata_, and _Plectoptera
rhabdota_ have been found living in trees between leaves or in
"butterfly-nests" of _Tetralopha scabridella_ in leaves of _Inga vera_,
or nests of _Pilocrocis secernalis_ (Möschler) in the leaves of _Petitia
domingensis_ in the mountains of Puerto Rico. Seín (_in_ Rehn and
Hebard, 1927) had collected _P. rhabdota_ in the nest of larvae of _T.
scabridella_.
Wolcott (1950) reported that _Nyctibora lutzi_ had been found in a large
rotten stump associating with "'comején' termites [_Nasutitermes
costalis_ (Holmgren)], yellow wood-ants and rhinoceros beetle grubs."
Rehn and Hebard (1927) found _Simblerastes jamaicanus_ in numbers in the
debris of an abandoned termites' nest in Jamaica: "To what extent the
species is dependent upon the protection of the termite or other
structures remains to be determined."
In Virginia _Cryptocercus punctulatus_ has been found living in the same
galleries with _Reticulitermes_ sp., and on the Pacific Coast it has
been found occupying the same log with _Termopsis_ sp. (Cleveland et
al., 1934).
Shelford (1909) found one male and one female of _Balta platysoma_ in a
nest of a spider of the genus _Phryganoporus_ and assumed a symbiotic
association. Chopard (1924) recorded _Mareta acutiventris_ from empty
nests of spiders on Barkuda Island, India; nothing is known of the
relationships, if any, between these cockroaches and spiders.
Chopard (1924c) found _Margattea_ sp. in the nest of the ant _Acropyga
acutiventris_ Roger; he also found _Margattea_ sp., _Periplaneta_ sp.,
_Polyphaga indica_, and _Temnopteryx obliquetruncata_ in deserted
termite mounds in India. However, he believed that none of these species
were more than accidental associates of the host insects; he considered
them hygrophilous cockroaches which had found a retreat in the nests.
McClure (1936) obtained a large nest of _Vespula maculata_ (Linnaeus)
[= _Vespa maculata_] in March in Illinois. In it were living 65 nymphs of
_Parcoblatta pensylvanica_, 3 spiders (_Philodromus pernix_ Blackwall),
2 immature spiders (_Drassus_ sp.), and 6 mites. Balduf (1936) observed
four individuals of _Parcoblatta pensylvanica_ in a nest of _Vespula
maculata_; he suggested that they probably fed on dead bodies and
organic wastes of the wasps. However, Rau (1940) has observed this
cockroach devour a _Polistes_ larva in its cell. Although we do not
imply that a commensal relationship exists between _Parcoblatta_ and the
wasp, it is well to recall a statement by Rothschild and Clay (1957): "A
commensal relationship is potentially even more dangerous than a merely
social tie, for by nature it is more intimate. The closer the
association, the more easily is the balance upset. One partner can then
suddenly take a mean advantage of the other."
Cockroach nymphs may enter bees' nests where, according to Imamura (_in_
Sonan, 1924), they do not feed on honey or pollen but presumably feed on
excreta of bees or anything scattered by bees in their nest; the bees
are not disturbed by the cockroaches.
Cockroaches that have been found in the burrows of vertebrates are
listed on pages 23-25.
Paulian (1950) found immature cockroaches in the nests of birds
(Ploceinae) in Madagascar and Ivory Coast. All nests of _Fondia_ sp.
examined in Madagascar contained many cockroaches, and Paulian believed
that the blattid was a species peculiar to the nests of birds. Three
nests of _Ploceus_ sp. in Ivory Coast yielded one or two cockroaches
each in association with more numerous mites, Psocoptera, Heteroptera,
beetles, and lepidopterous larvae (Delamare, Deboutteville and Paulian,
1952). These last cited workers also found four cockroaches in a nest of
_Estrildine_ sp., and two in a nest of an undetermined bird, all in
association with other arthropods. Moulton (1912) observed large numbers
of _Symploce cavernicola_ and _Periplaneta australasiae_ swarming in
soft bird guano on the floor of caves in Borneo. Abdulali (1942) found
in India many _Periplaneta americana_ in caves containing the
edible-nest swift; there was no indication of association of the
cockroaches with the birds. Danforth (_in_ Wolcott, 1950) reported
finding large numbers of _Aglaopteryx facies_ "in the nests of the grey
kingbird, in the region of the Cartagena Lagoon [Puerto Rico], 'living
among the twigs.'" In Trinidad, Kevan found a male of _Blaberus
discoidalis_ in a bird's nest (Princis and Kevan, 1955).
Davis (_in_ Rehn and Hebard, 1914a) stated that "At Punta Gorda
[Florida] there was a vacant house at the end of the town frequented at
night by a Nanny and Billy goat, and on warm evenings many _Periplaneta
australasiae_ would run about on the piazza floor and on the sides of
the house. They were seen feeding on the excrement of the goats and were
no doubt to a great degree dependent upon them." This is another example
of a coprophagous insect that has taken advantage of a particular
situation favorable to its survival. Similar associations exist in which
many of the domiciliary cockroaches feed on the feces of man and
domestic and other animals (Roth and Willis, 1957a).
XVI. COCKROACHES AS PREDATORS
INTERSPECIES PREDATION
Tepper (1893) made the broad statement that the majority of Australian
and Polynesian cockroaches appear to be wholly carnivorous, eating other
insects, eggs, and larvae. He stated that, because of their voracity and
cannibalistic tendencies, the carnivorous species lead more or less
solitary lives so that one rarely meets several in close proximity; they
are never very numerous at any time because the stronger devour the
weaker in the absence of other prey. Tepper stated that Australian
species of _Ischnoptera_ hunt for their prey among the foliage of
shrubs, and that Australian species of _Cutilia_ [= _Drymaplaneta_,
Hebard (1943)] run about actively on the surface, or ascend shrubs and
trees in quest of living insects and therefore are highly beneficial.
Tepper (1894) also stated that _Geoscapheus robustus_ ate earthworms,
grubs, and caterpillars. Froggatt (1906) and Marlatt (1915) attributed
to Tepper the statement that cockroaches, like _Epilampra notabilis_,
which are found out-of-doors in Australia, are carnivorous and feed on
caterpillars and other soft-bodied insects; but Froggatt (1907) believed
that this alleged behavior needed confirmation.
A number of observations have been recorded which indicate that
sometimes cockroaches may be predatory. According to Ealand (1915),
nymphs of the cockroach _Pseudomops cincta_ fed on the Argentine ant
_Iridomyrmex humilis_. In the laboratory, _Eurycotis floridana_ has been
observed to catch and devour the wasp _Anastatus floridanus_ which
parasitizes the eggs of _Eurycotis_ (Roth and Willis, 1954a).
_Parcoblatta pensylvanica_ was observed devouring a larva of _Polistes_
sp. in its cell in a deserted wasps' nest (Rau, 1940). Brigham (1866)
saw a cockroach kill and eat a centipede four or five inches long.
Annandale (1910) described the destruction in Calcutta of termites by
_Periplaneta americana_. During a heavy rain storm many termites flew
into the dining room and were set upon by the cockroaches which seized
them with their mandibles and began to gnaw their abdomens. If
disturbed, the cockroaches carried the termites away in their mandibles
without using their legs to seize, hold, or carry the prey. Sometimes
only the abdomen, but other times the whole body with the exception of
the wings, was devoured. Perhaps this observation led Allyn (Anonymous,
1937) to theorize that, first, cockroaches could eradicate termites from
houses, and then the blattids in turn could be eliminated. Falls (1938)
has pointed out the unfeasibility of this idea. _Blattella vaga_ has
shown some tendency to eat plant lice (Flock, 1941a). Certain small
cockroaches found beneath cane leaf-sheaths, in the Philippine Islands,
preyed in part upon leafhoppers (Uichanco, _in_ Williams et al., 1931).
Takahashi (1924) stated that the American cockroach will eat the eggs of
the hemipteron _Cantao ocellatus_ (Thunberg). Cunliffe (1952) observed
mite-infested cockroaches (_Blatta orientalis_, _Blattella germanica_,
and/or _Periplaneta americana_) dislodge and eat the mite _Pimeliaphilus
podapolipophagus_. Sonan (1924) reported that cockroaches (_P.
americana_ and _P. australasiae_) devoured the egg clusters and first
instar larvae of _Prodenia litula_ and the first instar larvae of
_Attacus atlas_ which were being reared in the laboratory. Lederer
(1952) stated that _Periplaneta americana_ ate reptile eggs in the
aquarium at Frankfurt am Main. Pettit (1940) stated that cockroaches
"are said to have destroyed a large colony of dermestids used to
skeletonize carcasses at the University of Kansas."
DeFraula (1780) believed that his silent "gryllon" [obviously _Blatta
orientalis_ from his drawings; see Willemet (1784)] was the enemy of the
chirping species of cricket, because after the cockroach became
established in his home he no longer heard crickets chirping. Gilbert
White (1905 ed.), writing in England in the late 18th century, stated
that "Poda says that these [_Blatta orientalis_] and house crickets will
not associate together; but he is mistaken in that assertion"; however,
in August 1792 White noted that "Since the _blattae_ have been so much
kept under, the crickets have greatly increased in number." For several
years Jolivet (1950) had observed changes in a mixed population of
_Blatta orientalis_ and _Acheta domesticus_ in an old kitchen in France.
He suggested that the cyclical fluctuations in the relative abundance of
the cockroaches and crickets might be caused by reciprocal predatism
with one species more susceptible than the other at certain stages.
Mallis (1954) has stated that crickets prey on other insects as well as
on one another. Lhéritier (1951) had also observed crickets becoming
rare in bakeries in France, having been superseded everywhere by _B.
orientalis_; however, he doubted that Jolivet's hypothesis was the
correct explanation and suggested that the higher optimum temperature
requirements of crickets might be the regulating factor. Lederer (1952)
stated that the number of crickets decreased in the aquarium buildings
at Frankfurt am Main as the population of American cockroaches
increased.
_Platyzosteria novae seelandiae_ was found under the bark of trees in
New Zealand devouring bugs (Walker _in_ Shelford, 1909b).
For years it has been believed that cockroaches feed on bedbugs (_Cimex
lectularius_ L.) and this statement has been repeated in many reference
works and articles. Ealand (1915) stated that cockroaches devour bedbugs
with avidity. Even today similar statements are to be found in the
literature. "In the old sailing ship days, they [cockroaches] were often
welcomed by crews because of the belief that they would eradicate a
population of bedbugs. This belief was based on scientific fact, as
cockroaches are known as predators of bedbugs" (Monro, 1951).
Cockroaches will often "help rid a house of bedbugs by devouring all the
little parasites they can capture" (Gaul, 1953). The basis for this
belief may have originated with a statement by Webster (1834) who wrote
that bedbugs disappeared aboard "H.M. Sloop Chanticleer" when
cockroaches made their appearance. Newman (1855) reported the
observations of a friend who claimed to have seen a cockroach seize a
bedbug in an infested boardinghouse in London. In 1920 Purdy
reintroduced cockroaches into a house from which they had been
exterminated, in order to control the bedbugs which had become
established. According to a popular account by Lillingston (1934)
African natives are said to ask sailors for a cockroach or two to be
used to hunt bedbugs.
In Siberia, Burr (1926, 1939) found _Blattella germanica_ and bedbugs
inhabiting the same room. Mellanby (1939) studied the populations of an
animal house in which bedbugs and cockroaches occurred in large numbers;
the bugs apparently were not attacked and their numbers increased
greatly over a period of a few weeks (Johnson and Mellanby, 1939). Wille
(1920) placed starved _B. germanica_ with bedbugs for 20 days, but the
cockroaches failed to attack the bugs. In India, captive adults and
nymphs of two species of house cockroaches would not touch living
bedbugs or their eggs (Cornwall, 1916). In laboratory experiments Gulati
(1930) found that _Periplaneta americana_ ate young bedbugs which had
soft, blood-filled abdomens; adult bedbugs with harder exoskeletons
sometimes were rejected. The maximum number of bedbugs eaten by a
cockroach was 3 out of 12 during a period of 48 hours. Johnson and
Mellanby (1939), also in laboratory experiments, were unable to show
that bedbugs can be controlled by _Blatta orientalis_ or that bedbugs
are eaten to any extent by them. The existing evidence indicates that
there is little basis for the often repeated statement that cockroaches
destroy bedbugs in nature. As Lorando (1929) pointed out, assassin bugs,
cockroaches, and red ants can hardly be considered as practical factors
in bedbug control, though he did recommend the use of spiders.
According to Martini (1952), cockroaches prey on mosquitoes and sand
flies but we have been unable to find any original sources for these
statements; the only reference we have found in which cockroaches and
_Phlebotomus_ are mentioned together is a paper by Whittingham and Rook
(1923); they fed ground-up cockroaches to larvae of _Phlebotomus
papatasii_. Wharton (1951) reported that cockroaches and other predators
attacked mosquitoes knocked down by insecticides and affected the number
recovered.
Cockroaches will on occasion attack and bite animals other than insects.
In an earlier paper (1957a) we discussed about 20 reports of cockroaches
biting man. The injury is usually confined to abrasion of the callused
portions of hands and feet but may result in small wounds in the softer
skin of the face and neck. We failed to include the following reference
in the above-mentioned paper. Sonan (1924) had his toes and breast
nibbled by cockroaches on Hiyakejima Island during sleep. He had
previously learned from a policeman that _Periplaneta americana_ and _P.
australasiae_ nibbled people on that island, but he had hardly believed
it before he experienced the biting himself.
INTRASPECIES PREDATION
Those who have reared cockroaches in the laboratory have undoubtedly
seen cannibalism occur in the cultures. Cannibalism has been observed
among the common domiciliary species of cockroaches as well as
laboratory colonies of _Leucophaea maderae_ (Scharrer, 1953), and
_Blaberus craniifer_[12] (Saupe, 1928). Edmunds (1957)
reported that cannibalism was common in a laboratory colony of
_Periplaneta brunnea_ and that egg capsules deposited by a female were
often eaten by the other cockroaches.
_Periplaneta americana_ occasionally ate other cockroaches and their
oöthecae and also attacked members of their own species (Lederer, 1952).
Griffiths and Tauber (1942) recorded the killing of male American
cockroaches by females of the species: "One female was especially
vicious and attacked each new male as he was introduced into the
container. Most of such males had molted less than 2 days previously.
Older males were more capable of defending themselves against attacks of
these cannibalistic females." Even though adequate food may be present,
females of _Periplaneta americana_ may eat their own eggs (Klein, 1933).
Some females may regularly eat their oöthecae as soon as they are
dropped (Griffiths and Tauber, 1942). To be completely eaten an oötheca
generally must be attacked before it has hardened. If a hole is eaten in
one side of the capsule, the cockroach may devour the eggs and leave a
portion of the oötheca. Frequently only the keel or a part of the keel
is eaten and when this occurs the eggs fail to hatch and usually do not
complete development because of the rapid loss of water (Roth and
Willis, 1955). When adults of _P. americana_ and _P. australasiae_ were
deprived of food, both males and females ate newly deposited eggs and,
finally, the females ate the males (Sonan, 1924).
_Parcoblatta virginica_ in laboratory cultures also may eat part of its
oöthecae; in this species only the soft end of the recently deposited
oötheca was eaten (Roth, unpublished data, 1957).
Cros (1942) observed oöthecae-bearing females of _Blatta orientalis_
attack and kill males of the same species which were attempting to mate;
these males were then eaten by the females. Cros also observed injured
and recently molted nymphs of _B. orientalis_ to be eaten by others of
the same species.
Pettit (1940) noted that cannibalism in his culture of _Blattella
germanica_ occurred only when the insects were molting. Adult insects
attacked the molting cockroaches more often than did the nymphs.
However, nymphs after the fourth instar occasionally set upon other
molting nymphs. First-to third-instar nymphs rarely victimized their
mates. The victims were all older than third instar; the later stadia
were progressively more subject to attack, and molting adults suffered
the greatest mortality. No direct correlation was noted between
population density and cannibalism.
German cockroaches may attack newly molted nymphs of their own kind and
cause them to deflate (Gould and Deay, 1938). Lhéritier (1951) has
observed the hatching nymphs of _B. germanica_ being devoured by their
congeners even before they have left the oötheca.
_Nauphoeta cinerea_ in laboratory cultures will eat newly hatched young
of the same species (Roth and Willis, 1954; Willis et al., 1958). In
Hawaii, in nature, _N. cinerea_ may kill and eat the cypress cockroach,
_Diploptera punctata_ (Illingworth, 1942; Fullaway and Krauss, 1945).
Bunting (1956) stated that species of _Neoblattella_ are omnivorous with
carnivorous and cannibalistic tendencies. An adult female _Panchlora_
sp. was killed and eaten by _Neoblattella_ sp. in captivity. A male,
provisionally identified as _N. celeripes_, was killed and partly eaten
by two other males of the same species.
The factors influencing the extent of cannibalism among cockroaches are
not completely known. According to Wille (1920) hunger was not the cause
of cannibalism in _Blattella germanica_. Wille claimed that the tendency
toward cannibalism increased at high temperatures and decreased at low
temperatures. Pettit (1940) also noted this effect. Gould and Deay
(1938) stated that under crowded laboratory conditions, when there was a
scarcity of food, cannibalism among _Periplaneta americana_ was common.
The injured cockroaches and those unable to molt were often eaten. Adair
(1923) made similar observations. Undoubtedly, conditions of crowding,
availability of food, temperature and other factors all influence
cannibalism, but practically no experimental work has been done on this
subject.
It is interesting, in comparison with the above positive examples of
cannibalism, that both Saupe (1928) and Roeser (1940) observed no
cannibalism during extensive studies with _Pycnoscelus surinamensis_. In
fact, Roeser stated that there was never a case of cannibalism in spite
of long hunger periods imposed on both nymphal and adult insects.
XVII. ASSOCIATIONS AMONG COCKROACHES
Besides preying on their own species or on other blattids, cockroaches
exhibit additional symbiotic relationships among themselves. These
relationships are (1) the familial associations of parent and offspring,
(2) gregariousness, (3) intraspecies fighting, (4) interspecies
compatibility, and (5) interspecies antagonism. There are some
inconsistencies between observations made on the same species by
different workers, which only further observation and experimentation
will explain. Some of the reported observations are unique; this is
especially true for the feral species. Because of the paucity of
information, it is impossible at this time to make valid generalizations
about some of these interesting relationships.
FAMILIAL ASSOCIATIONS
The females of many species of cockroaches insure varying degrees of
protection to the developing young in their ways of disposing of the
oötheca after it has been formed. The extent of this association between
the mother and her developing progeny varies from the minimum amount of
time spent by oviparous females in concealing their oöthecae, to the
duration of embryogenesis in the so-called viviparous species, a period
of over a month or more.
Haber (1920a) observed a female of _Periplaneta americana_ chew a groove
in a piece of pasteboard into which she attempted to deposit her
oötheca. The oötheca failed to adhere to the shallow hole and fell to
the floor. After several futile attempts to replace the oötheca in the
hole, the female finally left the egg case on the floor of the cage and
coated it with an oral secretion to which she attached bits of trash.
During this operation she chased other females away when they ventured
near the site. Qadri (1938) described the behavior of the female of
_Blatta orientalis_ in concealing her oötheca in a hole that she dug in
sand; she deposited the egg case in the hole, coated it with saliva and
sand, and then refilled the pit. Rau (1943) described in detail how
females of _P. americana_ and _B. orientalis_ covered their oöthecae
with wood dust or sand in holes they had prepared in the substrate. Both
species placed a sticky oral secretion in the holes and then deposited
their oöthecae therein. After coating the oöthecae with more sticky
secretion, the females adjusted the oöthecae so that the keels were
uppermost and then carefully concealed the oöthecae with the excavated
debris. Both females spent over an hour in the act. Rau (1924)
previously reported that of 90 oöthecae deposited by _B. orientalis_ in
jars containing earth and trash, 36 were placed in crevices or excavated
holes, and 38 were hidden by being covered with dirt stuck to them with
saliva; only 16 were left uncovered.
Edmunds (1957) described oviposition by _Periplaneta brunnea_. Some
females spent from 30 to 40 minutes secreting from the mouth a frothy
substance that was smeared on the substrate; the egg capsule was
deposited in the secretion and covered with additional froth, which
hardened into a very strong cement. Some females spent as long as two
hours coating the capsule after it was deposited. It was not stated
whether the oötheca was otherwise concealed. The female remained with
her body over the oötheca for several hours and drove away other
cockroaches which approached.
Sonan (1924) observed that _Periplaneta americana_ and _Periplaneta
australasiae_ spent from 40 minutes to an hour covering their oöthecae,
and that if the females were frightened away from this activity, they
returned again to complete it. As well as excavating holes in the
substrate in which to deposit its oöthecae, _P. americana_ also avails
itself of readymade crevices of appropriate size (Ehrlich, 1943).
Species of _Epilampra_ in Malaya were said by Annandale (1900) to
deposit their oöthecae in crevices in floating logs just above the water
line. However, Shelford (1906) stated that four genera (including
_Epilampra_) of the subfamily Epilamprinae are "viviparous," in which
event the females would carry their oöthecae within their bodies during
embryogenesis and would not place the oöthecae in crevices in logs.
The female of _Cryptocercus punctulatus_ was observed to make a groove
in a piece of wood, then carry her oötheca 6 inches from where she had
dropped it and place it in the groove; she covered the oötheca so that
only a portion was visible (Cleveland _in_ Cleveland et al., 1934). Dr.
W. L. Nutting (personal communication, 1954) collected a number of
oöthecae of _C. punctulatus_ in the field and found each one almost
completely sealed off with bits of wood in a deep groove in the roof of
a chamber in a log. The keel of the oötheca was visible but the rest was
well camouflaged. He stated that "The adult pair usually frequents the
chamber at this time, while their broods of previous years occupy
neighboring galleries."
Berland (1924) observed a female of _Loboptera decipiens_ filling a hole
(the abandoned nest of a hymenopteron) with earth that she carried in
her mouth; he later found her oötheca behind the earthen barricade which
she had erected.
In summary, the following species of oviparous cockroaches have been
observed concealing their oöthecae (only those references not previously
cited are given): _Blatta orientalis_; _Cryptocercus punctulatus_;
_Ectobius sylvester_ (Harz, 1956, 1957); _Epilampra_ sp.; _Eurycotis
floridana_ (Roth and Willis, 1954a); _Loboptera decipiens_; _Balta
scripta_, _Methana curvigera_, _Methana marginalis_, and _Methana
caneae_ (Pope, 1953a); _Pelmatosilpha marginalis_, _Pelmatosilpha
purpurascens_, and _Nauclidas nigra_ (Bunting, 1956); _Periplaneta
americana_ (Haber, 1919; Adair, 1923; Seín, 1923; Nigam, 1933; Gould and
Deay, 1938; Rau, 1940a); _Periplaneta australasiae_ (Girault, 1915b;
Spencer, 1943; Pope, 1953); _Periplaneta brunnea_ (Roth and Willis,
unpublished data, 1958); _Periplaneta fuliginosa_ (Gould and Deay,
1940); _Periplaneta ignota_ (Pope, 1953); _Supella supellectilium_
(Flock, 1941). Undoubtedly other oviparous species that drop their
oöthecae long before the eggs hatch also make some attempt to conceal
the oöthecae by placing them in crevices or covering them with debris.
Sometimes the oöthecae are deposited but not concealed. Hafez and Afifi
(1956) reported that in Egypt _Supella supellectilium_ attaches its
oötheca to a suitable substrate with a gummy oral secretion but leaves
the egg capsule otherwise exposed. We (1954) have noticed similar
behavior in laboratory colonies of this species and of _Blatta
orientalis_, as have Gould and Deay (1940). Cornelius (1853) stated that
the female of _B. orientalis_ takes care of the safety of her offspring
to the extent of usually dropping her oöthecae in places which are dry
and raised above the ground, although rarely one also may find some
oöthecae scattered on the ground. For lack of suitable material females
of _Periplaneta americana_ sometimes did not conceal their oöthecae
(Nigam, 1933). Frequently in laboratory colonies _P. americana_ merely
drops the oöthecae loosely in sand or food "in contrast to _P.
australasiae_, which almost always went to considerable trouble to
fasten their eggs securely and to conceal them with debris" (Pope,
1953). If conditions under which _Nauclidas nigra_ is kept are not
suitable, the female will drop her oötheca anywhere (Bunting, 1956). Rau
(1940) stated that the female of _Parcoblatta pensylvanica_ does not
conceal her oötheca. However, Gould and Deay (1940) stated that this
species deposits its oöthecae loosely behind bark. _Ellipsidion affine_
and _Ellipsidion australe_ attach their oöthecae to bark or the
underside of leaves but apparently make no attempt to conceal them
(Pope, 1953a).
The females of most of the above species have no further familial
association with their offspring. The eggs hatch with no attention from
the mother who is probably not even in the vicinity at that time. The
young apparently do not react to the presence of the parent, as such,
after hatching. This is not unexpected, as several additional oöthecae
may have been deposited by these oviparous females before the eggs of
the first oöthecae hatch. However, a different behavior is encountered
among species that do not form a second oötheca until after the eggs of
the first have hatched (see below) and in the so-called colonial
species.
Shaw (1925) reported that in Australia both _Panesthia australis_ and
_Panesthia laevicollis_ appear to live in families, and that one usually
finds a pair of adults associated with from 12 to 20 nymphs in
different stages of development; he continued, "it is only where the
molts are very abundant that one loses sight of this familial habit."
Tillyard (1926) also stated that the Australian species of _Panesthia_
live in burrows in soil in strict family communities of a pair of adults
and 10 to 20 nymphs. A related colonial species, _Cryptocercus
punctulatus_, lives in both sound and rotten logs in colonies consisting
of a pair of adults and 15 or 20 nymphs, probably representing two or
three broods (Cleveland et al., 1934; Cleveland, 1948). Chopard (1938)
has cited this association as an example of gregariousness, which it may
well be; however, the presence of only one pair of adults in each colony
suggests a more intimate relationship.
Among species of _Blattella_ and certain other genera with similar
reproductive habits the female carries her oötheca clasped in her
genital cavity with the posterior portion projecting behind her. Each
normal oötheca is carried for approximately the duration of
embryogenesis and is not dropped until, or shortly before, hatching. We
have seen (1954, fig. 65) newly hatched nymphs of _Blattella vaga_ crawl
over the body of the mother who stood quietly near the dropped oötheca;
this female raised her wings and some of the nymphs crawled under them
onto the dorsal surface of her abdomen. The nymphs seemed to feed on the
grease covering the mother's body. The association was short-lived,
however, and soon the nymphs scattered. Pettit (1940) stated that when
hatching of _Blattella germanica_ occurs in the open (on a table top),
the nymphs may remain near the capsule only a few minutes. Ledoux (1945)
found that newly hatched nymphs of _B. germanica_ remained together
without shelter in a single, sparse group. If the nymphs were separated
by blowing on them, the group quickly reassembled, usually in the same
spot. Ledoux showed that this gregarious grouping of first-instar nymphs
was not necessarily a familial association by placing nymphs from two
oöthecae together. In groups of 8 to 12 nymphs there was a perfect
intermingling of the offspring from the two different females.
It is among the so-called viviparous cockroaches that the greatest
number of observations have been made of postparturient associations
between female cockroaches and their offspring. The females of these
species carry their oöthecae in brood sacs within their bodies until
embryogenesis has been completed. This behavior ensures protection of
the young from desiccation and attack by parasites (Roth and Willis,
1955a). (See Roth and Willis, 1958a, for an analysis of oviparity and
viviparity in the Blattaria.) Shelford (1906, 1916) reported that he had
captured a female of _Pseudophoraspis nebulosa_ in Borneo with numerous
young nymphs clinging to the undersurface of her abdomen. He also
recalled that there was in the Hope Museum (Oxford) a female of
_Phlebonotus pallens_ to which the following label was attached:
"'Ceylon ... carries its young beneath its wing covers. 1878.'" Pruthi
(1933) found in South India another female of _P. pallens_ which was
carrying over a dozen young nymphs on her back beneath her wings. In his
paper Pruthi reproduced a photograph of this specimen with the
light-colored nymphs in place on the back of the female. Hanitsch (1933)
reported having seen a museum specimen from Luzon, Philippine Islands,
of the apterous female of _Perisphaerus glomeriformis_ with nymphs still
clinging to her undersurface; he also reported having seen a museum
specimen of a female of _Ellipsidion variegatum_ from Australia with
four young clinging to the upper side of the apex of her tegmina and six
to the oötheca which projected beyond her body. Presumably this specimen
was giving birth when captured. Gurney (1954; personal communication,
1958) stated that specimens of _Perisphaerus_ sp. from Mindanao and
Luzon have been found with young nymphs clinging to the middle and hind
coxae. The first-instar nymph has an elongate face and specialized
galeae. Karny (1925) also observed that at the slightest alarm the young
of some species of Phoraspidinae creep under the dome-shaped front wings
of the mother.
The newly hatched young of _Leucophaea maderae_ have also been seen
congregated under the mother on several occasions. Seín (1923) stated
that after being born, the nymphs of this species gather under the
mother and accompany her at night in her excursions in search of food.
Pessôa and Corrêa (1928) reported that "During the first days the free
larvae hide under the adult cockroach which becomes restless and active
in contrast to its usual slow gait." Wolcott (1950) stated that "They
are not only gregarious, but the mother broods over her young, and
together they sally forth at night in search for food, until they are of
such a size as to mingle with their elders."
The African mountain cockroach _Aptera fusca_ has been observed during
late summer and early winter in familial groups beneath loose bark,
under stones, and in dead leaves (Skaife, 1954): "Each party consists of
a number of black young ones, together with one, two or more adult
females and perhaps a winged male or two. Later on they scatter and live
more or less solitary lives." In Malaya Karny (1924) often found
phoraspidine females between leaves surrounded by about 20 young nymphs.
He stated that one also often found females of _Perisphaerus armadillo_
surrounded by pale, yellowish-white young; similarly he had observed
that _Archiblatta hoevenii_ was found mostly in colonies made up of
mothers and their young. The duration of these associations is not
known.
Saupe (1928) noticed that the newly hatched nymphs of _Blaberus
craniifer_ (see footnote II, p. 322) collected together under the body
of their mother and stated that this is as pronounced a case of brood
care as Zacher had observed with _Pycnoscelus surinamensis_. Nutting
(1953) stated that "A degree of maternal solicitude is exhibited by this
roach [_B. craniifer_], for many times I have observed the female to
remain motionless for an hour or more with her unpigmented brood
clustered around and beneath her body." We, too, have observed similar
behavior in laboratory colonies of _B. craniifer_ and _Leucophaea
maderae_.
Chopard (1950) noted that after hatching the young of _Gromphadorhina
laevigata_ remained grouped around the female for some time; the mother
stood motionless, high on her legs, with her thorax curved up to make
room for the brood which hid under her body. We (unpublished data, 1958)
have seen young nymphs of _Gromphadorhina portentosa_ also stay near
their mother for some time after birth; the mother at this time produced
a characteristic hissing sound when she was only slightly disturbed by
the movement of our hand near her and her brood. The sound is produced
as air is expelled through the second abdominal spiracle. We have seen
recently hatched nymphs of _Nauphoeta cinerea_ crawl beneath the mother,
even under her wings, where they remained about an hour (Willis et al.,
1958). Bunting (1956) observed a female of _Blaberus discoidalis_
collect a mound of debris into which she inserted the tip of her
abdomen; he found young in the mound later the same day. This female
showed no maternal care for the young after birth. Whole families of
cockroaches may be found in bromeliads in Brazil (Ohaus, 1900). Hebard
(1920) observed a colony of adults and young of _Dendroblatta sobrina_
on a tree trunk in the Panama Canal Zone.
Whether any of the above associations exemplify maternal care for the
newly hatched young is questionable. The behavior of the mother, beyond
placing her eggs in a suitable location, seems to be entirely passive.
The first-instar nymphs are the active partners in these associations,
and they may merely be seeking shelter under the nearest convenient
object rather than under the mother as such. More extensive studies of
some of these relationships will be needed before claims for maternal
care, as suggested by Scott (1929), can be substantiated.
GREGARIOUSNESS
Casual statements that cockroaches are gregarious are often encountered
in the literature. There has been some argument to the effect that
large numbers of these insects seeking the same environment in a limited
space would appear to be gregarious, whereas there is probably no true
social tendency (Rau, 1924). Reactions of cockroaches to certain stimuli
in the environment undoubtedly do result in aggregations of individuals.
However, as Chopard (1938) has pointed out, it is difficult to assign
the respective parts played in assembling by the attraction of the
milieu and by gregarious instincts. Chopard (1938) also stated that
Orthoptera with a gregarious tendency are found rarely isolated; one
finds them, on the contrary nearly always collected in the same
shelters, close together, as if conscious of a need for contact between
themselves. He continued further that one can be tempted to attribute
the assembling to taxes but that interattraction equally plays an
important role; for example, if one places a large number of cockroaches
in a container and offers them similar shelters composed of cardboard
tubes, one finds that nearly all the individuals will assemble in one of
the tubes, ignoring the others. Pettit (1940) claimed that in _Blattella
germanica_ gregariousness seemed to depend on the mutual attractiveness
of body secretions as well as a thigmopositive behavior and love of
warmth.
Ledoux (1945) has studied experimentally gregariousness and social
interattraction in _Blatta orientalis_ and _Blattella germanica_. He
also found that the cockroaches tended to collect in shelters containing
other cockroaches. He concluded that group formation is not the result
of chance, but is a social phenomenon, and that interattraction is
mainly olfactory, conditioned by (1) positive chemotaxis to odors
emitted by the cockroaches themselves, (2) positive hygrotaxis, and (3)
thigmotaxis. He found also that large groups are not stable and tend to
break into smaller groups.
Gregariousness in the Orthoptera varies in intensity according to the
species and within a species according to the age or physiological state
of the insects (Chopard, 1938). This is well exemplified by several of
the blattid species discussed below.
Gregarious groupings of cockroaches have been observed most frequently
among the domiciliary species. A few examples will suffice. Gal'kov
(1926) observed heavy infestations of undetermined cockroaches in
workers' living quarters in the Ural region: "In the corners near the
stove, the cockroaches covered the walls in a dense carpet." After
fumigating he collected about 135,000 dead cockroaches from one barracks
and about 475,000 from another. We have reviewed a few other examples of
heavy infestations in our 1957(a) paper.
_Periplaneta americana_ was observed by Gould and Deay (1938) in an old
meat-packing building in Indiana. Adult cockroaches were present in
large numbers between closely placed beams, but the nymphs were more
common in cracks between bricks. Clusters of several hundred cockroaches
were seen on the open walls of the cold, dark hide room. Gould and Deay
stated "American roaches of all sizes live together in perfect harmony.
Young nymphs have been noted in clusters underneath adults and crawling
over the adults as they wander about in rearing jars." In the monkey
house of the Hamburg zoo, _P. americana_ spent most of the day in the
cellars resting on the walls in groups of about 200 individuals
(Brecher, 1929). Lederer (1952) noted that in closed, dark, heated
spaces under the aquarium at Frankfurt am Main, _P. americana_ rested in
groups of 20 to 30 individuals; he stated that it was remarkable that
the "herd" divided itself into groups each of which usually contained
insects of the same age or stage of development. Eads (1954) found _P.
americana_ in 40 percent of 762 sewer manholes in Tyler, Tex.; 13
percent of 670 of these manholes were heavily infested with 100 or more
cockroaches in each. Other heavy sewer infestations have been reviewed
in our 1957(a) paper.
Ehrlich (1943) has stated that _Periplaneta americana_ exhibits social
behavior. For instance, cockroaches of various ages inhabit a fairly
large space jointly; the adults and older nymphs sense approaches with
their antennae and warn and protect the young by a beating of wings and
by body movements. There is complete utilization of the available living
space; the imagos drive older nymphs from their resting places, and the
older nymphs drive out the younger ones, until all cracks, depending on
their size, are occupied by various age groups of different sizes. In
his experiments Ehrlich observed that in cages with no hiding places the
cockroaches would group together; when given a choice of small and large
shelters, _P. americana_ hid only under the larger ones that could
shelter more insects. Finally, the cockroaches ceased to bite and fight
each other when they crowded together in the face of danger.
Of _Blatta orientalis_ Marlatt (1915) stated "This species is notably
gregarious in habit, individuals living together in colonies in the most
amicable way, the small ones being allowed by the larger ones to sit on
them, run over them, and nestle beneath them without any resentment
being shown." Haber (1919) also observed that this species is often
noticed "huddled together, the younger ones crawling over, around, and
beneath the older ones."
Wille (1920) observed that nymphs of _Blattella germanica_ remained
almost constantly in groups during the first and second instars, but
less so during the third instar. He believed that the aggregations of
young occurred because they could occupy narrow crevices where the
larger insects could not penetrate. At usual room temperatures the older
nymphs and adults lived completely isolated, but at certain temperatures
they gathered together in large, tightly pressed groups.
_Supella supellectilium_ is said to be gregarious (Gould and Deay,
1940). The smaller nymphs aggregate in small groups in rearing
containers, but the older ones remain separate from one another (Hafez
and Afifi, 1956). _Leucophaea maderae_ is sociable and rarely found
alone; in their favorite hiding places, hills of these cockroaches can
be seen hanging together (Seín, 1923). Wolcott (1950) also stated that
_L. maderae_ is gregarious. Annandale (1900) observed that in the
"Siamese Malay States" large colonies of _Periplaneta australasiae_
conceal themselves in hollows of bamboo logs from which houses are
built. Moulton (1912) stated that he was astonished at the large numbers
of _P. australasiae_ and _Symploce cavernicola_ that he saw swarming on
the sides of caves of Mt. Jibong, Borneo.
Rehn and Hebard (1905) stated that in Key West, Fla., _Eurycotis
floridana_ fairly swarmed under the coquina boulders in the woods, in
groups of a dozen containing both young and adults; _Pycnoscelus
surinamensis_ was very abundant in the same type of habitat. Caudell
(1905) also found the young of _E. floridana_ with the mature
individuals. Hebard (1917) in his discussion of _Lattiblattella rehni_
again mentioned finding frequent colonies of _E. floridana_ in Florida.
He also found many specimens of _Blaberus craniifer_ under boards on the
ground at Key West. He found _Parcoblatta lata_ numerous under bark of
dead pine trees in Alabama. However, Dowdy (1955), in an ecological
study of oak-hickory forest in Missouri, stated that "_Parcoblatta_
[sp.] were never recorded as being gregarious, in fact they were mostly
solitary. However, in some cases two were found together." Yet Blatchley
(1895) stated of _Parcoblatta pensylvanica_ that in the winter in
Indiana "One cannot pull the loose bark from an old log without
dislodging a colony of from ten to a hundred of the nymphs of various
sizes." Males of _Parcoblatta virginica_ were said to be often
gregarious beneath loose bark and under chunks and rubbish (Blatchley,
1920).
Rehn and Hebard (1927) quoted observations made earlier by Hebard on
_Byrsotria fumigata_ in Cuba: "I found the specimens under flat stones,
sometimes in colonies of 3 or 4 mature specimens and numbers of immature
individuals in all stages of development." These observers also reported
that _Aspiduchus borinquen_ was found in Puerto Rico in a limestone
cavern by thousands in the grass and on the walls. J. W. H. Rehn (1951a)
stated that a related species, apparently _Aspiduchus cavernicola_, was
seen in great numbers on the side walls and roof of a cave in Puerto
Rico, but it was not possible to collect any of these and, we infer,
confirm the species. Rehn and Hebard (1927) in their account of
_Simblerastes jamaicanus_ reported finding it in numbers in a termite
nest. Pemberton and Williams (1938) stated that _Diploptera punctata_ is
of gregarious habits in Hawaii. Saupe (1928) observed a strong "herd
instinct" in all age groups of _Blaberus craniifer_. Bunting (personal
communication, 1956) stated that large nymphs and adults of _Blaberus
discoidalis_ "congregate in narrow cracks or on the underside of some
low object. The younger nymphs keep in close communities of
approximately the same age." Sonan (1924) stated that in Formosa(?)
_Salganea morio_ is usually found in groups of six or seven in decayed
trees. Species of the genus _Litopeltis_ may be found in small groups as
they are somewhat gregarious (Rehn, 1928).
The physiological or psychological effects of gregariousness, or lack of
it, are interesting aspects of the basic phenomenon. Landowski (1937)
studied in _Blatta orientalis_ the effect on development and growth of
the transition from life in complete isolation to life in groups. He
kept nymphs in groups of 1, 2, 4, 8, and 16 in jars of identical size
and shape. Landowski found that (1) mortality increased with the size of
the group and with age, as each animal occupied more of the available
space. [Presumably these factors are less detrimental in nature where
the group is unconfined.] He further found that (2) life in complete
isolation extended the time required to produce an adult insect; and (3)
the mean weight of the adult insect was, generally, in inverse
proportion to the number of nymphs raised together; isolated insects
usually attained the greatest adult weight.
Similarly, Griffiths and Tauber (1942a) found that isolation extended
the period of nymphal development in _Periplaneta americana_. As most of
their isolates died before reaching maturity, these workers concluded
that the American cockroach does not thrive when individually isolated
and that several individuals must be together for optimum development to
occur. Pettit (1940, 1940a) observed that isolated nymphs of _Blattella
germanica_ take longer to mature than those reared in groups. Wallick
(1954) found indications in _B. germanica_ that there is an inverse
relationship between population density and individual weight; as the
population decreased the weight increased. He also noted an inverse
relationship between population density and life expectancy in this
species.
We (Willis et al., 1958) have confirmed the above observations that
_Blattella germanica_, _Blatta orientalis_, and _Periplaneta americana_
complete nymphal development in less time when reared in groups rather
than individually. We (loc. cit.) also found that nymphs of the
following additional species matured more quickly when reared in groups:
_Eurycotis floridana_, _Periplaneta fuliginosa_, _Supella
supellectilium_, _Nauphoeta cinerea_, and _Pycnoscelus surinamensis_;
only a very slight decrease in the average length of the developmental
period was found in grouped nymphs of _Leucophaea maderae_.
Wharton et al. (1954) observed that virgin adult males of _Periplaneta
americana_ that had been individually isolated upon emergence were
almost wholly unresponsive to the sexually stimulating, female odor for
a test period of four weeks. Similar males of comparable age that were
kept in groups reacted strongly from the sixth day on. Removal of
reactive males from the group inhibited the reaction in these isolates,
but the response returned when the insects were regrouped. We (1952) had
similarly observed that no isolated male of _Blattella germanica_ was
ever seen to give a courting response without having received some form
of external stimulation. Yet when numbers of males were kept together
isolated from females, on several occasions the males became active and
a few individuals gave a courting response. As the sexual stimulus is
received by the male of _B. germanica_ through contact rather than odor,
as in _P. americana_, presumably it was mutual contact between the
grouped males that released the courting activity.
Cloudsley-Thompson (1953a), in his studies of diurnal rhythms in
_Periplaneta americana_, observed a steady decline in total activity in
successive 24-hour cycles: "When two cockroaches, even of different
species (_P. americana_ and _P. australasiae_) were kept together, this
depression did not appear to set in so readily." The associates
apparently kept each other active.
Isolated females of _Periplaneta americana_ can be conditioned to run a
simple maze with less time and fewer errors per trial than when paired
or when a member of a group of three (Gates and Allee, 1933). There was
less activity, and accordingly fewer errors per minute, among
cockroaches tested as pairs and groups of three than as isolated
individuals. This observation should not be contrasted with that of
Cloudsley-Thompson (1953a), cited above, because the intervals during
which activity was observed were quite different.
In the above account we have presumed that aggregations of some species
are indications of gregariousness. However, until gregariousness has
been proved experimentally for each species, we concede that reactions
to environmental stimuli might be sufficient to bring about some of the
observed groupings without any interaction between individuals.
In concluding this section we note that Tepper (1893) stated that
carnivorous cockroaches in Australia lead more or less solitary lives,
and that one rarely meets several together in close proximity. Takahashi
(1940) observed that in Formosa _Blattella humbertiana_ does not have a
tendency to throng together. Rau (1947) stated that the adults of
_Ischnoptera deropeltiformis_ showed no tendency toward gregariousness,
but in the laboratory newly hatched young lived close together under
bark and remained together throughout the nymphal stages. We wonder
whether this gregariousness was not imposed by the restricted quarters
of the cage. As mentioned above, Dowdy (1955) did not find _Parcoblatta_
sp. to be gregarious in the field.
INTRASPECIES FIGHTING
Fighting occurs among cockroaches of the same species over food or
shelter or between males. Saupe (1928) observed late-instar nymphs of
_Blaberus craniifer_ attack each other and even adults. Additional
records cited in the section on intraspecies predation (p. 322) imply
fighting within a species. Rau (1924) saw a male of _Blatta orientalis_
attack another male in copula and bite away a large portion of its wing.
Two other males in the container had their wings badly torn overnight,
presumably as a result of fighting.
Ehrlich (1943) stated that individuals of _Periplaneta americana_ that
are feeding will ward off intruders by spreading their wings and pushing
with their hind legs. However, the intruder will approach again and
again biting the feeder in the legs and wings. Frequently the odor of
approaching food was sufficient to cause the cockroaches to fight and
bite each other. Biting and fighting also occurred when individuals of
this species defended their daytime hiding places. A position of attack
is assumed when two antagonistic individuals of _P. americana_ meet
(Ehrlich, 1943, fig. 14). The insects raise their bodies slightly above
the ground, by extending their legs, and they stretch their heads
forward horizontally so that their mouth parts protrude; when the
insects jump at each other, they may wound each other severely in the
soft parts of the body. Fighting between sexually excited males resulted
in injury to their legs, wings, cerci, and other parts of the body.
Frequently an insect that could no longer defend itself was killed.
Lederer (1952) also made similar but less extensive observations on
fighting in this species.
Pettit (1940) quoted Woodruff as stating that nymphs of _Blattella
germanica_, apparently healthy and perfectly normal, would do battle for
no apparent cause other than a chance meeting, and that occasionally the
fight was to the finish, the loser being eaten. Pettit could not
substantiate such voracious attacks, although he saw nymphs engage in
fights lasting about two seconds during which one would be driven off by
vigorous bites on legs or cerci. Small nymphs of _B. germanica_ tended
to ignore each other, but third-and later-instar nymphs would engage in
"quarrels" of short duration when two met. Pettit noted that males of
_B. germanica_ that were crowded together quickly set upon, but did not
always kill, other cockroaches introduced into their cage. When he
isolated a dozen males in a small cage, they became quarrelsome and
three of the group were killed and partly eaten. After several days the
surviving males had taken positions so that each was equidistant from
his neighbors. Some of these males attacked other males and a female
that were introduced, by biting their legs and cerci. Females under
similar conditions were much less aggressive, although Pettit saw some
females that roved about biting all large members of the group that were
within easy reach.
We have frequently observed aggressive behavior between males of
_Nauphoeta cinerea_, which resulted in torn wings. The males would
wrestle with each other rolling over and over.
INTERSPECIES COMPATIBILITY
We agree in essence with Chopard (1938) who stated that it is improper
to speak of associations apropos of the ecological distribution of
Orthoptera. He continued that it is clearly evident that different
species of Orthoptera, which are found grouped on a territory more or
less narrowly limited, have no interdependence among them. Their
grouping results uniquely from almost similar reactions to the different
factors which characterize this limited milieu. There is neither
interdependence nor interaction; the grouping is a false biocoenose,
born under the action of the environment, and does not survive a
modification of this milieu.
However, as there are numerous examples of mutual toleration between
different species as well as examples of incompatibility, the subject
has more than academic interest even if no true ecological significance.
On the other hand, further study may show that certain of these
associations are definitely ecological, particularly among the feral
species. As might be expected, most of the following examples pertain to
domiciliary cockroaches.
Dozier (1920) occasionally found _Periplaneta americana_ with _Eurycotis
floridana_ in decaying stumps, beneath loose bark of decayed trees, and
beneath corded wood. Adair (1923) stated that in his house in Egypt
_Periplaneta americana_, _Blatta orientalis_, and _Blattella germanica_
were found together in a cupboard. Sambon (1925) found _B. orientalis_
and _B. germanica_ side by side but not fraternizing in a home in Italy.
Gould and Deay (1938) observed that apartments over stores were infested
with both _B. germanica_ and _P. americana_, but did not indicate
whether these occupied the same microhabitat. Gould and Deay (1940)
observed that in the Purdue University greenhouse _Periplaneta
fuliginosa_ was found "under benches, boxes, pots and other objects in
association with the American roach." Dr. L. A. Hetrick (personal
communication, 1954) wrote us that several summers before he had had a
mixed infestation of cockroaches, which included _Periplaneta
australasiae_, _Periplaneta fuliginosa_, and _Pycnoscelus surinamensis_,
in his chicken shed.
Eads (personal communication, 1955), in response to our inquiry about
the mixed populations of cockroaches that he had reported infesting
sewers in Texas (Eads et al., 1954), stated that "Each of the ten
colonies of _B. orientalis_ found in Tyler manholes were associated with
larger colonies of _P. americana_. True breeding colonies of _B.
orientalis_ appeared to be present since all the developmental stages
were taken. The same situation existed with the _P. fuliginosa_ and the
two species of _Parcoblatta_. Larger colonies of _P. americana_ were
associated with the other species in each case. From our limited
observations the two species always appeared to be perfectly
compatible." Eads et al. (1954) had found _Periplaneta fuliginosa_ in
three manholes, _Parcoblatta bolliana_ in one manhole and _Parcoblatta
pensylvanica_ in one manhole. We assume that the groups of each species
were spacially discrete so that they were recognizable as colonies. Dr.
T. A. Olson (personal communication, 1958) has observed two or more
species of cockroaches in a single structure but never in mixed
colonies. Each species was separated physically from the others. Olson
concluded that cockroaches of different species do not mingle freely
unless forced to do so by some special environmental condition. Pettit
(1940) found _B. germanica_ and _P. americana_ similarly separated in
the same building or even in the same basement laboratory.
Perkins (1899) found _Lobopterella dimidiatipes_ generally living in
company with the young of _Periplaneta australasiae_ in Hawaii. Rehn and
Hebard (1914) in Florida found _P. australasiae_ abundant with
_Periplaneta americana_ on a quarter-boat. They also noted that the
forficulid _Marava_ [= _Prolabia_] _arachidis_ (Yersin) appeared in
numbers in a kitchen after dark accompanied by swarms of _P. americana_.
These workers also found _Leurolestes pallidus_ in a fruit store in Key
West "where the species was common in a pile of old burlap bags and in
cracks under the stands which it shared with one fairly large colony of
_Blattella germanica_, occasional specimens of _Holocompsa nitidula_, a
few specimens of _Periplaneta americana_, and one specimen of _Supella
supellectilium_." They also found _H. nitidula_ with _Blaberus
craniifer_ "between old boards in a woodshed, where nymphs were more
numerous than adults."
Rehn and Hebard (1914) stated of _Supella supellectilium_ in Florida
that "The females were all taken in cupboards where _Blattella
germanica_ was found in swarms." The association in human habitations of
_S. supellectilium_ and _B. germanica_ has been reported also by Sein
(1923), Puerto Rico; Shaw (1924), Australia; Mallis (1954): "German and
brown-banded roaches were often found in the same crevice."; Anonymous
(1958), Texas; and Anonymous (1958a), Georgia. Gould and Deay (1940)
stated that other species of cockroaches, especially _B. germanica_, may
be found with _S. supellectilium_ in the same part of a building. Yet
Shaw (1925) stated that "when _Supella supellectilium_ Serv. invades
places already occupied by _Blattella germanica_ L., it tends to oust
the latter."
_Blaberus discoidalis_ has been found in homes or in fruit debris in
Puerto Rico in company with the more common, domiciliary species
_Leucophaea maderae_, but never in abundance (Sein, 1923; Wolcott,
1950). Illingworth (1915) in Hawaii found _Symploce hospes_ associated
with _Nauphoeta cinerea_, _Graptoblatta notulata_, and _Diploptera
punctata_.
Hebard (1917) found _Aglaopteryx diaphana_ in a bromeliad on a forest
tree in Jamaica together with _Nyctibora laevigata_ and numerous
_Cariblatta insularis_. He also found numerous _Aglaopteryx gemma_ under
signs on longleaf pines in Alabama with occasional specimens of
_Parcoblatta lata_. In Virginia he found _Parcoblatta uhleriana_ in a
decaying chestnut log with _Cryptocercus punctulalus_. In Florida he
found _Latiblattella rehni_ with _Eurycotis floridana_ and, more rarely,
with _Periplaneta australasiae_ under bark of pine trees. In Key West he
found _Symploce hospes_ in the cupboard of a hotel with swarms of
_Blattella germanica_ and a few _Supella supellectilium_.
Rehn and Hebard (1927) in their study of West Indian blattids reported
finding _Neoblattella proserpina_ in epiphytic bromeliads in Jamaica in
company with _Neoblattella eurydice_ and _Neoblattella dryas_. They also
list most of the associations cited by Hebard (1917).
Ramme (1923) reported that he found in Germany four species of
_Ectobius_ (_lapponicus_, _lucidus_, _pallidus_, and _sylvester_) living
together in an area about 50 m. by 200 m. Although he had stated that
his specimens of _E. lucidus_ were a distinct species in 1923, Ramme
(1951) later decided that they were a form of _E. sylvester_, _E.
sylvester_ f. _lucidus_.
Dow (1955) reported trapping _Blattella germanica_, _Periplaneta
americana_, and _Periplaneta brunnea_ in houses and privies in south
Texas. At our request Dr. Dow (personal communication, 1958) analyzed
his records to determine whether there were indications of associations
between these species, with the following results:
As stated in my published note, the roaches were at first
classified to genus only. The 83 _Periplaneta_ subsequently
identified to species represented 28 different collections, 11 from
houses and 17 from privies, all in Pharr, Texas. Tabulation of the
data shows first that _P. americana_ was taken only once in a house
and that _P. brunnea_ was taken only 4 times in privies. Of course
this distribution greatly reduces the probability that they would
be caught together, and it is not surprising that _P. americana_
was trapped alone in the single house collection. _P. brunnea_,
however, was trapped with _P. americana_ 2 of the 4 times it
occurred in privy collections.
To investigate the occurrence of _Periplaneta_ with and without
_Blattella_, an analysis has been made of 560 trap collections
taken in 40 houses and 40 associated privies in Pharr, Texas, in
weekly intervals (from May 14 to June 23 [1948]). In the houses,
_Periplaneta_ and _Blattella_ were caught in the same jar 26
times, _Periplaneta_ alone 12 times, _Blattella_ alone 83 times,
and neither genus 159 times. In a fourfold table, the value of
chi-square (14.7) is significant and indicates that the
frequencies are not proportional. The number of times
_Periplaneta_ and _Blattella_ actually occurred together (26) is,
however, much larger than the expected number calculated from the
row and column frequencies (14.8). In the privies, _Periplaneta_
and _Blattella_ were caught in the same jar 9 times, _Periplaneta_
alone 50 times, _Blattella_ alone 18 times, and neither genus 203
times. In a fourfold table, the value of chi-square (1.95) is not
significant but the same type of disproportion is evident and the
expected frequency of both genera in one trap is 5.7, lower than
the actual frequency of 9. Both immature and adult roaches are
included in this analysis.
The above evidence would be more satisfactory if based on more
extensive data. There is also a possible objection in that the
traps were operated for at least overnight, during which time one
species could theoretically supplant another. Of course, it is
doubtful that there is anything involved here like territory (in
the ornithologists' sense). On the other hand, it is well to
consider that _Periplaneta_ and _Blattella_ are both likely to be
more abundant in the same type of favorable location and that this
factor might offset in part some direct antagonism between the
species.
The only known specimen of _Ischnoptera podoces_ was captured in company
with the type series of _Cariblatta nebulicola_, in dead leaf litter in
Jamaica (Rehn and Hebard, 1927). In Florida _Periplaneta australasiae_
was often taken in company with _Pycnoscelus surinamensis_ and
_Eurycotis floridana_ (Blatchley, 1920).
INTERSPECIES ANTAGONISM
In contrast to the presumably amicable associations mentioned above,
other observations in the literature seem to indicate that some species
of cockroaches are incompatible when they attempt to occupy the same
habitat niche. Marlatt (1915) stated "Rarely do two of the domestic
species occur together in the same house. Often, also, of two
neighboring districts one may be infested with one species, while in the
other a distinct species is the commoner one. The different species are
thus seemingly somewhat antagonistic, and it is even supposed that they
may prey upon one another, the less numerous species being often driven
out." Phelps (1924) stated "Roaches of different species are rarely
found together, although roaches of the same species live together on
very amicable terms."
In 1859 Darwin (1887) stated that "In Russia the small Asiatic cockroach
[_Blattella germanica?_] has everywhere driven before it its great
congener [_Blatta orientalis?_]." Yet in France Girard (1877) suggested
that the oriental cockroach be introduced into a restaurant infested
with the German cockroach as the best way to expel the latter, because
the more robust species drives away cockroaches of smaller size. Wille
(1920) in Germany found usually only one species of cockroach in a
house. Yet when he placed _B. orientalis_ and _B. germanica_ together,
there were no reciprocal attacks even by hungry individuals. Wille
concluded that because of their greater speed, smaller size, greater
number of eggs, and faster development, the German cockroaches eat the
available food and so make the environment unfavorable for the oriental.
However, he noted that cases may be seen in which the opposite is also
possible. Laing (1946; British Museum [Natural History], 1951) observed
that in the British Isles _B. orientalis_ seems to have lost its
dominant position to _B. germanica_ in recent years; it was stated that
these species are not as a rule found together and that the greater
rapidity of breeding and ability to climb of _B. germanica_, as well as
the layout of modern buildings, are some of the factors that favor the
spread of _B. germanica_. Ledoux (1945) found that first-instar nymphs
of _B. germanica_ and fourth-instar nymphs of _B. orientalis_, adults of
_B. germanica_ and sixth-instar nymphs of _B. orientalis_, as well as
adults of both species, did not form mixed groups. However, when he
combined fifth-and sixth-instar nymphs of _B. germanica_ with
fourth-and fifth-instar nymphs of _B. orientalis_, which are all
practically of equal size, sometimes he would find mixed groups, but
generally the groups were distinct. Lucas (1912) stated that Burr had
found _B. germanica_ and _B. orientalis_ swarming within a rubbish heap
in England; presumably both colonies were breeding and multiplying and
one species was not detrimental to the presence of the other.
Shaw (1925) claimed that _Supella supellectilium_ tended to oust
_Blattella germanica_, but Pope (1953) thought it doubtful in
Queensland. Wolcott (1950) stated that "The larger and more powerful
domestic cockroaches, _Periplaneta americana_ (L.), _P. australasiae_
(F.) and _P. brunnea_ Burmeister have very definitely fallen behind in
Puerto Rico in competition with the little German roach." Pessôa and
Corêa (1928) observed that other species of cockroaches were rare in
Brazil in houses that were infested with _Leucophaea maderae_. Lederer
(1952) noticed that in the reptile house of the aquarium at Frankfort am
Main _Blatta orientalis_ was obviously kept down by _Blattella
germanica_, even before the appearance of _P. americana_. However, _B.
germanica_ was not driven out of the reptile house by _P. americana_
although the populations of each fluctuated for about 22 years after the
American cockroach had settled there; both species occupied separate
resting places. Lederer further observed that within four years of the
introduction of _P. americana_ into the crocodile house, none of the
original infestation of _B. orientalis_ could be found; a small colony
of _Pycnoscelus surinamensis_ in the reptile house was apparently also
driven out by _P. americana_. Chopard (1932, 1938) stated that the
oriental cockroach does not exist in company with _P. americana_ which
very probably destroys it. Pettit (1940) kept _B. germanica_ and _P.
americana_ together in a cage for several weeks but neither species gave
any indication of feeding on the other.
Froggatt (1906) stated that "It is probable that the advent of the
larger and more formidable American cockroach into Australia has led to
the retirement or destruction of our indigenous species" [presumably
_Periplaneta australasiae_]. Tillyard (1926) noted that this statement
is incorrect as neither species is native to Australia. Yet Shaw (1925)
stated that in Australia "When both species live together in the same
places, _australasiae_ Fabr. will probably be found gradually to
displace _americana_ L." Local fluctuations in the relative abundance of
these species could be a basis for such dissimilar observations.
However, MacDougall (1925) observed that in the plant houses of the
Royal Botanical Garden, Edinburgh, the Australian cockroach seemed to
have overcome the American which had been more numerous in former years.
In conclusion, we emphasize that many of the above observations are
merely tentative impressions gathered by workers who have watched many
species of cockroaches in nature. Obviously, additional observations
coupled with appropriate experimentation will be needed to disclose the
true structure of each presumed association and to resolve apparent
discrepancies. Although we are greatly indebted to the cited authors for
their contributions to the known information, we anticipate that future
results of cleverly designed laboratory experiments will do much to
dispel the uncertainty that still surrounds our knowledge of the
relations of the Blattaria to each other.
XVIII. DEFENSE OF COCKROACHES AGAINST PREDATORS
Irritating or repellent secretions provide many animals belonging
to widely unrelated groups with a more or less potent means of
defence....
It will be seen that this method of defence does not rest merely
upon a passive unpalatable attribute, but upon an active emission
of the unpalatable substance which, since it occurs when the
animal is seized or threatened by an enemy, enforces its
effectiveness. In its highest development we find different forms
whose specialized habits and modified structure enables them to
_project_ secretion at the enemy, and thus to discourage attack.
COTT (1940)
There are very few records indicating that cockroaches are unaccepted as
food by other animals. Hutson (1943) found that the duck, guinea fowl,
and pigeon would not normally eat _Pycnoscelus surinamensis_, and in his
experiments with the chicken eye worm he had to force-feed his birds
with infected cockroaches. Lederer (1952) found that insectivorous birds
in the Zoological Garden, Frankfurt am Main, either refused hardened (as
opposed to teneral) American cockroaches or ate them unwillingly.
Carpenter (1925) reported that a monkey (_Cercopithecus_) failed to feed
on cockroaches and suggested that the insects' odor made them repugnant;
however, there are a number of positive records of monkeys feeding on
cockroaches (see pp. 284-286).
Cockroaches may escape capture by predators through evasive behavior,
concealment, protective coloration, mimicry, or secretion of malodorous
materials. Nocturnal cockroaches may avoid predators that are active
during the day (Crawford, 1934), but nocturnal predators are apparently
quite successful in capturing cockroaches. Some cockroaches may be
protected by their swiftness, others by their resemblance to vegetation
(Williams, 1928). The habit of squeezing into narrow cracks may afford
cockroaches some protection.
Burrowing forms such as _Pycnoscelus_ may spend much time in underground
cells (Roeser, 1940). Polyphagids rapidly burrow into sand (Fausek,
1906), where they may be protected from predators. Tepper (1893)
discovered that a very large Australian cockroach, _Geoscapheus
robustus_, had its fore legs, especially the tibiae, adapted for
digging. He observed this species in captivity and in 1894 reported that
it appeared to sink into the soil without raising any considerable
amount above the surface and that it did not form an unobstructed
tunnel. Another large Australian cockroach, _Macropanesthia rhinocerus_,
burrows about two feet below the surface of sandy soil; it also makes
nests among pine roots and the nymphs rarely appear above ground (Henson
_in_ Day, 1950). Tepper (1893) observed that Australian cockroaches of
the genera _Epilampra_ and _Oniscosoma_ buried themselves in loose soil
and dust. Baker (_in_ Rehn, 1930) observed that _Styphon bakeri_ is
found in humus and rubble in the Dutch West Indies where "It is sluggish
in the open, but wedges into the humus quite quickly."
_Therea nuptialis_, found in India, conceals itself at the roots of fig
trees, etc. The small hairs on its elytra retain sufficient dust to
conceal it, or at any rate to render it inconspicuous, when not on the
wing (Annandale, _in_ Chopard, 1924c). Rehn and Hebard (1914) observed
that the nymphs of _Blaberus craniifer_[13] at Key West, Fla., "were
usually found half buried in loose damp earth under boards, where they
remained motionless, looking much like lumps of earth (with which they
were usually much dusted) until disturbed." Hebard (1917) reported of
_Monastria biguttata_ from Brazil that "All of the juveniles are heavily
coated with foreign particles" which adhere "to a multitude of closely
placed, minute and usually curved spines, which cover the dorsal surface
and marginal portions of the ventral surface."
It is apparent from the numbers of predators reported herein that many
animals are not deterred by the odorous secretions of cockroaches; these
secretions, because they may seem repugnant to man, are often claimed to
be repellent to predators. However, Cott (1940) points out that "There
are many instances in which protective devices and associated warning
colours are known to be ineffectual against certain enemies. But this
does not necessarily imply that they are not on the whole beneficial to
the species attacked." Certain cockroach secretions may well be
repellent to many predators, but as this is a purely negative aspect of
the predator-prey relationship little thus far has been observed or
published. Potential prey that successfully defends itself against
attack is never found in a predator's stomach.
Cockroaches have a variety of glands which secrete odorous materials.
Certain secretions, produced by tergal or dorsal glands in males, are
involved in sexual behavior; the females feed on the secretion from
these glands prior to copulating (Roth and Willis, 1954). However, other
secretions which are produced by both sexes are ejected or given off
when the insect is disturbed; undoubtedly these are defensive weapons
that are used against predators. Very few experiments or observations
are on record to show how effective these secretions may be in
protecting the cockroach. Although the morphology of some of the glands
has been described, relatively little is known about the chemistry of
their secretions.
Many species of Australian cockroaches have been reported to emit
"disgusting" odors, though the glands producing these secretions have
not been described, nor is the chemistry of the compounds known.
_Cosmozosteria lateralis_ exposed two orange-red spots on the abdomen
while emitting a pungent odor which deterred a collector from capturing
it (Shelford, 1912). Another Australian species, _Platyzosteria
castanea_, when disturbed on barren ground tilts forward on the vertex
and straddles out the posterior legs, supporting itself in a vertical
position on the head and tarsi; in assuming this attitude it will squirt
a foetid fluid as a fine spray for a distance of 6 or 7 inches (Shaw,
1914). Spencer (1892) mentions the pungent odor given off by a cockroach
which had been accidentally cut in two. Rageau (1956) stated that in the
New Hebrides and New Caledonia _Cutilia nitida_ emits, when disturbed, a
corrosive liquid with an extremely disagreeable odor.
The adults of _Eurycotis floridana_ emit an odorous fluid when seized
(Rehn and Hebard, 1905). The fluid, which may irritate sensitive skin
areas, may be ejected as a spray for a distance of several inches. This
secretion has been identified as 2-hexenal (Roth et al., 1956), and the
ventral abdominal glands which produce it have been described (Stay,
1957). Eisner (personal communication, 1958) has found that the toad
_Bufo marinus_ and the frog _Rana pipiens_ invariably spit out adults of
_E. floridana_ which they have seized. The odor of 2-hexenal was
strongly apparent after these attacks, and the insect was never damaged.
However, the lizard _Anolis equestris_ seized and crushed _E.
floridana_ before releasing its hold and dropping the insect 5 to 10
minutes later. The blue jay _Cyanocitta cristata_ readily attacked
adults of _E. floridana_ and killed them but did not eat the insects
until after the odor had dissipated; however, the bird carried nymphs of
_E. floridana_ to its perch and ate them. Nymphs of this species do not
secrete 2-hexenal (Roth et al., 1956). Recently, 2-hexenal has been
tested for its antibacterial activity and has been found to be active
against seven species of pathogenic bacteria (Valcurone and Baggini,
1957). _Eurycotis decipiens_ from Trinidad also ejects a fluid which may
produce toxic symptoms such as vertigo and nausea (Bunting _in_ Roth and
Willis, 1957a).
Large reservoirs of glands similar in appearance and position to those
of _Eurycotis floridana_ are present in the adults of both sexes of
_Neostylopyga rhombifolia_ and _Platyzosteria novae seelandiae_. Walker
(1904) and Longstaff (_in_ Shelford, 1912) noted that the latter species
had a strong odor. Roth (unpublished data, 1957) found that the
secretion of _P. novae seelandiae_ when ejected is grayish or milky in
color. In the reservoirs of the ventral gland of this insect the
secretion is a milky liquid containing floating greenish globules. Both
infrared and mass spectrographic analyses show that the secretion is a
mixture containing 2-hexenal, the aldehyde that is found in _E.
floridana_. Eisner (personal communication, 1958) observed that the
lizard _Anolis carolinensis_ immediately released _Neostylopyga
rhombifolia_ without injury, but that _Bufo marinus_, _Anolis
equistris_, and _Cyanocitta cristata_ ate the insect despite the
secretion; several unidentified spiders and the ant _Pogonomyrmex
badius_ were not repelled by the secretion of _N. rhombifolia_.
Dorsal and ventral glands have been found in both sexes of _Blatta
orientalis_ and _Periplaneta americana_ (Minchin, 1888, 1890; Kul'vets,
1898; Oettinger, 1906; Harrison, 1906; Liang, 1956). The ventral glands
are found in the same general region as those of _Eurycotis_. We have
also found similar ventrally located glands in both _Periplaneta
australasiae_, and _P. brunnea_. The reservoirs which store the
secretion of the ventral glands are smaller in _Blatta_ and
_Periplaneta_ spp. than those found in _Eurycotis_, _Neostylopyga_, or
_Platyzosteria_.
In _Blatta orientalis_ the dorsal glands can be everted by pressure on
the abdomen; the secretion in these glands, according to Haase (1889),
has the typical oriental cockroach odor. Although the dorsal glands of
the oriental cockroach are usually given a defensive role (Haase, 1889,
1889a; Kul'vets, 1898; Oettinger, 1906; Konček, 1924), the functions of
secretions of these nonepigamic dorsal glands and the ventral glands are
still open to question. It is possible that some of the odors produced
by cockroaches have functions other than defense or sex attraction. For
example, Ledoux (1945) showed that the species odor is largely
responsible for the gregarious behavior shown by _Blatta orientalis_ and
_Blattella germanica_. The olfactory stimulus acts over a short distance
only, and the source of this odor in the insect is unknown. By washing
_Blattella germanica_ in warm chloroform Dusham (1918) extracted a wax
which had the odor of the German cockroach. However, there is no
evidence to show that cockroaches respond to the same cockroach odors
that are detected by man.
Certain cockroaches have recently been found to have odorous secretions
which are produced in tracheal glands. In _Diploptera punctata_ the
tracheae leading to the second abdominal spiracles of nymphs and adults
are modified into odoriferous glands which produce a mixture of
2-ethyl-1,4-benzoquinone; 2-methyl-1,4-benzoquinone; and _para_
benzoquinone; this material is ejected as a means of defense. The
offensive odor emitted by adults and nymphs of _Leucophaea maderae_ also
issues from the second abdominal spiracles (Roth and Stay, 1958).
_Diploptera_ is capable of ejecting its quinones from either its right
or left tracheal gland according to which side of the insect is attacked
(pl. 36, A-B). Eisner (1958) found that the secretion repelled the ant
_Pogonomyrmex badius_ (Latreille) (pl. 36, C) and the beetle _Galerita
janus_ Fabricius when they attacked the cockroach. The spider _Lycosa
helluo_ Walckenaer was repelled by large nymphs and adults of _D.
punctata_ but young nymphs were usually eaten promptly (Eisner, 1958).
Bordas (1901, 1908) believed that the "conglobate" gland (Miall and
Denny, 1886), found in males of _Periplaneta americana_ and _Blatta
orientalis_, was an odoriferous gland used for defense, but Gupta (1947)
has shown that in all probability this gland (the phallic gland)
secretes the outermost covering of the spermatophore.
What appears to be mimicry occurs in some species of Blattaria. The
nymphs of many Panchlorini and Blaberinae vaguely resemble sow bugs
(Chopard, 1938). Certain members of the Perisphaerini (e.g.,
_Perisphaerus glomeriformis_) from the Malayan region which resemble sow
bugs (Annandale, 1900; Hanitsch, 1915) can roll themselves up into a
ball thus hiding their antennae and legs (Lucas, 1862). Although these
cockroaches are found among dead leaves or under stones, in places in
which sow bugs are also found, the benefit to either or both forms is
questionable; Annandale (1900) believed that the crustacean and the
cockroach, living under similar conditions, developed the same general
body shape. Rolling up into a ball is nothing more than an exaggeration
of a reflex common to many young cockroaches, that is, an arched
position which these insects assume when they immobilize themselves in
response to certain stimuli (Chopard, 1938).
There are cockroaches that resemble various Coleoptera and Hemiptera
(Belt, 1874; Shelford, 1912; Hanitsch, 1915). Some look like
cerambycids, lampyrids, coccinellids, pentatomids, etc. Perhaps the most
striking examples are the resemblances of cockroaches in the genus
_Prosoplecta_ of the Epilamprinae to beetles of the family
Coccinellidae; Shelford (1912) has figured a number of species of
_Prosoplecta_ together with the species of beetles which they seem to
have taken for models. Williams (1928) mentioned diurnal cockroaches
which by a combination of markings, shape, posture, and active flight
about vegetation suggest certain wasps.
Unfortunately, practically nothing is known about the behavior of these
so-called mimics and models or their relationships with predators in the
field. For the most part, the examples are based on a comparison of
pinned insects from museum collections (Burr, 1899); for this reason
Chopard (1938) believed that not much value should be placed on
superficial resemblances of this kind. However, we believe that a lack
of knowledge of cockroach mimicry is not a valid reason for rejecting
the idea that mimicry, if it occurs, may be of some benefit in the
survival of mimetic species. Certainly Cott's (1940) voluminous
compilation of the literature on adaptive coloration should make the
most skeptic hesitate to conclude dogmatically that these instances of
mimicry are merely accidental and meaningless.
XIX. THE BIOLOGICAL CONTROL OF COCKROACHES
In the Navy [Japanese] a seaman who has captured 300 cockroaches
will be granted one day special shore leave. They call it "shore
leave for cockroaches." The purpose is to promote extermination of
cockroaches in a warship because, on the one hand, any warship
suffers from numerous cockroaches, and, on the other hand, any
seaman likes shore leave.... The formalities for a shore leave for
cockroaches are as follows. A seaman keeps cockroaches which he
captured (mainly _B. germanica_, because _P. americana_ and _P.
australasiae_ are seldom found in Japan) in a bottle or in a bag
until the number reaches 300. Then he brings them to the deck
officer to get the confirmation that he has actually captured more
than 300 cockroaches. If the deck officer confirms it, the seaman
goes to a cabin where a petty officer reports that the deck
officer confirmed the number of cockroaches. The petty officer
signs the seaman's name, name of division, rank, and date to be on
shore leave in the log book for cockroach shore leaves. The petty
officer brings the log book again to the deck officer to get his
approval and then goes to the commander for the final approval. In
the Navy, they have another special shore leave for rats. In this
system, a seaman gets one day shore leave for one rat. The
formalities for the latter are the same as for the former, and
there is a log book for the rat shore leave in the petty officer's
quarters. The author took advantage of these systems frequently.
SONAN (1924)
Little is known of the effects of predatism and parasitism on natural
populations of cockroaches. Many statements in the literature are very
general; yet there are a few data on egg parasites (e.g., _Tetrastichus
hagenowii_) which suggest that, in the absence of parasites, populations
of domestic cockroaches might be much larger than they are in certain
areas. We have summarized the literature on natural control and also
that on the use by man of predators and parasites in the biological
control of cockroaches. However, because of the paucity of information,
we have been unable to evaluate the effectiveness of biological control
in reducing the numbers of pest cockroaches. This is an area that might
reward further investigation.
INVERTEBRATES
_Scorpions._--In Puerto Rico, cockroaches are probably the principal
food of the scorpions which live in old houses, on tree trunks, etc.
(Seín, 1923). The staple diet of scorpions in Arizona is the small
cockroach commonly known as the water bug (Stahnke, 1949); in the part
of Arizona where he resides, Stahnke (personal communication, 1953) says
that the "water-bug" is most generally _Supella supellectilium_ although
_Blattella germanica_ is also found, but less abundantly.
_Spiders._--Jefferys (1760) mentioned a large spider which was protected
in the Antilles and especially on Guadeloupe because it hunted down and
fed on cockroaches; the spider was reputed to be common in every house.
Sir Hans Sloane (1725, _in_ Cowan, 1865) reported that residents of
Jamaica kept spiders in their houses to destroy cockroaches. Takahashi
(1924) reported that, in the Taihoku area of Formosa, human habitations
contained large numbers of spiders which caught and ate cockroaches.
Smith (_in_ Marlatt, 1915) reported that Brazilians encourage large
house spiders because they tend to rid the house of "other insect
pests." In British Guiana tarantulas were kept in a bungalow to control
_Periplaneta_ and _Pycnoscelus_ (Beebe, 1925a).
_Ants._--A Madam Merian noticed that ants cleared houses of cockroaches
(Kirby and Spence, 1822). A small reddish-yellow ant, called Pucchuçiçi
by Peruvian Indians, pursued and destroyed a cockroach called Chilicabra
which was a pest in native huts (Tschudi, 1847). Schwabe (1950b) found
swarms of ants attacking living _Pycnoscelus surinamensis_ and stated
that ants are probably the chief enemy of this cockroach in Hawaii.
Wallace (1891) stated that in Africa a band of driver ants may enter a
house and clear it of cockroaches and other arthropods. In British
Guiana, Beebe, (1925) found that several times a year army ants cleared
the laboratory of all cockroaches and tarantulas.
_Wasp egg parasites._--Matsumura (1917, _in_ Asano, 1937) proposed that
parasitic wasps such as _Evania_ and _Brachygaster_ be protected in
Japan as the natural enemies of cockroaches. In one area in France, 20
percent of the oöthecae of _Loboptera decipiens_ were parasitized by
_Zeuxevania splendidula_ (Genieys, 1924). Edmunds (1952a) found that 12
percent of 459 oöthecae of _Parcoblatta_ collected during December
through April of 1950-51 in Ohio were parasitized; evaniids accounted
for about 7 percent of the parasitization. Additional collection data in
1951-52 Edmunds (1953a) showed that 8.7 percent of 320 wood-cockroach
oöthecae were parasitized; 2.8 percent of these parasites were evaniids;
almost 13 percent of the egg capsules collected showed evidence of
previous parasite emergence. Cameron (1957) reported that oöthecae of
_Periplaneta americana_ collected in Saudi Arabia were 29 percent
parasitized in March and 25 percent parasitized in October by _Evania
appendigaster_. Sonan (1924) found 1 of 65 oöthecae of _P. americana_
and _P. australasiae_ parasitized by _E. appendigaster_ in Formosa.
Cottam (1922) stated that the increase of _Supella supellectilium_ in
Khartoum was checked by a wasp egg-parasite that was later identified as
_Anastatus tenuipes_ (see p. 246) (Ferrière, 1930, 1935). In this
country, this wasp seemed to be effective in decreasing the numbers of
_Supella_ in certain areas in Arizona (Flock, 1941).
In Formosa, _Tetrastichus hagenowii_ was an important parasite of
cockroach eggs (Maki, 1937). Sonan (1924) reported 30 percent
parasitization of 65 oöthecae of _Periplaneta americana_ and _P.
australasiae_ collected in Formosa. In Bangalore, India, the natural
parasitization of randomly collected oöthecae of _P. americana_ varied
from 21 percent (of 495 oöthecae), July 1947-June 1948, and 43 percent
(of 288 oöthecae), July-December 1948, to 57 percent (of 178 oöthecae),
July-October 1949 (Usman, 1949). Cameron (1955) obtained _T. hagenowii_
from oöthecae collected in Trinidad, B.W.I., and Saudi Arabia; some 15
percent of the oöthecae of _P. americana_ and _P. australasiae_
collected in October in Trinidad were parasitized; a later collection
(March) was 34 percent parasitized; a small sample of _P. americana_
oöthecae was 65 percent parasitized. The oöthecae collected in Saudi
Arabia in March were 20 percent parasitized. Plank (1947) found that the
eggs of the American cockroach in Puerto Rico (probably in laboratory
cultures) were so heavily parasitized by _T. hagenowii_ that he had to
use _P. australasiae_ for experimental purposes; in 1950 Plank stated
that more than 50 percent of American cockroach oöthecae were
parasitized.
Fahringer (1922) stated that _Prosevania punctata_ could be used to
eradicate cockroaches, but he did not test his hypothesis. Marlatt
(1902) felt that the usefulness of _Evania appendigaster_ in biological
control was impaired by _Tetrastichus_ acting as a hyperparasite (see
footnote 6, p. 236). However, Wolcott (1951) stated that in Puerto Rico
_E. appendigaster_ is quite abundant and is a factor of considerable
importance in controlling cockroaches. Kadocsa (1921) stated that
_Brachygaster minutus_ and _Evania appendigaster_ were not important in
the biological control of cockroaches. These general statements are not
supported by experimental evidence.
It is likely that the smaller wasp egg parasites are more effective than
the evaniids in controlling cockroaches. Only one evaniid develops in a
parasitized oötheca, but many individuals of the other wasps develop in
one oötheca and the number of females that emerge is usually large.
However, Cameron (1957) concluded that, with a parasitism rate of 25 to
29 percent and three to four generations a year, against one or less for
the host, _Evania appendigaster_ in the areas where it is established is
a valuable control agent.
The use of specific egg parasites to control cockroaches has not been
attempted extensively. Cros (1942) liberated a species of _Tetrastichus_
(=_Eulophus_ sp.; see p. 254) in his home in Algeria to control the
oriental cockroach; as far as we know, he did not report the parasite's
effectiveness in reducing the cockroach population. According to
Zimmerman (1948) _Comperia merceti_, when accidentally imported,
practically wiped out _Supella supellectilium_ in parts of Hawaii; he
claimed to have controlled the brown-banded cockroach in a store
building with this parasite. In some parts of Honolulu, almost 100
percent of the oöthecae of this cockroach were parasitized (Zimmerman,
1944). We (1954b) ran some simulated field tests in which we liberated
_Tetrastichus hagenowii_ in rooms artificially seeded with oöthecae;
from 28 to 83 percent of American cockroach oöthecae and 56 percent of
oriental cockroach oöthecae were parasitized during these tests.
_Evania appendigaster_ was introduced from Hawaii into Canton Island in
1940 against _Periplaneta americana_, and it has become established
(Dumbleton, 1957). This parasite was also successfully introduced into
Samoa (Dumbleton, 1957).
_Cockroach-hunting wasps._--An earnest attempt has been made to
establish in Hawaii wasps that prey on cockroaches. Just how effective
these wasps are in controlling cockroaches is still unknown. _Dolichurus
stantoni_ was introduced from the Philippines in 1917 and spread to
several of the Islands (Swezey, 1920, 1921; Williams, 1944). Bridwell
(1920) stated that as a result of this introduction there was a great
decrease in cockroaches of the genus "_Phyllodromia_." A number of
_Podium haematogastrum_ from Brazil were liberated in Honolulu
(Williams, 1925) but did not become established (Williams, 1928). The
effectiveness of _Podium_ was questioned by Williams (1928) who observed
that _Podium_ "destroyed innumerable Blattidae, which nonetheless
swarmed in their neighborhood, and I must confess from my observations
on the various cockroach-hunting wasps that the blattid more than holds
its own alongside its enemy."
Introductions of _Ampulex_ have proved more successful. _Ampulex
canaliculata_ was introduced into Hawaii from the United States
(Williams, 1928a, 1929). Williams also introduced _A. compressa_ into
Hawaii in 1940, and the species was reared in large numbers for
distribution (Pemberton, 1942). _A. compressa_ has since become
established on most of the Islands (Pemberton, 1945a, 1947; Williams,
1946; Van Zwaluwenburg, 1950). The thousand of _A. compressa_ now found
in the Hawaiian Islands are all descendants of three wasps captured in
Noumea, New Caledonia (Williams, 1944). According to Williams (1941),
the number of cockroaches was noticeably reduced at the University of
Hawaii poultry farm, where some _A. compressa_ were released. Pemberton
(1953) believed that this wasp has become sufficiently abundant to be of
definite value. Simmonds (1941) recommended importing _A. compressa_
into Fiji for cockroach control. _A. compressa_ was introduced from
Hawaii into Guam in 1954 against _Periplaneta americana_ and into the
Cook Islands in 1955 against _Periplaneta_ spp.; it is not yet known
whether the parasite became established in either place (Dumbleton,
1957).
VERTEBRATES
... on conserve avec soin les crapauds dans les maisons, et que les
dames les tolèrent, même sous leurs robes, en raison de leurs
continuels services, car ils se promènent sans cesse à la recherche
des Kakerlacs.
GIRARD (1877)
_Toads._--_Bufo marinus_ was first introduced into Puerto Rico from
Barbados in 1920 to reduce several major insect pests including
cockroaches (Leonard, 1933). It was introduced from Puerto Rico into
Hawaii by C. E. Pemberton in 1932 where it rapidly became established;
it has since been distributed throughout the Pacific area. _B. marinus_
is one of the world's largest toads; it attains a body length (exclusive
of the hind legs) of 7 to 9 inches (Oliver, 1949) and has been kept
alive for more than 11 years in captivity (Pemberton, 1945). Alicata
(1938) placed giant toads in a fenced area in Hawaii containing an
infestation of _Pycnoscelus surinamensis_; after 24 hours the toads were
dissected and each was found to have eaten from 11 to 25 cockroaches.
Illingworth (1941) found that 40 to 90 percent of 53 stools of this toad
in Hawaii contained remains of _P. surinamensis_. Alicata (1947)
recommended the maintenance of _B. marinus_ in poultry yards to reduce
the population of _P. surinamensis_, the vector of the chicken eye worm.
Toads have also been recommended for controlling cockroaches in houses
(Meech, 1889; Sweetman, 1936). Girard (1877) cited a note in a French
newspaper which stated that toads were kept in houses in Cuba to control
the American cockroach.
_Tree frogs._--Tree frogs enclosed in a room overnight were said to
effectively clear it of cockroaches (Marlatt, 1915); on sugar
plantations in Australia, these amphibians were encouraged in houses and
kept as pets because they hunted and devoured large brown cockroaches
(Froggatt, 1906).
_Birds._--In Guadeloupe, Dutertre (1654) claimed that all the fowls of
the country were fond of small cockroaches and lived on practically
nothing else. In Hawaii (Zimmerman, 1948) and in the Lesser Antilles
(Ballou, 1912) cockroaches are eaten by poultry whenever the birds can
find them. In Puerto Rico, Wetmore (1916) stated that owls kept in
houses feed extensively on cockroaches; the stomach of one owl which had
been kept in a native house was filled entirely with cockroaches. In
British Guiana, Beebe (1925) found that cockroaches were eaten by 27
species of birds.
_Reptiles._--H. (1800) claimed that two lizards cleared his house of the
"true brown cockroach" and suggested that lizards be used for cockroach
control because the reptiles are docile and harmless. On Arno Atoll
geckos and night-feeding skinks eat large numbers of cockroaches
(Usinger and La Rivers, 1953). According to Wolcott (1924) the number of
cockroaches eaten by lizards is surprisingly large considering the
nocturnal habits of these insects. Beebe (1925a) kept geckos in a
bungalow to help control _Periplaneta_ and _Pycnoscelus_.
_Mammalia._--Cowan (1865) stated that in England hedgehogs were often
kept domesticated in kitchens to destroy cockroaches. This writer also
stated that a lemur was kept on board ship to destroy cockroaches.
Large numbers of the American and Australian cockroaches were eaten by
the mongoose in Hawaii (Perkins, 1913).
ACKNOWLEDGMENTS
We would have been unable to complete this review without the help of
many people who have generously given us their time and the benefit of
their special experience. We are exceedingly grateful to these
individuals for they have contributed much to whatever merit this work
possesses; we alone are responsible for the deficiencies and
inaccuracies that remain in the text.
Dr. A. B. Gurney, Entomology Research Division, U. S. Department of
Agriculture, and J. A. G. Rehn, Academy of Natural Sciences of
Philadelphia, have given us much help and advice throughout the
preparation of this monograph. Both have patiently answered our many
queries, and Mr. Rehn allowed us free access to his large collection of
cockroach literature. We are especially pleased to thank them for their
many favors.
Many persons have determined at our request the identity of specific
organisms. These individuals are cited in the text and to them we extend
our thanks. We thank our colleagues, cited in the text, who have made
their unpublished observations available to us. We also thank the
individuals and organizations for the use of photographs and/or drawings
for which they are credited in the accompanying legends.
We thank the following individuals for supplying us with living
specimens of the species indicated: T. Campbell, Commonwealth Scientific
and Industrial Research Organization, Canberra, New South Wales
(_Panesthia australis_); Dr. L. R. Cleveland, Harvard University
(_Cryptocercus punctulatus_); Dr. N. T. Davis, University of Connecticut
(_Byrsotria fumigata_); Dr. F. Englemann, Albert Einstein Medical School
(_Gromphadorhina portentosa_); Dr. F. A. Lawson, Kansas State College
(_Comperia merceti_); Dr. K. D. Roeder, Tufts University (_Hierodula
tenuidentata_ (?)); Dr. E. O. Wilson, Harvard University (_Ischnoptera
deropeltiformis_).
We thank the following individuals for checking the taxonomy of the
following organisms or for reading the indicated sections of the
manuscript: _Fungi._--Dr. R. K. Benjamin, University of California; Dr.
E. G. Simmons, Quartermaster Research Laboratories. _Protozoa._--Dr. R.
R. Kudo, Professor Emeritus, University of Illinois. _Helminths._--Mrs.
May Belle Chitwood and Dr. J. T. Lucker, U. S. Department of
Agriculture. _Centipedes and whipscorpions._--Dr. R. E. Crabill, Jr., U.
S. National Museum. _Scorpions._--Dr. J. L. Cloudsley-Thompson,
University of London; Dr. R. E. Crabill, Jr. _Spiders._--Dr. B. J.
Kaston, University of Connecticut; Dr. R. E. Crabill, Jr. _Mites._--Dr.
J. H. Camin, Chicago Academy of Sciences; Dr. E. W. Baker, U. S.
National Museum. _Cockroaches._--Dr. A. B. Gurney and J. A. G. Rehn.
_Ants._--Dr. W. L. Brown, Jr., Museum of Comparative Zoology, Harvard
College. _Hymenopterous parasites of cockroach eggs._--Dr. B. D. Burks,
U. S. National Museum; Dr. H. K. Townes, University of Michigan.
_Cockroach-hunting wasps._--Dr. K. V. Krombein, U. S. National Museum.
_Lepidoptera._--Dr. J. F. G. Clarke, U. S. National Museum; Dr. E. L.
Todd, U. S. Department of Agriculture. _Miscellaneous insects._--Dr. R.
S. Beal, Dr. A. B. Gurney, C. W. Sabrosky, Dr. R. I. Sailer, and J. T.
Spilman, U. S. National Museum. _Fishes._--Dr. L. P. Schultz, U. S.
National Museum. _Amphibians and reptiles._--Dr. Doris M. Cochran, U. S.
National Museum. _Birds._--Dr. Herbert Friedmann, U. S. National Museum.
_Mammals._--Dr. D. H. Johnson, U. S. National Museum.
We thank the following individuals for reading the entire manuscript: J.
A. G. Rehn; Dr. A. B. Gurney; Maj. Gordon Field, U. S. Army; and Dr. H.
L. Sweetman, University of Massachusetts. The monograph has profited by
the friendly criticism of these entomologists.
We thank Dr. R. A. Howard, Harvard University, for checking lists of
plant names; Mrs. Maria E. W. Torok, formerly of the Quartermaster
Technical Library, for assistance in obtaining obscure literature; Miss
Louise Bercaw, U. S. Department of Agriculture Library, for identifying
the journal containing the paper by Vlasov and Miram; the individuals
who translated foreign language articles, for which they are credited in
the bibliography; and Miss G. Lillian Fede, Quartermaster Research
Laboratories, for typing the manuscript.
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