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SMITHSONIAN
MISCELLANEOUS COLLECTIONS

VOL. 141

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

(Pusiication 4470)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1961

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

ADVERTISEMENT

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

LEONARD CARMICHAEL,
Secretary, Smithsonian Institution.

(iii)

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 141 (WHOLE VOLUME)

THe BIOTIC: ASSOCIATIONS: OF
COCKROACHIES

(WitTH 37 PLATES)

By
LOUIS M. ROTH
AND

EDWIN R. WILLIS
Pioneering Research Division, United States Army

Quartermaster Research and Engineering Center
Natick, Mass.

(PuBLICATION 4422)

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

SMITHSONIAN MISCELLANEOUS COLLECTIONS VO aay Peed

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

SMITHSONIAN MISCELLANEOUS COLLECTIONS
VOLUME 141 (WHOLE VOLUME)

CRE PIGTIC ASSOCIATIONS OF
COCKROACHES

(Wit 37 PLATES)

By
LOUIS M. ROTH
AND

EDWIN R. WILLIS

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

(Pusiication 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 amaze-
ment 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 prede-
cessors, 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 in-
formation 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 in-
formation 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 coun-
tries, 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.

iii

iv SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

The present work is such a compilation, for it assembles what has
been gleaned from approximately 1,700 sources, including correspond-
ence with a large number of other workers. Original observations
during some eight years of concentrated effort in U. S. Army Quar-
termaster 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 in-
sects 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 re-
garded only as disagreeable pests; many species appear to be im-
portant, 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 con-
clusions 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 re-
view 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 Smith-
sonian 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

[Day eenordil Ueto Cin MnAinn cnbiolo bia oigob ogo UOOe DES DODUISSEe Hon Got Dotna iii
ep lntroductOomeusn soci serie ee cto et ers ieiscle te ele cieteiercs c c)steleterel sce I
le hvoratcal be ee hen yar b tomo Om OOOO DOCS SUGS aoe bone Ne oetos 2
IMetOdS eyes Sa aioe ae re et cle Sebati stators diets aceite tous 4
ate ODIC SA Re ae ree shoe arco aialaia eee crater e leche e cielo sists 5
TIS Er AHONS) Wa rare eRe Ae wich een re aa ae chet feat aie 7

Te eSpeciesi-of (cockroachess)avsd Sac Secioe nes ho boats clon ele es rie oe 7
IE Keolocical), relationships) fascias eels hats Wolens oreets pire ll ataimrniatet ss es 14
(Cave: Habitats). 3 cave cassie ow c.ae Hee On nee eee eters an eta eres eer 16
CGaveriicolousecockroachestaeeeeeee eee omer: 17
@ockroaches) from buLrOwS ei. \o sale Croletele sate! slarevstereeel stove ols 6 0)2 23

Desert: Hab itatsw icin cveicis si ciccuerot ie share SSR he are oP eo olle ata 25
DESertrCOCKMOAGCIES s aaa yt che tear te Ee P octane eater ets ele welts 27

Aveatic, Hapitats) sacs an wor’ 2 hat basa arora, Heer eae Me eiatein, e's ole 30
Amaplitbious), COCKEOaCHES y 49 a2 «ic Poneto Via elevabetetand ele) Hale aca we. 31

Outdoors habitats yc ey eo A heer cae atererait ue el Leaors eltay averaie 33
Cockroaches from) outdoor shabitats2e/je-eiesin-) cee eles ees oes 35
Stauctinal habitats escrito crate a tere crete retererc eet trevess ete 70
Wand=hasedistractunesu ste itee creel tec ce ereroterstetotey actors tots reks 73
Cockroaches associated with land-based structures....... 74

SAIS oh ccio ia wich aie Siere wis’ aaa > dialele MRNAS aeiaReTA, emia nals aa ese slaiaars &2
Cockroaches) associated: withhishipstetacciast «tie seit score ole 85

AICTE He oi Sa horas RTA ae ore a el Naha tee onahc matee tan eR P ana arcs 87
Cockroaches associated) withyaincralte. sce ost eee sle- 88

TiVen Classification ot uchewassOciatlOlS\tem aeicterasiniiote eters aeteretieieler =e QI
Ne INMst ITU LE STIR cits) 2 aiarala a ere cn easier etaels Shaver staiaa tis a mia wlan aehereRe a 96
RACHEL OLUS mavater terete calexorc ton srateuerrs ld iro avorer otal Mar clever ha sone siahet Aiatonetetee 96
Cockroaches in which bacteroids have been found........... 99

RACER Teas ere eee eee eer ea Seen rat aeTn ielsioharsrels stayarelerese 100

IP KOLOZOA aise Tee ee ee eat atalls oh Paterotete sear aiehel sheto-tasanehenctsiaions oye 101

Nise Virisesr associated, wathucocis Oa CheSmre iin aieiciraiieiiatc i tetorelele = elare 103
WIL Bacteria: associated wath) cocktoachesw.-\-isc0 seas 22 4 «cites = 104
NAlMnE ned alicaveAaStSeatmerercmieier rami ravereietcnectolskellslcrororshoveretegsloceie oretaletstats) = 127
Bunei associated, with (cOckraaches.. i... seen cane aeeccesaces 129

IX. Higher plants associated with cockroaches.............+++++04- 139
Damage to plants by cockroaches............+.eeeeeeeeeeeee 162

xe Protozoa associated) with’ Cockroaches: 1.5... ses 0 cle ele cele = = 1606
XI. Helminths associated with cockroaches.............-eeeeeeeeees 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.............seeeeeeees 210
DAtNGHaTii nee Ree a etete erento retell sei ayoroiseustoierstaicretene(oleieysucrausiefatersl's 211
Chilopoda:. \ovenesccess deen e EP Wemete Ine Arai Sart nu ate ce diese weatere he 222

vi SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI
Page

SEC tay ree cise sarees ate rar ere cia ralene oie losaycuetete ler hrs clare) satel ereeeteencsT stare terete 224
EVymienopterat ce. dice seh o's aires naive treelovele, stelraretom tates ean tarss 234

Predators and parasites of cockroach eggs............... 234

Host selection *by, eee: parasites sie lack oes «mtv ce 254
Cockroach-hunting wasps: ayesieicleisioriclsterseietielsieierciatieteye 255

Ants predaceous, On: COCKrOACHES oarcia ere) aye ctaie) Veieisiaiss sey ile 266

XIII. Vertebrata associated with cockroaches..............0- eee Colt SOS:
IPISCESH ciataitere a tlateiers, steterelsicveleraleranes asin: wet Shay elisa btoke etatensterarar e.evaters 268

7 -Xvanh 0) 061 YC PR eI A AS CO aAEES Oe OG oC Dob ao so lSocmpods 269

esata Tags Seatapeconers seine vate’ gusce avails: she! crevasse coheree tote re Chem systetenonerNeneke ctereratts 272

DANES sce ia ots esoaverte a: o evs eis.e,ev nkayevereefel ors /skeie eveveveraley sxcvepe te toueied enue rabereiceone 276
Mammatnial lita sis)c s sseveverss auc alesse oo wsevovelevayesiensoterclate ciare choke peeyeealerainee che 283

XIV. Checklist of cockroaches and symbiotic associates.............6. 290
DEV. ACOCKCGACHES, AS. COMMENSAIS << 5:.)s.c, <cs.cie sino clniers olaalerea arte aiece alors 308
flosts) of commensal ‘cockroaches. 14+ a atetieeioereers sites 309

Checklist of commensal cockroaches with their hosts......... 313

Obscure vassocsiations, a 4. dieda.cc esereioknararee eee i acals onan

Vel Cockroaches| vas) predators sje sciecieee ee ee eee events 317
Interspecies -PKECATIOM {s,.</c)s)s/oco1s) oiepstctovauctarentarsreteteererane otto craverene 317
Intraspecies predation’ cara gac dee seiea Tee eine eee 320

DOV Associations) aniong cockroaches 1s. sanen pomereine aloes 6 eine 322
Bamilial associationsisncm en seeds eee ee eee 323
GKEGAPIOUSHESS:: 5 a5)5)5;5 4's) asa wins dhevora saber Cena a aeRO Oe Sele eR 328
Intraspecies fighting, aisciep« sn occ ela ee aera ee eee eine 334
Interspecies compatibility i. .sm see ee oon iene 335
Interspecies: antagyonism 3.01 cisacia da coctmadeeeieen ee ee 339

XVIII. Defense of cockroaches against predators.............2+eeeeees 341
EX. he, biological ‘control. of cockroaches iiss sesame ewan nese 346
Invertebrates: \\s.35:/tadid.eweenrcnee taste rate ee oeeie eee & 347
SVICHEEI TA ECS fi ics, :'s(aivicia che, siete RSA Wee PRE doy senate tere Se sccle ae!) BO
BAUCLETIO WLC ETACTIES oc 5-95 io .5: 4:82, <; sos) b'5/- 6) os) caro) al Oren Toe ke meee he es 352
BR CREE CIIC OS) 5) aa) ohs: 6/6 oral nit e/eiosoro;e.acuinj sve, o\ ev aso) ofs\ syarard Cloke aR RET IeT: etnte 354
Pndex joes strere cle aloiata s]c?ehotalo evsje.s aisle ateractevelelaletatenn atte stenatateneReralevelicrer aie ie letar ais 439

THE BIOTIC ASSOCIATIONS OF
COCKROACHES:

By Louis M. Rory anv Epwin R. WILL Is ?

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 cock-
roaches 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 associa-
tions 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
‘nferred 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 satis-

1 Part of the cost of publication of this monograph was borne by the Office of
Naval Research, Department of the Navy (through the American Institute of
Biological Sciences), and by the Quartermaster Research and Engineering Cen-
ter, Department of the Army.

2 Present address of both authors, Central Research Laboratories, United Fruit
Co., Upland Road, Norwood, Mass.

SMITHSONIAN MISCELLANEOUS COLLECTIONS, VOL. 141 (WHOLE VOL.)

2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. TAT

factory 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 Orthoptéeres reviewed
much of the literature on cockroaches, but of the many biotic associa-
tions that exist he discussed only the commensal cockroaches, gregari-
ousness, 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 surinamensts) 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. Lin-
stow (1878, 1889) recorded 14 species of helminths from cockroaches.
Van Zwaluwenburg (1928) listed 33 names of roundworms which

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 3

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 rela-
tionships 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 im-
portant 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
4O 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 cock-
roaches. Of the arthropods there are about 2 species of scorpions,
4 spiders, 15 mites, 4 centipedes, and go insects. Of vertebrates there
are 4 species of fish, 16 amphibians, 12 reptiles, 20 birds, and 27 mam-
mals. Besides these there are many records of experimental associa-
tions 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

4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

synonyms), I 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 cock-
roaches 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 cock-
roaches are listed by the currently accepted name. Unidentified cock-
roaches 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 °
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

3 Names of organisms preceded by an asterisk (*) are known or suspected
pathogens of vertebrates. These records were presented with annotations in our
1957a paper on the medical and veterinary importance of cockroaches. For that
reason the annotations have not been repeated herein, although the records have
been included to make the listing of the biotic associates of cockroaches sub-
stantially complete.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 5

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 ap-
parently 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 cock-
roaches have been changed. Although it would have been compara-
tively simple to list the names as they appeared in the original ref-
erences, 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. How-
ever, 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 IJ. 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. Un-
fortunately, 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 rela-
tionships has bately 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

6 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI

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 para-
sites 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 informa-
tive 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 odthecae 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 (thig-
motaxis? 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 fre-
quently 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 7

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.

ih) SPEGibS, 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 illus-
trations 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) [Ceratinop-
tera 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 rec-
ords = Ceratinoptera diaphana
R. and H.; Hebard (1917) ]
vegeta Rehn
ypsilon Princis

Allacta similis (Saussure) [Phyllo-
dromia obtusata Brunner; Zim-
merman (1948) ]

Alluaudellina cavernicola (Shelford)
[Alluaudella cavernicola Shel-
ford; 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 [Acantho-
gyna deplanata Chopard;
Princis (1957) ]

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

Apteroblatia perplexa Shelford

Archiblatia hoeventi Vollenhoven

Archimandrita marmorata (Stoll)

tessellata Rehn

Arenivaga apacha (Saussure)

bolliana (Saussure)

erratica (Rehn)

floridensis Caudell

grata Hebard

rosem (Brancsik) [Heterogamodes
rosem; Bei-Bienko (1950).
“Polygamia”’ roseni is undoubt-
edly an erroneous citation of
Polyphaga roseni, as there is no
genus Polygamia (Gurney, per-
sonal communication, 1957) ].

tonkawa Hebard

Aristiger histrio (Burmeister) [Plu-
miger histrio (Burm.) ; Bruijn-
ing (1048). Hemithyrsocera
histrio Burm.; Hebard (10929) ]

Aspiduchus boriquen J. W. H. Rehn
[In Puerto Rico reccrds=
Aspiduchus deplanatus R. and
Hee Rehn, Jo We i. Crosta)]

8 SMITHSONIAN MISCELLANEOUS COLLECTIONS

cavernicola J. W. H. Rehn
deplanatus (Saussure)
Attaphila aptera Bolivar
bergi Bolivar
flava Gurney
fungicola Wheeler
schuppi Bolivar
sexdentis Bolivar
Atticola mortomi Bolivar
Audreia bromeliadarum Caudell
jamaicana Rehn and Hebard
Balta godeffroyt (Shelford)
patula (Walker)
platysoma (Walker)
teryx platysoma
Hebard (1943) ]
quadricaudata Hebard
scripta (Shelford)
torresiana Hebard
verticalis Hebard

[Temnop-
(Walker) ;

Bantua stigmosa (Krauss) [Dero-
calymma__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; He-
bard (1917) ]

(Shelfordella) lateralis (Walker)
[Shelfordella tartara (Saus-
sure); Princis (1957). Peri-
planeta tartara Saussure; Bei-
Bienko (1950) }

Blattella germanica (Linnaeus) (pls.
5, A, B; 31, F) [Blatella ger-
manica; Gurney (1952). Phyillo-
dromia germanica; Hebard
(1917). Ectobius germanicus;
Gurney (personal communica-
tion, 1957) ]

VOL. I4I

hwmbertiana (Saussure) [Blatta
humbertiana; Phyllodromia
humbertiana; Hebard (1929) ]
lituricollis (Walker) (fig. 7, A)
[Blattella bisignata (Brun-
ner); Bei-Bienko (1950) ]
schubotzi Shelford
vaga Hebard (pl. 5, C, D)
Buboblatta armata (Caudell) [Latin-
dia 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 (Beativois); Rehn
and Hebard (1927) J
hylaca Rehn
imitans Hebard
insularis (Walker)
landalet Rehn and Hebard
lutea lutea (Saussure and Zehnt-
ner)
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
parisht Rehn
specilliger Hebard
texensis Saussure and Zehntner
[Chorisoneura plocea Rehn;
Rehn and Hebard (1916) }
translucida (Saussure)

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 9

Choristima sp.
Choristimodes sp.
Chromatonotus infuscatus (Brunner)
notatus (Brunner)
Compsodes schwarzt (Caudell)
Comptolampra liturata (Serville)
[Compsolampra liturata; Comp-
tolampra 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 (Bolivar) (pl. 9)
[Determined by Dr. A. B.
Gurney from photographs. ]
Cyrtotria capucina (Gerstaecker)
Dendroblatta sobrina Rehn
Derocalymma cruralis (Stal) [Ho-
malodemas cruralis (Stal);
Gurney (personal communica-
tion, 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 (Ser-
ville) ; Zimmerman (1948) ]
Dryadoblatia scotti (Shelford) [Ho-
malopteryx scottt Shelford;
Rehn (1930) ]
Ectobius africanus Saussure
albicinctus (Brunner)
duskei Adelung
lapponicus (Linnaeus) [Ectobius
perspicillaris Herbst, as used by
Lucas (1920) ; Blair (1934) ]
lucidus Heb.
nicaeensis (Brisout)

pallidus (Olivier) (pls. 7, C; 20,
A) [Ectobius lividus (Fabri-
cius); Ectobius livens (Tur-
ton); Kevan (1952); Princis
(in Roth and Willis, 1957) ]

panzeri Stephens [Ectobius eri-
cetorum (Wesmael); Ramme
(1923) ]

panzeri var. nigripes Stephens

semenovi Bei-Bienko

sylvester (Poda) [Ectobius syl-
vestris (Poda) ; Ramme (1951) ]

tadzihicus Bei-Bienko

vittiventer (Costa) [Ectobius vit-
tiventris (Costa); Ramme
(1951) ]

Ellipsidion Saussure [Apolyta Brun-
ner; Hebard (1043) ]

affine Hebard

australe Saussure [Ellipsidion pel-

lucidum (Brunner); Hebard
(1943) ]

bicolor (Tepper)

simulans Hebard

variegatum (Fabricius) [Ellip-

sidion aurantium Saussure; He-
bard (1943) ]
Eptlampra 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
zwheeleri Rehn
Spx (ait. 7 iba C)
Eremoblatta subdiaphana (Scudder)
Ergaula capensis (Saussure) [Dys-
cologamia capensis Saussure;
Dyscologamia wollastoni Kirby ;
Princis (1957) ]
scarabaeoides Walker [Dyscolo-
gamia_ _ piolosa (Walker) ;
Princis (1957). Parapolyphaga

10 SMITHSONIAN MISCELLANEOUS COLLECTIONS

erectipilis Chopard; Princis
(1950). Dyscologamia chopardt
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 de-
castigmata WHebard; Princis
(1959) ]
Eurycotis bananae Bei-Bienko
biolleyi Rehn [Eurycotis carbo-
naria Biolley; Rehn (1918) ]
caraibea (Bolivar)
decipiens (Kirby)
dimidiata (Bolivar)
ferrum-equinum Rehn and Hebard
floridana (Walker) (pl. 11)
[Platyzosteria ingens Scudder ;
Platyzosteria sabalianus Scudder
(and hence, by inference, Eury-
cotis 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 (Stal) [Blatta
notulata Stal; 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) J

tristis Hanitsch

VOL. I41

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

Hemiblabera brunneri (Saussure)

Henicotyle antillarum (Brunner)

Heterogamodes kriigeri (Salfi)

rugosa (Schulthess)
Holocompsa asteca (Saussure)
cyanea (Burmeister )
fulva (Burmeister )
metallica Rehn and Hebard
nitidula (Fabricius)
gzapoteca Saussure
Hololampra bivittata (Brullé)
chavesi (Bolivar)
maculata (Schreber) [Aphlebia
maculata Schreber; Harz
(1957) ; Gurney (personal com-
munication, 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 dero-
peltiformis 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 Ger-
staecker ; Rehn (10932) ]

remota Hebard

Lamproblatta albipalpus Hebard

meridionalis (Brunner)

Latiblattella chichimeca (Saussure and
Zehntner) [Blattella chichimeca
S. and Z.; Hebard (1932) ]

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS II

lucifrons Hebard
rehni Hebard

vitrea (Brunner)
sapoteca (Saussure)

Leucophaea maderae (Fabricius) (pl.
13) [Rhyparobia maderae; He-
bard (1917). Panchlora ma-
derae; Kirby (1904). Very
probably “Blaberus” maderae is
a careless reference to this spe-
cies; Gurney (personal com-
munication, 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 (Bolivar)
[Loboptera dimidiatipes (Boli-
var); Princis (1957a). Lobop-
tera sakalava (Saussure) ; He-
bard (1933a). Loboptera ex-
tranea Perkins; Hebard (1922).
Princis (1957a) in erecting Lo-
bopterella pointed out that only
the nontypical variety of saka-
lava is identical with dimidia-
tipes.]
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 (Stal)

Nauclidas nigra (Brunner)
blatta nigra Brunner;
(1930) ]

Nauphoeta cinerea (Olivier) (pl. 14)
[Nauphoeta bivittata Burmeis-
ter ; Zimmerman (1948) ]

flexivitta (Walker) [Nauphoeta

[Poro-
Rehn

braszae (Bolivar) ; 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 Cau-
dell (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) J
Nesomylacris cubensis Rehn and He-
bard
relica Rehn and Hebard
Nocticola bolivarit Chopard
caeca Bolivar
decaryt Chopard
simom Bolivar
sinensis Silvestri
termitophila Silvestri

I2 SMITHSONIAN MISCELLANEOUS COLLECTIONS

Nothoblatta wasmanni (Bolivar)
Notolampra antillarum Shelford
Nyctibora azteca Saussure and Zehnt-
ner
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) [Pan-
chlora cubensis Saussure; Gur-

ney (1055). 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; He-
bard (1917) ]
caudelli Hebard [99 of Ischnoptera
insolita R. and H.; Ischnoptera
uhleriana fulvescens S. and Z.
(in part) ; Hebard (1917) J
desertae (Rehn and Hebard) (do
of Ischnoptera insolita R. and
H.; Hebard (1917) ]

VOL. I4I

divisa (Saussure and Zehntner)
[Ischnoptera divisa S. and Z.;
Hebard (1917) ]
fulvescens (Saussure and Zehnt-
ner) [Ischnoptera uhleriana
fulvescens S. and Z. (in part) ;
Hebard (1917) ]
lata (Brunner) [I[schnoptera
couloniana R. and H. (not Saus-
sure) ; 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 Saus-
sure; Hebard (1917) ]
virginica (Brunner) (pls. 17, B;
27, A; 33, C; fig. 6) [Ischnop-
tera 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 ameri-
cana; Blatta americana L.; He-
bard (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
Phliebonotus pallens (Serville)
Pholadoblatta inusitata (Rehn)
Phorticolea boliviae Caudell
testacea Bolivar
“Phyllodromia” treitliana Werner

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 13

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) [Poly-

sosteria analis Saussure; Kirby
(1904) ]

armaia Tepper

bifida (Saussure)

castanea (Brunner)

novae seelandiae (Brunner) (pl.
23) [Periplaneta fortipes
Walker; Shelford (1912) ; Pla-
tyzosteria novae-zealandiae |

scabra (Brunner)

Plectoptera dorsalis (Burmeister)
infulata (Rehn and Hebard)
lacerna Rehn and Hebard
perscita Rehn and Hebard
poeyt (Saussure) [Plectoptera

floridana Hebard ; Rehn and He-
bard (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 (per-
sonal communication, 1957).
“Polygamia”’ aegyptiaca; ac-
cording to Gurney (p. c.), there
is no genus Polygamia and al-
most surely the reference is to
Polyphaga aegyftiaca.]
indica Walker [Polyphaga pellu-
cida (Redtenbacher); Princis
(1957) ]
saussuret (Dohrn)
Polyzosteria limbata Burmeister
melanaria (Erichson)
Pseudoderopeltis aethiopica (Saus-
sure) [Blatta aethiopica Saus-
sure; Gurney (personal com-
munication, 1957) ]

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

laticorms Perty
septentrionalis Hebard
Pseudophoraspis nebulosa (Burmeis-

ter)
Pycnoscelus niger (Brunner)
striatus (Kirby) [Leucophaea

striata Kirby ; Gurney (personal
communication, 1957) ]
surinamensis (Linnaeus) (pl. 24)
[Leucophaea surinamensis (L.) ;
Hebard (1917). Blatia melano-
cephala 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 circwmducta (Walker) ;
Gurney (personal communica-
tion, 1057) ]
longiuscula (Walker) [Escala
longiuscula (Walker) ; Gurney
(personal communication, 1957) ]
Salganea morio (Burmeister)
Sibylloblatta panesthoides (Walker )
Simblerastes jamaicanus Rehn and
Hebard
Spelaeoblatta gestrot Bolivar
Sphecophila polybiarum Shelford
ravana Fernando
termitium Shelford
Steleopyga (?) sinensis Walker [Dr.
Gurney (personal communica-
tion, 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 rep-
resents. ]
Stictolampra
(Walker)
Styphon bakeri Rehn
Supella hottentotta (Saussure)
supellectilium (Serville) (pls. 25;
30, B-E; 31, A-E) [Phyllo-

buqueti concinula

I4 SMITHSONIAN MISCELLANEOUS COLLECTIONS

dromia supellectilium (Serv.) ;
Bei-Bienko (1950) ]
Symploce breviramis (Hanitsch)

cavernicola (Shelford) [Ischnop-
tera cavernicola (Shelford) ;
Phyllodromia nigrocincta Cho-
pard; Hebard (1929) ]

curta Hanitsch

flagellata Hebard

hospes (Perkins) [Symploce lita
Hebard; Hebard (1922) ]

jamaicana (Rehn)

kevani Chopard

parenthesis (Gerstaecker) [Phyl-

VOL. I4I

ruficollis (Fabricius) [Symploce
bilabiata Rehn and Hebard;
Princis (19492) ]
Tartaroblatta karatavica Bei-Bienko
Temnopteryx obliquetruncata Chopard
phalerata (Saussure)
Theganopteryx straminea Chopard
Therea nuptialis (Gerstaecker) [Co-
rydia nuptialis Gerstaecker ;
Princis (1950) ]
Tivia australica Princis
brunnea (Chopard)
fulva (Burmeister)
macracantha Chopard
obscura (Chopard)
Typhloblatta caeca (Chopard) [Spe-

lodromia parenthesis Ger- lacoblatta caeca Chopard; Cho-
staecker ; Rehn (1932) ] pard (1924b) ]

remyt (Hanitsch) [Ischnoptera Typhloblattodes madecassus Chopard
remyi _Hanitsch; Chopard Xestoblatta festae (Griffini)
(1938) ] immaculata Hebard

i; ECOLOGICAL RELATIONSHIPS

The ecology of extinct cockroaches is necessarily a highly specu-
lative 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 drift-
ing plant material to places where permanent deposits were accu-
mulating.

Some species of fossil cockroaches have long, well-developed ovi-
positors, very unlike present-day cockroaches whose ovipositors are
small and nonprotruding. Brongniart (1889) and Zalesskii (1939,
1953) have suggested that certain Permian and Carboniferous cock-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 15

roaches with long ovipositors may have inserted their eggs singly
into trees and other plants, rather than protecting the eggs with an
odtheca. However, Laurentiaux (1951), although conceding the
possibility of egg laying in vegetable material, suggested that ovi-
position 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 bio-
logical 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 cock-
roach 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 in-
cluded in ecological studies of the Orthoptera by Hubbell (1922),
Strohecker (1937), and Cantrall (1943). Fifteen species of cock-
roaches 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 arthro-
pods 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 clas-
sifications serve admirably to relate various species of cockroaches
within common bounds, but fail to give an integrated account of the

16 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 in-
cluded a checklist (p. 290) which serves as a convenient index to cer-
tain 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 ad-
verse 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 Peri-
planeta americana) may have accompanied man into caves and re-
mained 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 estab-
lished. 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) Troglox-
enes: Cockroaches that occur in caves in a sporadic fashion (the
domiciliary cockroaches and accidentals such as Ectobius and Hetero-
gamodes). (2) Troglophiles: Cockroaches found habitually in
caves (Symploce, Periplaneta cavernicola). (3) Guanobies: Cock-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 17

roaches that live in the guano of cavernicolous vertebrates (Gyna,
Acanthogyna, Dyscologamia, Pycnoscelus). (4) Troglobies: Cock-
roaches that apparently cannot live outside of caves and which show
very marked adaptive characters (Alluaudellina, Nocticola, Spelaeo-
blatta, Typhloblatta). For complete discussions of these groups in-
cluding 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 in-
habit mines in North America, but we have found no such records.
Have cockroaches been ignored in faunal 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 char-
acters. Collected in moist sand on floor of a sandstone grotto in-

18 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

habited by bats (Chopard, 1927, 19502). 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., 19514).

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 19

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 cavernico-
lous 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.—F ound 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, 1949).
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).

20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Gyna tristis

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

Heterogamodes kriigeri

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.—Bolivar (1892).

Nocticola decaryi

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

Nocticola simoni

Philip pine Islands.—Bolivar (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 cock-
roaches 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 en-
trance by man (Chopard, 1945, 19492).

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 21

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 feces (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 par-
ticularly 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, I immature male
under stone in complete darkness (Chopard, 1924b).

22 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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.—F ound 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 charac-
teristics of adaptation to a life in darkness (Bolivar, 1897; Annan-
dale, 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 Sara-
wak 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 23

Tivia macracantha
Belgian Congo.—A troglophile without well-marked adaptive char-
acters (Chopard, 1950a). At Haut-Katanga, troglophile and guano-
phile (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 ap-
pendages (Chopard, 1945).

Typhloblattodes madecassus
Madagascar.—Unpigmented integument and reduced eyes (Cho-
pard, 1945).
Xestoblatta immaculata

Panama.—Found under rocks on guano-covered floor of the Chili-
brillo 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;
19438).
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).

24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Pallastomys
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 bur-
rows 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 25

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 ab-
stracted 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 Polyphagi-
nae 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-inhabit-
ing 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 ex-
tremely 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 prob-

26 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

lem 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 bol-
liana 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 I to 15 percent during
the day and 15 to 40 percent at night ; but inside the burrows the rela-
tive 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 micro-
climates obtainable. Presumably whatever food these insects eat pro-
vides 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 informa-
tion 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, re-
lated 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 arbi-
trarily 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 27

DESERT COCKROACHES
Agis orientalis
Northern Kenya.—In desert-grass and thorn-bush country: scat-

tered, dry tufts of grasses interspersed among acacia bush and scat-
tered 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 Ari-
zona (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 debris in
desert-grass and thorn-bush country (Kevan and Chopard, 1954).

28 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Hetero-
gamodes females live more or less buried in the sand (Kevan and
Chopard, 1954).

Namablatta bitaeniata

Southwestern Africa—Limited in distribution to the more arid por-
tions, 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 29

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) indi-
cated 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).

30 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 distin-
guish 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 “pro-
tect” 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 habi-
tats. Strictly speaking, these cockroaches live on land at the edges of
streams or pools and spend relatively brief periods in the water. A

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 31

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,
waterfilled, 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 cock-
roaches will descend rapidly into the water when alarmed and prob-
ably 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 ovovivip-
arous.” (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 (Sein, 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 float-
ing 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 climb-
ing 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

32 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 sur-
face 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 terres-
trial 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 moun-
tain 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, 1920.)

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS BS

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 ab-
dominal 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 experi-
ments in which the insects’ abdomens were held immersed in water,
with the thorax exposed, the insects died in 6 to 12 hours or less. (Shel-

ford, 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,

19572).
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. Speci-
mens kept in jars lived several days with only a portion of their ab-
domens 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 con-

34 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

sidered. 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 unidenti-
fied nymphal cockroaches) in the litter of dead leaves, twigs, and other
plant products in over go 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.* 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 arach-
nids, 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 in-
habit abandoned cocoons and larval tents, wasp nests, ant nests, ter-
mite 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 sea-
weed, drift, and other debris on beaches ; burrowing in soil and under
clods of earth; in marshes and swamps; in dumps and rubbish heaps.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 35

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 substan-
tiate 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 ex-
pect 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 rela-
tively 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 move-
ment 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, addi-
tional research is needed on the bionomics of all species. Further con-
clusions 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 to feet above the
ground on Mona Island (Hebard, 1917). In trunks of trees under
bark and very often in abandoned cocoons of the “plumilla” (Sein,
1923). On rotten, wooden fence; in empty cocoons of Megalopyge
krugu on trunks of bucare trees, Erythrina glauca; on trunk of Inga
laurina; in larval tents of Tetralopha scabridella on Inga vera (Wol-
cott, 1936). In large numbers in nests of the gray kingbird (Wolcott,
1950).

36 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Sand-
hills 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

Hazwati—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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 37

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—tLives 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).

38 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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.

V enezuela.—Only taken in the forests of the Orinoco near the
trunks of rotten trees at night (Doumerc im 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 Feb-
ruary (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, Pali-
urus aculeatus, and Dictamnus fraxinelia; 19 specimens, apparently
breeding outdoors (Adelung, 1907).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 39

North-central U.S.—Observations since 1950 indicate a marked in-
crease 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 througout 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,
IQI2).

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).

40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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, 19372).

Cahita nahua

Honduras.—All beaten from foliage along roads or in thickets,
during rainy season (Rehn, 19374).

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. hylaeca 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, 19454).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 4!

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 under-
growth in pine and oak woods (Rehn and Hebard, 1916). In under-
growth 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,
EOE)

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 (domi-
nant), xeric hammock (infrequent), mesic hammock (dominant),
pond margin (infrequent), longleaf-pine flatwoods (frequent), bay-
head (occasional), low hammock (frequent), and alluvial hammock
(occasional). In the shrub stratum in these habitats: Scrub (fre-
quent), sandhills (dominant), and xeric hammock (infrequent). In
herbaceous stratum in these habitats: Sandhills (dominant), mesic
hammock (dominant), and black-pine flatwoods (infrequent). On

42 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI

bare soil or bare sand under vegetation in these habitats: Sandhills
(dominant), pond margin (infrequent), longleaf-pine flatwoods (fre-
quent), 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 (in-
frequent), 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 43

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.—F rom jungle undergrowth (Hebard, 1920).

Chorisoneura specilliger

Panama.—In grass (Hebard, 1920).

Chorisoneura texensis

Florida.—“The almost impenetrable jungle on Key Largo was ex-
amined, and in its depths the two specimens of this species were se-
cured 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 web-
worm 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, 1943).

Southeastern and southern U.S.—I\n undergrowth in pine woods;

44 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAT

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 eleva-
tions 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 45

Cutilia soror

Marquesas Islands—Males under stones and dead log (Hebard,
19332).

Hawaii—tIn 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

Hawau.—‘Crowds of these insects in various stages of development
sometimes gather in cypress trees, in suitable chinks, in old flower-
head 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. How-
ever, 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, 19334).

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 under-
growth (Rehn, 1931).

46 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Trebovie (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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 47

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. Odthecae 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
odthecae 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 asso-
ciation with forests (Bei-Bienko, 1950).

48 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 (Sein, 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 49

Epilampra tainana
Cuba.—Under dead leaves on stream bank (Rehn and Hebard,
1G27;)’.
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

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 (fre-
quent), 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 hack-
berry ; 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 51

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).

52 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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).

Missourt—Twenty to 30 males found resting on heads of wild
oats on successive evenings (Rau, 1947).

Texas.—It preferred damp, open woodlands (Hebard, 19432).

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 pre-
fer 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 (infre-
quent), scrub (infrequent), sandhills (occasional), xeric hammock

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 53

(frequent), mesic hammock (dominant), shrubby, longleaf-pine flat-
woods (infrequent), bayhead (dominant), and low hammock (dom-
inant). 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 (oc-
casional), 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 shrub-
bery 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).

54 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Moun-
tains” (Ball et al., 1942).

Latiblattella rehni

Florida.—Widely distributed throughout pine woods (Pinus cari-
baea) ; 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,
1900).

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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 55

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 con-
siderable altitude in the forests, under trash, stones, boards, etc. ( Wil-
liams et al., 1931). Often it is found with nymphs of Pertplaneta
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 sur-
face 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 fol-
lowing 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).

50 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Megamareta verticalis

Australia.—In sugarcane (Hebard, 1943).

Methana canae

Ausiralia.—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 ;
odthecae 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 brome-
liads; nearly all collected specimens were taken in these plants (Rehn
and Hebard, 1927).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 57

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 speci-
mens 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.—W idely 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).

58 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 (Tep-
per, 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 Litopeliis bispinosa
(Hebard, 1920).

Texas.—Lives in foliage and in the green sheaths of plants (He-
bard, 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 (Sein, 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 59

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, 19432).

Parcoblatta divisa

Eastern and southeastern U.S.—AIl 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 long-
leaf pines and pine stumps (Hebard, 1917). Widespread in south-
eastern U.S. in habitats as diverse as dry pine lands, oak scrub, moist
hammocks in northern Florida, and deep, cool ravines along Apala-
chicola River (Hebard, 19432).

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 bar-
rens 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

60 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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), sand-
hills (dominant), xeric hammock (dominant), mesic hammock, long-
leaf-pine flatwoods (infrequent), low hammock (infrequent), and
alluvial hammock (infrequent). In the shrub stratum in these habi-
tats: Scrub (dominant), sandhills (dominant), xeric hammock (dom-
inant), 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 ham-
mock habitat (Friauf, 1953).

Indiana.—Beneath rocks on sides and tops of high hills, in lime-
stone glades where cedar abounds (Blatchley, 1920).

Missouri.—lIn leaf stratum of oak-hickory forest (Dowdy, 1951).
Earlier, Dowdy (1947) reported finding numerous immature Pseudo-
mopinae [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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 61

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 oothecae com-
mon beneath loose bark of logs, especially shellbark hickory (Blatch-
ley, 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 oothecae were found
under bark of pine logs, where, in October and November, hibernating
nymphs were found (Strohecker, 1937). In nests of Vespula macu-
lata (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 under-
growth 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

62 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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,
LOL7):

Tennessee.—lIn 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 (Stro-
hecker, 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 63

piles of moist dead leaves and rotten logs in oak-hickory forest (Can-
trall, 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—tIn 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.—Odthecae under loose bark of fallen trees, where as many
as 184 odthecae 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 foetidissi-
mum (Wolcott, 1941). Common along the coast and in mountains.
“Very much at home” under the loose bark of Sideroxylon foetidissi-
mum (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

64 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

fumaroles where a railroad fill was burning internally (Davis, 1927).
Common n palm trees along the gulf coast of Texas, where they
often fly around street lights at night (Zimmern im 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).

Nthoa 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 encoun-
tered 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 im Blanchard, 1837).

Phyllodromica brevipennis

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 65

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).

66 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 scabri-
della in leaves of Inga vera, or of Pilocrocis secernalis in the leaves of
‘capa 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 grape-
fruit, 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 “Jucaro” (Gundlach in Rehn and
Hebard, 1927).
Plectoptera pygmaea

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 67

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 Plec-
toptera 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, ac-
cording 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 sun-
ning 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, 19432).

Pycnoscelus surinamensis

Florida.—Under stones and rubbish; very abundant under coquina
boulders in woods at Key West (Rehn and Hebard, 1905). “This

68 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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).

Hawati.—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 in-
dividuals bored into the soil, Puerto Rico. (Rehn and Hebard, 1927.)

Barbados.—F requents cane fields (Tucker, 1952).

Puerto Rico—‘Altho primarily a xerophytic species: collected
among dry stones on Mona Island, under dry cow dung at Boqueron,
and under boxes at Guanica, 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 eye-
worm, 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 69

Simblerastes jamaicanus

Jamaica.—Numerous in fragmentary debris of an abandoned ter-
mite 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 cavi-
ties 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

Hawaiu.—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 sugar-
cane (Wolcott, 1950).

70 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 under-
growth present (Bei-Bienko, 1950).

STRUCTURAL HABITATS

In this category we include all man-made structures, whether in-
habited 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 pos-
sibly started as casually as the short-lived association that Beebe
(1953) observed when he discovered three cockroaches in the newly
built couch of an orangutan. Obviously, when man came down from
the trees, his fellow travelers found his cave dwellings and other
abodes particularly favorable habitats. From such primitive begin-
nings, 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 in-
habited world. At least eight domiciliary cockroaches originated in
Africa (Rehn, 1945): Blatta orientalis, Blattella germanica, Leu-
cophaea maderae, Nauphoeta cinerea, Oxyhaloa buprestoides, Peri-
planeta americana, P. australasiae, and Supella supellectilium; and,
perhaps, Periplaneta brunnea as well; Neostylopyga rhombifolia was
probably of Indo-Malayan origin; Pycnoscelus surinamensis was of

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 71

oriental origin; and Lewurolestes 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 tem-
perate zones and some even in the Arctic. Bei-Bienko (1950) listed
the following Io species as sinanthropes in the Palearctic zone: Blatta
lateralis, B. orientalis, Blattella germanica, Leucophaea maderae, Peri-
planeta 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 cock-
roaches 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 tempera-
ture 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 indif-
ference) of Blatta orientalis is 20-29° C. The upper limit of the pre-
ferred 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 favor-
able 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 experi-
mentally that, unless the water content of the food is high, fluid water
is essential in the insects’ dietary. Ten species of domiciliary cock-
roaches 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

72 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

humidity (Willis and Lewis, 1957). Blatta orientalis, when in a state
or normal water balance, usually spent more time in the drier part of a
humidity gradient; but desiccated insects tended to become hygro-
positive (Gunn and Cosway, 1938). We presume that other domi-
ciliary 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 estab-
lished 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 micro-
climatic conditions of structural microhabitats will be needed before
meaningful conclusions can be drawn about the stratification of cock-
roaches 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 73

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 cock-
roaches 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 cock-
roaches (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, cock-
roaches 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 hun-
dreds) 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

74 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 fre-
quently 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 depart-
ments, 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 dwell-
ings 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 infre-
quently in structures and are not known to breed there, may possibly

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 75

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 in-
vader 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

Hawati—Found only indoors at Nauhi. Otherwise this is ap-
parently an outdoor species (Swezey and Williams, 1932).

Blaberus craniifer

Cuba.—Household pest (Deschapelles, 1939). Particularly abun-
dant 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 (Sein, 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

76 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 ger-
manica) ; 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). Good-
liffe (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 re-
frigerators ; 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 oothecae in cracks 1.6 mm. wide.

Shuyler (1956) has observed extensions into relatively new struc-
tural 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 ware-
houses, 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 77

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 col-
lected indoors in Texas (Riherd, 1953).

Chromatonotus notatus

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

Cutilia soror
Hawait.—Almost as common in houses as Neostylopyga rhombif olia
(Hebard, 1922).
Ectobius duskei

U.S.S.R.—Frequently occurs in living apartments in farming locali-
ties 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

Hawati.—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).

78 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 (Sein,
1923; Wolcott, 1950).

Reported as a domiciliary pest in Madeira (Heer, 1864) ; Wind-
ward Islands (Marshall, 1878); Tropics and sub-Tropics (Rehn,
1937) ; Philippine Islands (Uichanco, 1953) ; New York City (Anony-
mous, 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 79

Methana marginalis

Australha.—Reported entering houses (Pope, 19532).

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).

Hawai.—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 out-
door species.

80 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI

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 (Hub-
bell, 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 (Cam-
pos 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 81

Periplaneta fuliginosa

U.S.A.—Frequently encountered out of doors, but has been re-
ported 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

Iraqg—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 green-
house pest (Hebard, 1917; Zappe, 1918; Doucette and Smith, 1926;

82 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 odthecae 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; odthecae 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).

Hawati.—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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 83

the breeding populations of a few common domiciliary pests. Except
in the most rigorously disinsectized ships, this commerce in cock-
roaches has continued to the present day.

Cockroaches undoubtedly infested the first ships that sailed the
Mediterranean; of these we have no records. The earliest recog-
nizable record of cockroaches on shipboard is Moffett’s (1634) state-
ment that when Drake captured the ship Philip, he found it overrun
with cockroaches [ Blattarum alatarum]. Bligh (1792) described dis-
infesting 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 re-
sembled 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 resi-
dence.”

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. . . . Cock-
roaches soon became plentiful on board, and showed themselves when-
ever 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.”

84 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 cock-
roaches 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
1o-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 occasion-
ally 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 pec-
tinicornis (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 disin-
fecting ships: Canalis (1916), Pryor (1918), Brooke (1920), Rice
(1925), Williams (1931), Simanton (1946), Richardson (1947), and
Anonymous (1951, 1954).

' BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 85

COCKROACHES ASSOCIATED WITH SHIPS

In the following list we include some previously unpublished data
on cockroaches that were recovered from ships at the Miami, Fila.,
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 ship-
board and will probably never establish breeding colonies on ships or
become pests on shipboard or elsewhere; some were merely passen-
gers 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 communi-
cation, 1958).
Ischnoptera sp.

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

Latiblattella sp.
At sea.—One female alive in hold of S.S. Tenadores (Hebard,
LOL):

86 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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,
IQI7).
Neoblattella sp.

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 87

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 infesta-
tion 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.

88 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 south-
ern 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 air-
craft at Miami (Denning et al., 1947).

Blatta orientalis

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

Blattella germanica

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 89

Epilampra sp.

U.S.A.—One dead specimen, Miami (Denning et al., 1947). South-
ern 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 air-
craft, 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 communi-
cation, 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

go SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

(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 (19562).

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, Mi-
ami (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 Mendon¢a 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 odthecae 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS gI

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, Strepsip-
tera, 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), Coleop-
tera (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; Cam-
eron’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 im-
mediately 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 mu-
tually 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 cate-
gories 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 :

g2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Symbiosis is the living together in more or less intimate associa-
tion of organisms of different species; it includes virtually all rela-
tionships between cockroaches and other organisms, such as parasit-
ism, 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 sym-
biont or symbiote by authors.

Commensalism includes associations in which neither party ap-
pears 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 associa-
tion may be divided into interspecies predatism, in which the preda-
tor preys upon a different species, and intraspecies predatism (canni-
balism) in which the predator preys upon its own species.

Although we have attempted to adhere to these definitions through-
out 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 Thelastomati-
dae), 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 organ-
isms 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, in-
convenienced by the presence of Protozoa or worms or lice, or a
cuckoo in the nest, we can nevertheless assume that it would be better

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 93

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 com-
mensals 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.” TFaust’s suc-
cessful 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 odtheca, 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 odtheca; 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 pred-
atory 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

94 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 ootheca, 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 odtheca, 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, Eupelmi-
dae) of cockroach eggs many individuals develop in a single ootheca.
When a hundred or more wasps emerge from an odtheca which con-
tained less than 20 eggs, it is obvious that a single cockroach egg sup-
ported more than one wasp, yet it is possible that one particular wasp
larva may have fed upon more than one cockroach egg before be-
coming an adult. We consider all entomophagous wasps that develop
in cockroach oothecae to be parasites rather than predators.

On the other hand, even though Anastatus floridanus, A. tenupes,
and Tetrastichus hagenowti 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 95

given for each section, but all organisms with similar habits pre-
sumably 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. IOI).
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;
De2aT)s

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. TIT):

(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).

(£) Arthropods.
(1) Mites (Tyrophagus lintneri; p. 218).

96 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 I. 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 I. 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 97

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 cock-
roaches bacteroid free. Starvation, parasites, electromagnetic radia-
tion, heat or cold, or chemicals have all been used in attempts to
eliminate the bacteroids. Of these, only chemical treatment has pro-
vided 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 ob-
served 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, hexyl-
resorcinol, or metaphen. Bode (1936) reported that injection of
irritants such as lithium salts or quinine hydrochloride had no ap-
parent effect on the symbiotes.

Brooks (1957) reared Blattella germanica on diets containing dif-
ferent concentrations of inorganic ions. On a manganese-deficient
diet the cockroaches grew poorly and some of their progeny lacked
normal bacteroids; about Io percent of the aposymbiotic generation
grew and reproduced on a diet fortified with yeast. Varying the con-
centrations 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 cock-
roaches 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,

98 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 cock-
roaches 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) ques-
tioned 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 Blatiella
germanica. Every female whose bacteroids were reduced to the
vanishing point resorbed her ovaries and was incapable of reproduc-
tion. Brooks (1954; Brooks and Richards, 1955) found that adminis-
tration 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 bac-
teroids 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 prac-
tically 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). Ob-
viously, the intracellular symbiotes subserve the normal nutrition of
the cockroach. Whether the bacteroids produce only vitaminlike sub-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 99

stances, 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 aposym-
biotic cockroaches is much greater than the known vitamin require-
ments; 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) trans-
planted symbiote-containing fat body from Blatta orientalis into the
mealworm and larva of Ephestia kihniella, and from Blattella ger-
manica 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 in-
jected 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 im-
planted 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 (10945).

Blaberus giganteus. Blochmann (1892).

Blatta orientalis. Blochmann (1887, 1888, 1892) ; Bode (1936) ; Brooks (1954) ;
Buchner (1912); Cuénot (1896); Fraenkel (1921); Frank (1955, 10956) ;
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) ; Mencl (1907) ?; Mer-

100 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

cier (1906a, 1907, 1907b, 1907c); Ries (1932); Ronzoni (1949); Tacchini
(1946) ; Wolf (1924, 19244).

Blattella germanica. Blochmann (1887, 1888, 1892); Bode (1936); Borghese
(1946, 1948, 1948a) ; Brooks (1954) ; Brooks and Richards (1954, 1955, 1955a,
1956); Fraenkel (1921); Gier (1036, 1947) ; Glaser (1920, 1930a) ; Gropen-
giesser (1925); Haller (1055, 19055a); 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 (1036, 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 supellectiium. 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 de-
fined types of bacilli that he found in abundance in the digestive tract
of the Kiichenschabe, 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.

’

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS IOI

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 synthe-
sized 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. How-
ever, 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 Peri-
planeta americana) by oxygenation at 3.5 atmospheres. The ciliates
Nyctotherus and Balantidiwm, flagellates Lophomonas and Poly-
mastix, the amoeba Endamoeba blattae, and three unidentified pro-
tozoa 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, Lopho-
monas 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 punctu-
latus depends upon certain intestinal protozoa for survival; these
protozoa utilize as food the wood ingested by this cockroach. The

102 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Tricho-
nympha, 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 Crypto-
cercus and in termites.

The protozoa of cockroaches and termites are clues to the relation-
ship 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, im 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 similari-
ties 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 Clevelan-
dellidae (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 Pertplaneta and Blatta
(Kidder, 1937).

The protozoa of Cryptocercus can be transferred from one indi-
vidual to another (Nutting and Cleveland, 1954). They can also be

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 103

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, 19542).

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 foot-
note 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, Peri-
planeta americana and/or Periplaneta brunnea, and Supella supel-
lectilium, 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 Peri-
planeta 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, re-
moved 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 Io and

104 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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? (Find-
lay 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 7 are either not listed by Breed et al. or are
stated by them to be insufficiently characterized for definite classifica-
tion. Use of asterisk is explained in footnote 3, page 4.

Phylum SCHIZOPHYTA

Class, SOHIZOMY CETES
Order EUBACTERIALES
Family PPEUDOMONADACEAE

* Pseudomonas aeruginosa (Schroeter) Migula

Natural vectors——Blaberus cranifer, 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS I105

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

{ Spirillum periplaneticum Kunstler and Gineste

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

{ Spirillum a, B, and y 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).

Pertplaneta americana, Philippine Islands (Barber, 1914) ; Nether-
lands (Akkerman, 1933); Formosa (Morischita and Tsuchimochi,
1926).

Periplaneta australasiae, Formosa (Morischita and Tsuchimochi,
1920).

* Vibrio metschnikovii Gamaléia

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

Vibrio tyrogenus (Fliigge) Holland

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

106 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

+ 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 re-
sembled 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 Wed-
berg, 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.—Blatiella germanica, U.S.A. (Janssen and Wed-
berg, 1952).
+ Micrococcus nigrofaciens Northrup

Source.—Diseased June beetle larvae.

Experimental infection —Periplaneta americana, U.S.A. (North-
rup, 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 cranufer, U.S.A. (Wedberg et al.,
1949).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 107

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 meta-
thoracic 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 indi-
viduals. 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 hemo-
lymph cells (Tauber, 1940) ; these responses of the insect were in-
terpreted 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 Wed-
berg, 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) ].

108 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Natural vectors—Blatta orientalis, Italy (Spinelli and Reitano,
1932) ; Germany (Jettmar, 1935).

Blattella germanica, Germany (Jettmar, 1935).

Experimental vectors.——Blattella germanica, on shipboard (Mor-
rell, 1911) ; Germany (Vollbrechtshausen, 1953).

Micrococcus sp.

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

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

Sarcina aurantiaca Fligge
Habitat.—Air and water.
Experimental vectors.—Blatta orientalis, Italy (Cao, 1906): Intes-
tinal contents.
Sarcina lutea Schroeter
Habitat.—Air, soil, water, skin surfaces.

Experimental vectors——Blatta orientalis, Italy (Cao, 1906): From
intestinal contents.

{Sarcina symbiotica Pribram

Habitat—Oothecae 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 in-
jected.

+ Sarcina sp.

Synonymy.—Sarcina “blanche” of Sartory and Clerc.

Natural vectors——Blatta orientalis, France (Sartory and Clerc,
1908): Isolated from intestinal tract.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS I0Q

* + Sarcina sp.

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

* + Sarcina sp.

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

Family NEISSERIACEAE
* Neisseria meningitidis (Albrecht and Ghon) Holland
Experimental vectors Cockroaches, U.S.A. (Longfellow, 1913).

* Veillonella parvula (Veillon and Zuber) Prévot

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

Family LACTOBACTERIACEAE
* Diplococcus pneumoniae Weichselbaum

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

{ Enterococcus sp.

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

Lactobacillus fermenti Beijerinck

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

+ Pneumococcus Type I, No. 1231

Experimental vectors.—Blattella germanica, Germany (Vollbrechts-
hausen, 1953).

* Streptococcus faecalis Andrewes and Horder

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

Blattella germanica, U.S.A. (Steinhaus, 1941).

Pertplaneta americana? (“Blatella americana’), England (Shrews-
bury and Barson, 1948).

Cockroaches [presumably any or all of the above three species],
Egypt (El-Kholy and Gohar, 1945).

IIo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I
* Streptococcus liquefaciens Sternberg emend. Orla-Jensen

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

*+ Streptococcus microapoika Cooper, Keller, and Johnson

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

* + Streptococcus non-hemolyticus II Holman

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

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

* Streptococcus sp. (pyogenic group)

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

* Streptococcus sp. (viridans group)

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

* Streptococcus spp.
Natural vectors.—Blatta orientalis, Germany (Jettmar, 1935).
Blattella germanica, Germany (Jettmar, 1935); U.S.A. (Janssen
and Wedberg, 1952).
Cockroaches, U.S.A. (Longfellow, 1913).
Experimental vectors.—Blatta orientalis, Italy (Cao, 1808).

Family CORYNEBACTERIACEAE
* Corynebacterium diphtheriae (Fliigge) Lehmann and Neumann

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

Family ACHROMOBACTERIACEAE
} Achromobacter hyalinum (Jordan) Bergey et al.

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

Achromobacter sp.

Natural vectors.—Blattella germanica, U.S.A. (Janssen and Wed-
berg, 1952).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS II!

Alcaligenes faecalis Castellani and Chalmers

Synonymy.—Bacillus faecalis alkaligenes; Bacillus alcaligenes fae-
calis; B. alcaligenes faecalis.

Habitat.—Intestinal canal of man. Has been isolated from feces,
abscesses related to intestinal canal, and occasionally in the blood-
stream. However, it is generally considered nonpathogenic.

Natural vectors—Blaberus cranifer, 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): Iso-
lated 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 Wed-
berg, 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 craniufer, 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): Iso-
lated from intestinal tract.

Ii2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 odtheca.

Periplaneta americana, U.S.A. (Bitter and Williams, 1949): From
intestinal tract.

Aerobacter sp.

Natural vectors.—Reriplaneta americana, U.S.A. (Bitter and Wil-
liams, 1949): Isolated from intestines.

+ Eberthella oedematiens Assis

Habitat.—Intestinal canal.
Natural vectors ——Periplaneta americana, U.S.A. (Bitter and Wil-
liams, 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 (Jett-
mar, 1935) ; Poland (Nicewicz et al., 1946).

Blattella germanica, U.S.A. (Steinhaus, 1941).

Periplaneta americana, U.S.A. (Bitter and Williams, 1949, 19494).

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 (Vollbrechts-
hausen, 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS Las

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 Wil-
liams, 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 Wed-
berg, 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 Wed-
berg, 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, Blat-
tella germanica and/or Periplaneta americana], Egypt (El-Kholy and
Gohar, 1945).

Ii4 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

* Proteus mirabilis Hauser
Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Wil-
liams, 1949, 1949a).

* Proteus morganii (Winslow et al.) Rauss
Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Wil-

liams, 1949, 19492).
Cockroaches [presumably Blatta orientalis, Blattella germanica,
and/or Periplaneta americana], Egypt (EI-Kholy and Gohar, 1945).

* Proteus rettgeri (Hadley et al.) Rustigian and Stuart
Natural vectors.—Periplaneta americana, U.S.A. (Bitter and Wil-
liams, 1949).
* Proteus vulgaris Hauser
Natural vectors —Blaberus cranifer, U.S.A. (Wedberg et al.,

1949).
Blatta orientalis, Italy (Spinelli and Reitano, 1932).

Periplaneta americana, U.S.A. (Bitter and Williams, 1949, 19494).
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
Iudina in Pavlovskii, 1948).

Blattella germanica, U.S.A. (Olson and Rueger, 1950) ; U.S.S.R.
(Rozengolts and Iudina in Pavlovskii, 1948).

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS I15

*Salmonella morbificans (Migula) Haupt

Natural vectors—Periplaneta americana, Australia (Mackerras
and Mackerras, 1948, 1949).

Experimental vectors—Nauphoeta cinerea, Periplaneta australa-
siae, Periplaneta ignota, and Supella supellectilium, Australia (Mac-
kerras 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 Wil-

liams, 1949, 1949a).

Experimental vectors. Periplaneta americana, Gold Coast Colony
(Macfie, 1922).

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

* Salmonella sp. (Type Adelaide)

Experimental vectors—Nauphoeta cinerea, Periplaneta australa-
siae, Periplaneta ignota, and Supella supellectilium, Australia (Mac-
kerras 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 Wil-
liams, 1949, 1949a).

* Salmonella sp. (Type Derby)

Experimental vectors—Nauphoeta cinerea, Periplaneta australa-
siae, 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 (Mack-
erras and Pope, 1948).

116 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4}

* 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 Wil-
liams, 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 Mer-
tens, 1950).

Nauphoeta cinerea, Australia (Mackerras and Mackerras, 1948).

Experimental vectors —Blaberus craniufer, U.S.A. (Wedberg et al.,
1949). o

Blatta orientalis, U.S.S.R. (Rozengolts and Iudina 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 Iudina in Pavlovskii, 1948).

Nauphoeta cinerea, Australia (Mackerras and Pope, 1948).

Periplaneta americana, U.S.A. (Beck and Coffee, 1943; Jung and
Shaffer, 1952).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS TV,

Periplaneta australasiae and Supella supellectiliwm, Australia (Mac-
kerras 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 cranufer, U.S.A. (Wedberg et
al., 1949).

Blatta orientalis, Italy (Spinelli and Reitano, 1932) ; U.S.A. (Mc-
Burney and Davis, 1930) ; U.S.S.R. (Rozengolts and Iudina in Pav-
lovskii, 1948).

Blattella germanica, U.S.A. (Janssen and Wedberg, 1952) ; Ger-
many (Jettmar, 1927); U.S.S.R. (Rozengolts and Iudina im Pav-
lovskii, 1948).

Periplaneta americana, Gold Coast Colony (Macfie, 1922) ; Nether-
lands (Akkerman, 1933); Formosa (Morischita and Tsuchimochi,
1926) ; U.S.A. (Olson in Roth and Willis, 1957a).

Periplaneta australasiae, Formosa (Morischita and Tsuchimochi,
1920).

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

Cockroaches [presumably Blatia 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 mor-
tality in our laboratory colony of this insect.

Leucophaea maderae or Periplaneta americana, Philippine Islands
(Barber, 1912) : From hemolymph.

118 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 sus-
pensions of ground insects. In 1954 we received a culture of Peri-
planeta 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 sev-
eral moribund individuals. The Department of Public Health of Min-
nesota had at times in the past cultured S. marcescens but had dis-
carded 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, 19572): Iso-
lated 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 cranufer, 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 prac-
tically pure cultures of Serratia were recovered from the reddened
areas.

Blatta orientalis, Italy (Cao, 1898): Isolated from intestinal con-
tents. Passed unchanged through the gut.

Blattella germanica, Canada (Heimpel and West, 1959): Not nor-
mally pathogenic per os; LDso 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 Wil-
liams, 1949, 1949a).

* Shigella dysenteriae (Shiga) Castellani and Chalmers

Experimental vectors——Blatta orientalis, Italy (Spinelli and Rei-
tano, 1932).

Periplaneta americana, Formosa (Morischita and Tsuchimochi,
1926).

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 119g

* 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—Blatita orientalis, Germany (Kuster, 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) ; Ger-
many (Kuster, 1903).

Blattella germanica, Germany (Jettmar, 1927).

Leucophaea maderae and Periplaneta americana, Philippine Islands
(Barber, 1912).

Family BACTERIACEAE

+ Bacterium alkaligenes Nyberg

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

I20 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

{+ Bacterium delendae-muscae Roubaud and Descazeaux

Source.—Diseased fly larvae.
Experimental infection—Cockroach, France (Roubaud and Des-
cazeaux, 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 (Kuster, 1903).

+ Bacillus biitschlii 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° ml. of a 24-hour broth culture.

Bacillus circulans Jordan

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS I2I

+ 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 micro-
organism 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): Or-
ganism recovered, apparently unchanged, from intestinal contents.

+ Bacillus monachae (von Tubeuf) Eckstein

Synonymy.—Bacterium monache.

Source.—Diseased larvae of nun moth, Lymantria monacha.

Experimental infection—Blatta orientalis, Europe (Filatoff, 1904) :
Organism pathogenic to the cockroach when injected but not when fed.

} Bacillus periplanetae Tichomiroff

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

} Bacillus stellatus Hollande

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

+ Bacillus radiciformis

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

* Bacillus subtilis Cohn emend. Prazmowski
Natural vectors——Blaberus cranufer, U.S.A. (Wedberg et al.,
1949).

122 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI

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 (Shrews-
bury 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): Iso-
lated from oodthecae.

} 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,
1808).
* Clostridium perfringens (Veillon and Zuber) Holland

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

* Clostridium tetani (Fliigge) Holland
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898).

* Clostridium spp.

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 123

Order ACTINOMYCETALES
Family MYCOBACTERIACEAE
* Mycobacterium avium Chester

Experimental vectors.—Blatta orientalis, U.S.S.R. (Ekzempliar-
skaia 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,
im Roth and Willis, 1957a) : Organism isolated from batches of in-
testinal 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 (Ari-
zumi, 1934, 1934a).

Cockroaches, Venezuela (Tejera, 1926); Belgian Congo (Radna,
1939).

Experimental vectors.—Blatia orientalis, Europe (Paldrock in
Klingmuller, 1930) ; Nyasaland (Lamborn, 1940).

Blattella germanica, Europe (Paldrock in Klingmiller, 1930) ;
Southern Rhodesia and Keyna (Moiser, 1945, 1946, 1946a, 1947;
Anonymous, 1946).

Nauphoeta cinerea, Nyasaland (Lamborn, 1940).

Periplaneta americana, Gold Coast Colony (Macfie, 1922); For-
mosa (Arizumi, 1934, 1934a).

Periplaneta australasiae, Formosa (Arizumi, 1934, 19344).

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).

I24 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

* Mycobacterium piscium Bergey et al.

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

* Mycobacterium tuberculosis (Schroeter) Lehmann and Neumann
Experimental vectors.—Blatta orientalis, Italy (Cao, 1898) ; Ger-
many (Kiister, 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 Hammerschmudtiella diesingt
and Leidynema appendiculata in Periplaneta americana, U.S.A. (Hoff-
man, 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.—H ygrocrocis intestinalis.

Habitat—Blatta orientalis, Europe (Valentin, 1836; Robin, 1847,
1853; Peshkoff, 1940): Isolated from intestinal tract. The organ-
ism 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 125

Order SPIROCHAETALES
Family SPIROCHAETACEAE
+ Spirochaeta blattae Tejera

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

* + Spirochaeta periplanetae Laveran and Franchini

Habitat.—Blatta orientalis, France (Laveran and Franchini,
1920a).

Cockroaches, Venezuela (Tejera, 1926): Tejera reported finding
“Spirochaeta blatarum Laveran et Franchini” which may have been a
lapsus.

Family TREPONEMATACEAE
} Treponema parvum Dobell

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

+ Treponema stylopygae Dobell

Synonymy.—S pirochaeta 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).

126 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

*“Bacillo proteisimile” of Cao

Natural and experimental vectors.—Blatta orientalis, Italy (Cao,
1898, 1900).
*“Bacillo del barbone dei bufali” of Cao

Experimenal 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 808, 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.—(1) Diseased silkworm larvae. (II) Diseased Ocneria
dispar larvae and blood of Blatta orientalis.

Experimental infection.—(1) (11) Blatta orientalis, Europe (Fila-
toff, 1904) : Organism pathogenic when injected, nonpathogenic when
ingested.

(1) 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, Blat-
iella germanica, and/or Periplancta americana], Egypt (El-Kholy

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 127

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 (Vollbrechts-
hausen, 1953): Nonpathogenic to the insect when injected into the
mouth or anus.

“Silkworm disease bacillus”

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

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

“Spirochaetoid bacteria”

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

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

VII BUNGE ANDY 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,

128 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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).

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 con-
tact. 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,

Herpomyces
stylopygae

Speg.
ascus- filled ane
perithecium

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

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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 129

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 debili-
tant. 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 reac-
tions 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 cul-
tured 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 odthecae of P. americana,
Griffiths and Tauber (1942a) autoclaved their rearing containers and
dipped the oothecae 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 Saccharo-
mycetes 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.—Ootheca of Blatta orientalis, Italy (Ronzoni, 1949).

130 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Torulopsis sp.

Natural host.—Odtheca 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—Odthecae of Blattella germanica and Eurycotis
floridana, U.S.A., Pennsylvania (Roth and Willis, unpublished data,
1952): On outer surface. Determination by Miss Mary Downing.

Oothecae of Periplaneta americana, U.S.A., Pennsylvania (Roth
and Willis, unpublished data, 1952): Inside odthecae. 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) : Organ-
ism passed unchanged through the gut of the insects.

Aspergillus sydowi (Bainier and Sartory) Thom and Church

Natural host.—Odtheca of Eurycotis floridana, U.S.A., Pennsyl-
vania (Roth and Willis, unpublished data, 1952): On outer surface.
Determination by Miss Mary Downing.

Aspergillus tamarii Kita

Natural host.—Odothecae of Blattella germanica, U.S.A., Pennsyl-
vania (Roth and Willis, unpublished data, 1952): On exterior sur-
face. Determination by Miss Mary Downing.

Aspergillus sp.?

Natural and experimental vector.—Blattella germanica, on ship-
board (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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 131

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 suspen-
sion 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) : Organ-
ism retained its pathogenicity after passing through the insect’s gut.

Penicillium sp.

Natural vector.—Blaberus crantfer, 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, un-
published 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 odthecal formation.

Cockroach, Puerto Rico (Johnston, 1915): From a “‘small roach”
in the pathological collection at Rio Piedras (no data).

132 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141

Family DEMATIACEAE
Memnoniella echinata (Rivolta) Galloway

Natural host.—Odotheca of Blattella germanica, U.S.A., Pennsyl-
vania (Roth and Willis, unpublished data, 1952): On material that
had oozed from a damaged ootheca. 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 odthecae. 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 craniufer, 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 "PHY COMYCEGES
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—Odtheca of Periplaneta americana, U.S.A., Pennsyl-
vania (Roth and Willis, unpublished data, 1952): Inside oodtheca.
Determination by Miss Mary Downing.

Pycnoscelus surinamensis, Germany (Bode, 1936): Isolated from
fat body which it had stained red.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS) 133

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. ort-
entalts.

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 craniufer, U.S.A. (Wedberg et al.,
1949): In feces.
Saccharomyces sp.
Natural vector——Blattella germanica, U.S.A. (Janssen and Wed-
berg, 1952): Found consistently in alimentary tract of B. germanica
fed sucrose solutions.

134 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Order HY POCREALES
Family HY POCREACEAE
Cordyceps amazonica Hennings

Natural host.—Cockroaches, British Honduras (Mains, 1940).

Cordyceps blattae Petch
Natural host.—Blattella germanica, Ceylon (Petch, 1924): Col-
lected at Hakgala twice. A slight covering of brown mycelium over-
ran 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 iden-
tified 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 /schnoptera) were spe-
cies of Parcoblatta. Hebard (1917) has shown that all the species

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 135

referred to Jschnoptera in the United States, except /. 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 Parcoblatia.

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 experi-
mentally (Richards and Smith, 1954).

Cockroach, Fiji (Thaxter, 1931).

Herpomyces ectobiae Thaxter

Natural hosts —Blattella germanica, U.S.A., Massachusetts (Thax-
ter, 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 Is-
lands (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.

136 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 an-

tennae.
Herpomyces macropus Spegazzini

Natural host.—Blaberus sp.?, Argentina (Spegazzini, 1917).

Cockroaches, Peru, Puerto Rico, Ecuador, and Haiti (Spegazzini,
I915, 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, per-
sonal 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 an-
tennae.

Cockroaches, Trinidad and Argentina (Thaxter, 1931).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 137

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 Tan-
gier (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 den-
sity 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, per-
sonal 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).

138 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Experimental hosts——wNeostylopyga rhombifolia (Richards and
Smith, 1954): A few plants matured.

Pycnoscelus surinamensis (Richards and Smith, 1954): Some de-
velopment 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 Stig-
matomyces 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 cranifer, 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, Min-
danao (Thaxter, 1931).

Experimental hosts—Blaberus cranifer, 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. tri-
cuspidatus. FE-xperiments 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 139

INCERTAE SEDIS

According to Dr. R. K. Benjamin (personal communication, 1957)
and Dr. E. G. Simmons (personal communication, 1957), the phylo-
genetic 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 cock-
roaches that feed on living plants (p. 162) and those allegedly cap-
tured 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 com-
munications, 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.

I40 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 rehmi, and Parco-
blatta 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 Wil-
liams, 1938; Zimmerman, 1948).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS I4I

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 Pember-
ton (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.—H ololeptoblatta 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 cock-
roach was rather abundant in clumps of Rhodes grass.

Panicum purpurascens Raddi

Synonymy.—Panicum barbinode | Hitchcock, 1936].
Associate —E pilampra abdomen-nigrum, Puerto Rico (Sein, 1923 ;
Wolcott, 1936): Abundant in “malojillo” meadow.

Saccharum officinarum Linnaeus

Associates —Balta quadricaudata, Balta scripta, Balta torresiana,
Balta verticalis, Ellipsidion simulans, and Megamareta verticalis, Aus-
tralia, Queensland (Hebard, 1943): All collected by J. F. Illingworth
on sugarcane.

142 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Blattella humbertiana, Ischnoptera schenklingi, and Pycnoscelus
surinamensis, Formosa (Box, 1953).

Cariblatta stenophrys, Puerto Rico (Sein, 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, per-
sonal 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 (Sein, 1923; Wol-
cott, 1936).

Ellipsidion bicolor, Australia, Queensland (Hebard, 1943).

Lophoblatta arawaka, Trinidad (Princis and Kevan, 1955).

Phoraspis sp., Brazil and Guiana (Doumerc im Blanchard, 1837).

Supella supellectilium, New Caledonia (Cohic, 1956).

Family CYPERACEAE
Cyperus sp.
Associate —Maretina uahuka, Marquesas Islands, Uahuka (He-
bard, 19332).
Family PALMAE
Acrocomia aculeata (Jacq.) Lodd.

Associate —Pycnoscelus surinamensis, Trinidad (Princis and
Kevan, 1955): On “gru-gru” fruits.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 143

Cocos nucifera Linnaeus
Associates—Aglaopteryx gemma, Florida (Rehn and Hebard,

IQI2).
Cariblatta lutea minima, Florida, and Cariblatta delicatula, San

Domingo (Hebard, 1916a).
Eurycotis floridana, Florida (Rehn and Hebard, 1912; Hebard,

HOU)’.
Periplaneta australasiae, Jamaica (Rehn and Hebard, 1927).
Pycnoscelus surinamensis, Florida (Rehn and Hebard, 1912; He-

bard, 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).

144 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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.—E pilampra 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 145

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).

Eurycottis 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 epi-
phytes 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).

146 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I41

“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 —Latiblatiella 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 ad-
ventitious 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 ob-
served. 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 147

Archimandrita marmorata, Denmark (Henriksen, 1939): Adven-
tive (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 tessellata, 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 speci-
mens 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): Ad-
ventive.

Eurycotis lixa, New York (Rehn, 1930): Adventive, on banana
ship from Jamaica.

Graptoblatta notulata, Marquesas Islands, Uahuka (Hebard,
1933a): In banana leaves.

148 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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. Eng-
land (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 (Sein, 1923): Sein 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.

Nauchidas 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) : Ad-
ventive.

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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 149

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, Ne-
braska (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): Ad-
ventive, from Jamaica.

Nyctibora obscura, Trinidad (Princis and Kevan, 1955): In
banana bunch.

Nyctibora sericea, Canada (Stevenson, 1905; Walker, 1912): Ad-
ventive ; Hebard (1917) synonymized Walker’s specimen under JN.
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): Adven-
tive, from Brazil.

Oxyhaloa deusta, U.S.S.R., Leningrad (Bei-Bienko, 1947): Ad-
ventive, from Colombia.

Panchlora antillarum, England (Bunting, 1955): Adventive, from
Dominica,

150 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Panchlora exoleta, Scotland (Distant, 1902): Adventive. Great
Britain (Shaw, 1902): Adventive. England (Coney, 1918) : Adven-
tive. Sweden, Norway (Princis, 1947): Adventive, Norwegian speci-
men from Brazil. Germany (Zacher, 1917): Adventive, from Ja-
maica.

Panchlora nivea, Colorado, Nebraska, New Jersey, New York (2),
Utah (Hebard, 1917): Adventive. Nebraska (Hauke, 1949): Ad-
ventive. Washington (Hatch, 1938): Adventive. Massachusetts
(Roth and Willis, 1958): Adventive. England (Bunting, 1955):
Adventive, from Dominica. U.S.S.R. (Bei-Bienko, 1947): Adven-
tive, from Colombia. Sweden (13), Norway (3), Finland (3) (Prin-
cis, 1947): Adventive, mostly females; origin (where known) Ja-
maica.

Panchlora fraterna(?) and Panchlora peruana(?), Denmark (Hen-
riksen, 1939): Adventive; origin (where known) Danish West In-
dies 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 communica-
tion, 1959).

Panchlora sp., Canada (Walker, 1912) : Adventive. England (Tul-
loch, 1939): Adventive, from Brazil.

Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1936).

Pelmatosilpha marginalis and Pelmatosilpha purpurascens, Eng-
land (Bunting, 1955, 1956) : Adventive, from Dominica; both species
common.

Pelmatosilpha vagabunda, New Zealand (Princis, 1954): Adven-
tive, probably from South America.

Periplaneta americana, Belgium (Schepdael, 1931): Adventive,
on bananas from the American Tropics.

Periplaneta americana and Periplaneta brunnea, England (Bun-
ting, 1955, 1956): Adventive, from Dominica.

Periplaneta americana and Periplaneta australasiae, England (Wat-
son, 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. Den-
mark (Henriksen, 1939): Adventive (9); origin mostly Jamaica.
England (Tulloch, 1939): Adventive, from Brazil. England (Bun-
ting, 1955, 1956): Adventive, from Dominica; common.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS I5I

Platyzosteria bifida, Nebraska (Hebard, 1917) : Adventive.

Plectoptera dorsalis, Puerto Rico (Rehn and Hebard, 1927): Cap-
tured 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 con-
siderable damage to banana plants growing in a conservatory.

Sibylloblatta panesthoides, Massachusetts (Rehn, 19372): Adven-
tive, 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, 1940).
Periplaneta australasiae, England (Lucas, 1918).

Undetermined orchids

Associates —Blaberus discoidalis, Blatta orientalis, Periplaneta
americana, Hawaii (Swezey, 1945).

Blatta orientalis, americana, cinerea, maderae, unidentified cock-
roaches, England, in bulb from Ecuador (Westwood, 1876).

Graptoblatta notulata, Hawaii (Swezey, 1945): On orchid from
India.

Homalopteryx laminata and Hormetica apolinari, New York (Heb-
ard, 1912c): In orchids shipped from Colombia.

Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1936).

Periplaneta americana, Germany (Tashenberg, 1884).

Periplaneta australasiae, England (Wainwright, 1898). Pennsyl-
vania (Skinner, 1905). Massachusetts (Morse, 1920).

152 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 pensyl-
vanica, North Carolina (Rehn and Hebard, 1916): Under signs on
red oak.

Parcoblatia 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 153

Quercus spp.

Associates.—A glaopteryx gemma, Alabama, Georgia, Florida, Lou-
isiana, 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). Massa-
chusetts (Flint, 1951): Under loose lichens and bark.

Parcoblatta divisa, Georgia, Louisiana, and Parcoblatta pensyl-
vanica, 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, 19454).

Family CHENOPODIACEAE
Beta maritima Linnaeus

Associate.—Ectobius panzeri, England (Lucas, 19204).

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 communi-
cation, 1959].

Associate-—Chorisoneura texensis, Florida (Rehn and Hebard,
1912).

154 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 ap-
parently 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 Sarracemia pitchers.

Sarracenia purpurea Linnaeus

Natural prey.—Cariblatta lutea lutea, North Carolina (Wray and
Brimley, 1943).
Sarracenia minor Walter

Synonymy.—Sarracema variolaris Michx. | Howard, personal com-
munication, 1958].

Natural and experimental prey.—Periplaneia 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 155

Family CUNONIACEAE
Weinmannia sp.

Associates —Aneurina viridis, Marquesas Islands, Nukuhiva and
Fatuhiva (Hebard, 19334).
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, particu-
larly 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).

156 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Ceratonia siliqua Linnaeus

Associate.—Diploptera punctata, Hawaii (Pemberton and Wil-
liams, 1938) : Damaging algarroba.

Erythrina glauca Willd.

Associates.—Aglaopteryx absimilis, Puerto Rico (Wolcott, 1950) :
In abandoned cocoon.
Aglaopteryx facies, Puerto Rico (Wolcott, 1936): In empty co-
coons.
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 rhab-
dota, Puerto Rico (Wolcott, 1950): In “butterfly nests” in leaves.

Mesquite
Associate.—N yctibora 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 157

Family RUTACEAE
Citrus aurantifolia Swingle

Associates.—Plectoptera dominicae and Plectoptera perscita, Do-
minica (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.

158 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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.—Illex 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—Graptoblatia 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——Amagonina emarginata, Cariblatta antiguensis, Eury-
cotis kevani, and Rhytidometopum dissimile, Trinidad (Princis and
Kevan, 1955).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 159

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.

160 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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, 19332).
Aneurina tahuata, Marquesas Islands, Tahuata (Hebard, 19332).
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.—E pilampra 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 panzert, England (Lucas, 1927) : “numerous imagines of
both sexes were swept from heather.”

Vaccinium meridionale Sw.

Associates.—Chorisoneura formosella, Neoblattella dryas, Neo-
blattella proserpina, Jamaica (Rehn and Hebard, 1927).

Family SAPOTACEAE
Sideroxylon foetidissimum Jacq.

Associate-—Pelmatosilpha coriacea, Puerto Rico (Wolcott, 1941):
Under bark.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS I61

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 communica-
tion, 1958].

Associates —Cariblatta antiguensis, Ischnoptera rufa rufa, Supella
supellectiium, Symploce ruficolis 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,
IQI2).
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 (He-
bard, 1933a).
Family RUBIACEAE
Canthium barbatum (Forst.) Seem.

Associate-—Graptoblatta notulata, Marquesas Islands, Uahuka
(Hebard, 1933a).

162 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Cinchona pubescens Vahl.

Associate —Periplaneta americana, Puerto Rico (Plank and Win-
ters, 1949): In greenhouse.

Coffea sp.

Associate —Plectoptera porcellana, Puerto Rico (Sein, 1923).

Family COMPOSITAE
Goldenrod

Associate —Eurycotis floridana, Florida (Hebard, 1917): “Climb-
ing 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 ma-
terial. 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, [Peri-
planeta?| americana, [Nauphoeta?] cinerea, [Leucophaea?] maderae,
and two others unknown to him. Fullaway (1938) stated that cock-
roaches 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 163

orchids as well as the aerial roots and flower spikes of Vanda orchids
(Rau, 1940a). Wainwright (1898) stated that Periplaneta austra-
lasiae 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. awstralasiae 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 or-
chids. Swezey (1945) in Hawaii stated that the following cock-
roaches have been reported as occasional minor pests on orchids:
Blatta orientalis, Blaberus discoidalis, P. americana, and P. surinam-
ensis; he further stated that Graptoblatta notulata had been inter-
cepted 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, Eu-
charis, 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 Peri-
planeta 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 (Zimmer-
man, 1948). On St. Kitts, B.W.I., young cotton plants were severely
attacked by P. australasiae; this caused loss of the stand on a con-
siderable area and necessitated replanting (Ballou, 1916). P. austra-
lasiae 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

164 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

where it may do considerable damage to the plants. Periplaneta fulig-
inosa 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 (Cro-
well, 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, pa-
paya fruits.”

In the reports of damage to plants by cockroaches, Pycnoscelus
surinamensis has been implicated most often. This species is un-
doubtedly 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 al-
sophilas 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. surinamen-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 165

sis 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 in-
festation 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-men-
tioned damage caused by P. surinamensis and added damage to chrys-
anthemums 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 be-
came 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 Lin-
naeus was attacked by Supella supellectilium. Wolcott (1924a) re-
ported 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 ap-
peared in large numbers in gardens where it gnawed at the under-
ground parts of vegetables and ornamental plants. Lever (1947)
listed it as a pest on the leaves of pineapple.

166 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141

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. Golden-
berg (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 ma-
jority 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): Otkomonas sp. and Monas sp. were found in the intestines of
83 percent of 124 cockroaches.

Cockroach, Venezuela (Tejera, 1926).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 167

Family TRYPANOSOMATIDAE
Leptomonas blaberae Tejera

Natural host.—Blaberus sp., Venezuela (Tejera, 1926).

Leptomonas sp.

Natural hosts—Parcoblatta lata, Parcoblatta pensylvanica, Parco-
blatta 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 Fran-
chini, 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 cock-
roaches examined. L. G. Feo (im Wenrich, 1932) successfully cul-
tured this protozoan (fig. 2, F).

Retortamonas sp.?

Natural host—Leucophaea maderae, Philippine Islands (Hegner
and Chu, 1930).

168 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Fic. 2.—Representative Protozoa associated with cockroaches. A, Monocercomonoides
melolonthae, X 3094 (after Grassé). B, Coelosporidium periplanetae, X 1310 (after Sprague) ;
trophozoite with spores and chromatoid bodies. C, Endamoeba blattae, X 273 (after Kudo);
trophozoite. D, Lophomonas striata, X 330 (after Kudo). E, Lophomonas blattarum, X 660
(after Kudo). F, Retortamonas blattae, X 3004 (after Wenrich). G, Nyctotherus ovalis,
X175 (after Kudo). H, Gregarina rhyparobiae, c. X52: mature trophozoite attached to
intestinal wall of Leucophaea maderae. (Redrawn from J. M. Watson [1945].) I, Diplocystis
schneideri, c. X 14.4 (after Kunstler). J, Gregarina blattarum, c. X 57 (after Kudo). K,
Protomagalhaesia serpentula, X 36 (after Pinto). L, Gamocystis tenax, magnification not
papwn iter Schneider). (AIl figures except H redrawn from Kudo [1954] after sources
indicated.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 169

Order: (POLY MASTIGINA
Family CHILOMASTIGIDAE
Chilomastix mesnili (Wenyon)

Experimental vectors——Blatta orientalis and Periplaneta ameri-
cana, 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 in-
fected 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 cock-
roaches, and of other species of insects.

Monocercomonoides orthopterorum (Parisi)

Synonymy.—Trichomonas (Trichomastix) orthopterorum Parisi;
Monocercomonas orthopterorum [Bélar, 1916]; Trichomastic or-
thopterum? [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).

“Kuchenschaben,” Austria (Bélar, 1916).

The protozoan is found in the hind gut. Zasukhin (1930) found

170 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 (Kid-
der, 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 OX YMONADIDAE
Oxymonas doroaxostylus (Cleveland et al.)

Synonymy.—Saccinobaculus doroaxostylus Cleveland et al. [Cleve-
land, 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 171

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 (Bélar)

Synonymy.—Octomitus periplanetae Belar [| Kudo, 1954].

Natural hosts—Blatta orientalis, U.S.S.R. (Zasukhin, 1930): Or-
ganism 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).

“Kitichenschaben,” Austria (Belar, 1916).

Hexamita sp.?

Natural host—Leucophaea maderae, Philippine Islands (Hegner
and Chu, 1930): The flagellates were present in large numbers.

* Giardia intestinalis (Lambl)

Experimental veciors—Blatta orientalis, South Africa (Porter,
1918).

Blattella germanica, Brazil (Pessoa and Corrca, 1927).

Eurycotis floridana, U.S.A. (Young, 1937).

Leucophaea maderae, Brazil (Pessoa and Correa, 1927).

Periplaneta americana, South Africa (Porter, 1918) ; Gold Coast
Colony (Macfie, 1922); Brazil (Pessoa and Corréa, 1927); U.S.A.
(Young, 1937).

Periplaneta brunnea, U.S.A. (Young, 1937).

Cockroaches, Venezuela (Tejera, 1926); Argentina (Bacigalupo,
in Tejera, 1920).

172 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

* 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., northern Cali-
fornia (Cleveland et al., 1934).

Family LOPHOMONADIDAE
Lophomonas blattarum Stein

Natural hosts—Blatta orientalis, Czechoslovakia (Stein, 1860) ;
Germany (Butschli, 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 (De-
Coursey and Otto, 1956, 1957).

Pertplaneta americana, England (Schuster, 1898); Europe (Ja-
nicki, 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).

“Kuchenschaben,” Austria (Belar, 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 173

percent of 124 B. orientalis infested. Zasukhin (1930) found Io per-
cent 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 con-
sidered a commensal (Kudo, 1926).

Lophomonas striata Biitschli

Synonymy.—Lophomonas sulcata Schuster is most probably identi-
cal with L. striata (Kudo, 1926b).

Natural hosts—Blatta orientalis, Germany (Bitschli, 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 Is-
lands (Hegner and Chu, 1930); U.S.A. (Kudo, 1926b; McAdow,
1931; Armer, 1944).

Cockroach, Venezuela (Tejera, 1926); England or U.S.A.?
(Lucas, 1928).

“Kiichenschaben,” Austria (Bélar, 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 Fusifornus lopho-
monadis.

Prolophomonas tocopola Cleveland et al.

Natural host.—Cryptocercus punctulatus, California, Oregon, Vir-
ginia, 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).

174 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 ex-
amined 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 TRICHON YMPHIDAE
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.

Fic. 3.—Protozoa from the gut of the wood-feeding cockroach Cryptocercus
punctulatus. A, Eucomonympha imla, female above, male below, c. X 375. (From
Cleveland [1950c].) B, Barbulanympha sp. (From Cleveland [1953].) C, Uri-
nympha talea, c. X 712. (From Cleveland [1951a].) D, Rhynchonympha tarda,
c. X 450. (From Cleveland [1952].) E, Trichonympha okolona or T. algoa, c.
X 390. (From Cleveland [1949].) (All drawings reproduced through the cour-
tesy of Dr. L. R. Cleveland.)

175

176 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 EUCOMON YMPHIDAE
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 in-
dividuals 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 mal-
pighian tubules.
Class SARCODINA
Order MYCETOZOA
INCERTAE SEDIS
Peltomyces periplanetae (Léger)

Synonyny.—Peltomyces blattellae. Sprague (1940a) synonymizes
Peltomyces periplanetae, with Coelosporidium periplanetae.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 177

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 myceto-
zoan, but they may have erred in synonymizing Plistophora peri-
planetae with the organism they studied. Debaisieux found intracel-
lular stages of Peltomyces periplanetae that have not been found in
Plistophora periplanetae or Coelosporidium periplanetae.

Order AMOEBINA
Family AMOEBIDAE
Hartmannella blattae Ivanic

Natural host—Blatta orientalis, Yugoslavia (Ivani¢, 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 Pertplaneta ameri-
cana, Egypt (DeCoursey and Otto, 1956, 1957): Thirty out of 261
cockroaches examined contained this protozoan.

Endamoeba blattae (Biitschli)

Synonymy.—Amoeba blattae, Entamoeba blattae, Entamoeba blat-
tarum.

Natural hosts.—Blatta orientalis, Germany (Bitschli, 1878; Schu-
botz, 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? (Ja-
nicki, 1908, 1909) ; U.S.S.R. (Yakimov and Miller, 1922; Zasukhin,
1929, 1930); England (Thomson and Lucas, 1926; Lucas, 1927,
1927a, 1928); Yugoslavia (Ivanic, 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).

178 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Periplaneta australasiae, U.S.A. (Morris, 1936).

Cockroaches, Paraguay ? (Elmassian, 1909); Austria (Bélar,
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 per-
cent of the examined cockroaches to be infested; Yakimov and Miller
(1922) found 4 percent of 124 oriental cockroaches infested; Za-
sukhin (1930) found up to 50 percent of over 3,000 B. orientalis in-
fested ; 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 (19072) observed a fungus, Nucleophaga sp., hyper-
parasitic in the nucleus of Endamoeba blattae.

Endamoeba javanica Kidder

Natural hosts—FPanesthia 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 en-
doplasm of this amoeba contains large amounts of wood and cellulose
fibers.

Endamoeba philippinensis Kidder

Natural host.—Panesthia angustipennis, Philippine Islands (Kid-
der, 1937) : Occurred in about ro 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. colli.

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 179

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 (De-
Coursey and Otto, 1956, 1957).

Cockroaches, Venezuela (Tejera, 1926).

Experimental vectors.—Blatta orientalis, Italian Somaliland (Mari-
ani and Besta, 1936).

Periplaneta americana, Gold Coast Colony (Macfie, 1922) ; U.S.A.
(Frye and Meleney, 1936).

Cockroaches, Venezuela (Tejera, 1920).

Entamoeba pitheci Prowazek?

Experimental vector—Periplaneta americana, Formosa (Mori-
schita 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 communica-
tion, 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 cock-
roach. Smith and Barret (1928) developed a synthetic medium in
which cultures of E. thomsoni were carried through successive trans-
fers for 24 months.

Entamoeba sp.

Natural vector —Periplaneta americana, Gold Coast Colony (Mac-
fie, 1922): Under the heading “Entamoeba histolytica and E. coli”

180 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Macfie (p. 445) stated, ““The cockroaches used in these experiments
had previously been carefully examined for amoebic infections a pre-
caution 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 EF. coli, from feces of the
monkey [Erythrocebus patas patas (Schreber)] were fed to cock-
roaches, 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, 19274) ;
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 ex-
amined.

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 patho-
genic.

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 cock-
roaches examined.

Blattella germanica and/or Periplaneta americana, Egypt (De-
Coursey and Otto, 1956, 1957) : Seventy-four out of 261 cockroaches
examined harbored this protozoan.

Iodamoeba sp.

Natural vectors.—Blattella garmanica and/or Periplaneta ameri-
cana, Egypt (DeCoursey and Otto, 1956, 1957): Fifty-nine of 261
cockroaches examined contained this protozoan. Jodamoeba sp. was
common in human feces in villages in which the cockroaches were
collected.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 181

Undetermined species of Amoeba

Natural host—Panesthia angustipennis, Philippine Islands. (Kid-
der, 1937): Found in only one specimen.

Class SPOROZOA
Order GREGARINIDA
Family DIPLOCYSTIDAE
Diplocystis schneideri Kunstler

(Fig. 2, I)

Natural hosts—Blatia 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 cranufer, U.S.A. (Nutting, 1953): From
I to 12 or more paired trophozoites or cysts may be found in the he-
mocoele and occasionally in the thorax.

Family STENOPHORIDAE
Stenophora sp.

Natural host—Blatta orientalis, India (Bal and Rai, 1955): Or-
ganism found in the midgut of the cockroach.

Family GREGARINIDAE
Gregarina blattarum von Siebold

Synonymy.—Gregarina blattae orientalis; Clepsidrina blattarum.
Natural hosts—Blatta orventalis, Germany (Siebold, 1837, 1839;

182 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Stein, 1848; Biitschli, 1881; Wolters, 1891; Marshall, 1892; Well-
mer, 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 (Magalhaes, 1900; Pinto, 1919); U.S.S.R. (Zasukhin, 1929,
1930).

Blattella germanica, U.S.A. (Crawley, 1903) ; South Africa (Fan-
tham, 1929; Porter, 1930: these appear to be the same record).

Blattella germanica and/or Periplaneta americana, Egypt (De-
Coursey and Otto, 1956, 1957).

Periplaneta americana, Brazil (Magalhaes, 1900) ; U.S.A. (Craw-
ley, 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 or1-
ental 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. Occa-
sionally, 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. german-
ica, collected in restaurants in Egypt, infested with this species. Wat-
son (1917) found a dozen or more in one specimen of Blatta ori-
entalis. 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 per-
cent were infected and only to 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 183

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,
1g18a, 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 uhle-
riana, 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.

184 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Gregarina thomasi Semans

Natural host—Parcoblatta pensylvanica, U.S.A., Ohio (Semans,
1939): Enteric caeca and midgut.

Protomagalhaesia serpentula (de Magalhaes)
Synonymy.—Gregarina serpentula [Pinto, 1918a, 1919; Semans,
1943].
Natural host.—Periplaneta americana, Brazil (Magalhaes, 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 palidus, 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 cranufer, U.S.A. (Roth and Willis, un-
published 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 185

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 some-
times generally distributed through the body (head, legs, antennae,
etc.) of C. punctulatus; this parasite was probably the species de-
scribed by Yarwood.

Order HAPLOSPORIDIA
Haplosporidium periplanetae Georgévitch
Natural host—Blatta orientalis, Yugoslavia (Georgévitch, 1953) :
This organism was described from the malpighian tubules of the cock-
roach where it apparently occurred in a mixed infection with the
microsporidian Plistophora periplanetae. See synonymy under Plisto-
phora periplanetae.

Coelosporidium periplanetae (Lutz and Splendore)

Synonymy.—N osema periplanetae, Coelosporidium blattellae, Ber-
tramia 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 (Ivanié, 1926).

Blattella germanica, U.S.A. (Crawley, 1905) ; Germany (Wellmer,
IQI0, IQII).

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 troph-
ozoite 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 epithe-

186 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

lial 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 Micro-
sporidia, Haplosporidia, and Mycetozoa, and that some of the dis-
crepancies 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, Haplospo-
ridium periplanetae, and Peltomyces periplanetae.

Natural hosts.——Blatta orientalis, France (Mercier, 1906a; De-
baisieux, 1927); England (Perrin, 1906, 1906b); U.S.S.R. (Zhi-
vago, 1909); Yugoslavia (Georgévitch, 1925, 1926, 1926a, 1927);
Germany (Wellmer, I910, I9I1).

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 cock-
roaches. 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 col-
lected in South Africa. It may or may not have been a species of
Plistophora.

Class CILIA TA
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 de-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 187

partment 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): In-
testinal 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., Hlinois (Kudo and Meg-
litsch, 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 (Magalhaes, 1900) :
These organisms were numerous in the intestine.

188 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Family SPIROSTOMIDAE
Nyctotherus buissoni Pinto

Natural host—‘Barata sylvestre,’ Brazil (Pinto, 1926): Organ-
ism found in the cockroach’s intestine.

Nyctotherus ovalis Leidy

Synonymy.—Bursaria blattarum; Plagiotoma blattarum.

Natural hosts —Blatta orientalis, U.S.A. (Leidy, 1850, 1853, 1853,
1879a; Kudo, 1922, 1926, 1936; McAdow, 1931; Kudo and Meg-
litsch, 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) ; Portu-
gal (Lima Ribiero, 1924); Brazil (Pinto, 1926); Venezuela (Te-
feta, O26).

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 (Lom, 1956).

“Barata sylvestre,” Brazil (Pinto, 1926).

“Kuchenschaben,” Austria (Bélar, 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; Lom, 1956).

Nyctotherus uichancoi Kidder

Natural hosts—Panesthia angustipennis, Philippine Islands, and
Panesthia spadica, Japan (Kidder, 1937) : About go percent of all P.
angustipennis harbored this ciliate in their hindguts.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 189

Nyctotherus viannai Pinto

Natural host.—“Barata sylvestre,” Brazil (Pinto, 1926) : In the in-
testine 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 pre-
occupied; 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 (Kid-
der, 1937).
Clevelandella hastula (Kidder)

Natural host.—Panesthia angustipennis, Philippine Islands (Kid-
der, 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 (Kid-

der, 1937).

Igo SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 demon-
strate 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 ap-
parent pathological effect on their cockroach hosts. Galeb (1878) has
shown experimentally that oxyurids eat the same food as the host in-
sect 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 commen-
sals. 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-para-
sitised specimens.”

Very few papers have dealt with the ecology of the oxyurid para-
sites 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 Ham-
merschmidtiella 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS IQr

cockroaches were infected with both of the above species of nema-
todes. 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 cock-
roaches infected by only one species (see pp. 195 and 197). Dobrov-
olny 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 cock-
roach 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 cock-
roaches damage the insect to varying degrees depending upon the ex-
tent 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 Oryspirura 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 pro-
thoracic 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 (Ali-
cata, 1937). Cysts of similar appearance have been found in cock-
roaches infected with Physaloptera rara, P. maxillaris, P. hispida
(Petri, 1950; Hobmaier, 1941; Schell, 1952), and Gongylonema

Ig2 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 pig-
mented cysts in Blatta orientalis and Periplaneta americana that were
experimentally infected with Rictularia coloradensis.
Our classification of the helminths follows Hyman (1951, 19514).

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, Massachu-
setts (Roth and Willis, 1957) : This mermithid lies coiled in the body
cavity of the host and one end may extend into the thorax. Ap-
parently, 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 193

chresima Steiner but apparently is a new species. Nauphoeta cinerea
was very susceptible to infection; B. germanica and P. americana
were less susceptible.

Order OX YUROIDEA
Family THELASTOMATIDAE

These nematodes are found in the intestinal tract of cockroaches.

Aorurus philippinensis Chitwood and Chitwood, 1934

Natural host—Panesthia angustipennis, Philippine Islands (Chit-
wood and Chitwood, 1934).

Binema mirzaia (Basir, 1940) Basir, 1956
Synonymy.—Periplaneticola mirgaia 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; O-xyuris blatticola
Galeb, 1878; Blatticola blatticola (Galeb, 1877) Schwenck, 1926
[ Chitwood, 1930, 1932].

Natural hosts—Blattella germanica, Brazil (Pessoa and Corréa,
1926; Schwenck, 1926); U.S.A. (Chitwood, 1930; Bozeman,
1Q42).> Meypt 2) (Galeb, 1877; 1878); U.Sis: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.

194 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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, ap-
parently accepted it as a valid species in describing the variety javanica.

Natural host——Panesthia laevicolis [Cobb recorded the host as
Panesthia brevicollis, but no such cockroach exists. Van Zwaluwen-
burg (1928) and Caudell (in Chitwood, 1932) believed that Cobb
meant Panesthia laevicollis. According to Gurney (personal com-
munication, 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 (Chit-
wood 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 magalhaesi (Schwenck, 1926) Basir, 1956
Synonymy.—Bulhéesi magalhadesi Schwenck, 1926; Thelastoma
magalhaesi (Schwenck, 1926) Travassos, 1929 | Basir, 1956].
Natural host—“Barata selvagem,” Brazil, Sao Paulo (Schwenck,
1926).
Euryconema paradisa Chitwood, 1932

Natural host——Eurycotis floridana, U.S.A., Florida (Chitwood,
1932).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS I95

Galebia aegyptiaca (Galeb, 1878) Chitwood, 1932
Synonymy.—O-xyuris 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 diesingt
(Hammerschmidt, 1838) Travassos, 1929; Streptostomum gracile
Leidy, 1850; O-xyuris diesingi Hammerschmidt, 1838; O-xyurts
blattae orientalis Hammerschmidt, 1838 [ Chitwood, 1932]. Oryurtis
macrura of Lankester (1865).

Natural hosts.—Blatta orientalis, Europe (Hammerschmidt 1838,
1847; Bitschli, 1871); Egypt? (Galeb, 1878); England
(Lankester, 1865 ;. Lee, 1958); U.S.A. CLeidy, 1850a) ;, U.S.S.R.
(Yakimov and Miller, 1922; Sobolev, 1937; Sondak, 1935) ; Brazil
(Travassos, 1929) ; China (Chitwood, 1932) ; Czechoslovakia (Gro-
schaft, 1956).

Leucophaea maderae, Brazil (Pessoa and Corréa, 1926).

Periplaneta americana, Brazil (Magalhaes, 1900; Pessoa and Cor-
rea, 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 (Chit-
wood, 1932). India (Basir, 1940). Czechoslovakia (Groschaft,
1956). England (Lee, 1958).

Periplaneta australasiae, Brazil (Pessoa and Corréa, 1926).

Polyphaga aegyptiaca (Linstow, 1878).

Cockroaches (Blatta orientalis, Blattella germanica, and/or Peri-
planeta 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. Butschli (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

196 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL air

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 ap-
pendiculata.

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 di-
gestive 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 1s 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 ap-
pendiculata.

Hammerschmidtiella neyrai Serrano Sanchez, 1945

Synonymy.—Hammerschnmudtiella neyrae Serrano Sanchez, 1947.
[ According to M. B. Chitwood, personal communication, 1957, Ser-
rano Sanchez’s emendation is apparently an error. ]

Natural host.—Blatta orientalis, Spain, Grenada (Serrano Sanchez,
1947): Of 2,943 specimens examined, 1,143 were parasitized by
oxyurids and of these 45 percent contained H. neyrat.

Leidynema appendiculata (Leidy, 1850) Chitwood, 1932

Synonymy.—Oxyuris blattae orientalis Hammerschmidt, 1847, of
Bitschli, 1871, and Oxyuris blattae-orientalis of Magalhaes, 1900) ;
Oxyuris blattae Hammerschmidt, 1847, of Galeb, 1878; Aorurus
(Thelastoma) appendiculatus Leidy, 1850. [Chitwood, 1932.] Ser-
rano Sanchez (1947) has divided this species into three geographical
varieties as follows: L. appendiculata (Leidy, 1852) (Dobrovolny

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 197

and Ackert, 1934) var. indiana; L. appendiculata (Leidy, 1852)
(Chitwood, 1932) var. americana; L. appendiculata (Serrano San-
chez, 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 (Butschli, 1871) ;
U.S.S.R. (Sobolev, 1937; Sondak, 1935); U.S.A., Nebraska (Todd,
1944) ; Spain (Serrano Sanchez, 1947): Recorded as var. hispana.
Czechoslovakia (Groschaft, 1956). England (Lee, 1958a).

Blatta orientalis or Periplaneta americana, Brazil (Magalhaes,
1900).

Periplaneta americana, U.S.A.: Texas (Todd, 1943); Nebraska
(Todd, 1944) ; Kansas (Dobrovolny, 1933; Dobrovolny and Ackert,
1934) ; North Carolina (Hatcher, 1939) ; lowa, North Dakota, Michi-
gan (Hoffman, 1953). Czechoslovakia (Groschaft, 1956). Eng-
land (Lee, 1958a).

Cockroach, Venezuela (Tejera, 1926).

Cockroaches (Blatta orientalis, Blattella germanica, and/or Peri-
planeta 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 em-
bryonated 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, inarticu-
late, amorphous filaments, growing from nematodes infecting B. ort-
entalis. Bitschli (1871) and Magalhaes (1900) described similar
filaments adhering to the surface of oxyurids from cockroaches.

198 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 cock-
roach 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, 10932

Natural hosts—Blaberus cranufer, 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. craniufer
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, 10932

Natural host—Leucophaea maderae, Cuba, Havana (Chitwood,
1932).

Leidynema nocalum Chitwood and Chitwood, 1934

Natural host.—Panesthia angustipennis, Philippine Islands (Chit-
wood 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 Chit-
wood, 1934).

Leidynemella panesthiae (Galeb, 1878) Chitwood and Chitwood, 1934

Synonymy.—Oxyuris panesthiae Galeb, 1878, in part; Thelastoma
panesthiae (Galeb, 1878) Travassos, 1929. [Chitwood, 1932; Chit-
wood and Chitwood, 1934. |]

Natural host—FPanesthia 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 (Chit-
wood and Chitwood, 1934).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 199

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 question-
able (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
Snyonymy.—Protrellina aurifluus Chitwood, 1932.
Natural hosts —Parcoblatta lata, U.S.A., North Carolina, Maryland
(Chitwood, 1932).
Parcoblatta uhleriana, North Carolina (Hatcher, 1939).

Protrellus australasiae (Pess6a and Corréa, 1926) Travassos, 1929

Synonymy.—O-xyuris australasiae Pessoa and Corréa, 1926; Pro-
trellina australasiae (Pessoa and Corréa, 1926) Chitwood, 1932 [Chit-
wood, 1933].

Natural host.—Periplaneta australasiae, Brazil (Pessoa and Corréa,
1926, 1927))).

Protrellus galebi Schwenck, 1926

Synonymy.—Protrellina galebi (Schwenck, 1926) Chitwood, 1932
[ Chitwood, 1933].

Natural host—“Barata selvagem,” Brazil (Schwenck, 1926).

Protrellus kiinckeli (Galeb, 1878) Travassos, 1929
Synonymy.—Oxyuris kiinckeli Galeb, 1878; Protrellina kiinckelt
(Galeb, 1878) Chitwood, 1932 [Chitwood, 1933].
Natural hosts——Periplaneta americana, Egypt? (Galeb, 1877,
1878. [Chitwood (1932) questioned the determination of this host

200 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

because he failed to find this nematode in a large number of specimens
from U.S.A. and China.] Brazil (Pessoa and Corréa, 1926).
Periplaneta australasiae, Brazil (Pessoa and Corréa, 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.—Bulhdesia icemi Schwenck, 1926; Thelastoma icemi
(Schwenck, 1926) Travassos, 1929; Thelastoma aligarhica Basir,
1940. [Basir, 1956. ]

Natural hosts —“Barata selvagem,” Brazil, Sao 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.—Bulhdesia severianoi Schwenck, 1926 [Travassos,
1929].
Natural hosis—“Baratas de pau podre,” Brazil (Schwenck, 1926).
Pycnoscelus surinamensis, U.S.A., Florida (Chitwood, 1932).

Suifunema caudelli Chitwood 10932
Natural host.—Steleopyga? sinensis, Asia: Suifu, Szchuen, China
(Chitwood, 1932).

Thelastoma pachyjuli (Parona, 1896) Travassos, 1929
Synonymy.—Oxyuris bulhdesi de Magalhaes, 1900; Bulhdesia bul-
hoesi (Magalhaes, 1900) Schwenck, 1926 [Travassos, 1929; Chit-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 20I

wood, 1932]; Thelastoma bulhdesi (Magalhaes, 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 (Magalhaes, 1900) ; North America
(Chitwood, 1932); U.S.A., North Carolina (Hatcher, 1939).

Thelastoma palmettum Chitwood and Chitwood, 1934

Natural host—Panesthia angustipennis, Philippine Islands (Chit-
wood 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) ; Pennsyl-
vania, 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.

202 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Ransom (in Pierce, 1921) states that G. aquaticus may be an acci-
dental parasite of man. Faust (1955) summarizes the few reported
cases of human parasitism. Dorier (1930) reported that the re-
gurgitated liquid of Blatta orientalis had no effect on cysts of G. aquat-
icus 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 (Mobius, 1855) Diesing, 1861

Synonymy.—Chordodes pilosus Mobius, 1855 [Diesing, 1861].
Natural host—Blaberus giganteus, Venezuela (Mobius, 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 20, B.

Parahormetica bilobata, Brazil (Pessoa and Corréa, 1929): Worm
referred to as “‘gordiaceo.”’

Cockroaches, Australia (E. F. Riek, personal communication,
1953): Three undescribed gordian worms were found in undeter-
mined cockroaches of the subfamily Blattinae.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 203

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, I915

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 (Magalhaes, 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, 10931

Natural host—Blattella germanica, India (Meyer, 1931, 1932).

4 The following helminths also have been stated in the literature to pass their
intermediate stages in cockroaches: Hymenolepis diminuta (Rudolphi, 1819)
[Blanchard (1891)]; Inermicapsifer madagascariensis (Davaine in Grenet,
1870) [Baer (1956)]; Spirocerca sanguinolenta (Rudolphi, 1819) [Seurat
(1913) ]. These doubtful records are discussed in Roth and Willis (1957a).

204 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

* Moniliformis moniliformis (Bremser in Rudolphi, 1819) Travassos, 1915

Natural hosts——Periplaneta americana, Argentina (Bacigalupo,
1927, 1927a, 1928); Brazil (Pessoa and Corréa, 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
Lariviére, 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,,1920;
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 re-
produce parthenogenetically. The parthenogenetic strain is undoubt-
edly 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 in-
volved in most Pacific areas. However, from internal evidence in his

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 205

papers, we concluded that Schwabe, in Hawaii, may well have been
working with the bisexual strain and possibly also with the partheno-
genetic 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, Parco-
blatta pensylvanica, Parcoblatta virginica, Periplaneta americana, and
Supella supellectilium, U.S.A. (Oswald, 1958): The larvae under-
went 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 de-
veloped in each infected Parcoblatta; fewer than Io per insect devel-
oped 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 Pessoa and Correa, 1929
Natural host—Parahormetica bilobata, Brazil (Pessoa and Corréa,
1929).
* Gongylonema ingluvicola Ransom, 1904

Experimental host.—Blattella germanica, U.S.A. (Cram, 1935).

206 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I41

* 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); For-
mosa (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 (Magalhaes, 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, Leuco-
phaea maderae, France (Brumpt, 1931).

* Protospirura columbiana Cram, 1926

Experimental host.—Blattella germanica, U.S.A. (Cram, 1926).

* Protospirura muricola Geodoelst, 1916

Natural host.—Leucophaca maderae, Panama (Foster and Johnson,

1938, 1939).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 207

* Seurocyrnea colini (Cram, 1927) Cram, 1931
Experimental host.—Blattella germanica, U.S.A. (Cram, 1931,
1931a, 1933a).

* Spirura gastrophila (Miiller, 1894) Seurat, 1913

Natural hosts —Blatta orientalis, Europe? (Deslongchamps, 1824,
in Seurat, 1911); Italy (Grassi, 1888) ; Algeria (Seurat, 1916).

Periplaneta americana, Brazil (Pessoa and Corréa, 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, per-
sonal 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).

208 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 HY MENOLEPIDIDAE
* 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 209

Phylum ASCHELMINTHES
Class NEMATODA
Order OX YUROIDEA
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 (Ak-
kerman, 1933).

* Ancylostoma ceylanicum (Looss, 1911) Leiper, 1015

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).

210 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 austra-
lasiae, Neostylopyga rhombifolia, Formosa (Morischita and Tsuchi-
mochi, 1926).

Family TRICHOSTRONGYLIDAE
* Trichostrongylus sp.

Natural vector—Blatta orientalis, Italian Somaliland (Mariani
and Besta, 1936).

Order TRICHUROIDEA
Family TRICHURIDAE
* Capillaria hepatica (Bancroft, 1803) 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 Hymen-
optera follows the classification of Peck (1951).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 2II

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) longe-
manus seems to prefer blattids to other insects (Schultze, 1927).

Order PEDIPALPIDA
Family THELYPHONIDAE
Mastigoproctus giganteus (Lucas)

Synonymy.—Thelyphonus giganteus Lucas [Dr. R. E. Crabill, per-
sonal 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 prin-
cipal food for scorpions at the Poisonous Animals Research Labora-
tory 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 communica-
tion, 1957].

212 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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, un-
published data, 1953): Scorpion collected in Florida by Roth and
identified by Dr. M. H, Muma.

Centruroides hentzi (Banks)

Experimental prey.—FPeriplaneta 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, Peri-
planeta 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.—Euscor pius carpathicus [ Cloudsley-Thompson,
19552].

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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 213

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 cock-
roach.

Family SCORPIONIDAE
Heterometrus longimanus (Herbst)

Synonymy.—Palamnaeus longimanus [Cloudsley-Thompson, per-
sonal 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.

214 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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. austra-
lasiae in Formosa; this spider also occurs on Hiyakejima Island and
Okinawa. Passmore (1936), who has produced some excellent photo-
graphs 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 cock-
roaches 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 215

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,
19302); 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 ; Clouds-
ley-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 fe-
male 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 im 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 THERIDITDAE
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 ac-
cidental contact of the insect with sticky threads surrounding the en-
trance. This spider apparently attacks any insect or arachnid that be-

216 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

comes 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 pre-
sumptive 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 prob-
ably 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 (Womer-
sley, 1956).
Coleolaelaps (?) sp.

Natural host—Gromphadorhina portentosa, the hosts were im-
ported into U.S.A. from Madagascar via Europe (Roth and Willis,

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 217

unpublished data, 1958): The mites (pl. 12, C) were tentatively de-
termined 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). Urop-
odids 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 sub-
family Diplogyniinae, and is most closely related to the genus
Lobogynioides. Mites of this family live as ectoparasites and com-
mensals 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.

218 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Caloglyphus sp.

Natural hosts—Blatiella 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 (per-
sonal 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 in-
fested with this species, but it did not affect the health or mortality of
the cockroaches in his culture.

Tyrophagus noxius A. Z.

Natural host.—FPeriplaneta americana, U.S.A. (Roth and Willis,
unpublished data, 1953) : Mite determined by Dr. E. W. Baker (per-
sonal communication, 1953). Mites were found in the oothecal 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 ap-
peared 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 cock-
roaches.

Family GLYCIPHAGIDAE
Chaetodactylus sp.

Synonymy.—Trichotarsus sp. [Baker and Wharton, 1952].

Natural host—Leucophaea maderae, Puerto Rico (Sein, 1923):
Mites found on cockroach’s thorax and particularly among the folds
of the wings (Sein, 1923). Mites of this genus are found infesting
organic matter (Baker and Wharton, 1952).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 219

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 NV. 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 be-
tween certain predaceous and phytophagous mites (Pritchard, 1956).

Family PTERYGOSOMIDAE
Pimeliaphilus podapolipophagus Tragardh

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 secre-
tion 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

220 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

unless live cockroaches were supplied. Parasitism was proved by de-
tecting 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 (Sein, 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 99].”

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 odthecae, 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 dis-
persed. 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). Ha-
waii, in laboratory (Schwabe, 1950) : Some of the cockroaches appar-
ently died from mite infestations.

A OY ee: I>
ZB SS FN
ESA \ MY WiLZ
\\
(

i Ve
Le id \ x

WS GE SSS
%

i
a

a
Se

Fic. 4.—The cockroach mite, Pimeliaphilus podapolipophagus. (From Baker
et al. [1956] ; reproduced through the courtesy of Dr. E. W. Baker and the Na-
tional Pest Control Association. )

221

222 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 podapoli-
pophagus (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 cock-
roach 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 (Sein,
1923). In Hawaii, centipedes preyed on insects generally but espe-
cially 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 (Cra-
bill, personal communication, 1953).

Experimental prey.—Blattella germanica, newly hatched nymphs

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 223

and adult female, U.S.A. (Snodgrass, 1930; Roth and Willis, un-
published 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 cock-
roaches 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 em-
braced 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.

224 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 re-
ported to be a great enemy of cockroaches.

Scolopendra sp.

Natural prey—Ectobius panzeri, England (Lucas, IgII, 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

XVI
Order ORTHOPTERA
Family MANTIDAE
Hierodula tenuidentata (Saussure) (?) (Serville)
(P1432)

Experimental prey.—Blatta orientalis, Diploptera punctata, Eury-
cotis floridana, Leucophaea maderae, Nauphoeta cinerea, Neostylo-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 225

pgya rhombifolia, and Periplaneta americana, U.S.A. (Rilling, per-
sonal communication, 1957): Mrs. Rilling wrote us that with the
exception of NV. rhombifolia, all the above cockroaches were readily
eaten. All the mantids initially rejected N. rhombifola after grasping
and making a brief attempt to chew the cockroaches. However, if
specimens of NV. rhombifolia were left in the jars with the mantids, the
cockroaches were usually eaten within the next 24 hours. N. rhombi-
folia ejects an odorous substance when seized and the mantids prob-
ably ate these insects after most of this secretion had been depleted.
It is highly probable that the secretion of NV. rhombifolia may deter
the mantid’s attack, but it should be pointed out that, with the possible
exception of NV. 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 ef-
fectiveness 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 ameri-
cana, U.S.A. (Rilling, personal communication, 1957).

Metallyticus semiaeneus Westwood

Experimental prey.—Cockroaches, Borneo (Shelford, 1916).

Sphodromantis viridus (Forskal)

Synonymy.—S phodromantis bioculata Burmeister [Gurney, per-
sonal 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 ameri-
cana, U.S.A. (Breland, 1941): The mantids were fed 1-2 German
cockroaches daily. One female mantid consumed to adult German
cockroaches plus one odtheca and part of another in 2.5 hours. An

226 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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. vy. 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 oothecae which ap-
peared to have been eaten into. He assumed from the condition of the
oothecae 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 (ob-
tained from an o6theca being carried by a female) and eggs of P.
japanica (presumably in odthecae) 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): Ac-
cording to Chopard, Brisout de Barneville in 1848 indicated that ear-
wigs in captivity can be fed small cockroaches.

Order HEMIPTERA

Family LYGAEIDAE

Clerada apicicornis Signoret
Natural prey.—Cockroach, Hawaii (Illingworth, 1917): This pre-
daceous bug is commonly found about buildings. Illingworth says that

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 227

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 saussuret, Turk-
men 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 (So-
meren, 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).

228 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. EAT

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 communica-
tion, 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) : Ob-
tained complete evolution of the parasite in this host. This may have
been an experimental host.

Calodexia spp.

Hosts.—Cockroaches, Panama (Rettenmeyer, personal communica-
tion, 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 suc-
cessfully reared.

Undetermined tachinids

Hosts—Eurycotis floridana, from Florida (Roth, unpublished
data, 1953): Three larvae (det. by W. W. Wirth) were found ina
living adult male.

Panesthia australis, from Australia (Roth, unpublished data,
1957): Reared from a wild-caught cockroach that was maintained in
a laboratory colony.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 229

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 communica-
tion, 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 at 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. punctu-
Jatus in nature. In the laboratory it killed and devoured cockroaches
as large as itself.

230 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 ®
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 circum-
ducta, 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 Pseudo-
mopinae, 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 circum-
ducta, Australia Capital Territory (Riek, 1955).

5 Barber (1939) has shown that the correct spelling of the name of the type
genus is Ripidius and not Rhipidtus, as it is frequently written, and that, conse-
quently, the family should be Ripiphoridae and not Rhipiphoridae.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 231

Rhipidioides mollis Riek

Natural host.—Robshelfordia longiuscula or Robshelfordia circum-
ducta, 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 Ripi-
diini 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 Ecto-
biinae 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 > boissyi Abeille

Balduf (1935) lists Ripidius boissyi as parasitic on nymphs of
Ectobius pallidus giving Abeille de Perrin (1909) as a source for this

232 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 /tipidius 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 steam-
ship “Samui” (Stamm, 1936); on cruiser ‘““Duguay-Trouin” (Bar-
bier, 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 identifi-
cation.

Ectobius pallidus? Abeille de Perrin (1909) stated that R. pectint-
cornis was first described by Sunders [sic] as blattarwm 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. pectini-
cornis apparently attacks only adult females and nymphs of B. ger-
manica. Barbier (1947) found only B. germanica parasitized, al-
though 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 fe-
male 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 233

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 ab-
domen (Barbier, 1947). Chobaut (1892) first suggested this method
of attack by the ripiphorid larva. As the parasites develop, the abdo-
men 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 odtheca 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 ro 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 biologi-
cal 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 cock-
roaches in baskets or sacks of provisions.

Ripidius scutellaris Heller

Natural hosts—Blattidae, Philippine Islands (Schultze, 1925).

234 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Family DERMESTIDAE
Dermestes ater De Geer

Common name.—Black larder beetle.

Natural prey.—Blatta orientalis, U.S.A. (Roth and Willis, unpub-
lished 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 kill-
ing them; the beetles probably attack only the weakened or injured
cockroaches in a culture. This was a natural infestation of a labora-
tory culture by a predator.

Experimental prey.—Blattella germanica, odthecae, U.S.A. (Roth
and Willis, 1950) : The beetle larvae can penetrate unhatched oothecae
of the German but not those of the American or oriental cockroaches.

Dermestes sp.

Natural prey.—Blatta orientalis, odthecae, 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 cock-
roaches 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 Halicto-
phagidae.”” 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 re-
corded as developing on cockroach eggs. All the Evaniidae are pre-
sumed 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 dwell-
ings indicates the presence of cockroaches (Gross, 1950). At times

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 235

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 sum-
maries of their biology.

Family EVANITIDAE
Acanthinevania princeps (Westwood)

Synonymy.—Evania princeps |Dr. H. Townes, personal communi-
cation, 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 attrib-
uted 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 re-
ports, 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 odtheca attached to the abdomen until or just
before the eggs hatch, it would seem that the female of B. minutus Git
the host records are valid) must oviposit into the odtheca of this
species while it is still being carried by the female; this would not

236 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

necessarily be true for the other hosts which drop the egg case long
before the eggs hatch.

Distribution.—Europe: Sweden, Russia, England, France, Ger-
many, Austria, Hungary, Switzerland, Italy (Kieffer, 1920).

Evania appendigaster (Linnaeus)

Synonymy.—Evama desjardinsu Bordage, Evania laevigata La-
treille [Dalla Torre, 1901-1902].

Natural hosts —Blatta, “exotic species” (Westwood, 1854, 1954a).

Blatta orientalis, Europe? (Schletterer, 1886; Howard, 1888,
Kieffer; 1912): Eeyptr "(Alferi, -ror4;” Adair, 1023). ~ [Girault
(1907, 1914) erroneously attributed another record to Marlatt (1902) ;
see footnote 6. See also notes under Brachygaster minutus with re-
spect to Kadocsa. ]

Blattella germanica? (Girault 1907, 1914). [This record is obvi-
ously 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 cock-
roach incubates its eggs internally (Roth and Willis, 1954). Later,
after finding that L. maderae is ovoviviparous, Bordage (1913) ad-
mitted having misidentified a parasitized ootheca from some other

6 Page 11 of Marlatt (1902) has been cited erroneously so many times in
support of host records for T. hagenowii and Evania appendigaster that we are
quoting the pertinent section below. In the section preceding the quoted material
Marlatt discusses the American, Australian, oriental, and German cockroaches.
There is nothing in the paper to connect any of these cockroaches specifically
with the parasites mentioned below:

NATURAL ENEMIES AND PARASITES

“In Europe the egg capsules of the cockroach are often parasitized by an
ichneumon fly (Evania appendigaster). This insect has become widely dis-
tributed over the world following its host insect, and has been redescribed under
a great many different names. It was found in Cuba as early as 1829, and has
been several times collected in the United States. Unfortunately, its usefulness
as a means of keeping the roach in check by destroying the egg capsules is
greatly impaired by the occurrence of another ichneumon fly (Entedon hage-
nowt), which is parasitic upon the first. This is also a European species which
has been brought over with its host parasite. If the true egg capsule parasite of
the roach could have been introduced into this country without this secondary
parasite, its usefulness would doubtless have been very much greater. The sec-
ondary parasite, however, seems to have been introduced early, and has been
found in Cuba and Florida, and probably occurs as widely as its host and pre-
vents the latter from multiplying very greatly.”

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 237

species ; he concluded that the developing eggs of this species are pro-
tected 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 (Sein,
1923) ; Jamaica (Gowdey, 1925); Hawaii (Swezey, 1929); Pal-
estine (Bodenheimer, 1930); U.S.A., Florida (Ashmead, 1900) ;
Maryland (Piquett and Fales, 1952); Saudi Arabia, Jedda (Cam-
eron, 1957) ; Canton Island and Samoa (Dumbleton, 1957).

Periplaneta americana or P. australasiae, Formosa (Sonan, 1924).

Periplaneta australasiae, U.S.A., Florida (Ashmead, 1900) ; Ha-
wali (Swezey, 1929; Zimmerman, 1948). [Girault (1914) errone-
ously 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 dis-
covered, 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 odthecae of cockroaches. Westwood
(18542) found the larvae, pupae, and adults of E. appendigaster in
egg cases of an unidentified species of cockroach found on orchids re-
ceived from Calcutta.

Adult behavior—Adult wasps visited flowers of parsley, Petro-
selium crispum, and fennel, Foentculum vulgare (Margretti in Schlet-
terer, 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 odtheca at
the crista and then deposited her egg or eggs on the eggs of the cock-
roach. Haber (1920) observed and described oviposition. The female

238 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

wasp crawled over the surface of the odtheca, 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 odtheca 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)

Fic. 5.—Evania appendigaster. Left, dorsal view, X 8. Right, side view, X 5.
(Reproduced with permission. British Museum [Natural History], 1951,
figs. IA and 1B.)

stated that the female deposits her eggs before the walls of the ootheca
harden.

Development.—Kieffer (1912) stated that the larvae in this family
eat the cockroach eggs and pupate in the odtheca 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 odtheca. 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 239

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 cock-
roaches 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 oothecae.

Hyptia dorsalis of Ashmead

Synonymy.—Dr. H. Townes, (personal communication, 1956) be-
lieves 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 (Ash-
mead, 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.: Ootheca collected by
L. Roth, May 17, 1956; wasp emerged June 12, 1956 (pl. 33, B) ; de-
termined by Dr. H. Townes. The keel region of the odtheca 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 cock-
roach odtheca (Edmunds, 1954). Five odthecae 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).

240 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Hyptia reticulata Say
Natural host.——FParcoblatta pensylvanica, U.S.A., Missouri (Rau,
1940).
Adult wasps have been taken on parsnip, Pastinaca sativa (Robert-
son, 1928).
Distribution.—U.S.A.: Pennsylvania to Florida and Louisiana.
Mexico. Upper Austral to Tropical Zones (Townes, 1951).

Hyptia thoracica (Blanchard)

Natural host.—Parcoblatia 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 (Ed-
munds, 1954).

Development.—Entire contents of odtheca are eaten by the single
larva. Last instar larva overwinters inside the odtheca. 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 odtheca near one end. Adult took about 65 minutes
to emerge from the time its mandibles first broke through the oothecal
wall. (Edmunds, 1954.)

Distribution —Canada, Ontario, U.S.A.: Connecticut to Wiscon-
sin, south to Florida and Texas. Upper Austral to Tropical Zones.
(Townes, 1951.)

Hyptia sp.

Natural host—Cariblatta delicatula, Cuba (Hebard, 1916a) ; Para-

site identified by Ashmead.

Hyptia sp. (undescribed)
Natural host—Parcoblatta sp., U.S.A., Ohio (Edmunds, 19522).

Prosevania punctata (Brullé)

Synonymy.—Evania punctata Brullé [Townes, 1949].

Natural and experimental hosts—Blatta orientalis, Istrian Penin-
sula (Fahringer, 1922) ; Algeria (Cros, 1942); U.S.A., Ohio (Ed-
munds, 1954).

Blattella germanica? Europe? (Girault 1907, 1914) ; Europe (Fahr-
inger, 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 241

seven female parasites from odthecae of Blattella germanica. He
placed female parasites with adults of B. germanica in a glass cage. As
soon as odthecae could be seen between folds of a woolen rag, he re-
moved all the larger cockroaches and held the oothecae until the para-
sites emerged. Fahringer may have been dealing with a different
species of cockroach, because placing oothecae in crevices (or between
folds of rag) is a habit foreign to B. germanica, the female of which
usually carries her ootheca until hatching or until about a day before.
Edmunds (1953b) could not induce this wasp to parasitize eggs of B.
germanica. |

Pertplaneta americana, Istrian Peninsula (Fahringer, 1922); Pal-
estine (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 abdo-
men moves up and down like a waving flag; because of this behavior,
these insects are commonly known as ensign-flies. Cros (1942) main-
tained 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 odthecae of P. ameri-
cana for oviposition and ignored those of B. orientalis and Parcoblatta
pensylvanica in the same cage. Oviposition was accomplished as de-
scribed for Evania appendigaster. One odtheca 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 odtheca, but she also oviposited from the opposite side or,
rarely, directly down into the side of the odtheca. 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 odthecae
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 cock-
roaches have reached the final stages of preemergence development.

242 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Edmunds (1954) placed females of Periplaneta americana that were
carrying odthecae, into cages with Prosevania; some of the female
wasps showed considerable interest in the attached odthecae, but he
observed oviposition only into egg cases that had been dropped by the
cockroaches.

Cros (1942) described an interesting reaction that he called “in-
stinctive hostility” of the oriental cockroach toward Prosevamia. A
wasp was placed in a jar in which a cockroach had just deposited its
ootheca. The wasp tried to oviposit into the egg case but was upset and
pursued by the cockroach. The cockroach placed herself over the
odtheca, 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 odtheca 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 de-
veloped in 45-53 days (Edmunds, 1954). In Blattella germanica:
Almost 4 weeks spent in development (Fahringer, 1922). In Peri-
planeta americana: Three wasps developed in 127 days (Edmunds,
1952). Only one parasite develops in each ootheca. 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
odtheca through a hole 4 mm. in diameter. (Cros, 1942.) Partheno-
genesis 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, Ha-
wali (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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 243

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 re-
lationship by dissecting two specimens of Z. splendidula from odthecae
of L. decipiens.

The following information is taken from Genieys (1924): Ovi-
position.—Wasp egg is introduced into the still-soft odtheca before the
wall hardens. Some oodthecae had four oviposition scars but never
contained more than two parasite eggs. Development.—Larva com-
mences development in July or August. Only one larva completes
development, but it eats all the eggs in the odtheca. 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 odthecae of Loboptera decipiens that were parasitized
by Z. splendidula were also hyperparasitized by an eulophid (see
Ss yntomosphyrum 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,
Ig15a).
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. Gur-
ney the odtheca 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 pre-
occupied by Blatticida Ashmead, 1904. In 1923 Gahan and Fagan re-
named Blatticida Girault, Blatticidella. [Burks, p. c., 1956.]

Natural host.—Cockroach, Australia, Queensland (Girault, 1915).

244 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Cheiloneurus viridiscutum (Girault)

Synonymy.—Cristatithorax Girault = Cheiloneurus Westwood
[ Mercet, 1921 |.

Natural host—Ellipsidion australe, Australia, Queensland (Dodd,
IQI7).

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. How-
ever, in the body of the paper, he states that the supposed origin of
the parasite was the odtheca of B. germanica. Burks (personal com-
munication, 1956) does not believe that this wasp parasitizes the eggs
of B. germanica. We (unpublished data, 1957) exposed six oothecae
of B. germanica to C. merceti. In order to retard water loss the
odthecae were removed from the females by cutting the insects in two
so that each odtheca remained attached to the posterior pars of the
abdomen. No wasps developed in these odthecae.

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 odtheca
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 oothecae
less than a week old and in embryos in the green band stage. There
were I-50 oviposition punctures per odtheca. (Lawson, 1954a.)

Development.—lf enough wasp larvae were present, they ate all
eggs in an odtheca. Occasionally wasps developed in one end of an
odtheca while cockroaches developed in the other ; when this occurred,
the cockroach nymphs always emerged last. The developmental pe-
riod was 30-41 days at room temperature. There were 5-25 parasites
per odtheca. The single exit hole in the ootheca varied from 0.6 to 0.9
mm. in diameter. (Lawson, 1954a.)

Distribution—U.S.A.: New Jersey south to Florida, west to Illi-
nois, Kansas, and Arizona. West Indies; Central and South America ;
Hawaii. (Burks, personal communication, 1956.)

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 245

Dicarnosis alfierii Mercet

Natural hosts —“Phyllodromia’ sp., Egypt (Mercet, 1930) : Ac-
cording to Mercet, Dr. Alfieri claimed that this wasp parasitized one
of the species of “Phyllodromia”’ found in Egypt, namely, Phyllo-
dromia [=Blattella] germanica, Phyllodromia [=Supella] supel-
lectilium and/or Phyllodromia treithana. 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*
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, 19542).

Experimental hosts.—Blatta orientalis, Eurycotis floridana, and
Periplaneta americana, U.S.A. (Roth and Willis, 19544).

Adult behavior—Female wasps are sexually receptive almost im-
mediately on leaving the odtheca. 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 ootheca with her
sheathed ovipositor until she finds an acceptable spot; she then drills
through the wall of the odtheca with her ovipositor. One female ovi-
posited for 5 hours, but briefer periods were more usual. We have
seen six or more females ovipositing simultaneously into an ootheca
of Eurycotis floridana. One female was seen to feed on material that
oozed from the oviposition puncture. The wasp (pl. 34, A) may ovi-
posit into the odtheca of E. floridana while it is still being carried by
the female, as well as in odthecae that have been dropped and which
have hard walls. Eggs 36 days old were successfully parasitized.

Development.—In Eurycotis floridana: In the laboratory, de-
velopment was completed in 34-36 days at about 85° F. This time was
regulated to some extent by the number of parasites in the ootheca.

7 Brues et al. (1954) include this family in the Encyrtidae.

246 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

There is evidence that larvae eat unhatched wasp eggs or other larvae.
In 34 odthecae 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 odthecae 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 some-
times developed but usually failed to hatch. Adult wasps made one
to six emergence holes in the odtheca; the average number in 42
oothecae was two holes.

Number of parasites per odtheca.—In Blatia orientalis: One of
III odthecae exposed to female wasps yielded 48 parasites. In Eury-
cotis floridana: One oodtheca parasitized in the field yielded 68 para-
sites; 8 odthecae exposed to single wasps for their entire lifespan
yielded an average of 50+6 parasites (range 23-81); 34 oothecae
exposed to many wasps for their entire lifespan yielded an average of
198+8 parasites (range 93-306). In Periplaneta americana: Nine
odthecae of 152 exposed to the wasps were found to be parasitized
when dissected; 11 adults emerged from one odtheca; no parasites
emerged from the other 8 oothecae.

Sex ratio—4 92:1 ¢ from ovipositions by isolated females. In
the one ootheca collected in the field, the ratio was 21.6 99:1 ¢
Parthenogenesis exists ; the unfertilized eggs produced only males.

Anastatus tenuipes Bolivar y Pieltain

Synonymy.—Anastatus blattidarum Ferriere. Dr. C. Ferriere (per-
sonal 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 de-
scription agrees with A. blattidarum. I had not yet knowledge of
Bolivar’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 (Fer-
riére, 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 cock-
roach host. The wasp rarely flies but hops proficiently; when dis-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 247

turbed, it can hop from several inches to several feet. The female licks
up the drop of fluid that oozes from the oviposition puncture. Fe-
males die in a few days, but if fed honey and water may live two
weeks. Oviposition—The female selects an odtheca 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 ovi-
posited simultaneously into a single odtheca; a single female re-
peatedly oviposited into one odtheca at intervals. Development.—
Completed in an average of 32.6 days at a constant temperature of
82> F.

Number of parasites per odtheca.—Average about 10.7 (range 4-
16) (Flock, 1941); 15 (Ferriére, 1935).

Sex ratio—4 92:1 # (Ferriére, 1935); average of 6 99:1 ¢
(Flock, 1941). Parthenogenesis occurs; the unfertilized eggs pro-
duced 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
odtheca which contained 96 unidentified chalcids had 16 dentations at
the edge; the description fits the odtheca 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 Entomo-
logical Society in 1838 Mr. Sells exhibited 94 specimens of a small

248 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Pteromalus (apparently identified by Westwood) obtained from one
cockroach odtheca. This same record of Sells was published posthu-
mously in 1842, although in this paper he identified the host odtheca as
“Blaberus” maderae. Cameron (1955) lists a European record of
Pteromalus sp. from Pertplaneta 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 oodtheca of P. pensylvanica yielded 14 parasites with a sex ratio
of 2.5 92: 1 & (Judd, 1955). The average number of parasites in
11 odthecae of Parcoblatta sp. collected in 1950-51 was 27 wasps (Ed-
munds, 1952a, 1953a). The adults made two to three emergence holes
in the odtheca (Edmunds, 1953a; Judd, 1955).

Family EULOPHIDAE
Melittobia chalybii Ashmead

Natural host.—Periplaneta americana, U.S.A., Missouri (Rau,
1940a): MM. chalybu 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,
I9I7).
Syntomosphyrum blattae Burks
Natural hosts——Parcoblatta sp., U.S.A., Ohio (Burks, 1952; Ed-
munds, 1952a, 1953a): Ten odthecae yielded an average of 92 wasps
(Edmunds, 1952a). Five odthecae, collected a year later, yielded an
average of 74 wasps; adults sometimes made two to three exit holes
in the odtheca (Edmunds, 1953a).
Cockroach, U.S.A., West Virginia (Burks, 1952).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 249

Syntomosphyrum ischnopterae (Girault)

Synonymy.—E pomphaloides ischnopterae Girault [Peck, 1951].

Parker and Thompson (1928) called their hyperparasite Tetrasti-
chus sp. However, Dr. B. D. Burks (personal communication, 1955)
has examined the teneral specimens which Parker and Thompson de-
posited in the U.S. National Mueseum; he stated that the species is
apparently Syntomosphyrum ischnopterae. In view of the experi-
mental work by Parker and Thompson (see below), this wasp may
prove to be a hyperparasite on evaniids in cockroach oothecae rather
than a primary parasite on cockroach eggs. (See Zeuxevania splen-
didula, 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 odthecae of Lobop-
tera 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 odthecae that were parasitized by
Zeuxevania, never into normal, nonparasitized oothecae. Oviposition.—
Oviposition occurred two days after mating. The female wasp stroked
the odtheca with her antennae, selected a site, and bored into the
ootheca 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 odthecae. Sex ratio.—5 29:1 f (from 3 odthecae).

Distribution—uvU.S.A., District of Columbia, Maryland (Burks,
1952).

Tetrastichus australasiae Gahan

Natural host—Periplaneta australasiac, Sumatra (Gahan, 1923).

Tetrastichus hagenowii (Ratzeburg)

Synonymy.—Entedon hagenowti Ratzeburg, Blattotetrastichus hag-
enowi (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 hagenowti and concluded that T.
asthenogmus is only a weakly developed specimen of T. hagenowitt.

250 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

He stated, “The longitudinal dorsal grooves of the scutellum, which
are strongly developed in normal hagenowi, 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 ge-
nerically 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.
hagenowwti 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 rec-
ord is based on one parasitized odtheca. We have exposed, at three
different times, groups of 10 to 20 odthecae 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 glos-
sinae Wtstn., a parasite of tsetse fly pupae (Jordan, 1956), Formosa
(Takahashi, 1924; Sonan, 1924); Palestine (Bodenheimer, 1930) ;
Puerto Rico (Sein, 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 odthecae 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 com-
munication, 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 odtheca of P. americana collected on shipboard.
Burks (personal communication, 1955) stated that the wasp was prob-
ably T. hagenowit.

Periplaneta australasiae, Australia (Shaw, 1925) ; India (Usman,
1949) ; Saudi Arabia, Trinidad (Cameron, 1955) ; Formosa (Sonan,
1924).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 251

Periplaneta brunnea, U.S.A., Florida (parasitized odthecae were
collected near Orlando, by members of the Orlando Laboratory, En-
tomology 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 ootheca.

“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. hagenowit was a primary parasite
of eggs of P. americana because the parasitized odtheca 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. hag-
enowi 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 acci-
dental, the eulophid happening to oviposit into an odtheca already con-
taining 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 ac-
complished in from “several” to 20 seconds (Takahashi, 1924; Ed-
munds, 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 (Sein, 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).

252 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

In Hawaii, Severin and Severin (1915) caught 571 T. hagenown in
10 kerosene traps that were set up to sample populations of Mediter-
ranean fruitfly. Apparently the parasite is attracted by the odor of
kerosene.

Oviposition—The female wasp explores the surface of the ootheca
with vibrating antennae (Edmunds, 1955). She bends her abdomen
ventrad and repeatedly touches the surface of the ootheca with her
valvae; when she finds an acceptable oviposition site, the wasp un-
sheathes her ovipositor and bores through the wall of the ootheca
(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 odtheca and more than one wasp oviposited
into the same odtheca (Roth and Willis, 1954b; Edmunds, 1955).
We found freshly laid wasp eggs in 34 empty but previously para-
sitized odthecae from which the wasps had emerged (Roth and Willis,
1954b).

Development.—In Periplaneta americana: Development is com-
pleted 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 odtheca; the
larger the number of wasps (up to an average of about 70 wasps per
ootheca), the shorter the time required to complete development.
Wasps in odthecae 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 o6theca, 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 odtheca; 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 odtheca (Usman, 1949; Roth and
Willis, 1954b).

Number of offspring per female.—In Blatta orientalis: In the
laboratory, 5 odthecae were left with each of 25 female wasps for their

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 253

entire lifespans; of the 125 odthecae, 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 odthecae
was exposed to a single female wasp for 24 hours; the average
number of offspring per female was 103 (range 50-139). Five
oothecae were left with each of 38 females for their entire lifespans ;
of the 190 oothecae, 81 were parasitized. The average number of off-
spring per female was 94 (range 45-168 [from original data]) (Roth
and Willis, 1954b).

Number of parasites per ootheca.—In Eurycotis floridana: In the
laboratory, 3 oothecae 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 ootheca yielded 73 parasites (Pemberton, 1941). In Parcoblatta
sp.: Two oodthecae yielded an average of I00 parasites (Edmunds,
1953a). In Pertplaneta americana: 100 parasites per odtheca (Sein,
1923); 140 (Schmidt, 1937); 25 (Rau, 1940a); 7-38, average 33
(Usman, 1949); 71 (Wolcott, 1951); 4 oothecae exposed to 20 fe-
male 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 oothecae yielded an average of 93 parasites
(range 12-187) (Edmunds, 1955). In Periplaneta australasiae: Oo-
thecae yielded an average of 40-50 adult parasites (Cameron, 1955) ;
about 50 (Shaw, 1925).

Sex ratio —3 92:1 g (Usman, 1949); 4 92:1 g (Cameron,
1955); 2-8 99:1 ¢ (Roth and Willis, 1954b); 1.2 99:1 ¢ (Ed-
munds, 1955). Parthenogenesis exists; the unfertilized eggs pro-
duced 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; For-
mosa; Hawaii.

Tetrastichus periplanetae Crawford

Natural hosts—Periplaneta americana, Mozambique (Crawford,
1910); Union of South Africa (parasites reared from odthecae
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).

254 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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. hagenowu.

Natural hosts—Periplaneta americana, Union of South Africa
(parasites reared from odthecae collected in Durban, Natal, by the
City Health Department [ Burks, p. c., 1956]).

Periplaneta australasiae, Manila, Philippine Islands (Burks, p. c.,
1950).

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. hagenowti because of its brilliant
steel-blue color.

Natural host.—Blatia orientalis, Algeria (Cros, 1942): Adult be-
havior.—Mating began as soon as wasps emerged from an ootheca.
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 odthecae 6, 22, 40,
and 43 days old, and the parasites developed successfully. More than
one female oviposited into the same odtheca. Oviposition was of long
duration. Development.—From egg to eclosion took an average of 34
days in summer (range: 30-38 days, 5 odthecae), and an average of
67 days in fall (range: 58-73 days, 3 odthecae). An average of 55
parasites developed per odtheca (range 21-105, 5 odthecae) ; over 130
wasps emerged from a sixth odtheca. Sex ratio.—10-20 99:1 ¢.

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 odthecae that had been cemented to
the substrate than in clean odthecae that had simply been dropped.
Cros (1942) experimented with two females of P. punctata to see if
the wasps could find odthecae 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 mis-
took the location of the odthecae. 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 odtheca.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 255

The extent of host selection varies among these parasites; some
species will oviposit into the eggs of more than one species of cock-
roach, but others show some degree of host specificity. Positive selec-
tion 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
odtheca, recently removed from Eurycotis floridana, to C. merceti; no
wasps developed ; we had similar negative results with C. merceti and
odthecae of B. germanica. We (1954b) could not induce Tetrastichus
hagenowii to parasitize eggs of Blattella germanica, B. vaga, or Parco-
blatta virginica in the laboratory. In our experiments, T. hagenowu
oviposited into eggs of Supella supellectilinm, but the wasp eggs either
failed to hatch, or if they hatched, the larvae died before completing
development. Neither would T. hagenowti parasitize eggs of N.
rhombifolia (Roth and Willis, unpublished data, 1957). Anastatus
tenuipes would not parasitize the eggs of Latiblattella lucifrons He-
bard, 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
ootheca, 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 com-
munication, 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 com-
munication, 1956). Marshall (1866) suggested that the braconid
Paxylomma buccata Bréb., which he found frequenting cockroach

256 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 im 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 Am-
pulex have been described, the prey of only a small number of species
have been discovered, but the known prey are all cockroaches.

Ampulex amoena Stal

Synonymy.—Ampulex novarae Saussure [| Krombein, personal com-
munication, 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 ; Koh-
riba, 1957).

Kamo (1957) observed that in the field both males and females
sucked juices from wounds they made in the stems of Clerodendron

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 257

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 cock-
roach 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 (per-
sonal 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., Mis-
souri (Williams, 1928a, 1929): figure 6.

258 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL TAT

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).

Fic. 6.—Ampulex canaliculata attacking Parcoblatta virginica. A, Female
wasp stinging her prey, c.X 4.8. B, Wasp’s egg attached to the coxa of the
mesothoracic leg of the cockroach. C, Larva of A. canaliculata (about three-
quarters grown) feeding on the internal organs of the host from the exterior,
c.X 4. (Reproduced from F. X. Williams [1929], through the courtesy of Dr.
F, X. Williams and F. A. Bianchi, Hawaiian Sugar Planters’ Association. )

Distribution.—U.S.A.: Connecticut south to Georgia; Ohio, Wis-
consin, Missouri, Kansas; in open woods (Krombein, 1951).

Ampulex compressa (Fabricius)

(PI. 35)

Synonymy.—Gueépe ichneumon of Réaumur [| Williams, 1929] ;
Chlorion (Ampulex) compressum.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 259

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—WNeostylopyga rhombifolia, Perplaneta amert-
cana, 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. as-
similis. 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 99, supplied with a
cockroach per day, stored an average of 57+14 cockroaches; 8 99
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, maxi-
mum 140 days (Williams, 1942). About 6 weeks (Swezey, 1944).
Longevity of adults—13 99 lived an average of 110+11 days (min-
imum 31, maximum 159); several ¢¢ 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 im 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

260 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 mis-
identification 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 con-
sidered 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 stbirica Fab.
Natural hosts—Cockroaches, West Africa (Perkins im 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 ready-made 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 cornic-
ulus. 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]. Ber-
land (1925) listed D. haemorrhous separately but stated that it is per-
haps a color variety of D. corniculus.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 261

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—FParcoblatta 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 (Ar-
nold, 1928): The wasp is “usually seen running up and down the

262 ' SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141

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—tThe 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. ( Wil-
liams, 1918, 1919; Williams et al., 1931.)

Dolichurus sp.

Natural hosts—Cockroaches, nymphs, South Africa (Bridwell,
1917). Adult female cockroach carrying an odtheca, France (Deleur-
ance, 1943).

Nesting site——Plant stem, or in ground possibly an old abandoned
nest of Ammophile. Behavior —bBridwell 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 263

ee

J

Bf
Tey

y, hf tH Vif

Y444'2,/),
Me,

44,

Uy
las si
be VY by
ify Vf,
Yi Mp ff
< 4 VY ,
Yj Y)

fa

a
¢
ype
“4
Nfl?

iN
VN

GM Ui, 7
fry ie

ify is
ip Ye
Ni

Fic. 7.—Cockroach-hunting wasps. A, Dolichurus stantomi leading a nymph
of Blattella lituricollis to her nest, c.X 4. (Reproduced from F. X. Williams
[1919].) B, Podium haematogastrum attaching her egg to an Epilampra sp.
while on the side of a termite mound that contains the wasp’s nest, c. X 1.6. C,
Epilampra sp. parasitized by P. haematogastrum showing the wasp’s egg at-
tached to the right fore coxa, c. X 3.2. (B and C reproduced from Williams
[1928], through the courtesy of Dr. F. X. Williams and F. A. Bianchi.)

2604 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Trirhogma sp.

Natural hosts——Cockroaches, Oriental region (Williams, 1918,
1928): As far as is known species of this genus of wasps hunt cock-
roaches.

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 Grap-
toblatta 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 265

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 uro-
sternum.

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 cock-
roaches are stored in each nest. The prey is not immobilized as a re-
sult of the sting, and its antennae are left intact.

Podium carolina Rohwer

Natural host——Parcoblatia 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—uvU.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. haemato-
gastrum.

Podium flavipenne Lepeletier

Natural host.—Epilampra abdomen-nigrum, British Guiana (Wil-
liams, 1928): Nesting site—Burrows, about 2 inches deep and len-
ticular in cross section, are dug in the ground in well-drained, partly
sheltered areas; also old Podiwm 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 cock-
roaches 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

266 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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, Para (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 col-
lected, 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)

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 267

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. [un-
doubtedly 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. sjéstedi 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 cock-
roaches 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.

268 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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,
IQI5).

Lasius alienus (Forster)

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 (Illing-
worth, 1914, 1942): The ants followed and killed N. cinerea and P.

surinamensis as they burrowed in moist soil and attacked and de-
stroyed JN. cinerea in breeding cages.

XIII. VERTEBRATA
GlassJPISCES

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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 269

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.2 Captain William Owen (in Web-
ster, 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®
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.

8archy (Marquis, 1931) was living in a dream world when he typed:

“there is always
something to be thankful
for you would not
think that a cockroach
had much ground
for optimism
but as the fishing season
opens up i grow
more and more
cheerful at the thought
that nobody ever got
the notion of using
cockroaches for bait”

9 Classification of Amphibia and Reptilia follows Hegner (1936).

270 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI

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 cock-
roaches. 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS) 271

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 (Giinther)

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): Cock-
roach wings were found in the stomach of a frog caught around hu-
man 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 cock-
roaches.

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 con-
tained 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.

272 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Neostylopyga rhombifolia, U.S.A. (Dr. T. Eisner, personal com-

munication, 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 speci-
men 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 pre-
ferred 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 REP TELA
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 tor-
toise house, Zoological Gardens, Regent’s Park, and the terrapin ate it.

Order SAURIA
Family GEKKONIDAE
Gekko gecko (Linnaeus)
Natural prey.—Cockroaches, Philippine Islands, Laguna ( Villa-
dolid, 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 (Villa-
dolid, 1934): Bulk of stomach contents of 22 lizards consisted of
Orthoptera, mostly cockroaches.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 273

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 Sym-
ploce flagellata, Puerto Rico (Wolcott, 1924) : The last-named cock-
roach 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 ex-
amined, 16 contained Orthoptera, including cockroaches. Puerto Rico
(Wolcott, 1924): One hundred A. cristatellus had eaten 8 cock-
roaches, 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—WNeostylopyga rhombifolia, U.S.A. (Eisner,
personal communication, 1958.)

274 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Anolis grahami Garman

Natural prey.—Periplaneta spp. and Blattidae, Bermuda (Sim-
monds, 1958): Stomachs of 176 lizards yielded 6 cockroaches.

Anolis leachi Duméril and Bibron

Natural prey.—Periplaneta spp. and Blattidae, Bermuda (Sim-
monds, 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 Giinther

Common name.—Spiny lizard.
Natural prey.—Cockroaches, Philippine Islands, Laguna (Villa-
dolid, 1934) : Food of this species was mostly Blattidae.

Unidentified skinks

Natural prey.—Cockroaches, Arno Atoll (Usinger and La Rivers,
1953).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 275

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 kingit
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 Peri-
planeta 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 con-
tents of 18 out of 4o 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.

276 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL.) LAT:

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, per-
sonal 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 in-
sectivorous 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 cock-
roaches. 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 cock-
roaches.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 277

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 I 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 dimidia-
tipes, Hawaii (Schwartz and Schwartz, 1949).

278 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Domestic chicken

Natural and experimental prey.—Blaberus cranitifer, U.S.A., Key
West, Florida. J.A.G. Rehn in 1912 (personal communication) ob-
served 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 cock-
roaches that were caught in traps.

Hebardina concinna, Japan (Asano, 1937): Experimental feeding
to white Leghorn chickens.

Periplaneta americana, Surinam (Stage, 1947): Several cock-
roaches 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.

Pynoscelus surinamensis, Australia (Fielding, 1926); Formosa,
experimental feeding (Kobayashi, 1927); Australia, experimental
feeding (Fielding, 1927, 1928) ; U.S.A., Florida, experimental feed-
ing (Sanders, 1928); Antigua (Hutson, 1943); Hawaii (lIlling-
worth, 1931; Schwabe, 1949, 1950a, 1950b). This cockroach is the
intermediate host of Oxyspirura mansom, 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).

' BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 279

Pigeon

Experimental prey—Pycnoscelus surinamensis, Australia (Field-

ing, 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
Dendrocopos mahrattensis (Latham)

Synonymy.—Liopicus mahrattensis [Friedmann, p. c. 1957].
Common name.—Y ellow-fronted pied woodpecker.
Natural prey.—Cockroaches, India, Central Provinces (D’Abreu,

1920).

280 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 odthecae 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.—bBlue 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 re-
leased by the insect on being attacked by the bird, had dissipated.

Family PARADISEIDAE

Paradisea papuana Bechstein

Experimental prey.—Cockroaches, Malaya and on shipboard (Wal-
lace, 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.”

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 281

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 any-
where else in Hawaii. The birds followed tractors that were destroy-
ing cactus and kept close to the chain that turned over the stumps.
The following species were collected: Allacta similis, Blattella ger-
manica, Cutilia soror, Diploptera punctata, Leucophaea maderae, Peri-
planeta americana, Periplaneta australasiae, Pycnoscelus surinamensis.
Illingworth did not state whether the birds ate all these species in-
discriminately.

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.

282 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Family ICTERIDAE
Agelaius xanthomus (Sclater)

Common name.—Y ellow-shouldered blackbird.
Natural prey.—Cockroaches, Puerto Rico (Wetmore, 1916) : Oo-
thecae 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 (odthecae) 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): Ani-
mal food was found in 5 of 72 stomachs examined; one bird had
eaten two cockroaches among other insects.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 283

Class MAMMALIA”

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 Pros-
thenorchis 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.

10 Classification of Mammalia follows Simpson (1945).

284 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4T

Family LORISIDAE
Loris tardigradus (Linnaeus)
Synonymy.—Lemur tardigradus [Dr. D. H. Johnson, personal com-

munication, 1957].
Natural prey.—Cockroaches, on board ship (Cowan, 1865).

Perodicticus potto (P. L. S. Miller)

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 John-
son, 1939): Captive monkeys became naturally infested with Proto-

spirura 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 John-
son, 1939): See comment under Aotes zonalts.

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 John-
son, 1939): Favorite item of food in the laboratory. See comment
under Aotes zonalis.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 285

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 communica-
tion, 1957].

Natural prey.—Blattella germanica, France (Brumpt and Urbain,
1938) : The monkey apparently became infested naturally with Pros-
thenorchis 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 cock-
roaches 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 chrys-
oleucos.

Leontocebus ursulus (E. Geoffroy)

Natural prey.—Blattella germanica, France (Brumpt and Urbain,
1938) : See comments under Callithrix chrysoleucos.

286 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Ur-
bain, 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, 19382): 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 Wie-
gand Bruss, 1946) : Indirect evidence for this relationship was shown
by these workers who found two animals heavily infected with Pros-
thenorchis 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—Oothecae of Blatta orientalis and Neostylopyga
rhombifolia, Thailand (Bristowe, 1932).
Periplaneta americana, Formosa (Takahashi, 1924).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 287

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 communica-
tion, 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 cock-
roach which is the intermediate host of the worm.

Periplaneta americana, Brazil (Magalhaes, 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).

288 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 14!

Family CAVIIDAE
Cavia sp.

Experimental prey—Blattella germanica, U.S.A. (Hobmaier,
1941): Guinea pigs were fed cockroaches infested with Physaloptera
maxilaris.

Order CARNIVORA
Family CANIDAE
Canis familiaris Linnaeus

Experimental prey.—Blattella germanica, U.S.A. (Hobmaier,

1941) : Dogs were fed cockroaches infested with Physaloptera maxil-

laris. 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 cock-
roaches 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 Pros-
thenorchis 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 289

Nasua sp.

Natural prey.—Cockroaches, British Guiana (Beebe, 19254).

Family MUSTELIDAE
Meles sp.

Experimental prey.—Blattella germanica, France (Brumpt and Ur-
bain, 1938a): A badger was successfully infected when fed cock-
roaches infested with Prosthenorchis elegans and P. spirula.

Family VIVERRIDAE
Herpestes javanicus auropunctatus Hodgson

Natural prey—Epilampra wheeleri, Eurycotis improcera, Pan-
chlora 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 Pi-
mentel).

Pimentel (personal communication, 1958) has given us the follow-
ing percentage occurrence of cockroach species in the total number
of mongoose stomachs that he examined in Puerto Rico: Epilampra
wheelert 1.8, Ischnoptera rufa rufa 3.6, Panchlora nivea 1.8, Peri-
planeta 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.—FPeriplaneta 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 ma-xil-
laris. U.S.A. (Petri and Ameel, 1950): Cockroaches infested with
Physaloptera rara were fed to a kitten. France (Brumpt and Ur-

290 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

bain, 1938a): A young cat was fed cockroaches infested with Pros-
thenorchis elegans and P. spirula.

Felis pardalis mearnsi J. A. Allen

Natural prey.—Cockroaches, Panama (Dr. H. L. Sweetman, per-
sonal communication, 1958) : An ocelot was seen collecting and feed-
ing on cockroaches, possibly Blaberus sp. ““The ocelot was quite effi-
cient and seemed to relish the roaches.”

XIV., CHECKLIST {OF |} COCKROACHES AND: SYMBIOTIC
ASSOCIATES

Only naturally occurring associations are included in this list. Com-
mensal 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 in-
cluded 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 stantont, p. 262. Solindenia picticornis, p. 247.

Anaplecta sp.
Fungus: Herpomyces anaplectae,
p. 134.
Aptera fusca

Protozoan: Gregarina fastidiosa,
p. 182.

Arenivaga bolliana

Insect: Sarcophaga oman, p. 220.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 29QI1

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.
Nematodes :

Leidynema appendiculata, p. 197.

Entamoeba coli, p. 178.

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.

Fungi:
Herpomyces tricuspidatus, p. 138.
Penicillium sp., p. 131.
Protozoan: Diplocystis (?) sp.,
pp. 181, 184.
Nematodes:
Leidynema cranifera, p. 108.
Mite: Jolina nana, p. 219.
Undetermined, p. 220.
Bird :? *Ghieken}p.'278:

Escherichia freundii, p. 113.

Micrococcus pyogenes var. albus,
p. 106.

Micrococcus pyogenes var. aureus,
p. 107.

Proteus vulgaris, p. 114.

Pseudomonas aeruginosa, p. 104.

Rhizopus nigricans, p. 133.
Saccharomyces cerevisiae, p. 133-

Protrelleta floridana, p. 199.

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.
Protozoan: Leptomonas blaberace,
p. 167.

Herpomyces periplanetae, p. 137.
Herpomyces tricuspidatus, p. 138.

to

Ne)
No

Protozoan:

Bacterium:

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Blaptica dubia

Pileocephalus blaberae,
p. 184.

Blatta (Shelfordella) lateralis

Shigella paradysenteriae,

p. 119.
Insect: Ampulex assimilis, p. 257.

Blatta orientalis

Bacteria:

Aerobacter aerogenes, p. III.

Alcaligenes faecalis, p. 111.

Alcaligenes recti, p. III.

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 biitschli, p. 120.

Bacillus periplanetae, p. 121.

Bacillus stellatus, p. 121.

Bacillus subtilis, p. 122.

Bacillus tritus, p. 122.

Bacteroides uncatus, p. 110.

Clostridium lentoputrescens, p. 122.

Clostridium novyi or Clostridium
Sporogenes, p. 122.

Enterococcus sp., p. 100.

Escherichia coli, p. 112.

Lactobacillus fermenti, p. 109.

Micrococcus pyogenes var. albus,
D: Loz:

Micrococcus pyogenes var. aureus,
p. 107.

Fungi:

Aspergillus fumigatus, p. 130.

Blastocystis hominis, p. 133.

Blastocystis sp., p. 133.

Candida zeylanoides, p. 129.

Coccidioides periplanetae, p. 133.

Protozoa:

Balantidium praenucleatum, p. 187.
Bodo blattae, p. 167.
Coelosporidium periplanetae, p. 185.

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, 1009.
Sarcina symbiotica (host may have
been B. germanica), p. 108.
Sarcina ventriculi, p. 108.
Serratia marcescens, p. 117.
Spirillochaeta blattae, p. 127.
Spirillum a, B and y, p. 105.
Spirillum sp., p. 105.
Spirochaeta periplanetae, p. 125.
Streptococcus faecalis, p. 109.
Streptococcus liquefaciens, p. 110.
Streptococcus microapotka, p. 110.
Streptococcus pyogenes, p. 110.
Streptococcus sp., p. 110.
Treponema parvum, p. 125.
Treponema stylopygae, p. 125.
Vibrio sp., p. 106.

Herpomyces periplanetae, p. 137.
Herpomyces stylopygae, p. 137.
Torula gropengiesseri, p. 132.
Torulopsis sp., p. 130.

Diplocystis schneideri, p. 181.
Endamoeba blattae, p. 177.
Entamoeba thomsoni, p. 179.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 293

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.

Helminths :

Ascaris sp., p. 200.

Enterobius vermicularis, p. 200.

Gongylonema neoplasticum, p. 206.

Gordius blattae orientalis, p. 202.

Hammerschmidtiella diesingi, p. 195.

Hammerschmidtiella neyrai, p. 1096.

Mite: Undetermined, pp. 220, 222.

Insects :

Dermestes ater, p. 234.
Dermestes sp., p. 234.
Evania appendigaster, p. 236.
Evania dimidiata, p. 230.

Nyctotherus ovalis, p. 188.
Oikomonas sp., p. 166.
Peltomyces periplanetae, p. 177.
Plistophora kudot, p. 185.
Plistophora periplanetae, p. 186.
Plistophora sp., p. 186.
Retortamonas blattae, p. 167.
Stenophora, sp., p. 181.
Tetratrichomastix blattidarum,

p. 170.

Leidynema appendiculata, p. 197.
Spirura gastrophila, p. 207.
Thelastoma pachyjuli, p. 201.
Trichostrongylus sp., p. 210.
Trichuris trichiura, p. 210.

Prosevamia punctata, p. 240.
Systellogaster ovivora, p. 248.
Tetrastichus hagenowitt, 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 hagenowt, p. 250.

Blattella germanica and/or Blattella vaga

Viruses: Unspecified strains of polio-
myelitis virus, p. 103.

Blattella germanica

Bacteria:
Achromobacter sp., p. 110.
Aerobacter aerogenes, p. 111.

Micrococcus epidermidis, p. 106.
Micrococcus pyogenes var. albus,

Aerobacter cloacae, p. 112. p. 107.
Alcaligenes faecalis, p. 111. Micrococcus pyogenes var. aureus,
Alcaligenes viscosus, p. 111. p. 107.

Bacillus circulans, p. 120.
Escherichia coli, p. 112.
Escherichia freundii, p. 113.
Micrococcus aurantiacus, p. 106.

Micrococcus ureae, p. 107.
Micrococcus sp., p. 108.
Mycobacterium leprae, p. 123.
Paracolobactrum aerogenoides,

pi 113:

204

Paracolobactrum coliforme, p. 113.

Paracolobactrum sp., p. 113.

Pseudomonas aeruginosa, p. 104.

Sarcina symbiotica (host may have
been B. orientalis), p. 108.

Fungi:

Aspergillus flavus, p. 130.

Aspergillus tamari, p. 130.

Aspergillus sp., p. 130.

Cordyceps blattae, p. 134.

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.

Helminths:

Blattelicola blattelicola, p. 193.
Blatticola blattae, p. 193.
Enterobius vermicularis, p. 200.
Galebia aegyptiaca, p. 195.
Gongylonema neoplasticum, p. 206.

Mites:

Blattisocius tineivorus, p. 216.
Caloglyphus sp., p. 218.

Insects :

Brachygaster minutus, p. 235.
Eupelmus atriflagellum, p. 247.

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.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Salmonella typhimurium, p. 116.
Serratia marcescens, p. 117.
Streptococcus faecalis, p. 109.
Streptococcus sp., p. 110.

Herpomyces ectobiae, p. 135.
Memnoniella echinata, p. 132.
Saccharomyces sp., p. 133.

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.

Moniliformis kalahariensis, p. 203.
Prosthenorchis elegans, p. 203.
Prosthenorchis spirula, p. 203.
Tetrameres americana, p. 207.
Trichuris trichiura, p. 210.

Undetermined, p. 220.

Dolichurus corniculus, p. 261.
Ripidius pectinicornis, p. 232.

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.

BIOTIC ASSOCIATIONS OF COCKROACHES—-ROTH & WILLIS 295

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:
Sarracemia 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 wenyom, 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,
Dp. 172.

Monocercomonoides globus, p. 169.

Notila proteus, p. 170.
Oxymonas doroaxostylus, p. 170.
Amphibian: Plethodon glutinosus,

p. 2609.

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.

296 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Cutilia soror

Insects :
Dolichurus stantoni, p. 262. sepligetella sericea, p. 242.
Evania appendigaster, p. 236.

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,
poer7:
Fungus: Herpomyces diploptcrac,
p. 135.
Mites:
Tolina 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. 160.
Nematode: Blatticola blattae, p. 193.
Insects :
Brachygaster minutus, p. 235. Tachysphex lativalvis, p. 264.

Dolichurus corniculus, p. 261.

Ectobius pallidus

Protozoan: Gamocystis tenax, p. 184.

Helminths:
Blatticola blattae, p. 193. Undetermined mermithid, p. 192.
Insects :
Ampulex fasciata, p. 250. Ripidius denist (presumptive record),
Aphaenogaster picea, p. 267. pyeses
Dolichurus corniculus, p. 261. Tachysphex lativalvis, p. 264.

Lasius alienus, p. 268.
Ripidius boissyt (presumptive rec-
ord), p. 231.

Ectobius panzeri

Centipede: Scolopendra sp., p. 224.
Insects :
Dolichurus corniculus, p. 261. Tachysphex lativalvis, p. 264.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 297

Ectobius sp.

Insects :
Brachygaster minutus, p. 235. Tachysphex lativalvis, p. 264.
Dolichurus corniculus, p. 261.

Ellipsidion affine

Insect: Rhipidioides fuscatus, p. 230.

Ellipsidion australe
Insects :
Agamerion metallica, p. 243. Mestocharomyia oophaga, p. 248.
Cheiloneurus viridiscutum, p. 244.

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. 2606.

Bird: Gymnasio nudipes, p. 279.

Epilampra wheeleri
Reptiles :
Ameiva exsul, p. 275. Anolis cristatellus, p. 273.
Mammal: Herpestes javanicus auro-
punctatus, p. 280.
Escala (?) sp.

Insect: Rhipidioides adynatus, p. 230.

Eurycotis floridana

Fungi:

Aspergillus flavus, p. 130. Aspergillus sydowt, p. 130.
Helminths :

Euryconema paradisa, p. 194. Protrelloides paradoxa, p. 200.

Leidynema appendiculata(?), p. 108. Undetermined gordian worm, p. 202.
Insects :

Anastatus floridanus, p. 245. Undetermined tachinid, p. 228.

Eurycotis improcera

Mammal: Herpestes javanicus auro-
punctatus, p. 289.

298 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Eurycotis manni

Fungus: Herpomyces szanzibarinus,
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 sanzibarinus,
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 auro-
punctatus, p. 280.

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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 299

Leucophaea maderae

Bacterium: Serratia marcescens,
powr7e
Fungus: Herpomyces tricuspidatus,
p. 138.
Protozoa:
Gregarina rhyparobiae, p. 183. Retortamonas (?) sp., p. 167.
Hexamita (?) sp., p. 171. Undetermined gregarine, p. 184.
Nematodes:
Hammerschmidtiella diesingi, Protospirura bonnet, p. 206.
p. 195. Protospirura muricola, p. 206.
Leidynema delatorrei, p. 108.
Mite: Chaetodactylus sp., p. 218.

Mammals:
Aotes zonalis, p. 284. Cebus capucinus, p. 284.
Ateles dariensis, p. 284. Rattus norvegicus, p. 287.

Leucophaea sp.
Nematode: Cephallobellus brevicauda-
tum, Pp. 194.
Leurolestes pallidus

Fungus: Herpomyces leurolestis,

p. 136.
Loboptera decipiens
Insects :
Dolichurus corniculus, p. 261. Zeuxevama splendidula, p. 243.

Loboptera sp.

Fungus: Herpomyces lobopterae,
p. 136.

Lobopterella dimidiatipes

Insect: Ampulex canaliculata, p. 257.
Bird: Coturnix coturnix japonica,

Dp. 277.
Melanosilpha capensis
Protozoa:
Gregarina impetuosa, p. 183. Gregarina sandont, p. 183.

Monastria sp.

Insect: Triatoma arthurneivai, p. 227.

Nauphoeta cinerea

Bacteria :
Salmonella typhimurium, p. 116. Serratia marcescens, p. 117.

300 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Fungus: Herpomyces tricuspidatus,

p. 138.
Mites:

Blattilaelaps nauphoetae, p. 216. Locustacarus sp., p. 219.
Insect: Pheidole megacephala, p. 268.

Neostylopyga rhombifolia

Bacterium: Serratia marcescens,
pe nn7.
Insects :
Evania appendigaster, p. 237. Tetrastichus hagenowitt, p. 250.
Szepligetella sericea, p. 242.
Mammal: Homo sapiens, p. 286.

Nyctibora obscura
Fungus: Herpomyces amazonicus,
p. 134.
Nyctibora sp.
Fungus: Herpomyces nyctoborae,
p. 136.
Nyctibora tomentosa
Fungus: Herpomyces nyctoborae,
p. 136.
Oniscosoma granicollis

Insect: Paranephrites xents, p. 230.

Panchlora exoleta

Protozoan: Gregarina panchlorae,
p. 183.

Panchlora nivea

Bacterium: Serratia marcescens,
Dp. TL7:

Fungus: Herpomyces panchlorac,
p. 136.

Mammal: Herpestes javanicus auro-
punctatus, p. 280.

Panesthia angustipennis

Protozoa:
Clevelandella constricta, p. 189. Clevelandella hastula, p. 189.
Clevelandella contorta, p. 189. Clevelandella panesthiae, p. 189.

Clevelandella elongata, p. 180. Clevelandella parapanesthiae, p. 189.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 301

Endamoeba javanica, p. 178. Nyctotherus uichancoi, p. 188.
Endamoeba philippinensis, p. 178. Paraclevelandia brevis, p. 190.
Hexamita cryptocerci, p. 171. Paraclevelandia simplex, p. 190.
Monocercomonoides panesthiae, Undetermined amoeba, p. 181.
p. 170.
Nematodes:
Aorurus philippinensis, p. 193. Leidynemella fusiformis, p. 198.
Blattophila sphaerolaima var. ja- Leidynemella paracranifera, p. 198.
vanica, p. 104. Thelastoma palmettum, p. 201.

Leidynema nocalum, p. 108.

Panesthia australis
Fungus: Metarrhizium anisopliae,
Ds 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 fustformis,

p. 198.
Panesthia lobipennis

Fungus: Herpomyces panesthiae,

p. 136.
Panesthia spadica
Protozoa:
Clevelandella constricta, p. 189. Endamoeba javanica, p. 178.
Clevelandella contorta, p. 189. Nyctotherus uichancoi, p. 188.
Clevelandella nipponensis, p. 189. Paraclevelandia brevis, p. 190.
Clevelandella panesthiae, p. 189. Paraclevelandia simplex, p. 190.

Panesthia sp.

Nematode: Leidynemella panesthiae,

p. 198.
Parahormetica bilobata
Helminths :
Agamospirura parahormeticae, Undetermined gordian worm, p. 202.
p. 205.

Parcoblatta lata

Plant: Sarracenia flava, p. 154.
Protozoan: Leptomonas sp., ,. 167.
Nematode: Protrellus aurifluus, p. 199.

302 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Parcoblatta pensylvanica

Protozoa:
Gregarina blattarum, p. 182. Gregarina thomasi, p. 184.
Gregarina illinensis, p. 183. Leptomonas sp., p. 167.
Gregarina parcoblattae, p. 183. Nyctotherus ovalis, p. 188.
Nematode: Rictularia coloradensis,
p. 205.
Scorpion: Centruroides vittatus (?°),
Dp. 2t2:
Insects :
Camponotus pennsylvanicus, p. 267. Hyptia thoracica, p. 240.
Hyptia dorsalis, p. 239. Podium carolina, p. 265.
Hyptia harpyoides, p. 239. Systellogaster ovivora, p. 248.

Hyptia reticulata, p. 240.
Amphibian: Rana (?) sp., p. 272.

Parcoblatta uhleriana
Fungus: Herpomyces arietinus,

p. 134.

Protozoan: Gregarina parcoblattae,
p. 183

Nematode: Protrellus aurifluus,
p. 190.

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. 210.

Insects :
Coenosia basalis, p. 220. Syntomosphyrum blattae, p. 248.
Dolichurus greenei, p. 261. Systellogaster ovivora, p. 248.
Hyptia sp., p. 240. Tetrastichus hagenowti, p. 250.

Megaselia sp., p. 227.

Periplaneta americana

Viruses: Unspecified strain(s) of
poliomyelitis virus (host may have
been P. brunnea), p. 103.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 303

Bacteria:

Achromobacter hyalinum, p. 110.
Aerobacter aerogenes, p. III.
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 freundu, 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,
D. E13:
Paracolobactrum coliforme, p. 113.
Paracolobactrum sp., p. 113.
Proteus mirabilis, p. 114.
Proteus morganti, p. 114.
Proteus rettgeri, p. 114.
Proteus vulgaris, p. 114.
Proteus sp., p. 114.
Pseudomonas aeruginosa, p. 104.
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.

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.

Pseudomonas fluorescens, p. 105.

Salmonella anatis, p. 114.

Salmonella morbificans, p. 115.

Salmonella schottmuelleri, p. 115.

Salmonella sp. (Type Bareilly),
p. II5.

Salmonella sp.
p./ 115:

Salmonella sp. (Type Kentucky),
p. II5.

Salmonella sp. (Type Meleagris),
p. 116.

Salmonella sp.
p. 116.

Salmonella sp. (Type Oranienburg),
p. 116.

Salmonella sp.
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. 1009.

Streptomyces leidynematis, pp. 196,
197.

Tetragenous sp., p. 127.

Veillonella parvula, p. 1009.

(Type Bredeny),

(Type Newport),

(Type Panama),

Mucor guilliermondii, p. 132.
Mucor sp., p. 132.
Penicillium sp., p. 131.
Rhizopus sp., p. 133.
Syncephalastrum sp., p. 133.
Torula acidophila, p. 132.

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.

304 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Eutrichomastix sp. (host may have

been B. germanica), p. 160.
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.
Helminths :
Ancylostoma duodenale, p. 209.

Ascaris luwmbricoides or Ascaris

suum, p. 209.
Binema mirsaia, p. 193.

Gongylonema neoplasticum, p. 206.

Gongylonema sp., p. 206.

Gordius sp., p. 202.

Hammerschmidtiella diesingi,
p. 195.

Hymenolepis sp., p. 208.
Mites:

Caloglyphus sp., p. 218.

Pimeliaphilus podapolipophagus,

p. 210.
Spiders:

Avicularia avicularia, 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.
Amphibian:

Hyla cinerea, p. 270.
Reptiles:

Ameiva exsul, p. 275.

Birds:

Chicken, p. 278.
Mammals:

Felis catus, p. 280.

Herpestea javanicus auropunctatus,

p. 289.
Herpestes sp., p. 280.

Lophomonas blattarum, p. 172.
Lophomonas striata, p. 173.
Monocercomonoides orthopterorum,
p. 160.
Nyctotherus ovalis, p. 188.
Plistophora periplanetae, p. 186.
Protomagalhaesia serpentula, p. 184.
Tetratrichomastix blattidarum,
p. 170.

Leidynema appendiculata, p. 197.
Moniliformis dubius, p. 203.
Moniliformis moniliformuis, p. 204.
Necator americanus, p. 210.
Protrellus kiinckeli, p. 199.
Schwenkiella icemi, p. 200.
Spirura gastrophila, p. 207.
Thelastoma pachyjuli p. 201.
Trichuris trichiura, p. 210.

Rhizoglyphus tarsalus, p. 218.
Tyrophagus noxius, p. 218.
Undetermined, p. 220.

Avicularia sp., p. 214.

Ripidius pectinicornis, p. 232.
Spiniger domesticus, p. 227.
zepligetella sericea, p. 242.
Tetrastichus hagenowiti, p. 250.
Tetrastichus periplanetae, p. 253.
Tetrastichus sp., p. 254.
Trirhogma caerulea, p. 262.

Anolis cristatellus, p. 273.
Sparrow, p. 282.
Homo sapiens, p. 286.

Rattus norvegicus, p. 287.
Rattus rattus, p. 287.

Periplaneta australasiae

Bacteria:
Mycobacterium leprae, p. 123.

Serratia marcescens, p. 118.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 305

Fungus: Herpomyces periplanetae,
p. 137.
Plant: Sarracenia minor, p. 154.
Protozoa:
Endamoeba blattae, p. 177. Endolimax blattae, p. 180.
Nematodes:
Gongylonema neoplasticum, p. 206. Protrellus australasiae, p. 199.
Hammerschmidtiella diesingi, p. 195. Protrellus kiinckeli, p. 200.
Moniliformis dubius, p. 203.
Centipedes: Undetermined, p. 222.

Insects:
Ampulex amoena, p. 256. Tetrastichus australasiae, p. 249.
Ampulex compressa, p. 259. Tetrastichus hagenowii, p. 250.
Evania appendigaster, p. 237. Tetrastichus, sp., p. 254.
Ssepligetella sericea, p. 242. Trirhogma caerulea, p. 262.
Reptiles :
Anolis cristatellus, p. 273. Chrysemys picta, p. 272.
Bird: Gallus sp., p. 277.
Mammals:
Herpestes sp., p. 280. 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,
ps 072:
Helminths: Thelastoma riveroi, p. 201. Moniliformis moniliformis, p. 204.
Insects :
Ampulex compressa, p. 259.
Evania subspinosa (presumptive rec- Szepligetella sericea, p. 242.

ord?), p. 239.
Amphibian: Bufo marinus, p. 270.

306 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 melo-
lonthae, p. 160.

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. Hammerschmudtiella diesingi, p. 195.
Galebia aegyptiaca, p. 195. Oxyuris (?) heterogamiae, p. 199.
Polyphaga saussurei

Helminths :
Hymenolepis sp., p. 208. Undetermined tapeworm ova, p. 208.
Insects: Undetermined reduviids,
Dp. 227.

Polyzosteria melanaria (?) or Platyzosteria analis

Nematode: Protrellus aureus, p. 190.

Pycnoscelus surinamensis

Bacterium: Serratia marcescens,
p. 117.
Fungus: Mucor sp., p. 132.
Protozoa:
Undetermined ciliate, p. 190. Undetermined gregarine, p. 184.
Undetermined flagellate, p. 176.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 307

Nematodes :

Oxyspirura mansoni, p. 204. Severianoia severianoi, p. 200.
Mites:

Caloglyphus spinitarsus, p. 217. Tyrophagus lintneri, p. 218.

Histiostoma feroniarum, p. 217. 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. Passer domesticus, p. 282.
Ducks, p. 277. Phasianus sp., p. 277.
Melanerpes portoricensis, p. 280. Streptopelia chinensis, p. 278.

Meleagris gallopavo, p. 278.
Mammal: Herpestes javanicus auro-
punctatus, p. 280.

Robshelfordia circumducta or Robshelfordia longiuscula

Insects :
Neorhipidius neoxenus, p. 230. Rhipidioides mollis, p. 231.
Rhipidioides helenae, p. 230.

Steleopyga (?) sinensis

Nematode: Suifunema caudelli, p. 200.

Supella supellectilium

Viruses: Unspecified strain(s) of
poliomyelitis virus, p. 103.

Bacterium: Serratia marcescens,
Ptr.

Fungus: Herpomyces supellae, p. 138.

Nematode: Blattophila supellaima,

p. 104.
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.

Mite:

308

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

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. 100.
Micrococcus pyogenes var. aureus,
p. 107.
Mycobacterium leprae, p. 123.
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.

Plants:

Nepenthes ampularia, p. 154.
Nepenthes gracilis, 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.

Helminths:

Cephalobellus magalhdaesi, 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.

Scorpion: Heterometrus longimanus,

p. 213.

Spiders :

Ctenid, p. 214.

Heteropoda venatoria, p. 215.

Pimeliaphilus podapolipopha-

gus, p. 219.

Centipedes :

Allothereua maculata (circumstantial
evidence), p. 223.

Scolopendra morsitans, p. 223.

Paracolon bacilli, p. 113.

Proteus morgant, p. 114.

Proteus vulgaris, p. 114.

Spirillum sp., p. 105.

Spirochaeta periplanatae, p. 125.

Streptococcus sp., p. 110.

Vibrio Types I and II Heiberg,
p. 1006.

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 sanzibarinus, p. 138.
Metarrhizium anisopliae, p. 131.

Nepenthes sp., p. 154.

Monocercomonoides orthopterorum,
p. 169.

Nyctotherus buissoni, p. 188.

Nyctotherus ovalis, p. 188.

Nyctotherus viannai, p. 180.

Oikomonas blattarum, p. 166.

Oikomonas sp., p. 166.

Paramecium sp., p. 186.

Trichomonas sp., p. 171.

Schwenkiella icemi, p. 200.

Severianoia magna, p. 200.

Severianoia severianoi, p. 200.

Spirura gastrophila, p. 207.

Undetermined gordian worms,
p. 202.

Latrodectus mactans p. 216.

Scolopendra subspinipes, p. 224.
Scutigera coleoptrata (circumstantial
evidence), p. 222.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 309

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 japanica, p. 226.
Dolichurus bicolor, p. 260.
Dolichurus corniculus, p. 261.
Dolichurus gilberti, p. 261.
Dolichurus ignitus, p. 201.
Dolichurus sp., p. 262.

Dorylus nigricans sjéstedi, p. 267.
Dorylus sp., p. 267.

Dorylus wilverthii, p. 267.
Eciton burchelli, p. 268.

Fish:

Chalceus macrolepidotus, p. 260.

Cyrtocharax magdalenae essequiben-
Sis, p. 269.

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.

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.

Birds:

Acridotheres tristis, p. 281.
Agelaius xanthomus, p. 282.

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 picticormts, 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 hagenowni, p. 251.
Tetrastichus periplanetae, p. 253.
Trirhogma sp., p. 264.
Undetermined tachinid, p. 220.

Potamotrygon humboldti, p. 268.
Rhamdia sebae, p. 268.

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.

Geckos, p. 273.

Gekko gecko, p. 272.
Hemuidactylus 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.

Bambusicola thoracica, p. 277.
Chickens, p. 278.

310 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Coturnix coturnix japonica, p. 277. Partridge, p. 278.
Dendrocopos mahrattensis, p. 279. Phasianus sp., p. 277.
Ducks, p. 277. Sparrow, p. 282.
Gymnasio nudipes, p. 279. Tiaris bicolor omissa, p. 282.
Gymnopithys leucaspis, p. 280. Tockus birostris, p. 270.
Holoquiscalus brachypterus, p. 282. Troglodytes aedon, p. 281.
Icterus portoricensis, p. 280. Troglodytes audax, p. 281.
Owl, p. 351.

Mammals:
Bassariscus astutus, p. 288. Homo sapiens, p. 287.
Callithrix jacchus, p. 285. Loris tardigradus, p. 284.
Erinaceus sp., p. 283. Molossus sp., p. 283.
Felis catus, p. 280. Monodelphis sp., p. 283.
Felis paradalis mearnsi, p. 290. Nasua nasua, p. 288.
Herpestes javanicus auropunctatus Nasua sp., p. 289.
Ferpestes 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 com-
mensal 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 bene-
fit 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.—S phecophila ravana, Ceylon (Fernando, 1957): Six
females, 50 males, and nymphs of both sexes were found among de-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 3II

caying timber in the ground in association with a colony of this ter-
mite. The antennae of most specimens were mutilated unsymmetri-
cally.

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.—S phecophila termitium, Kibonoto, East Africa (Shel-
ford, 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.—N octicola sinensis, Repulse Bay, Australia(?) (Sil-
vestri, 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 Fsperanza
(Caudell, 1923): Three males collected in the joint nests of C.
femoratus and Crematogaster limata.

312 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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.—M yrmecoblatta 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 Phor-
ticolea testacea, Brazil, San Leopoldo (Bolivar, 1905) : Found in the
formicaries of C. rufipes.

Formica rufibarbis Fabricius
and
Formica subcyanea Wheeler

Commensal.—M yrmecoblatta 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,
Huasan, Argentina? (Bruch, 1916).

Acromyrmex lundi (Guérin)

Synonymy.—Atta lundi [ Brown, personal communication, 1957].

Commensal.—Attaphila bergi, Argentina and Uruguay (Bolivar,
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. lundt.

Acromyrmex niger (F. Smith)

Synonymy.—Atta nigra Schupp [ Brown, p.c., 1957].

Commensal.—Attaphila schuppi, Brazil, Porto Alegre (Bolivar,
1905): Found outside the nest of the ant and mixed in the columns
of ants on the march.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 313

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 (Bolivar, 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 en-
countered in Huasan 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 (Bolivar,
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 uninten-
tional, it is peculiar that Bolivar (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.

314 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI

Crematogaster limata parabiotica Forel

Commensal.—Phorticolea boliviae, Bolivia, Cachuela Esperanza
(Caudell, 1923): Collected in joint nests of C. limata and Compo-
notus femoratus.

Solenopsis geminata (Fabricius)

Commensal.—M yrmecoblatta 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 myr-
mecine ant.

Subfamily PONERINAE
Odontomachus affinis (Guérin)

Commensal.—M yrmeblattina longipes, Brazil, Rio de Janeiro
(Chopard, 1924, 1924a; Hancock, 1926): Originally described as
Phileciton longipes by Chopard (1924) from the nest of an ant mis-
takenly identified as Eciton sp.

Family VESPIDAE
Polybia pygmaea Fabricius

Commensal.—S phecophila 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 be-
tween the cockroach and the bees. [See comment by Sonan (1924) on

page 318.]

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS

315

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: Acromyrmesx lobicornis,
D5, Bre)
Acromyrmex lundi, p. 312.
Acromyrme.x silvestrit, p. 313.
Attaphila flava
Host unknown, presumably an ant,
p. 314.
Attaphila fungicola

Ants: Acromyrmex octospinosus,

Dp. 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(?),
pu Bre.

Formica rufibarbis, p. 312.

Formica subcyanea, p. 312.

Myrmecoblatta wheeleri

Ant: Solenopsis geminata, p. 314.

Nocticola sinensis

Termites: Macrotermes barnecyi,
jo Spite

Odontotermes sp., p. 311.
Nocticola termitophila

Termites: Macrotermes malaccen-

SIS; DP. 3IL.
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,
pashi
Crematogaster limata para-
biotica, p. 312.
Phorticolea testacea

Ant: Camponotus rufipes, p. 312.

Sphecophila polybiarum
Wasp: Polybia pygmaca, p. 314.

Sphecophila ravana
Termite: Coptotermes ceylonicus,
p. 310.
Sphecophila termitium

Termite: Macrotermes bellicosus or
Macrotermes natalensis,

p. Sit.

316 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

OBSCURE ASSOCIATIONS

Cockroaches that are sometimes found in the nests of, or in asso-
ciation 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 myrme-
cophilous (Donisthorpe, 1900). Mann (1911) found an “/schnop-
tera’ 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 Vero-
messor andrei (Mayr) [=Stenamma andrei]. Hebard (1917) re-
ported that W. M. Wheeler collected Eremoblatta subdiaphana in
Arizona as an ant guest. Rehn (1906a; Rehn and Hebard, 1927) re-
ported 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 Myrmucaria nata-
lensis 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. (He-
bard, 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 ootheca of Aristiger histrio (sp. ?) between leaves (Costus sp.)

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 7,

that had been stuck together by a thysanopteron, Anaphothrips sp.
He pointed out that the odtheca would not adhere to leaves that were
not stuck together but would fall to the ground.

Sein (im Rehn and Hebard, 1927) in Puerto Rico found Aglaop-
teryx facies in abandoned cocoons of Megalopyge krugiu (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. krugw 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 absimulis also living in the abandoned cocoon
of M. krugit on bucare trees in Puerto Rico. Wolcott (1950) re-
ported that Plectoptera dorsalis, Plectoptera infulata, and Plectoptera
rhabdota have been found living in trees between leaves or in “‘butter-
fly-nests” of Tetralopha scabridella in leaves of Inga vera, or nests of
Pilocrocis secernalis (Moschler) in the leaves of Petitia domingensis
in the mountains of Puerto Rico. Sein (im 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 jamatcanus 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 sym-
biotic 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., Peri-
planeta 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 re-
treat in the nests.

318 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. TAT

McClure (1936) obtained a large nest of Vespula maculata (Lin-
naeus) [=Vespa maculata] in March in IIlinois. 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. Al-
though we do not imply that a commensal relationship exists between
Parcoblatta and the wasp, it is well to recall a statement by Roth-
schild 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 up-
set. One partner can then suddenly take a mean advantage of the
other.”

Cockroach nymphs may enter bees’ nests where, according to Ima-
mura (im Sonan, 1924), they do not feed on honey or pollen but pre-
sumably 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 be-
lieved that the blattid was a species peculiar to the nests of birds.
Three nests of Ploceus sp. in Ivory Coast yielded one or two cock-
roaches each in association with more numerous mites, Psocoptera,
Heteroptera, beetles, and lepidopterous larvae (Delamare Deboutte-
ville and Paulian, 1952). These last cited workers also found four
cockroaches in a nest of Estrildine sp., and two in a nest of an un-
determined bird, all in association with other arthropods. Moulton
(1912) observed large numbers of Symploce cavernicola and Peri-
planeta australasiae swarming in soft bird guano on the floor of caves
in Borneo. Abdulali (1942) found in India many Periplaneta ameri-
cana in caves containing the edible-nest swift ; there was no indication
of association of the cockroaches with the birds. Danforth (in Wol-
cott, 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).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 319

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 Pert-
planeta 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 ASuEREDATORS
INTERSPECIES PREDATION

Tepper (1893) made the broad statement that the majority of
Australian and Polynesian cockroaches appear to be wholly carnivo-
rous, 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 ro-
bustus 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 con-
firmation.

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 Ar-
gentine ant Iridomyrmex humilis. In the laboratory, Eurycotis flori-
dana 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

320 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

long. Annandale (1910) described the destruction in Calcutta of ter-
mites by Periplaneta americana. During a heavy rain storm many ter-
mites flew into the dining room and were set upon by the cockroaches
which seized them with their mandibles and began to gnaw their abdo-
mens. 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 observa-
tion 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, im 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, Blat-
tella germanica, and/or Periplaneta americana) dislodge and eat the
mite Pimelaphilus 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 cock-
roaches “‘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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 21

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 hy-
pothesis 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 im Shelford, 1gogb).

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 [cock-
roaches] 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. New-
man (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 cock-
roach or two to be used to hunt bedbugs.

In Siberia, Burr (1926, 1939) found blaitella germanica and bed-
bugs inhabiting the same room. Mellanby (1939) studied the popula-
tions 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 cock-
roaches would not touch living bedbugs or their eggs (Cornwall,
1916). In laboratory experiments Gulati (1930) found that Peri-

322 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

planeta americana ate young bedbugs which had soft, blood-filled ab-
domens; 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 bed-
bugs 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 cock-
roaches and Phlebotomus are mentioned together is a paper by Whit-
tingham 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 in-
sects. 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 Peri-
planeta 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 un-
doubtedly 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! (Saupe, 1928). Edmunds (1957)

11 The West Indian “Blabera fusca Brunner” of Saupe (1928) is obviously
Blaberus craniifer as can readily be seen from a comparison of Saupe’s figures of
the pronotal shields of his species with the descriptions by Hebard (1917) of
B. craniifer and Blaberus atropos (Stoll). The Chilean B. fusca Brunner is a
junior synonym of B. atropos (Stoll), a South American insect (Hebard, 1917).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 323

reported that cannibalism was common in a laboratory colony of Pevi-
planeta brunnea and that egg capsules deposited by a female were
often eaten by the other cockroaches.

Periplaneta americana occasionally ate other cockroaches and their
odthecae 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 Pertplaneta americana may eat their
own eggs (Klein, 1933). Some females may regularly eat their 00-
thecae as soon as they are dropped (Griffiths and Tauber, 1942). To
be completely eaten an odtheca generally must be attacked before it
has hardened. If a hole is eaten in one side of the capsule, the cock-
roach may devour the eggs and leave a portion of the odtheca. Fre-
quently 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
oéthecae; in this species only the soft end of the recently deposited
odtheca was eaten (Roth, unpublished data, 1957).

Cros (1942) observed odthecae-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 popu-
lation density and cannibalism.

324 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141

German cockroaches may attack newly molted nymphs of their
own kind and cause them to deflate (Gould and Deay, 1938). Lhéri-
tier (1951) has observed the hatching nymphs of B. germanica being
devoured by their congeners even before they have left the ootheca.

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 cock-
roaches 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 simi-
lar 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 surinamen-
sis. 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, cock-
roaches exhibit additional symbiotic relationships among themselves.
These relationships are (1) the familial associations of parent and
offspring, (2) gregariousness, (3) intraspecies fighting, (4) inter-
species compatibility, and (5) interspecies antagonism. There are
some inconsistencies between observations made on the same species

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 325

by different workers, which only further observation and experimenta-
tion will explain. Some of the reported observations are unique; this
is especially true for the feral species. Because of the paucity of in-
formation, 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
ootheca after it has been formed. The extent of this association be-
tween the mother and her developing progeny varies from the mini-
mum amount of time spent by oviparous females in concealing their
oothecae, 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 ootheca. The odtheca failed to adhere to the shallow hole and fell
to the floor. After several futile attempts to replace the odtheca 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 ootheca 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
oothecae 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 oothecae therein. After coating the odthecae with
more sticky secretion, the females adjusted the oothecae so that the
keels were uppermost and then carefully concealed the odthecae with
the excavated debris. Both females spent over an hour in the act.
Rau (1924) previously reported that of go odthecae deposited by B.
orientalis in jars containing earth and trash, 36 were placed in crey-
ices 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 cap-
sule was deposited in the secretion and covered with additional froth,
which hardened into a very strong cement. Some females spent as

326 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. IAI

long as two hours coating the capsule after it was deposited. It was
not stated whether the odtheca was otherwise concealed. The female
remained with her body over the ootheca 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 odthecae,
and that if the females were frightened away from this activity, they
returned again to complete it. As well as excavating holes in the sub-
strate in which to deposit its odthecae, 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 oothecae in crevices in floating logs just above the water line.
However, Shelford (1906) stated that four genera (including Epilam-
pra) of the subfamily Epilamprinae are “viviparous,” in which event
the females would carry their odthecae within their bodies during
embryogenesis and would not place the oothecae in crevices in logs.

The female of Cryptocercus punctulatus was observed to make a
groove in a piece of wood, then carry her ootheca 6 inches from where
she had dropped it and place it in the groove; she covered the ootheca
so that only a portion was visible (Cleveland im Cleveland et al.,
1934). Dr. W. L. Nutting (personal communication, 1954) collected a
number of odthecae 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 ootheca 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 odtheca behind the earthen
barricade which she had erected.

In summary, the following species of oviparous cockroaches have
been observed concealing their oothecae (only those references not
previously cited are given): Blatta orientalis; Cryptocercus punctu-
latus; Ectobius sylvester (Harz, 1956, 1957) ; Epilampra sp.; Eury-
cotis floridana (Roth and Willis, 1954a) ; Loboptera decipiens; Balta
scripta, Methana curvigera, Methana marginalis, and Methana caneae
(Pope, 1953a); Pelmatosilpha marginalis, Pelmatosilpha purpura-
scens, and Nauclidas nigra (Bunting, 1956) ; Periplaneta americana
(Haber, 1919; Adair, 1923; Sein, 1923; Nigam, 1933; Gould and
Deay, 1938; Rau, 1940a) ; Periplaneta australasiae (Girault, 1915b;
Spencer, 1943; Pope, 1953) ; Periplaneta brunnea (Roth and Willis,

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 327

unpublished data, 1958) ; Periplaneta fuliginosa (Gould and Deay,
1940); Periplaneta ignota (Pope, 1953); Supella supellectilium
(Flock, 1941). Undoubtedly other oviparous species that drop their
oothecae long before the eggs hatch also make some attempt to con-
ceal the oothecae by placing them in crevices or covering them with
debris.

Sometimes the odthecae are deposited but not concealed. Hafez
and Afifi (1956) reported that in Egypt Supella supellectilium at-
taches its odtheca 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 odthecae in places
which are dry and raised above the ground, although rarely one also
may find some odthecae scattered on the ground. For lack of suitable
material females of Periplaneta americana sometimes did not con-
ceal their oothecae (Nigam, 1933). Frequently in laboratory colonies
P. americana merely drops the odthecae 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 ootheca anywhere (Bunting,
1956). Rau (1940) stated that the female of Parcoblatta pensylvanica
does not conceal her odtheca. However, Gould and Deay (1940)
stated that this species deposits its odthecae loosely behind bark.
Ellipsidion affine and Ellipsidion australe attach their odthecae 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
odthecae may have been deposited by these oviparous females before
the eggs of the first odthecae hatch. However, a different behavior is
encountered among species that do not form a second ootheca 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

328 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 punctu-
latus, 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 ootheca clasped in her
genital cavity with the posterior portion projecting behind her. Each
normal o6theca is carried for approximately the duration of embryo-
genesis 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
odtheca; 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 neces-
sarily a familial association by placing nymphs from two oothecae to-
gether. 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 odthecae 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 ovi-
parity and viviparity in the Blattaria.) Shelford (1906, 1916) re-
ported that he had captured a female of Pseudophoraspis nebulosa in
Borneo with numerous young nymphs clinging to the undersurface of

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 329

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 speci-
men from Luzon, Philippine Islands, of the apterous female of Peris-
phaerus glomeriformis with nymphs still clinging to her undersur-
face; 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 ootheca 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 en 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. Sein (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. Pessoa and Corréa (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 fe-
males of Perisphaerus armadillo surrounded by pale, yellowish-white
young ; similarly he had observed that Archiblatta hoevenii was found

330 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 ob-
served similar behavior in laboratory colonies of B. crantifer and
Leucophaea maderae.

Chopard (1950) noted that after hatching the young of Grompha-
dorhina 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 (unpub-
lished data, 1958) have seen young nymphs of Gromphadorhina por-
tentosa 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 Nau-
phoeta 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) ob-
served 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 exten-
sive 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 en-
countered in the literature. There has been some argument to the

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 331

effect that large numbers of these insects seeking the same environ-
ment in a limited space would appear to be gregarious, whereas there
is probably no true social tendency (Rau, 1924). Reactions of cock-
roaches 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 as-
sembling by the attraction of the milieu and by gregarious instincts.
Chopard (1938) also stated that Orthoptera with a gregarious tend-
ency are found rarely isolated; one finds them, on the contrary
nearly always collected in the same shelters, close together, as if con-
scious 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 gre-
gariousness 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 so-
cial 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 re-
sult 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 fre-
quently among the domiciliary species. A few examples will suffice.
Gal’kov (1926) observed heavy infestations of undetermined cock-
roaches in workers’ living quarters in the Ural region: “In the cor-
ners near the stove, the cockroaches covered the walls in a dense
carpet.” After fumigating he collected about 135,000 dead cock-
roaches from one barracks and about 475,000 from another. We
have reviewed a few other examples of heavy infestations in our

1957(a) paper.

332 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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.” Inthe monkey house of the Ham-
burg 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 cock-
roaches ceased to bite and fight each other when they crowded to-
gether 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 resent-
ment 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 re-

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 333

mained 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 (Sein, 1923). Wolcott (1950) also
stated that L. maderae is gregarious. Annandale (1900) observed that
in the “Siamese Malay States” large colonies of Periplaneta australa-
siae 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., Ewrycotis
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 in-
dividuals. Hebard (1917) in his discussion of Lattiblattella rehm
again mentioned finding frequent colonies of E. floridana in Florida.
He also found many specimens of Blaberus cranufer 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 “Par-
coblatta [sp.] were never recorded as being gregarious, in fact they
were mostly solitary. However, in some cases two were found to-
gether.” 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 rub-
bish (Blatchley, 1920).

Rehn and Hebard (1927) quoted observations made earlier by He-
bard on Byrsotria fumugata 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 ob-
servers also reported that Aspiduchus borinquen was found in Puerto

334 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Aspi-
duchus 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
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
cranifer. 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 Lito-
peltis 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 in-
creased with the size of the group and with age, as each animal occu-
pied 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 in-
sect 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 ex-
tended 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 in-
dividually 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 335

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 com-
plete 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 supel-
lectiium, 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 Peri-
planeta 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. Re-
moval 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 re-
sponse. 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 court-
ing 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 ap-
parently 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

336 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

species are indications of gregariousness. However, until gregarious-
ness 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 in-
dividuals.

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. Taka-
hashi (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. Ad-
ditional 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 337

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 occasion-
ally 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 im-
proper 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 dif-
ferent 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 be-
tween different species as well as examples of incompatibility, the sub-
ject 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

338 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 de-
cayed trees, and beneath corded wood. Adair (1923) stated that in
his house in Egypt Periplaneta americana, Blatta orientalis, and Blat-
tella 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 Pycno-
scelus surinamensis, in his chicken shed.

Eads (personal communication, 1955), in response to our inquiry
about the mixed populations of cockroaches that he had reported in-
festing 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 ob-
servations the two species always appeared to be perfectly compati-
ble.” Eads et al. (1954) had found Periplaneta fuliginosa in three
manholes, Parcoblatta bolliana in one manhole and Parcoblatta pensyl-
vanica 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 339

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. amert-
cana. 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 Holo-
compsa 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 ger-
manica 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 brownbanded roaches were often found in the same
crevice.”; Anonymous (1958), Texas; and Anonymous (1958a),
Georgia. Gould and Deay (1940) stated that other species of cock-
roaches, especially B. germanica, may be found with S. supellectilium
in the same part of a building. Yet Shaw (1925) stated that “when
Supella supellectiium Serv. invades places already occupied by Blat-
tella 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 asso-
ciated 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 long-leaf 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

340 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 £. lucidus were a distinct species in 1923, Ramme
(1951) later decided that they were a form of E. sylvester, E. syl-
vester 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 differ-
ent collections, 11 from houses and 17 from privies, all in Pharr, Texas. Tabu-
lation 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 as-
sociated 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 terri-
tory (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 /schnoptera podoces was captured in
company with the type series of Cariblatta nebulicola, in dead leaf

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 34I

litter in Jamaica (Rehn and Hebard, 1927). In Florida Periplaneta
australasiae was often taken in company with Pycnoscelus surina-
mensis 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. ger-
manica together, there were no reciprocal attacks even by hungry in-
dividuals. 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 un-
favorable 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. ori-
entalis 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

342 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Queens-
land. 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 be-
hind in Puerto Rico in competition with the little German roach.”
Pessoa and Corea (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. ameri-
cana. 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” [pre-
sumably 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 343

seemed to have overcome the American which had been more numer-
ous 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 re-
solve 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 ex-
periments 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 effective-
ness. 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.

Cotr (1940)

There are very few records indicating that cockroaches are unac-
cepted as food by other animals. Hutson (1943) found that the duck,
guinea fowl, and pigeon would not normally eat Pycnoscelus surina-
mensis, 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 sug-
gested 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 be-
havior, concealment, protective coloration, mimicry, or secretion of
malodorous materials. Nocturnal cockroaches may avoid predators
that are active during the day (Crawford, 1934), but nocturnal pred-
ators are apparently quite successful in capturing cockroaches. Some

344 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

cockroaches may be protected by their swiftness, others by their re-
semblance 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 re-
ported that it appeared to sink into the soil without raising any con-
siderable amount above the surface and that it did not form an unob-
structed 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) ob-
served 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 cranufer'* 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 cock-
roaches; 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

12 The species was recorded by these authors as Blaberus atropos. (Hebard,
1917 p. 204, footnote 327.)

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 345

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 de-
fends itself against attack is never found in a predator’s stomach.

Cockroaches have a variety of glands which secrete odorous ma-
terials. Certain secretions, produced by tergal or dorsal glands in
males, are involved in sexual behavior; the females feed on the secre-
tion 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 mor-
phology 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, Platy-
zosteria 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 dis-
agreeable 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 in-
variably 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

346 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 EF. 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 (Val-
curone 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 secre-
tion 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 im-
mediately 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 Pogo-
nomyrmex badius were not repelled by the secretion of N. rhombi-
folia.

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 Peri-
planeta 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; Koncek,
1924), the functions of secretions of these nonepigamic dorsal glands
and the ventral glands are still open to question. It is possible that

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 347

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 punc-
tata the tracheae leading to the second abdominal spiracles of nymphs
and adults are modified into odoriferous glands which produce a mix-
ture 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 them-
selves 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

348 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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 Hemip-
tera (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 Coc-
cinellidae; Shelford (1912) has figured a number of species of Pro-
soplecta 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 compari-
son 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 bene-
fit in the survival of mimetic species. Certainly Cott’s (1940) volu-
minous compilation of the literature on adaptive coloration should
make the most skeptic hesitate to conclude dogmatically that these in-
stances 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 ex-
termination 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

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 349

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 hagenowti) which suggest that, in the absence of para-
sites, 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 para-
sites in the biological control of cockroaches. However, because of the
paucity of information, we have been unable to evaluate the effective-
ness 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.
(Sein, 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 communica-
tion, 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 pro-
tected 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 cock-
roaches. Takahashi (1924) reported that, in the Taihoku area of
Formosa, human habitations contained large numbers of spiders which
caught and ate cockroaches. Smith (im 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).

350 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4t

Ants.—A Madam Merian noticed that ants cleared houses of cock-
roaches (Kirby and Spence, 1822). A small reddish-yellow ant,
called Pucchucici by Peruvian Indians, pursued and destroyed a cock-
roach 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 cock-
roaches 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 odthecae of Loboptera decipiens were parasitized by
Zeuxevania splendidula (Genieys, 1924). Edmunds (1952a) found
that 12 percent of 459 odthecae of Parcoblatta collected during De-
cember through April of 1950-51 in Ohio were parasitized ; evaniids
accounted for about 7 percent of the parasitization. Additional col-
lection data in 1951-52 Edmunds (1953a) showed that 8.7 percent of
320 wood-cockroach oothecae were parasitized; 2.8 percent of these
parasites were evaniids; almost 13 percent of the egg capsules col-
lected showed evidence of previous parasite emergence. Cameron
(1957) reported that odthecae 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 odthecae 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) (Ferriére, 1930, 1935).
In this country, this wasp seemed to be effective in decreasing the num-
bers of Supella in certain areas in Arizona (Flock, 1941).

In Formosa, Tetrastichus hagenowi was an important parasite of
cockroach eggs (Maki, 1937). Sonan (1924) reported 30 percent
parasitization of 65 odthecae of Periplaneta americana and P. austra-
lasiae collected in Formosa. In Bangalore, India, the natural para-
sitization of randomly collected odthecae of P. americana varied from
21 percent (of 495 odthecae), July 1947—June 1948, and 43 percent
(of 288 odthecae), July-December 1948, to 57 percent (of 178 00-
thecae), July—October 1949 (Usman, 1949). Cameron (1955) ob-
tained T. hagenowii from odthecae collected in Trinidad, B.W.L., and

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 351

Saudi Arabia; some 15 percent of the odthecae 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 odthecae was 65 percent parasitized. The odthecae
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 para-
sitized by T. hagenowii that he had to use P. australasiae for experi-
mental purposes; in 1950 Plank stated that more than 50 percent of
American cockroach odthecae 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 de-
velops in a parasitized odtheca, but many individuals of the other
wasps develop in one odtheca 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 odthecae of this cockroach were parasitized (Zimmer-
man, 1944). We (1954b) ran some simulated field tests in which we
liberated Tetrastichus hagenowii in rooms artificially seeded with
odthecae ; from 28 to 83 percent of American cockroach odthecae and

352 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

56 percent of oriental cockroach odthecae 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 effec-
tive these wasps are in controlling cockroaches is still unknown. Doli-
churus 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 “P/yllo-
dromia.”’ 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 (Wil-
liams, 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; Wil-
liams, 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 notice-
ably 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. Sim-
monds (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 Is-
lands in 1955 against Periplaneta spp. ; it is not yet known whether the
parasite became established in either place (Dumbleton, 1957).

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 353

VERTEBRATES

. On Conserve avec soin les crapauds dans les maisons, et que les
dames les tolérent, méme sous leurs robes, en raison de leurs
continuels services, car ils se proménent sans cesse a la recherche
des Kakerlacs.

GrrarpD (1877)

Toads— Bufo marinus was first introduced into Puerto Rico from
Barbados in 1920 to reduce several major insect pests including cock-
roaches (Leonard, 1933). It was introduced from Puerto Rico into
Hawaii by C. E. Pemberton in 1932 where it rapidly became estab-
lished; 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 I1 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 II
to 25 cockroaches. Illingworth (1941) found that 40 to go percent
of 53 stools of this toad in Hawaii contained remains of P. surina-
mensis. 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 plan-
tations in Australia, these amphibians were encouraged in houses and
kept as pets because they hunted and devoured large brown cock-
roaches (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 An-
tilles (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 cock-roach” and suggested that lizards be used for

354 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I41

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 Pycno-
scelus.

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. Depart-
ment 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.

BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 355

Lawson, Kansas State College (Comperia merceti); Dr. Kee:
Roeder, Tufts University (Hierodula tenmdentata (2) ) De Ea:
Wilson, Harvard University (/schnoptera deropeltifornus).

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. Hel-
minths—Mrs. May Belle Chitwood and Dr. J. T. Mucker, 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. Anis.—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.
Noe Ge Glance, US 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. Spil-
man, U. S. National Museum. Fishes=Dri 1 Ps Schultz, U.S:
National Museum. Amphibians and reptiles——Dr. Doris M. Cochran,
U. S. National Museum. Birds——Dr. Herbert Friedmann, Ures:
National Museum. Mammals—Dr. D. H. Johnson, U. S. National
Museum.

We thank the following individuals for reading the entire manu-
script: J. A. G. Rehn; Dr. A. B. Gurney ; Maj. Gordon Field, LOM Se
Army; and Dr. H. L. Sweetman, University of Massachusetts. The
monograph has profited by the friendly criticism of these entomol-
ogists.

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 manu-
script.

350 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

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Journ. Bombay Nat. Hist. Soc., vol. 43, pp. 446-451.
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1909. Etude d’un Rhipidius nouveau de Provence, R. Boissyi Abeille.

Compt. Rend. Soc. Biol., Paris, vol. 67, pp. 854-858.
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1938. Une curieuse €pizootie vermineuse a acanthocéphales, devenue
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1939. Prophylaxie et traitement utilisés pour combattre les acanthocephales
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1942. A note on Timulla (Timulla) eriphyla Mickel (Hym., Mutillidae),
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1920. A naturalist on Lake Victoria with an account of sleeping sickness
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1921. Experiments on the relative edibility of insects, with special reference
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1925. A naturalist in East Africa. 187 pp., 8 pls. 1 map. Oxford, England.

CASTELLANI, A., and CHALMERS, A. J.
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1905. Notes on some Florida Orthoptera. Ent. News, vol. 16, pp. 216-219.

1914. Some bromeliadicolous Blattidae from Mexico and Central America.
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1923. Phorticolea boliviae, a new myrmecophilous cockroach from South
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1924. Some insects from the Chilibrillo bat caves of Panama. Insecutor
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1925. Pycnoscelus surinamensis Linnaeus (Orthoptera) ; on its nymphs and
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1931. Notes on Blattidae, adventive to the United States (Orthop.). Ent.
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1955. Cycle évolutif d’un ascaride: Subulura jacchi (Marcel 1857) para-
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1942. Notes diverses sur Leucophaea surinamensis (L.) et Nauphoeta

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1949. Insects of agricultural and household importance in Alaska with sug-
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1926. Some factors affecting the propagation of hookworm infections in
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1941. The specific status of Moniliformis (Acanthocephala) of Texas rats,
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1949. Introduction to parasitology. 756 pp. New York.

CHARLES, VERA K.

1941. A preliminary check list of the entomogenous fungi of North

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1933. Ziichtungsversuche an parasitischen Protozoen von Periplaneta
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1934. Nematodes parasitic in Philippine cockroaches. Philippine Journ.

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1919. Zoological results of a tour in the far east. Les Orthoptéres
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1933. The generic names Cephalobellus Cobb, 1920 and Scarabanema
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1930. The symbiosis between the wood-feeding roach, Cryptocercus punctu-
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1931. The effects of molting on the protozoa of Cryptocercus. Anat. Rec.,
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1947. Sex produced in the protozoa of Cryptocercus by molting. Science,
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1947a. The origin and evolution of meiosis. Science, vol. 105, pp. 287-289.

1948. An ideal partnership. Sci. Month., vol. 67, pp. 173-177.

1949. Hormone-induced sexual cycles of flagellates. I. Gametogenesis,
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1950. Hormone-induced sexual cycles of flagellates. II. Gametogenesis,
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1951. Hormone-induced sexual cycles of flagellates. VI. Gametogenesis,
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1952. Hormone-induced sexual cycles of flagellates. VIII. Meiosis in
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1953. Hormone-induced sexual cycles of flagellates. IX. Haploid gameto-
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1954. Hormone-induced sexual cycles of flagellates. X. Autogamy and
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1956. Hormone-induced sexual cycles of flagellates. XIV. Gametic meiosis
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1934. The wood-feeding roach Cryptocercus, its protozoa, and the symbiosis
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1954. Protozoa as indicators of developmental stages in molting of the
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1951. Notes on Arachnida. 16. The behavior of a scorpion. Ent. Month.
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1953. Notes on Arachnida. 19. Biological observations, etc. Ent. Month.
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1933. Observations on the life history of Tetrameres pattersoni. Journ.
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BIOTIC ASSOCIATIONS OF COCKROACHES—ROTH & WILLIS 439

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B, Dead individual showing the bright red coloration (very dark areas) characteristic
of infection with Serratia marcescens. C, Living male. D, Living female.

Pela

VOL. 141,

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female.

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Periplaneta brunnea, c. X 2.9.

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SMITHSONIAN MISCELLANEOUS COLLECTIONS

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 24

Cc D

Pycnoscelus surinamensis, c. X 3.7. A, Male from Hawaii. B, Macropterous par-
thenogenetic female from Florida. C, Brachypterous nonparthenogenetic female from
Hawaii. D, Late instar nymph. (Photograph of nymph D, by Jack Salmon.)

5

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VOL. 141,

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 26

Bacteroids from Blattella germanica. A, Part of abdomen showing mycetocytes in fat body, X 225.
B, Lobe of fat body showing 3 mycetocytes, X 750. C, Single mycetocyte; bacteroids appear hollow
as result of fixation in Carnoy’s fluid, X 1725. D, Smear of fat body showing bacteroids in various
stages, X 1800. (All preparations and photographs through the courtesy of Dr, Marion A. Brooks.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOLES D4 Rs 27,

Fungi parasitic on cockroaches. A, Herpomyces arictinus growing on antennae, legs,
body, and cerci of a nymph of Parcoblatta virginica, X 7. B, Herpomyces stylopygae
on antenna of Blatta orientalis, X 35. (Reproduced from Richards and Smith [10955,
1956].) C, Herpomyces sp. [probably H. stylopygac]| on antenna of B. orientalis, X 132.
(Photographs B and C through the courtesy of Dr. A. G. Richards.)

VOL. 141, PL. 28

MITHSONIAN MISCELLANEOUS COLLECTIONS

S

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 29

A, Undetermined mermithid that parasitizes Ectobius pallidus, X 9. The worm has partly emerged
from the neck region of the cockroach. (Reproduced from Roth and Willis [1957].) B, Undeter-
mined gordian worm that parasitized Eurycotis foridana shown beside its host, X 1.8. (Specimen
courtesy of Dr. T. Eisner.)

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 30

D | \ | Beh

A, Heteropoda venatoria, a cockroach-hunting spider, slightly less than natural size, on
bananas. (Reproduced from a Kodachrome transparency through the courtesy of Dr. B
Kaston.) B to E, Lycosa sp. (avida?) capturing and feeding on a nymph of Supella supel-
lectilium in the laboratory, X 1.4.

PL. 31

VOL. 141,

SMITHSONIAN MISCELLANEOUS COLLECTIONS

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SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 142; PE 33

Cc

A, Prosevania punctata & beside an odtheca of Periplaneta americana, X5. B,
Hyptia harpyoides with ootheca of Parcoblatta uhleriana from which it had emerged,
X 5. C, Larva of a lampyrid beetle feeding on Parcoblatta virginica in the laboratory,

X 4.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 34

ie

C 8 SSVSsSSSESBED

Chalcid parasites of cockroach eggs. A, Anastatus floridanus OVipositing into an oOtheca which is
still being carried by Eurycotis floridana, c.X 4. B, Comperia merceti ovipositing into an oOtheca of
Supella supellectilium, c. X 13. C, Tetrastichus hagenowtt ovipositing into an odtheca of Periplaneta
americana, c. X 10. (C from Roth and Willis [1954b].)

VOES 241; RES 35

SMITHSONIAN MISCELLANEOUS COLLECTIONS

eriplaneta sp. (presumably americana), about natural size.

B. She stings the prey in the thorax. C, She then leads the
disabled cockroach (antennae clipped) to her nest. D, The wasp’s egg was placed on the coxa of the
cockroach’s right mesothoracic leg where it hatched. E, Portion of the host’s abdomen removed to
show feeding larva. F, New adult wasp emerging from dead host. (Reproduced from F. X. Williams
[1942] from the color paintings of the late W. Twigg-Smith, through the courtesy of F. A. Bianchi.)

Ampulex compressa attacking P
A, The wasp finds a cockroach.

SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. 141, PL. 36

Chemical defense of Diploptera punctata against predators; the spray pattern is displayed
on KlI-starch indicator paper. .\, Spray pattern after right mesothoracic leg was pinched.
3, Cumulative spray pattern after left mesothoracic leg of the same insect was pinched.
C, The defensive glands of the cockroach on the left had been excised, and it is under per-
sistent attack by ants from a laboratory colony of Pogonomyrmex badius (Latreille). The intact
cockroach on the right was also attacked by the ants, but it discharged a spray of quinones
and repelled the attackers. (From Eisner [1958], through the courtesy of Dr. T. Eisner.)

INDEX

Plate and page numbers in boldface type indicate illustrations. In general,
entries are placed in the index as unmodified substantives except where a modifier
contributes significantly to the identification of the item (e.g., blue heads, prairie
dog). This index should be used in conjunction with the indexed check list of
natural associations (pages 290 to 310) because these are not repeated here.
All experimental associations are indexed below except those involving Blatta
orientalis, Blattella germanica, and Periplaneta americana that are cited on pages
that also contain references to the natural associations of these three species.

Abies sp., 257
Acacia, 56, 69
Acacia farnesiana, 155, 164
harpophylla, 55
Sp.) 155
Acanthinevania princeps, 235
Acanthocephala, 203
Acanthogyna deplanata, 7
Acaridae, 217
Acarina, 210, 216
Acheta domestica, 320
Achromobacter hyalinum, 110
sp., 110
Achromobacteriaceae, I10
Acridotheres tristis, 281
Acrocomia aculeata, 142
Acromyrmex lobicornis, 312, 313
lundi, 312
niger, 312
octospinosus, 312
silvestrui, 312
Acropyga acutinventris, 317
Actinocephalidae, 184
Actinomycetaceae, 124
Actinomycetales, 123
Adeleidae, 184
Adelina cryptocerci, 184-185
Aechmaea porteoides, 144
Aerobacter aerogenes, 111
cloacae, 112
sp., I12
Aeschnidae, 224
Agamerion metallica, 243
Agamidae, 275
Agamospirura parahormeticae, 205

Agaricus, 164
Agave, 43, 57
Agelaius xanthomus, 282
Agis orientalis, 7, 27
Aglaopteryx absimilis, 7, 35, 156, 317
devia, 7
diaphana, 7, 35, 36, 145, 146, 290,
339
facies, 7, 35, 156, 200, 317, 318
gemma, 7, 36, 140, 143, 144, 153,
158, 159, 339
vegeta, 7, 146
ypsilon, 7, 75
Aircraft, 87-90
Alcaligenes faecalis, 111
rectt, III
ViSCOSUS, III
Algaroba, 45, 156, 164
Allacta similis, 7, 36, 75, 276, 290
Allothereua maculata, 223
Alluadellina cavernicola, 7, 17
Alluaudella cavernicola, 7
Alpinia, 163
Alsophila sp., 139
Alsophilas, 164
Amazonina emarginata, 7, 36, 146, 158
Ameiva exsul, 275
Sp., 275
Amitermes, 102
Amoeba blattae, 177
coli, 178
undetermined sp., 181
Amoebidae, 177
Amoebina, 177
Amphibia, 269-272

442

Amphibians, 3, 353
Amphibolurus barbatus, 275
Amphoromorpha blattina, 139
sp., 139
Ampulex, 2, 256
amoena, 250-257
assimilis, 257
canaliculata, 94, 257-258, 352
compressa, 258-259, 352; pl. 35
compressiventris, 260
fasciata, 259
novarae, 256
ruficornis, 250
siberica, 260
sibirica, 250
sonnerati, 260
Ampulicidae, 256
Anacardiaceae, 157
Ananas comosus, 144
Anaphothrips sp., 317
Anaplecta asema, 7, 36
azteca, 7, 145
decipiens, 7, 36
fallax, 7, 36
hemiscotia, 7, 36
lateralis, 7, 36
mexicana, 7, 145
spp., 88, 145, 290
Anastatus blattidarum, 246
blattidifurax, 245
floridanus, 94, 245, 255, 319; Pl. 34
tenuipes, 94, 246, 255, 350
Anatidae, 277
Anax strenuus, 224
Ancylostoma caninum, 209
ceylanicum, 209
duodenale, 209
Ancylostomidae, 209
Androctonus australis, 211
Aneurina tahuata, 7, 160
viridis, 7, 155, 160, 161
Anguillula macrura, 195
Anisogamia, 26
Annato, 159
Annetta, Steamship, 149
Anoetidae, 217
Anolis carolinensis, 273, 346
cristatellus, 273
equestris, 273, 345, 346

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Anolis—Continued.
grahami, 274
leachi, 274
pulchellus, 273
sagret, 274
Sp., 274
stratulus, 274
Anseriformes, 277
Antagonism, interspecies, 341-343
Antbird, 280
Antibiotics, 97-98, 346
Ants, 57, 266-268, 316, 317, 350
Argentine, 268, 319
army, 228, 267, 268, 280, 350
big-headed, 268
carpenter, 267
hosts of cockroaches, 311-314
predaceous, 266
safari, 267
wood, 317
Aorurus (Thelastoma) appendiculatus,
196
diesingt, 195
philippinensis, 193
sphaerolaima, 194
Aotes zonalis, 284
Aotus, 284
A phaenogaster picea, 267
Aphelocoma coerulesens, 280
Aphids, 62
Aphlebia maculata, 10
punctata, 10
Apocynaceae, I61
Apolyta, 9
Apotrogia angolensis, 7, 17
Apple, 166
Aptera cingulata, 7
fusca, 7, 290, 329
Apteroblatta perplexa, 7, 18
Aquifoliaceae, 158
Arachnida, 211
Arachnids, 34
Araceae, 143
Aranea, 214
Araneida, 214
Arbor vitae, 44
Archiacanthocephala, 203
Archiblatta hoevenii, 7, 329
Archimandrita marmorata, 7, 147
tessellata, 7, 147

INDEX

Arenivaga apacha, 7, 23, 26, 27
bolliana, 7, 23, 26, 27, 37, 200
erratica, 7, 23, 26, 27
floridensis, 7, 24, 37
grata, 7, 17, 18, 37
rosem, 7, 24, 20, 27, 291
tonkawa, 7, 24

Aristida pennata, 65, 141

Aristiger histrio, 7, 37, 150, 316

Armadillo, 287

Army transports, 84

Arthroleptis variabilis, 270

Arthromitaceae, 124

Arthromitus intestinalis, 124

Arthropoda, 210-268

Arum sp., 143

Ascalaphidae, 227

Ascaridae, 209

Ascaris lumbricoides, 209
sp., 209
suum, 209

Ascaroidea, 209

Aschelminthes, 192, 204, 209

Ascomycetes, 133

Asmorrhiza longistylis, 240

Asparagus officinalis, 235

Aspen, 46, 152, 164

Aspergillaceae, 120

Aspergillus flavus, 130
fumigatus, 130
niger, 130
Sp:; E30; 131
sydowit, 130
tamarit, 130

Aspiduchus borinquen, 7, 18, 37, 333-

334
cavernicola, 8, 18, 334
deplanatus, 7, 8, 37

Asplenium nidus, 44

Assembling of cockroaches, 331-334

Associations, 6, 15, 91-96
among cockroaches, 324-343
biotic, 2, 91, 94-96
classification of, 91, 95-96
ecological, 14-90, 324-343
facultative, 95-06
familial, 325-330
gregarious, 330-330
interspecies, 337-343
intraspecies, 336-337

443

Associations—C ontinued.
mutualism, 96-102
number of, 3, 5
obligate, 95
obscure, 96, 316-319
with man, 70-90
Astata, 255
Ateles dariensis, 284
Athel, 39
Atta cephalotes, 313
fervens, 313
lundi, 312
nigra, 312
octospinosa, 313
sexdens, 313
texana, 313
Attacus atlas, 320
Attaphila aptera, 8, 313
bergi, 8, 312, 313
flava, 8, 314
fungicola, 8, 313
schuppi, 8, 312
sexdentis, 8, 313
SP., 313
Atticola mortont, 8, 312
Attraction, intraspecies, 328-334
to decaying material, 53, 59-63
to honeydew, 62
to lights, 52, 61, 64, 66, 79, 81
Audreia bromeliadarum, 8, 31, 37, 145
jamaicana, 8, 37
marginata, II
Aves, 276-282
Avicularia avicularia, 214
sp., 214

“B. aerobia del pseudoedema maligno,”
125
B. alcaligenes beckeri, 125
faecalis, 111
recti, III
“B, del pseudoedema maligno,” 125
B. violaceus, 106
Baboon, 286
Bacillaceae, 120
Bacilli, colon, 126
paracolon, 113
“Bacillo del barbone dei bufali,” 126
“proteisimile,” 126
“similcarbonchio,” 126

444 SMITHSONIAN MISCELLANEOUS COLLECTIONS

“Bacillo”—Continued.
“similtifo,” 126
“tifosimile,” 126

“Bacillus,” 126
“subtilis group,” 122

Bacillus acidi lactici, 112
albolactis, 120
alcaligenes faecalis, 111
anthracis, 120
biitschlu, 120
cereus, 120
circulans, 120
cloacae, 112
faccalis alkaligenes, 111
flacheriae, 121
lactis acrogenes, 111
megaterium, 121
monachae, 121
periplanetae, 121
prodigiosus, 117
radiciformis, 121
stellatus, 121
subtilis, 121
tritus, 122

Bacteria, 2-3, 100-101, 104-127, 346
“spirochaetoid,” 127

Bacteriaceae, I19

“Bacterium,” 126

Bacterium alkaligenes, 119
delendae-muscae, 120
haemophosphoreum, 120
monache, 121
prodigiosum, 117

Bacteroides uncatus, 119

Bacteroids, 96-100; pl. 26

Badger, 289

Balantidium, 101
blattarum, 187
coli, 187
ovatum, 187
praenucleatum, 187
sp., 187

Ballota nigra, 38

Balta godeffroyi, 8, 37
patula, 8, 201
platysoma, 8, 317
quadricaudata, 8, 37, 141
scripta, 8, 37, 141, 326
torresiana, 8, 38, 141
verticalis, 8, 38, 141

VOL. I4I

Bamboo, 44, I41
Bambusicola thoracica, 277
Banana, 36, 40, 53, 54, 58, 66, 82, 146-
I5I, 163, 165, 215
blossoms, 36, 41, 56, 57, 146-148
Bantua stigmosa, 8, 99
“Barata selvagem,” 194, 199, 200
“sylvestre,” 188, 189
“Baratas de pau podre,” 200
Barbulanympha, 102
coahoma, 173
estaboga, 173
laurabuda, 174
Sp., 175
ufalula, 174
wenyont, 174
Bark, 36-38, 42, 43, 45-65, 141, 152, 153,
160, 164, 165
Bass, large-mouth black, 269
Bassariscus astutus, 288
Bat, 77, 283
caves, 17-18, 21-23
Batrachians, unidentified, 272
Bayberry, 43, 44, 152
Beauveria bassiana, 131
Bedbug, 321-322
Beech, 47, 61, 62
Bees, 314, 318
Beetles, 26, 34, 57, 63, 91, 229-234, 318,
347, 348
black (see Blatta orientalis)
black larder, 234
rhinocerus, 317
Bertramia blatellae, 185
Beta maritima, 47, 153
vulgaris cicla, 153
Binema mirzaia, 193
Birds, 3, 4, 276-282, 353
nests, 35, 318
Bites, cockroach, 83, 322, 336-337
mite, 220
spider, 215
Bixa sp., 159
Bixaceae, 159
Blabera fusca, 8, 322
Blaberus atropos, 8, 18, 38, 125, 147,
203, 204, 201, 322, 344
boliviensis, 8, 147

INDEX

Blaberus—Continued.
clarazianus, 8
craniifer 8, 18, 75, 97, 99, 116-1 18,
132, 138, 291, 322, 330, 333, 336,
330, 344; Pls. 1, 2, 28
cubensis, 8
discoidalis, 8, 38, 75, 85, 147, 151,
163, 291, 318, 330, 334, 339
giganteus, 8, 18, 38, 99, 291; pl. 3
“maderae,’ 11, 248
spp., 38, 290, 291
Blackbird, 282
Puerto Rican, 282
yellow-shouldered, 282
Blaptica dubia, 8, 292
Blastocystidaceae, 133
Blastocystis hominis, 133
Sp 133
“Blatella americana,” 108, 110, 119,
12%, 122
Blatta aegyptiaca, 13
aethiopica, 13
americana, 12
caraibea, 79
concinna, 10
humbertiana, 8
(Shelfordella) lateralis, 8, 18, 38,
71, 75, 202
melanocephala, 13
notulata, 10
orientalis, 8, 18, 38, 70-72, 75-76,
84, 85, 88, 96, 99, 100, 118, 123,
128, 120, 131, 151, 153, 162, 163,
190, 204, 205, 212, 219, 222, 224-
226, 232, 235, 237, 245, 255, 269,
275, 276, 292-203, 320-323, 325-
327, 331, 332, 334-336, 338, 341,
342, 346, 347, 352; PIs. 4, 27
sp., 292
Blattaria, 3, 224
Blattarum alatarum, 83
Blattelicola blattelicola, 193
Blattella bisignata, 8
brunneriana, 11
chichimeca, 10
delicatula, 8
germanica, 8, 17, 18, 39, 70-72, 70,
83-85, 87, 88, 96-101, 103-105,
114, 118, 119, 124, 126, 131, 143,
I7I, 192, 204, 205, 211, 212, 219,

445

Blattella—Continued.
germanica—C ontinued.
222, 225-227, 234-236, 240, 241,
255, 272, 276, 277, 283, 203-294,
320, 321, 323, 324, 328, 331, 332,
334, 335, 337-342, 347-349; Pls.
5, 26, 31
humbertiana, 8, 39, 142, 144, 294,
336
lituricollis, 8, 263, 295
nahua, 11
schubotzi, 8, 76
spp. 88, 145, 295
vaga, 8, 27, 39, 77, 135, 141, 166,
255, 293, 320, 328; pl. 5
Blatticida ashmeadi, 243
pulchra, 243
Blatticidella ashmeadi, 243
Blatticola, 199
aegyptiaca, 195
blattae, 193-194
blatticola, 193
“Blattidae sylvestres,” 200
Blattilaelaps nauphoetac, 216
Blattina concinna, 10
Blattisocius tineivorus, 216
triodons, 216
Blattophila sphaerolaima, 194
sphacrolaima var. javanica, 194
supellaima, 194
Blattotetrastichus hagenowt, 249
Bluegills, 269
Blue heads, 269
Blue jay, 280, 346
Bobolink, 282
Bodo blattae, 167
sp., 167
Bodonidae, 167
Bog, peat, 46
Boraginaceae, 161
Bounty, H. M. S., 83
Brachygaster, 350
minutus, 235-236, 351
Bracken, 45, 47
Brambles, 51, 155
Breadfruit, 162
Bream, 269
Bromeliaceae, 144
Bromeliads, 31, 36, 37, 41, 42, 45, 49,
50, 54, 56, 57, 68, 144-146, 320

446

Brucella abortus, 119
Buboblatta armata, 8, 145
Bucerotidae, 279
Bufo funereus, 270
ictericus, 270
marinus, 270, 345, 346, 353
valliceps, 270
Bufonidae, 270
Bulhéesia bulhéesi, 200
icemi, 200
magalhaesi, 194
severianot, 200
Bullfrog, 271
Burrows, 27
cockroach, 55, 58, 67, 68, 328, 344
relative humidity in, 26
vertebrate, 23-24, 26, 27, 29
Bursera simaruba, 157
Burseraceae, 157
Busaria blattarum, 188
Busariidae, 187
Buthidae, 211
Buthus australis, 211-212
Butterfly nests, 66, 156, 317
Byrsotria cabrerae, 8, 40

fumigata, 8, 19, 40, 225, 333, 3543
pl. 6

Cacao, 43, 52, 53, 55

Cacomistle, 288

Cafards, 207

Cahita borero, 8, 40
nahua, 8, 40

Caladium, 66, 143

Callithricidae, 285

Callithrix chrysoleucos, 285
chrysolevea, 285
jacchus, 285

Callitris sp., 55

Calluna vulgaris, 160

Calodexia venteris, 228
spp., 228

Caloglyphus spinitarsus, 217
sp., 218

Calyptracordia alba, 161

Camouflage, 344

Camponotus femoratus, 311, 314
maccooki, 316
maculatus, 312

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I41

Camponotus—Continued.
pennsylvanicus, 267
rufipes, 312

Candida seylanoides, 129

Canidae, 288

Canis familiaris, 288
latrans, 288

Canna, 151, 163

Cannaceae, I5I

Cannibalism, 322-324

Cantao ocellatus, 320

Canthium barbatum, 161

Capa blanco, 66

Capillaria hepatica, 210

Capucinella delicatula, 147

Carabidae, 229

Carboniferous, 14

Care, maternal, 325-330

Cariblatta antiguensis, 8, 40, 158, 161
cuprea, 8, 40
delicatula, 8, 40, 143, 147, 295
hylaea, 8, 40, 147, 153
imitans, 8, 41
insularis, 8, 41, 145, 147, 339
landalei, 8, 41, 147
lutea, 23
lutea lutea, 8, 41, 153, 158, 295
lutea minima, 8, 42, 143; pl. 7
nebulicola, 8, 42, 145, 340
orestera, 8, 151
punctipennis, 8, 147
punctulata, 8, 143
reticulosa, 8, 42
spp., 42, 88
stenophrys, 8, 42, 142, 156

Cariblattoides instigator, 8, 43
suave, 8, 43

Carica papaya, 159

Caricaceae, 159

Carnations, 163

Carnivora, 288

Carpinus orientalis, 38

Caryophanales, 124

Casuarina, 152, 164

Casuarinaceae, 152

Cat, 4, 289
ring-tailed, 288

Caterpillars, 319

Catopsis fulgens, 144

Cattleya, 151, 162-163
Caudata, 269
Caves, 16-17
Cavia sp., 288
Caviidae, 288
Cebidae, 284
Cebus apella, 284
capucinus, 284
Cecropia sp., 40, 153
Cedar, Japanese, 164
Centipedes, 3, 222-224, 319
house, 222
Centruroides gracilis, 212
hentz1, 212
vitattus, 212
Cephalobelius brevicaudatum, 194
magalhdesi, 194
Cephalosporium sp., 131
Ceratinoptera diaphana, 7
picta, 8, 43, 1590
Ceratonia siliqua, 156
Cercopithecidae, 286
Cercopithecus sp., 286, 343
Cestoda, 208
Chactidae, 212
Chaetochloa verticillata, 142
Chaetodactylus sp., 218
Chalceus macrolepidotos, 269
Challenger, H. M. S., 83
Chamaeleon chamaeleon, 275
oustaleti, 275
Chamaeleontidae, 275
Chameleon, 275
Channel cat, 269
Characidae, 269
Cheiloneurus viridiscutum, 244
Chelonia, 272
Chemotaxis, 331
Chenopodiaceae, 153
Chestnut, 44, 60-61
Chicken, 68, 82, 276, 278
Chilicabra, 281
Chilomastigidae, 169
Chilomastix mesnili, 169
Chilopoda, 222
Chimpanzee, 286
Chiroptera, 283
Chlamydosaurus kingit, 275
Chlorion compressum, 258
Chloris gayana, 141

INDEX 447

Chordodes morgani, 201
pilosus, 202
puerilis, 201
Chordodidae, 201
“Chordotes” puerilis, 201
Chorisoneura barbadensis, 8, 147
flavipennis, 8, 43
formosella, 8, 43, 160
parishi, 8, 43
plocea, 8
Sp., 295
specilliger, 8, 43
texensis, 8, 43, 152, 153, 157, 161,
316
translucida, 8, 44
Choristima sp., 9, 205
Choristimodes sp., 9, 295
Chromatonotus tnfuscatus, 9, 44
notatus, 9, 44, 77
Chromobacterium violaceum, 106
Chrysanthemums, 165
Chrysemys picta, 272
Cicadas, 257
Ciliata, 186
Ciliate, unidentified, 190
Cimex lectularius, 321
Cinchona pubescens, 162, 163
Ciniflo, 214
Citharexylum villosum, 43, 161
Citrus aurantifolia, 157
maxima, 157
sinensis, 157
Sp., 157, 164
Cleonymidae, 243
Clepsidrina blattarum, 181
Clerada apicicornis, 226
Clerodendron trichotomum, 256-257
Clevelandella constricta, 189
contorta, 189
elongata, 189
hastula, 189
nipponensis, 189
panesthiae, 189
parapanesthiae, 189
Clevelandellidae, 102, 189
Clevelandia, 189
Clevelandiidae, 189
Clostridium feseri, 122
lentoputrescens, 122

448

Clostridium—C ontinued.
novyl, 122
perfringens, 122
sporogenes, 122
spp., 122
tetami, 122
Clubiona, 214
Cnemidophorus sp., 275
Coati, 288
Coccidia, 184
Coccidioides periplanetac, 133
Coccinellidae, 348
“Coccobacillus,” 126
Coccobacillus cajae, 120
Cockroach, American (see Periplaneta
americana)
amphibious, 30-33, 48
aposymbiotic, 97-99
Australian (see Periplaneta ais-
tralasiae )
beetle (see Difloptera punctata)
brown (see Periplaneta brunnea)
brown-banded (see Supella supel-
lectilium)
burrowing, 27, 29, 30, 55, 58, 67,
68, 328, 344
carnivorous, 6, 14, 319-324, 336
cavernicolous, 16-25, 37
checklists of, 290-310, 315
cinereous (see Nauphocta cine-
rea)
colonies, 75, 327-334
commensal, 310-315
control, 1, 6, 84, 162, 331, 348
Cuban (see Panchlora nivea)
cypress (see Diploptera punctata)
dendricolous, 51
defenses, 343-348
desert, 25-30
disease relations, I, 3, 74
domiciliary, 51, 70-90, 322, 33I,
338, 349
enemies, 2-4, 91, 95-96
field (see Blattella vaga)
Florida (see Eurycotis floridana)
German (see Blattella germanica)
harlequin (see Neostylopyga
rhombifolia)
Key West (see Blaberus craniifer)

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I41

Cockroach—Continued.
Madeira (see Leucophaea made-
rae)
myrmecophilous, 310, 311-314
odor, 6, 84, 280, 344-347
oriental (see Blatta orientalis)
oviparous, 325-327
phytophagous, 68, 162-166
predaceous, 319-324
predators, escape from, 343
ship, 82-87
smoky brown
fuliginosa)
species, list of, 7-14
number of known, iii, 5
spotted Mediterranean (see Ecto-
bius pallidus)
structural habitats of, 70-90
Surinam (see Pycnoscelus suri-
namensis )
undetermined, 308-310
viviparous, 3I, 326, 328-330
“wood” (except here, usually Par-
coblatta spp.), 214, 266, 274
Coconut, 35, 38, 49, 143
Cocos nucifera, 143
Coelosporidium blattellae, 185
periplanetac, 168, 176, 177, 185
Coenosia basalis, 229
Coffea sp., 162
Coffee, 66, 67
Coleolaelaps sp., 216; pl. 12
Coleoptera, 229-234, 348
“Colon bacilli,” 126
Colonies, cockroach, 75, 327-334
Colubridae, 276
Columbidae, 278
Columbiformes, 278
Combretaceae, 159
Commensalism, 92
Commensals, 2-3, 95, 173, 310-314
cockroach, check list of, 315
Compatibility, interspecies, 337-341
Comperia mercett, 244, 255, 351, 355;
pl. 34
merceti var. falsicornis, 244
Compositae, 162
Compsodes schwarzi, 9, 27, 316
Compsolampra, 9

(see Periplaneta

INDEX

Comptolampra liturata, 9, 44, 140, 141
Conocarpus erectus, 159
Conopidae, 228
Control, biological, 1, 6, 348-354
Convolvulaceae, 161
Coptotermes ceylonicus, 310
Coraciformes, 279
Cordaites, 14
Cordia dentata, 161
Cordyceps amazonica, 134
blattae, 134
Corn, 42, 48, 57, 58
Cornus mas, 38
Corvidae, 280
Corydia nuptialis, 14
Corynebacteriaceae, 110
Corynebacterium diphtheriae, 110
Cosmozosteria lateralis, 9, 345
Costus sp., 316
Cotton, 66, 67, 158, 163
Coturnix coturnix japonica, 277
Coxsackie viruses, 103-104
Coyote, 288
Crappies, 269
Crataegus sp., 155
Crickets, 320-321
stone, 226
Crinum, 163
Cristatithorax, 244
Cryptocercus punctulatus, 9, 44, 62, 67,
100-103, 175-220, 295, 317, 326, 328,
339, 354; Pl. 8
relictus, 9, 44
spp., 6, 34
Cryptomeria, 45, 140, 164
Cucarachero, 281
Cucumber, 164
Cunoniaceae, 155
Cupressaceae, I41
Cupressus, 164
macrocarpa, 141, 104
Cutilia nitida, 9, 345
soror, 9, 45, 77, 296
sp. near sedilloti, 9; pl. 9
Spp., 45, 296, 319
Cyanocitta cristata, 280, 346
Cyatheaceae, 139
Cycads, 166
Cynopotamus essequibensis, 269

449

Cyperaceae, 142

Cyperus sp., 142

Cypress, 45, 63, 164

Cypress pine, 55

Cyrilla, 42

Cyrtandra sp., 161

Cyrtocharax magdalenae essequibensis,
269

Cyrtotria capucina, 9, 27

Dasypodidae, 287
Dasypus novemcinctus, 287
Dates, 39
DDT, 278
Defenses, cockroach, 343-348
Dematiaceae, 132
Dendroblatta sobrina, 9, 45, 316, 330
Dendrocopus mahrattensts, 279
Dermaptera, 226
Dermestes ater, 234
Sp., 234
Dermestidae, 234
Dermestids, 320
Derocalymma cruralis, 9, 100
lampyrina, 9, 27
porcellio, 9, 28
stigmosa, 8
Deropeltis autraniana, 9, 28
erythropeza, 9, 19
melanophila, 9, 28
nigrita, 9, 28
Deserts, 25-30
Dicarnosis alfierit, 245
Dictamnus fraxinella, 38
Didelphidae, 283
Diestrammena apicalis, 226
japanica, 226
Digenea, 208
Dinenymphidae, 170
“Diphtheroid I and II,” 127
“Diplococci,” 127
Diplococcus pnewmoniae, 109
Diplocystidae, 181
Diplocystis schneideri, 168, 181
sp., 181; pl. 28
Diplogaster, 192
Diplogasteridae, 192
Diplogyniidae, 217
Diplogyniid, undetermined, 217

450

Diploptera dytiscoides, 9
punctata, 9, 45, 140-142, 152, 155-
157, 159, 164, 216, 224-226, 259,
273, 280, 287, 296, 324, 334, 339,
3475 Pls. 10, 36
Dipodomys spectabilis spectabilis, 23
Diptera, 227
Disease organisms transmitted by cock-
roaches, 3
Dobellina, 177
Dog, 288
Dolichonysx orysivorus, 282
Dolichurus bicolor, 260
corniculus, 260-261
gilberti, 261
greenei, 261
haemorrhous, 260
ignitus, 261-262
stantont, 262, 263, 352
sp., 262
Dorylaea rhombifolia, 11
Dorylus (Anomma) nigricans subsp.
sjostedt, 267
sp., 267
wilverthi, 267
Dove, Chinese, 278
Dragonfly, giant Hawaiian, 224
Drake (Francis), 83
Drassodes, 214
Drassus sp., 318
Dryadoblatta scotti, 9, 31, 45, 144, 145
Drymaplaneta, 319
Ducks, domestic, 277, 343
Duguay-Trouin, Cruiser, 232
Dyscologamia capensis, 9
chopardi, 10
pilosa, 9
wollastont, 9
Dytiscidae, 230

Earthworms, 319

Earwigs, undetermined, 226

Eberthella oedematiens, 112

Echinococcus granulosis, 208

Eciton burchelli, 268, 280
sp., 228

Ecology, 14-96, 324-343

“Ectobia” spp., 135

Ectobia livida, 232

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Ectobius africanus, 9, 45
albicinctus, 9, 46
duskei, 9, 46, 77
ericetorum, 9
germamncus, 8
lapponicus, 9, 46, 83, 100, 152, 160,
164, 296, 340
livens, 9
lividus, 9
lucidus, 9, 340
nicaeensis, 9, 46
nigripes, 256
pallidus, 9, 19, 47, 77, 100, 140, 153,
161, 2906, 340; pls. 7, 29
panseri, 90, 47, 153, 160, 296
panzeri var. nigripes, 9
perspicillaris, 9
semenovi, 9, 47, 152
Spp., 19, 214, 207
sylvester, 9, 47, 326, 340
sylvestris, 9
tadzihicus, 9, 48
vittiventer, 9, 19, 48
vittiventris, 9
Edentata, 287
Eleagnus, 48
Eleutheroda dytiscoides, 9
Ellipsidion affine, 9, 48, 297, 327
aurantium, 9
australe, 9, 48, 155, 160, 207, 327
bicolor, 9, 48, 142
pellucidum, 9
simulans, 9, 48, 141
spp., 35, 48
variegatum, 9, 329
Elm, 38
Embadomonas blattae, 167
Embryophyta siphonogama, 140
Emydidae, 272
Encephalomyelitis virus, mouse, 104
Encyrtidae, 94, 243, 245
Endamoeba blattae, 101, 102, 168, 177-
178
coli, 178
javanica, 178
philippinensis, 178
thomsont, 179
versus Entamoeba, 177
Endamoebidae, 177

INDEX

Endolimax blattae, 101, 180
nana, 180
sp., 180
Endomycetales, 133
Entamoeba blattae, 177
blattarum, 177
coli, 178, 179
histolytica, 179
nana, 180
pithect, 179
sp., 179-180
thomsoni, 179
Entedon hagenowii, 236, 249
Enterobacteriaceae, III
Enterobius vermicularis, 209
Enterococcus, 109
Entomophthorales, 133
Ephestia kiihniella, 99
Epilampra abdomen-nigrum, 9, 31, 48,
77, 141, 147, 160, 164, 297
annandalei, 9, 31-32
azteca, 9, 48
conferta, 9, 207
conspersa, 9, 145
grisea, 9, 100
maya, 9, 85, 145, 147
mexicana, 9, 147
mona, 9, 48, 144
notabilis, 9, 319
sodalis, 9, 145
Spp., 9, 32, 85, 80, 145-147, 263,
297, 326, 344
tainana, 9, 49
wheeleri, 9, 49, 207
Epilamprinae, 30, 326, 348
unidentified, 33
Epimys norvegicus, 287
Epiphytes, 34, 44 (see also Brome-
liads)
Epomphaloides ischnopterae, 249
Eremoblatta subdiaphana, 9, 28, 316
Ergaula capensis, 9, 49, 77, 311
scarabaeoides, 9, 19
Ericaceae, 160
Erinaceidae, 283
Erinaceus europaeus, 283
sp., 283
Erythrina glauca, 35, 156
Erythrocebus patas patas, 180

451

Escala circumducta, 13
longiuscula, 13
sp., 10, 297
Escherichia coli, 112
coli var. acidilactici, 112
coli var. communior, 113
freundii, 113
intermedium, 113
“Espino rubial,” 66, 155
Estrildine sp., 318
Euandroblatta palpalis, 10, 28
Eubacteriales, 104
Eublaberus posticus, 10, 77, 89
Eucalyptus, 48, 231
Eucalyptus sp., 160
Eucharis, 163
Eucomonympha, 102
imla, 175, 176
Eucomonymphidae, 176
Eudromiella bicolorata, 10, 49
calcarata, 10, 147
Eugenia aromatica, 160
Euglena sp., 166
Euglenidae, 166
Euglenoidina, 166
Eulophidae, 94, 248
Eulophus, 250, 351
Eupelmidae, 94, 210, 245
Eupelmus atriflagellum, 247
Sp., 247
Euphorbiaceae, 157
Euphyllodromia angustata, 10, 147
liturifera, 10, 49
decastigmata, 10
Euryconema paradisa, 194
Eurycotis bananae, 10, 47
biolleyt, 10, 49, 145, 158
caraibea, 10, 147
carbonaria, 10
decipiens, 10, 49, 346
dimidiata, 10, 49, 147
ferrum-equinum, 10, 49
floridana, 6, 10, 49-50, 77, 100, 140,
143, 144, 152, 162, 166, 171, 224,
255, 280, 297, 319, 326, 333, 334,
338, 339, 341, 345, 346; pls. 11,
29, 34
galeoides, 10, 50
improcera, 10, 297

452

Eurycotis—Continued.

kevani, 10, 50, 158

lixa, 10, 147

mann, 10, 208

opaca, 10, 50

sabalianus, 10
Euscorpius carpathicus, 212

germanus, 212

italicus ttalicus, 213
Euthlastoblatta abortiva, 10, 24, 50, 143
Euthyrrhapha nigra, 10, 19

pacifica, 10, 77
Eutrichomastix sp., 169
Eutrichosomella blattophaga, 245
Evania, 237, 350

abyssinica, 239

appendigaster, 235, 236-237, 238,

239, 350-352

desjardinsui, 236

dimidiata, 239

impressa, 242

laevigata, 236

minuta, 235

princeps, 235

punctaia, 240

sericea, 242

SD:fi 2375. 251

subspinosa, 239
Evaniidae, 93, 234-243
Exothea paniculata, 36, 158
Eyeworm, chicken, 68, 164, 204-205,

277, 278, 343

Fagaceae, 152
Felidae, 289
Felis catus, 289
pardalis mearnsi, 290
Fennel, 237
Fern, 44, 140, 164
Fig, 344
Fighting, intraspecies, 323, 336-337
Filarien, 202
Fir, Scotch, 140
Fish, 3, 268-269
Flacourtiaceae, 159
Flagellate, unidentified, 176
Flies, conopid, 228
ichneumon, 236
muscid, 229

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I41

Flies—Continued.
phorid, 227
sand, 322
sarcophagid, 229
tachinid, 228
Foeniculum vulgare, 237
Fondia sp., 318
Food, cockroaches as human, 286-287
contamination of, 83
of cockroaches, 16, 21, 31, 32, 45,
68, 72, 77, 83, 87, 162-166, 314,
318, 319-324
Forficulid, 339
Forficulidae, 226
Formica, 267
omnivora, 268
rufa, 316
rufibarbis, 312
Sp., 316
subcyanea, 312
Formicariidae, 280
Formicas, 316
Formicidae, 267, 311
Formicinae, 311
Fossils, 14
Fowl, guinea, 343
jungle, 277
Fox, 288
Freycinetia sp., 141
Fringillidae, 282
Frogs, 4, 270-272
tree, 270, 353
Fungi, 3, 127-139
hyperparasitic in protozoa,
188
Imperfecti, 129

178,

Fungus gardens, 313
Fusiformis lophomonadis, 119, 173

Galebia aegyptiaca, 195
Galerita janus, 347
Galliformes, 277

Gallus sp., 277
Gamocystis tenax, 168, 184
Geckos, 273, 354

Gekko gecko, 272

INDEX

Gekkonidae, 272
Geomys sp., 24
Geoscapheus robustus, 10, 319, 344
Geotrichum candidum, 131
Geraniaceae, 156
Geraniums, 156, 164
Gesneriaceae, I61
Giardia intestinalis, 171
Sp., 172
Ginger, wild, 151
Glands, secretory, 345-347
Glomeropitcairma erectiflora, 31, 144
Glyciphagidae, 218
Goat, 319
Goldenrod, 162
Gongylonema ingluvicola, 205
neoplasticum, 191, 206, 287
pulchrum, 192, 206
sp., 206
Gopher, pocket, 24
Gordiaceo, 202
Gordian worms, undetermined, 202; pl.
29
Gordiidae, 201
Gordioidea, 201
Gordius, 192
aquaticus, 201-202
blattae orientalis, 202
orientalis, 202
pilosus, 202
raphaelis, 202
sp., 202
Gossypium spp., 158
“Gram positive rods,” 127
Gramineae, 141
Grapefruit, 66, 67
Graptoblatta notulata, 10, 50-51, 141,
147, 151, 158-161, 163, 208, 339
Grass, 37, 40, 42-53, 55, 59-62, 64-66,
69, 141-142
bunch, 64
desert, 27-30
feather, 46
marram, 47
Rhodes, 39, 141
saw, 42, 50
wire, 41, 52
Grassquit, Carib, 282

453

Greenhouses, 68, 80, 81, 155, 157, 162-
165, 342
Gregarina blaberae, 184
blattae orientalis, 181
blattarwm, 168, 181-182
fastidiosa, 182
gibbsi, 182
illinensis, 183
impetuosa, 183
légeri, 183
neo-brasiliensis, 183
ohioensis, 183
panchlorae, 183
parcoblattae, 183
rhyparobiae, 168, 183
sandoni, 183
serpentula, 184
thomasi, 184
Gregarine, 181; pl. 28
Gregarinida, 181
unidentified, 184
Gregarinidae, 181
Gregariousness, 328, 330-336
physiological effects, 334-335
Grevisia, 144
Gromphadorhina laevigata, 10, 330
portentosa, 10, 298, 330, 354; pl. 12
Gru-gru, 142
Grubs, 317, 319
Gryllacrididae, 226
Guano, 16, 18-23
Guanobies, 16-17, 19
Guava, 54, 63, 67
Guépe ichneumon, 258
Guinea pigs, 288
Gum tree, 36, 53, 59, 61, 63, 155
Gumbo limbo, 43, 44, 157
Gymnasio nudipes, 279
Gymnopithys leucaspis, 280
Gyna kasungulana, 10, 19
maculipennis, 10, 19
sp., 298
tristis, 10, 20
vetula, 10

Habitats, 15
aquatic, 30-33, 48
beach, 36, 42, 47, 52, 53, 58, 63, 69
Cave, 16-17

454

Habitats—C ontinued,

cultivated areas, 38, 30, 53, 58

desert, 25-30, 39, 59, 65, 67, 69

dumps and rubbish, 38-40, 50, 53,
55, 67, 69

flatwoods, 37, 41, 42, 53, 60

forest, 36-38, 40-47, 49-57, 59-66

grassland, 41, 42, 52, 53, 64

hammock, 41-43, 50, 52, 53, 59, 60,
68

marsh, 42, 50
outdoor, 33-70
rotten wood, 34, 35, 38, 44, 45, 47,
49-52, 56-63
sand dune, 47, 61
sandhills, 36, 41, 42, 47, 50, 52, 60
scrub, 36, 37, 40-42, 48, 50, 52, 55,
57-60, 62, 66, 67
soil, 37, 39, 49, 53, 55, 58, 60, 67,
68, 344
steppe, 46, 47
stones and rock, 29, 31-33, 36-40,
43, 45-57, 59, 60, 62, 64, 67-70
structures, 70-90
aircraft, 87-90, 232
land-based, 73-82
ships, 82-87, 232-233
swamp, 41, 42, 46, 51, 53, 58, 61,
63, 66, 68
winter, 38-30, 61, 71
Hackberry, 50
Hadrurus arizonensis, 213
Hamamelidaceae, 155
Hammerschmidtiella diesingi, 124, 190,
IQI, 195-106
neyrae, 196
neyrai, 196
Haplosporidia, 185
Haplosporidium periplanetae, 185
Harpalus pennsylvanicus, 229
Hartmannella blattae, 177
Heather, 46, 47, 160
Hebardina concinna, 10, 278
Hedgehog, 4, 283, 354
Hedychium, 163
Helianthus sp., 162
Heliconia, 40
Helminths, 2-3, 190-210
cockroaches as intermediate hosts,
203-207

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Helminths—Continued.
cockroaches as primary hosts, 192-
202
ova carried by cockroaches, 208-
210
Hemuiblabera brunneri, 10, 51, 156
Hemidactylus frenatus, 272
Hemiptera, 226, 348
Hemithyrsocera histrio, 7
Hemlock, 44
Henicotyle antillarum, 10, 51
Herpestes javanicus auropunctatus, 289
sp., 289
Herpetomonas periplanetae, 167
Herpomyces, 93, 127-129
amazonicus, 134
anaplectae, 134
appendiculatus, 134
arietinus, 128, 134-135; pl. 27
chaetophilus, 135
chilensts, 135
diplopterae, 135
ectobiae, 135
forficularis, 135
gracilis, 135
grenadinus, 136
leurolestis, 136
lobopterae, 136
macropus, 136
nyctoborae, 136
panchlorae, 136
panesthiae, 136
paranensis, 136
periplanetae, 137, 138
phyllodromiae, 137
platyzosteriae, 137
stylopygae, 128, 137-138; pl. 27
supellae, 138
tricuspidatus, 138
sansibarinus, 138
Heterodon contortrix, 276
platyrhinos, 276
Heterogamia aegyptiaca, 13
Heterogamodes kriigeri, 10, 20
rosem, 7
rugosa, 10, 28
Heterometrus longemanus, 211, 213
Heteropoda regia, 215
venatoria, 215; pl. 30
Heteroptera, 318

INDEX

Hexamita, 102
cryptocerci, 171
periplanetae, 171
Speer7i
Hexamitidae, 171
2-hexenal, 6, 280, 345-346
Hibiscus, 36, 40, 68, 158
Hibiscus rosa-sinensis, 68, 158
Hickory, 53, 60, 61, 63
Hierodula tenuidentata, 224-225, 355;
pl. 32
Histiostoma feroniarum, 217
Holocompsa azteca, 10, 77
cyanea, 10, 78
fulva, 10, 208
metallica, 10, 51
nitidula, 10, 78, 148, 339
sapoteca, 10, 20
Hololampra bivittata, 10, 51
chavesi, 10, 51, 155
maculata, 10, 51
marginata, 10, 51
punctata, 10, 208
Sp., 51
Hololeptoblatta sp., 51, 141
Holomastigotidae, 172
Holoquiscalus brachypterus, 282
Holotricha, 186
Homalodemas cruralis, 9
Homalopteryx laminata, 10, 52, 151
scotti, 9
Hominidae, 286
Homo sapiens, 286
Honeydew, 237, 242
Hookworm, ova, 210
Hoplonymphidae, 173
Hoplosphoropyga babaulti, 10, 20
Hormetica apolinari, 10, 51
laevigata, 10, 52, 143, 145, 148
spp., 148
ventralis, 10, 148
Hormurus caudicula, 213
Hornbill, gray, 279
Huckleberry, 43, 57
Humidity reactions, 35, 72, 120
Hygrocrocis intestinalis, 124
Hygroreceptors, 72
Hygrotaxis, 72, 331
Hyla cinerea, 270

455

Hylidae, 270
Hymenolepididae, 208
Hymenolepis diminuta, 203
sp., 208
Hymenoptera, 2, 234, 311
cockroach-hunting wasps, 255-266
host selection by egg parasites,
254-255
hosts of commensal cockroaches,
311-314
predators and parasites of cock-
roach eggs, 234-254
Hypermastigina, 172
Hyperolius picturatus, 271
Hypoaspis sp., 217
Hypocreaceae, 134
Hypocreales, 134
Hyporhicnoda, 146
Hyptia dorsalis, 239
harpyoides, 239; pl. 33
reticulata, 239, 240
sp., 240
thoracica, 239, 240

Icteridae, 282
Icterus portoricensis, 280
Idionympha perissa, 174
Ignabolivaria bilobata, 10, 52
Iguanidae, 273
Ilex cassine, 66, 158
coriacea, 41, 158
lucida, 41, 158
rotunda, 257
Incertae sedis, fungi, 139
protozoa, 176
Inermicapsifer madagascariensis, 203
Infestations of cockroaches, 331
Inga laurina, 35, 156
vera, 35, 66, 67, 156, 317
Insecta, 224-268
Insectivora, 283
Inuus sylvanus, 286
Invertebrates in biological control, 349-
352
Iodamoeba sp., 180
Tolina nana, 219
Tolinidae, 219
Ipomoea tiliasea, 66, 161
Iridomyrmex humilis, 268, 319

456

Ironwood, 164
Ischnomyrme.x sp., 316
Ischnoptera, 134-135, 319
borealis, 12
cavernicola, 14
couloniana, 12
deropeltiformis, 10, 52, 142, 164,
298, 316, 336, 355; pl. 12A
divisa, 12
imsolita, 12
major, 12
panamae, 10, 53
pennsylvanica, 12
podoces, 10, 53, 340
remyt, 14
rufa occidentalis, 10, 78, 145
rufa rufa, 10, 53, 78, 89, 148, 161,
208
rufescens, 82
schenklingt, 10, 142
spp., 85, 89, 267, 298, 316
uhleriana, 12
uhleriana fulvescens, 12
Ischnuridae, 213
Isoptera, 310
Isotricha caulleryi, 187
Isotrichidae, 187

Jasmine, 45
Jasminum pubescens, 45
Jay, blue, 280
Florida, 280
Jucaro, 66
Juniper, 59
Juniperus sp., 65, 141
Jussiaea natans, 160, 164

Kakerlac schaeffert, 12
Kakkerlac, 260
Karnyia discoidalis, 10, 298
Kiawe tree, 164
Kingbird, grey, 35, 318
Klebsiella pneumoniae, 113
Kojukei, 277
Kuchinga hemerobina, 10, 298
remota, 10, 141, 148
Kiichenschaben, 100, 169, 171-173, 188

Laboulbeniaceae, 127, 134
Laboulbeniales, 134

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Lactobacillus fermenti, 109
Lactobacteriaceae, I09
Laelaptidae, 216
Lamproblatta albipalpus, 10, 53, 148
meridionalis, 10, 53
Lampyrid, undetermined, 230; pl. 33
Lampyridae, 230
Laniidae, 281
Lanius ludovicianus, 281
Larvaevoridae, 228
Lasius alienus, 268
Lastrea aristata, 163
Latiblattella chichimeca, 10, 54, 145
lucifrons, 11, 54, 146, 255
rehm, 11, 54, 140, 144, 339
spp., 85, 148
vitrea, 11, 143
sapoteca, II, 54
Latindia armata, 8
Latrodectus indistinctus, 215-210
mactans, 216
Lauraceae, 153
Leafhoppers, 320
Legumes, 165
Leguminosae, 155
Leidynema appendiculata, 124, 190, 191,
196-198
appendiculata var. americana, 197
appendiculata var. hispana, 197
appendiculata var. indiana, 197
cranifera, 198
delatorrei, 108
nocalum, 198
Leidynemella fusiformis, 198
panesthiae, 108
paracranifera, 198
Leiolopisma laterale, 274
Lemur coronatus, 283
fulvus, 283
tardigradus, 284
Lemuridae, 283
Leontocebus oedipus, 285
rosalia, 285
ursulus, 285
Lepidoptera, 316-318
Lepomis pallidus, 269
Leptodactylus albilabris, 271
pentadactylus, 271

INDEX

Leptomonas blaberae, 167
sp., 167
Leptopelis calcaratus, 271
rufus, 271
Leptospironympha, 102
eupora, 172
rudis, 172
wachula, 172
Lesser apple worm, 65
Leucophaea maderae, 11, 54, 70, 71, 78,
86, 89, 110, 148, 162, 168, 171, 203,
204, 213, 224, 236, 247, 200, 322, 320,
330, 333, 335, 339, 342, 347; pls. 13,
28
sp., 299
striata, 13
surinamensis, 13
Leurolestes circumvagans, 11, 78
pallidus, 11, 71, 78, 299, 339
Lice, plant, 320
Lichens, 47, 51, 56, 64, 153
Liliaceae, 146
Lily, Brig, 84
Lily, Easter, 146, 164
Lime, 45, 66, 157, 164
Liopicus mahrattensis, 279
Liquidambar styraciflua, 155
Lissoblatta fulgida, 13, 157
Litopeltis biolleyi, 11, 54, 145
bispinosa, 11, 54, 58, 145, 148
deiantra, 11, 54
musarum, II, 54, 148
SPP-, 334
Lizards, 272-276, 353-354
Lobogynioides, 217
Lobolampra subaptera, 11, 55
Loboptera decipiens, 11, 55, 100, 299,
326, 350
dimidiatipes, 11
extranea, 11
sakalava, 11
Sp., 209
thaxtert, 11, 55
Lobopterella dimidiatipes, 11, 55, 299,
338
Locustacarus sp., 219
Lophoblatta arawaka, 11, 55, 142
Lophocerus birostris, 279
Lophomonadidae, 172

457

Lophomonas, 101
blattarum, 101, 168, 172-173
striata, IOI, I19, 168, 173
sulcata, 173
Loris tardigradus, 284
Lorisidae, 284
Lycolaimus, 192
Lycosa avida, 216
helluo, 216, 347
Sp) 2l6enpl 30
Lycosidae, 216
Lygaeidae, 226
Lymantria monacha, 121

Macaca cyclopis, 179
mulatta, 286
rhesus, 286
sylvanus, 286
Macaque, 286
Macropanesthia rhinocerus, 11, 55, 344
Macrospironympha, 102
xylopletha, 172
Macrotermes barneyi, 311
bellicosus, 311
malaccensis, 311
natalensis, 311
Maize, 55, 64, 142
Malleomyces mallet, 119
Malojillo, 31, 48, 141
Malpighian tubules, parasites in, 177,
185, 186
Malvaceae, 158
Mamestra oleracea, 120
Mammalia, 283, 354
Mammals, 3, 283
Mangifera indica, 157
Mangoes, 45, 157, 164
Mangrove, 36, 42, 53, 66
Mantidae, 224
Mantis, Carolina, 225
Chinese, 226
European, 225
Mantis religiosa, 225
Maple, 61-63
Marava arachidis, 339
Mareta acutiventris, 11, 317
Maretina uahuka, 11, 142, 155
Margattea sp., 317
Marsupialia, 283

458

Mastigophora, 166
Mastigoproctus giganteus, 211
sp., 211
Maze, 335
Mediterranean fruitfly, 252
Megachilidae, 314
Megalixalus fornasinti, 271
Megaloblatta blaberoides, 11, 55
rufipes, 11
Megalopyge krugit, 35, 317
Megamareta verticalis, 11, 56, 141
Megaselta sp., 227
Melanerpes portoricensis, 280
Melanosilpha capensis, 11, 299
Meleagrididae, 278
Meleagris gallopavo, 278
Meles sp., 289
Melipona nigra, 314
Melittobia chalybiu, 248
Memnoniella echinata, 132
Mermis, 192, 202
Mermithidae, 192
Mermithids, undetermined, 192; pl. 29
Mermithoidea, 192
Mesquite, 37, 58, 156
Mestocharomyia oophaga, 248
Metallyticus semiaeneus, 225
Metarrhizium anisopliae, 131
Metrosideros collina, 160
Methana canae, 11, 56, 326
curvigera, II, 56, 155, 326
marginalis, 11, 56, 79, 326
Methionine, 101
Micrococcaceae, 106
Micrococcus aurantiacus, 106
citreus, 106
epidermidis, 106
migrofaciens, 106
pyogenes var. albus, 106-107
pyogenes var. aureus, 107
spp., 107, 108
ureae, 107
Microhabitats, 16-17, 25-26, 30-31, 33-
35, 70-74
Microsporidia, 185
Microtetrameres helix, 206
Midas rosalia, 285
Mimicry, 6, 347-348
Mimosa, 57

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Mines, 16, 19-21, 23
Miroblatta silphoides, 10
Mirotermes, 102
Mites, 2, 3, 211, 216-222, 318, 320
cheese, 222
cockroach, 219, 221
control, 222
mushroom, 218
unidentified, 220
Molossidae, 283
Molossus sp., 283
Moluchia (?) dahli, 11, 56
Monadidae, 167
Monas sp., 167
Monastria biguttata, 11, 344
sp., 299
Mongoose, 280, 354
Monilaceae, 130
Moniliales, 129
Moniliformidae, 203
Moniliformis dubius, 203, 270, 271, 276
kalahariensis, 203
moniliformis, IQ1, 204
Monkey, 4, 284-286, 343
Canal Zone night, 284
Darien black spider, 284
white-faced, 284
Monocercomonas orthopterorum, 169
Monocercomonoides, 102
globus, 169
melolonthae, 168, 169
orthopterorum, 169-170
panesthiae, 170
Monodelphis sp., 283
Mononychoblatta, 26
Moraceae, 153
Mosquitoes, 322
Moss, 46, 47, 56, 64
Spanish, 54, 144
“Mouche bleue, La,” 260
Mouse, 103-104
sand, 24
white-footed, 24, 205
Mucor guilliermondi, 132
sp., 132
Mucoraceae, 132
Mucorales, 132
Muridae, 287
Murorinae, 129

INDEX

Mus decumanus, 287
musculus, 287
Musa (see banana)
Musaceae, 146
Muscidae, 229
Mustelidae, 289
Mutant, white-eyed, 21
Mutualism, 92, 96-102
Mutuals, 95
Muzoa madida, 11, 56
Mycetozoa, 176
Mycobacteriaceae, 123
Mycobacterium aviunt, 123
friedmannii, 123
lacticola, 123
leprae, 123
lepraemurium, 123
phlei, 123
piscium, 124
spp., 124
tuberculosis, 124
Myna (mynah), 281
Myrica cerifera, 43, 152
Myricaceae, 152
Myrmeblattina longipes, 11, 314
Myrmecoblatta rehni, 11, 312
wheeleri, 11, 314
Myrmecophiles, 311-314, 316
Myrmicaria natalensis, 316
Myrmicinae, 312
Myrtaceae, 160

Namablatta bitaeniata, 11, 28
Nasua narica, 288
nasua, 288
sp., 289
Nasutitermes costalis, 317
Nauclidas nigra, 11, 56, 148, 326, 327
Nauphoeta bivittata, 11
brazzae, 11
cinerea, 11, 70, 79, 86, 89, 100, 115,
116, 123, 162, 163, 192, 224-226,
259, 209, 324, 330, 335, 337, 339;
pl. 14
flexivitta, 11, 148, 163
punctipennis, 11, 28
Necator americanus, 210
Nectandra coriacea, 43, 153
Neisseria meningitidis, 109

459

Neisseriaceae, 109
Nelipophygus ramsdeni, 11, 56
Nematoda, 192, 204, 209
Nematodes, 2-3, 190-192, 204, 209
undetermined, 201
Nematomorpha, 201
Neoaplectana chresima, 192-193
sp., 192
Neoblatiella brunneriana, 11, 145
carcinus, 11, 148
celeripes, 11, 148, 324
detersa, 11, 56, 148
dryas, 11, 56, 145, 160, 340
eurydice, II, 57, 145, 340
fratercula, 11, 86, 145, 148
fraterna, 11, 86
grossbecki, 11, 57, 145
laodamia, 11, 148
nahua, 11, 86, 145
proserpina, 11, 57, 145, 160, 339
semota, 11, 57, 148
spp., 79, 86, 89, 148, 149, 324
tridens, 11, 148
vatia, II, 148
Neonephrites partiniger, 230
Neorhipidius neoxenus, 230
Neostylopyga rhombifolia, 11, 70, 77,
79, 86, 138, 200, 210, 224, 255, 259,
272, 273, 280, 300, 346; pl. 15
Neotoma sp., 23-25, 37, 50
Nepenthaceae, 154
Nepenthes ampularia, 154
gracilis, 154
Sp. 154
Nephrites australis, 231
nitidus Riek not Shuckard, 231
sp., 231
Nepidae, 227
Nesomylacris cubensis, 11, 57
relica, 11, 57, 145
Nests, ant, 27, 311-314, 316
bird, 35, 318
insect, 35, 43, 66, 67, 69, 156, 317,
318
rodent, 23-25, 37, 50
spider, 215-216, 317
Neuroptera, 227
Nicotiana sp., 161
Nocardia sp., 124

460

“Nocticola” azteca, 149
Nocticola, 17, 149

bolivart, 11, 17, 57

caeca, II, 20

decaryi, 11, 20

simoni, 12, 20

sinensis, 12, 311

ternutophila, 12, 311
Nosema periplanetae, 185, 186
Nosematidae, 185
Nothoblatta wasmanni, 12, 312
Notila proteus, 170
Notolampra antillarum, 12, 144, 159
Nucleophaga sp., 178
Nyctibora azteca, 12, 149

brunnea, 12, 145

holosericea, 12, 149

laevigata, 12, 57, 145, 149, 339

latipennis, 12

lutet, 12, 57, 145, 317

mexicana, 12, 149

noctivaga, 12, 86, 100, 145, 149

obscura, 12, 57, 149, 300

sericea, 12, 149

spp., 86, 140, 300

stygia, 12, 58, 156

tomentosa, 12, 300
Nyctotherus, 92, 101, 102

buissoni, 188

ovalis, 101, 102, 168, 188

uichanci, 188

viannai, 189
Nymphytria, 26

Oak, 36-38, 41, 43, 44, 46, 47, 52, 53,
59-66, 152-153
Oats, wild, 52, 142
Ocelot, 290
Ocneria dispar, 126
Ocotea catesbyana, 43, 153
Octomitus periplanetac, 171
Odonata, 224
Odontomachus affinis, 314
clarus, 316
haematodes insularis var. pallens,
316
Odontotermes sp., 311
Oikomonadidae, 166

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 141

Oikomonas blattarum, 166
sp., 166, 167
Oligacanthorhynchidae, 203
Onagraceae, 160
Oniscosoma granicollis, 12, 300
spp., 58, 344
OGdthecae, concealment of, 47, 61, 63,
325-327
Ophion luteus, 261
Opisthoplatia maculata, 12, 30, 32-33
orientalis, 12, 33, 58
Orange, 45, 157, 164
Orangutan, 70
Orchidaceae, I51
Orchids, 151, 162-163, 164, 237
Origins of domiciliary cockroaches, 70-
71
Oriole, Puerto Rican, 280
Oriolidae, 280
Orthoptera, 2, 15, 16, 25, 34, 85, 88, 224
Oulopteryx meliponarum, 12, 314
Ovipositor, cockroach, 15
Owls, 276, 353
bare-legged, 279
Oxyhaloa buprestoides, 12, 70, 79, 86
deusta, 12, 149
Oxymonadidae, 170
Oxymonas, 102
doroaxostylus, 170
nana, 170
Oxyspirura mansoni, 68, 191, 204-205,
278
Oxyuridae, 209
Oxyurts aegyptiaca, 195
australasiae, 199
blattae, 193, 196
blattae orientalis, 195, 196
blatticola, 193
bulhéest, 200
diesingi, 105
(2?) heterogamiae, 199
kiinckeh, 199
macrura, 195
panesthiae, 198
Oxyuroidea, 193, 204, 209

Palamnacus, 211
Palissota, 34
Paliurus aculeatus, 38

INDEX

Pallasiomys meridionalis pennicilliger,
24
Palmae, 141
Palmetto, 37, 40, 50, 57, 62, 64, 67, 166
Palms, 45, 50, 52, 58, 64, 143, 150, 164
coconut, 36, 40, 42, 50, 53, 56, 58,
64, 68, 143
date, 143
royal, 54, 143
sago, 166
Pan sp., 286
Panchlora antillarum, 12, 58, 143, 149
cubensis, 12
exoleta, 12, 150, 300
fraterna, 12, 150
maderae, 11
nivea, 12, 58, 79, 86, 89, 142, 150,
300; pl. 16
peruana, 12, 150
sagax, 12, 58, 150
SPP., 150, 324
virescens, 12, 150
Panchlorinae, 231
Pandanaceae, 141
Pandanus, 51, 141, 150
Panesthia, 102
angustipennis, 12, 100, 300
australis, 12, 58, 301, 327, 3543
pl. 8
“brevicollis,” 194
javanica, 12
laevicollis, 12, 58, 301, 327
lobipennis, 12, 301
Spp., 301, 328
Spadica, 12, 301
Panicum barbinode, 141
purpurascens, 141
Papayas, 45, 159, 164
Papio papio, 286
Parabuthus capensis, 212
Parachordodes raphaelis, 202
Paraclevelandia brevis, 190
simplex, 190
Paracolobactrum aerogenoides, 113
coliforme, 113
Spp., 113
Paradisea papuana, 280
Parahormetica bilobata, 12, 301
Parameciidae 186

461

Paramecium sp., 186-187
Paranephrites xenus, 230
Parapolyphaga erectipilis, 9-10
Parasites, 91, 95-90; pls. 16, 27, 29, 35
cockroach egg, 2, 234-254, 350-352;
pls. 33, 34
host selection by, 254-255
Parasitism, 91-04
Parcoblatta, 350
americana, 12, 267
bolliana, 12, 59, 338
caudelli, 12, 59
desertae, 12, 25, 20, 59, 316
divisa, 12, 59, 140, 152, 153, 155
fulvescens, 12, 24, 59-60, 79, 140
lata, 12, 60, 79, 100, 140, 152, 301,
333, 339
notha, 12, 80
pensylvanica, 12, 15, 61, 80, 100,
152, 153, 257, 302, 318, 319, 327,
333, 339; Pl. 17
spp., 17, 20, 63, 144, 206, 302, 316,
333, 336
uhleriana, 12, 15, 61-62, 100, 302,
339; pls. 18, 33
virginica, 12, 15, 62-63, 100, 230,
255, 257, 258, 302, 316, 323, 333;
Pls. 17, 27, 33
zebra, 12, 63, 140, 155
Parinarium, 34
Parsley, 237
Parsnip, 240
Partridge, 278
Parvobacteriaceae, 119
Passer domesticus, 282
Passeriformes, 280
Passiflora sp., 37, 159
Passifloraceae, 159
Pasteurella multocida, 119
pestis, 119
Pastinaca sativa, 240
Paxylomma buccata, 255
p-benzoquinone, 216, 347
p-chlorophenyl, p-chlorobenzene sulfo-
nate, 222
Pedipalpida, 211
Pelmatosilpha coriacea, 12, 63, 142, 150,
151, 160
kevani, 12, 63

462

Pelmatosilpha—Continued.
marginalis, 12, 150, 326
purpurascens, 12, 63, 150, 326
rotundata, 12, 145
vagabunda, 12, 150
Peltomyces blattellae, 176
periplanetae, 176-177
Penicillium sp., 131
Periplaneta americana, 12, 17, 20, 63-
64, 70, 71, 80, 83, 84, 87, 80, 98-101,
103, 104, 106, 126, 129, 143, 150, I5I,
156, 162, 163, 166, 190, I9I, 205, 21I-
213, 223, 225, 226, 220, 245, 255, 271,
272, 276, 280, 302-304, 318, 320, 321,
323-327, 332, 334-336, 338-340, 342,
346-348, 350-352, 354; Pls. 19, 33;
34, 35
australasiae, 12, 21, 55, 64, 70, 71,
80, 87, 89, 100, 105, 115, 117, 123,
143, 150, 151, 153, 158, 159, 163,
209, 210, 214, 225, 304-305, 318-
320, 323, 326, 327, 333, 335, 338,
339, 341, 342, 348, 350, 351, 354;
pls. 20, 32
brunnea, 12, 64, 70, 80, 150, 153,
I7I, 305, 323, 325-326, 340, 346;
pl. 21
cavernicola, 12, 16, 21
fortipes, 13
fuliginosa, 12, 64, 81, 158, 164, 251,
277, 305, 327, 335, 338; pl. 22
ignota, 12, 81, 115, 327
lata, 12, 21
orientalis, 8
picea, 12
SPP., 2I, 90, 305-306, 317, 352
tartara, 8
Periplaneticola mirzaia, 193
Perisphaerus armadillo, 12, 320
glomeriformis, 12, 320, 347
Sp., 21,..320
Periwinkle, 47
Permian, 14
Perodicticus potto, 284
Peromyscus leucopus noveboracensis,
205
polionotus rhoadsi, 24
Petitia domingensis, 66, 317
Petroselium crispum, 237

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. 141

Phaetalia pallida, 12, 81
Phasianidae, 277
Phasianus calchicus karpowi, 277
Sp., 277
Pheasants, 277
Pheidole megacephala, 268
Phidon (?) dubius, 13, 64
Phileciton longipes, 314
Philip, Ship, 83
Philodromus pernix, 318
Phlebonotus pallens, 13, 329
Phlebotomus papatasii, 322
Phoenix dactylifera, 143
Pholadoblatta inusitata, 13, 316
Phoraspidinae, 329
Phoraspis sp., 64, 142
Phoridae, 227
Phormictopus cancerides, 214
Phorticolea boliviae, 13, 311, 314
testacea, 13, 312
Phryganoporus, 317
Phycomycetes, 132
Phyllodromia germanica, 8
hemerobina, 10
hieroglyphica, 10
humbertiana, 8
nigrocincta, 14
obtusata, 7
parenthesis, 14
spp., 306, 352
supellectilium, 13-14
treitliana, 13, 245
Phyllodromica brevipennis, 13, 64
graeca, 13, 65
trinae, 13, 65
maculata, 13, 65
megerlet, 13, 65, 153
polita, 13, 65
pygmaea, 13, 65, 141
tartara, 13, 65
tartara nigrescens, 13, 65, 141, 162
Physaloptera hispida, 191, 207
ma-illaris, 191, 207, 288, 289
praeputialis, 207
rara, 191, 207, 288, 289
turgida, 191, 207
Physalopteridae, 207
Physignathus lesueurti, 275
Phytoseiidae, 216

INDEX

Picidae, 279
Piciformes, 279
Pigeon, 279, 343
Pileocephalus blaberae, 184
Pilocrocis secernalis, 66, 317
Pimeliaphilus podapolipophagus, 219-
221, 320
Pimelodidae, 268
Pinaceae, 140
Pine, 36, 41-44, 46, 47, 49-54, 57, 59-67,
140
Pineapple, 39, 45, 68, 144, 165
Pinus australis, 140
caribaea, 50, 52, 53, 59, 64, 140
clausa, 53, 140
echinata, 140
Spp., 40
sylvestris, 140
Pisces, 268
Plagiotoma blattarum, 188
Plants, carnivorous, 154
damage to, by cockroaches, 162-
166
higher, 139-162
pitcher, 154
Platyhelminthes, 208
Platyzosteria analis, 13, 300
armata, 13, 100
bifida, 13, 151
castanea, 13, 65, 306, 345
ingens, 10
novae seclandiae, 13, 65, 306, 321,
346; pl. 23
novae-zealandiae, 13
sabalianus, 10
scabra, 13, 306
sp., 306
Plectoptera dominicae, 13, 66, 157
dorsalis, 13, 66, 67, 81, 142, 143,
I5I, 155-158, 160, 161, 317
floridana, 13
infulata, 13, 66, 142, 155, 156-158,
317
lacerna, 13, 66, 140, 153
perscita, 13, 66, 157
poeyi, 13, 66, 158, 159
porcellana, 13, 66, 157, 162
pygmaea, 13, 66

463

Plectoptera—Continued.
rhabdota, 13, 60, 67, 142, 155-158,
160, 317
sp., 306
vermiculata, 13, 67, 140
Pleistophora periplanetae, 186
Plethodon glutinosus, 269
Plethodontidae, 269
Plistophora kudot, 185-186
periplanetae, 177, 185, 186
sp., 186
Ploceinae, 318
Ploceus sp., 318
Plumiger histrio, 7
Plumilla, 35
Pneumococcus, 109
Podapolipodidae, 219
Podium abdominale, 265
carolina, 265
dubium, 265
flavipenne, 265-266
haematogastrum, 263, 266, 352
luctuosum, 266
rufipes, 266
sp., 266
Podocarpus, 42
Pogonomyrmex badius, 346, 347; pl.
36
Poinsettia sp., 157
Poinsettias, 164
Polecat, 4
Poliomyelitis viruses, 3, 103
Polistes, 318, 319
Polybia pygmaea, 314
“Polygamia” aegyptiaca, 13
rosent, 7
Polymastigidae, 169
Polymastigina, 169
Polymastix, 101
Polyphaga aegyptiaca, 13, 21, 24, 25,
29, 67, 81, 100, 190, 306
indica, 13, 24, 20, 317
pellucida, 13
Saussuret, 13, 24, 29, 67, 71, 81,
114, I15, 117-119, 306
Sp., 21
Polyphagids, 344
Polyphaginae, 25
Polypodiaceae, 140

464

Polystichum aristatum, 163
Polyzosteria analis, 13
limbata, 13, 67
melanaria, 13, 306
Pomarrosa, 160
Pompilidae, 256
Pompilus bracatus, 256
sp., 256
Ponerinae, 314
Pongidae, 286
Ponies, pit (mine), 16, 23
Populus diversifolia, 152
euphratica, 47, 152
sp., 152
Poroblatta nigra, 11
spp., 67
Potamotrygon humboldti, 268
hystrix, 268
Potamotrygontidae, 268
Potato, 165
Potto, 284
Poultry, 276-278, 353
Prairie dog, 24
Predation, interspecies, 92, 319-322
intraspecies, 92, 322-324
Predatism, 91-95
Predators, 91, 96, 319
defense against, 343-348
of cockroach eggs, 234
Prey, experimental, 4
natural, 4
Primates, 283
Pritchardia, 64, 163
Privies (latrines), cockroaches in, 74,
80, 81, 178, 340
Procyonidae, 288
Prodenia litula, 320
Prolabia, 339
Prolophomonas tocopola, 173
Prosevania punctata, 240-242, 254-255,
351; pl. 33
Prosoplecta, 348
Prosthenorchis elegans, 203, 283, 285,
286, 288-290
Spirula, 203, 283, 286, 288-290
Proteus mirabilis, 114
morganit, 114
rettgeri, 114
spp., 114
vulgaris, 114

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Protomagalhaesia serpentula, 168, 184
Protomonadina, 166
Protospirura bonnei, 206
columbiana, 206
muricola, 191, 206, 284
Protozoa, 2, 92, 101-103, 166-190
cultured in vitro, 170, 178, 179, 188
Protrelleta floridana, 199
Protrellina, 199
aurifluus, 199
australasiae, 199
galebi, 199
kiinckeli, 199
manni, 200
phyllodromi, 200
Protrelloides paradoxa, 200
Protrellus aureus, 199
aurifluus, 199
australasiae, 199
galebi, 199
kiinckeli, 199
mann, 200
phyllodromi, 200
Pseudoderopeltis aethiopica, 13, 100
Pseudomonadaceae, 104
Pseudomonas aeruginosa, 104
etsenbergil, 105
fluorescens, 105
Pseudomopinae, 60, 230, 231
Pseudomops cincta, 13, 319
laticornis, 13, 145
septentrionalis, 13, 67, 162
Pseudophoraspis nebulosa, 13, 81, 328
Pseudosaccharomycetaceae, 120
Psidium guajava, 67, 160
Psocoptera, 318
Pteridophyta, 139
Pteromalidae, 247
Pteromalus sp., 247, 248
Pterygosomidae, 219
Pycnosceloides aporus, 12, 58
Pycnoscelus niger, 13, 22
striatus, 13, 22
surinamensis, 2, 13, 22, 25, 67-68,
70, 71, 81, 87, 90, 100, 138, 139,
142-146, 151, 152, 155-157, I61-
165, 191, 212, 220, 259, 279, 280,
282, 306-307, 324, 330, 333, 335,

338, 341-343, 350, 353; pl. 24
Pyrus sp., 62

Quail, Japanese, 277

Quercus alba, 152
pubescens, 38
rubra, 152
Spp., 153
virginiana, 152

Quinones, 347

Rana catesbeiana, 271
mascareniensis, 271
pipiens, 271, 345

Ranatra sp., 227

Ranidae, 270

Rats, 4, 287
kangaroo, 23, 20
Norway, 287
wood, 23, 26

Rattus norvegicus, 287
rattus, 287
spp., 287

Reduviidae, 227

Reduviids, undetermined, 227

Reduvius christophi, 227
fedtschenkianus, 227

Renealmia sp., 151

Repellents, 345-347

Reptiles, 3, 272-276, 353
eggs of, 320

Reptilia, 272-276

Reticulitermes sp., 317

Retortamonas blattae, 167, 168
orthopterorum, 169
sp., 167

Rhabditoidea, 192

Rhamdia sebae, 268

Rhantus pacificus, 230

Rhicnoda natatrix, 13, 33
sp., 146

Rhinopsis caniculatus, 257

Rhinotermitidae, 310

Rhipidioides ableptus, 230
adynatus, 230
fuscatus, 230
helenae, 230
mollis, 231
rubricatus, 231

Rhipidius, 230

Rhipiphoridae, 230

Rhizobiaceae, 106

INDEX 465

Rhizoglyphus tarsalus, 218
Rhizophora mangle, 30, 159
Rhizophoracea, 159
Rhizopus nigricans, 133
sp., 133
Rhombomys opimus, 24
Rhynchonympha, 102
tarda, 174, 175
Rhyparobia maderae, 11
Rhythms, diurnal, 335
Rhytidometopum dissimile, 13, 68, 158
Riatia fulgida, 13, 157
orientis, 13, 68, 158
Riboflavin, 101
Rictularia coloradensis, 192, 205
Riekella australis, 231
nitidioides, 231
sp., 231
Ripidiini, biology of Australian, 231
Ripidius, 230
boissyi, 231-232
denisi, 232
pectinicornis, 84, 232-233
scutellaris, 233
sp., 232
Ripiphoridae, 230
Roach (see cockroach)
Robshelfordia circumducta, 13, 307
longiuscula, 13, 307
Rodentia, 287
Rosa sp., 155
Rosaceae, 155
Rose, 155, 163-165
Roundworms, 2 (see Helminths)
Roystonea regia, 143
Rubiaceae, 161
Rubus spp., 155
Rushes, 46
Rutaceae, 157

Sabal palmetto, 50
Saccharomyces cerevisiae, 133
sp., 133
Saccharomycetaceae, 133
Saccharomycetes, 129
Saccharum officinarum, 141-142
Saccinobaculus, 102
ambloaxostylus, 170
doroaxostylus, 170

466 SMITHSONIAN MISCELLANEOUS COLLECTIONS VOL. I4I

Saccinobaculus—C ontinued.
lata, 170
minor, 170
undescribed spp., 170

Saimiri sciurea, 285

Salamander, 269

Salganea morio, 13, 334

Salicaceae, 152

Salientia, 270

Salius verticalis, 256

Salix sp., 152

Salmonella anatis, 114
choleraesuis, 114
enteritidus, 114
morbificans, 115
paratyphi, 115
schotimuelleri, 115
typhimurium, 116
typhosa, 117

Salmonella sp.
(Type Adelaide), 115
(Type Bareilly), 115
(Type Bredeny), 115
(Type Derby), 115
(Type Kentucky), 115
(Type Kottbus), 115
(Type Meleagris), 116
(Type Montevideo), 116
(Type Newport), 116
(Type Oranienburg), 116
(Type Panama), 116
(Type Rubislaw), 116
(Type Tennessee), 116

Salsola kali, 65

Samanea saman, 35, 156

Samut, Steamship, 232

Sapindaceae, 158

Sapotaceae, 160

Sarcina alba, 108
aurantiaca, 108
lutea, 108
spp., 108-109
symbiotica, 108
ventriculi, 108

Sarcodina, 176

Sarcophaga lambens, 2, 229
omani, 229
sternodontis, 229

SPP-, 229

Sarcophagidae, 229
Sarracenia flava, 154
minor, 154
purpurea, 154
variolaris, 154
Sarraceniaceae, 154
Sauria, 272
Scale, diaspine, 237
Sceliphron caementarium, 248
Schistosoma haematobium, 208
Schistosomatidae, 208
Schizomycetes, 104
Schizophyta, 104
Schwenkiella icemi, 200
Scincidae, 274
Scolopendra, 222
cingulata, 223
morsitans, 223
subspinipes, 224
Sp., 224
Scolopendridae, 223
Scolopendromorpha, 223
Scorpionida, 211
Scorpionidae, 213
Scorpions, 3, 2II, 349
Scorzonera acanthoclada, 65, 162
Scutigera coleoptrata, 222-223; pl. 31
forceps, 222
maculata, 223
Scutigeridae, 222
Scutigeromorpha, 222
Sea grape, 43, 69
Seaweed, 47, 55
Secretions, repellent, 6, 216, 225
sexual, 235
Sedge, 66
Serpentes, 276
Serratia marcescens, 117-118; pl. 16
Setaria verticillata, 142
Seurocyrnea colini, 207
Severianoia magna, 200
severianoi, 200
Sewers, cockroaches in, 74, 332, 338
Shelfordella tartara, 8
Shigella alkalescens, 118
dysenteriae, 118
paradysenteriae, 119
Ships, 82-84
cockroaches on, 79, 82-87
slave, 79, 86, 87

INDEX

Shrike, 281
Sibylloblatta panesthoides, 13, 151
Sida, 64, 159, 163
Sideroxylon foetidissimum, 63, 160
Silkworm, bacterium from, 126
“disease bacillus,” 127
Simblerastes jamaicanus, 13, 69, 317,
334
Sumia jacchus, 285
Sinanthropes, 71, 81
Skinks, 274, 354
Snakes, 276
Sodium fluoride, 278
Solanaceae, 161
Solanum tuberosum, 161
Solenopsis geminata, 314
Solindenia picticornis, 246, 247
Sound, cockroach, 330
Sow bugs, 347
Sparassidae, 215
Sparrow, 282
English, 282
Spartan, Ship, 84
Spartina marsh, 42
Spelaeoblatta caeca, 14
gestrot, 13, 22
Spermophilopsis leptodactylus, 24
Sphaerita, 189
Sphaerodactylus sp., 273
Sphagnum, 52
Sphecidae, 264
Sphecophila polybiarum, 13, 314
ravana, 13, 310-311
termitium, 13, 311
Sphodromantis bioculata, 225
viridis, 225
Spicaria prasina, 130
Spiders, 3, 211, 214-216, 317, 318, 347,
349; pl. 30
banana, 215
big, brown house, 215
bird-eating, 214
black widow, 216
button, 215
huntsman, 215
nests, 215-216
Spiniger domesticus, 227
Spirillochaeta blattae, 127

467

Spirillum a, B, and y, 105
periplaneticum, 105, 127
Sp., 105
Spirocerca sanguinolenta, 203
Spirochaeta blattae, 125
blattarum, 125
periplanetae, 125
stylopygae, 125
Spirochaetaceae, 125
Spirochaetales, 125
Spirochaetes, unidentified, 125
“Spirochaetoid bacteria,” 127
Spirorbis pusillus, 14
Spirostomidae, 188
Spirotricha, 187
Spirura gastrophila, 207
Spiruridae, 205
Spiruroidea, 204
Spondias, 67
mombin, 158
purpurea, 158
Sporozoa, 181
Spruce, 44
Squirrel, ground, 23, 24, 26
Stagmomantis carolina, 225-226
Staphylococci, 107
Staphylococcus albus, 107
Staurojoeninidae, 174
Stegobium paniceum, 99
Steinernematidae, 192
Steleopyga sinensis, 13, 307
Stenamma andrei, 316
Stenobothrus vagans, 46
Stenophora sp., 181
Stenophoridae, 181
Sterculiaceae, 159
Stictolampra buqueti concinula, 14, 33
Stigmatomyces sp., 129, 138
Stilpnoblatta minuta, 316
Stipa lessingiana, 46
Strepsiptera, 234
Streptococcus faecalis, 109
liquefaciens, 110
microapoika, 110
non-hemolyticus, 110
pyogenes, 110
Spp., IIO
Streptomyces leidynematis, 124, 1096,
197
Streptomycetaceae, 124

468

Streptopelia chinensis, 278
Streptostomum gracile, 195
Strigidae, 279
Strigiformes, 279
Strongyloidea, 209
Sturnidae, 281
Stylogaster spp., 228
stylata, 228
Stylopyga americana, 12
Styphon bakeri, 14, 60, 344
Subulura jacchi, 204
Subuluridae, 204
Sugarcane, 37-39, 45, 48, 51, 53-56, 58,
64, 66-60, 141-142
Suifunema caudelli, 200
Sulfur, flowers of, 222
Sunfish, 269
Sunflower, 67
Supella hottentotta, 14, 29
supellectilium, 14, 60, 70, 71, 76,
82, 87, 89, 100, 115, 117, 142, 161,
165, 205, 216, 223, 232, 255, 307,
327, 333, 335, 339, 342, 349-351;
pls. 25, 30, 31, 34
sp., 90
Swift, edible-nest, 318
Swiss chard, 47, 153
Symbiosis, 25-26, 92
Symbius blattarum, 232
Symploce bicolor, 14, 82
bilabiata, 14
breviramis, 14, 22
cavernicola, 14, 21, 22, 318, 333
curta, 14, 22
flagellata, 14, 60, 307
hospes, 14, 82, 161, 330
jamaicana, 14, 69
kevani, 14, 29
lita, 14
parenthesis, 14, 307
remyt, 14, 22
ruficollis, 14, 69, 142, 161, 307
Sp., 90
Syncephalastrum sp., 133
Synonyms, 5
Syntomosphyrum blattae, 248
glossinae, 250
ischnopterae, 249

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOL. I4I

Systellogaster ovivora, 248
Syzygium aromaticum, 160
sepligetella sericea, 242

Tachinid, undetermined, 228
Tachysphex blatticidus, 264
coriaceus, 264
fanuiensis, 264
lativalvis, 264-265
Taenia echinococcus, 208
saginata, 208
Taeniidae, 208
Taenioidea, 208
Tamarind, 51
Tamarindus indica, 156
Tapeworm, undetermined, 208
Tarachodes maurus, 226
Tarantula, 214
Tarantula, 214
Tarsiidae, 284
Tarsius, 284
Tartaroblatta karatavica, 14, 70
Tatu novemcinctum, 287
Taxodiaceae, 140
Tegenaria, 214
Teiidae, 275
Temnopteryx, 134-135
deropeltiformis, 10
obliquetruncata, 14, 317
phalerata, 14, 307
platysoma, 8
Temperature preference, 25, 71
Tenadores, Steamship
Tenodera aridifolia sinensis, 226
Termes bellicosus, 311
flavipes, 316
malaccensis, 311
Sp., 311
Termites, 57, 60, 102, 310-311, 317, 320
comején, 317
Termitidae, 311
Termopsis sp., 317
Testudo horsfieldi, 24
Tetragenous sp., 127
Tetralopha scabridella, 35, 66, 67, 317
Tetrameres americana, 207
pattersont, 207
Tetrastichodes asthenogmus, 249

INDEX

Tetrastichus australasiae, 249
hagenowti, 94, 249-253, 255, 349-
351; Pl. 34
periplanetae, 253
SP., 249, 254, 351
Tetratrichomastix blattidarum, 170
Tettix kraussi, 46
Thallophyta, 129
Thecadactylus sp., 273
Theganopteryx straminea, 14, 30
Thelastoma aligarhica, 200
brevicaudatum, 194
bulhdesi, 201
heterogamiae, 199
icemi, 200
indiana, 194
magalhaesi, 194
pachyjuli, 200-201
palmettum, 201
panesthiae, 108
riverot, 201
Thelastomatidae, 190, 193
Thelaziidae, 204
Thelyphonidae, 211
Thelyphonus giganteus, 211
Theobroma cacao, 159
Theraphosidae, 214
Therea nuptialis, 14, 344
Theridiidae, 215
Thistle, 60
giant, 51
Thyrsocera cincta, 13
Tiaris bicolor omissa, 282
Tillandsia fasciculata, 144
usneoides, 144
uttriculata, 50, 144
sp., 31, 36, 144
Timulla eriphyla, 264
Tivia australica, 14, 316
brunnea, 14, 30
fulva, 14, 28, 30
macracantha, 14, 23
obscura, 14, 30
Sp., 23
Toads, 270, 353
Tobacco, 165
Tockus birostris, 279
Todidae, 279
Todus mexicanus, 279

469

Tody, 279
Torula acidophila, 132

gropengiessert, 132

rosea, 132
Torulopsis, 130
Toxoplasma gondii, 190
Trapping, 53, 59-63
Treeferns, 36, 42, 50, 56, 57, 139
Trees (see under common or scientific

name)

in general, 35-68
Trematoda, 208
Treponema parvum, 125

stylopygae, 125
Treponemataceae, 125
Triatoma arthurneivat, 227
Tribulus sp., 63, 156, 163, 165
Trichomastic orthopterum, 169
Trichomastix, 169
Trichomonadidae, 171
Trichomonas hominis, 171

orthopterorum, 169

Sp. 171
Trichonympha, 102

acuta, 174

algoa, 174, 175

chula, 176

grandis, 176

lata, 176

okolona, 175, 176

parva, 176
Trichonymphidae, 174
Trichostrongylidae, 210
Trichostrongylus sp., 210
Trichotarsus, 218
Trichuridae, 210
Trichuris trichiura, 210
Trichuroidea, 210
Trigonopsis abdominalis, 265
Trirhogma caerulea, 262

sp., 264
Troglobies, 17, 20
Troglodytes aedon, 281

audax, 281
Troglodytidae, 281
Troglophiles, 16, 17, 19, 20, 23
Trogloxenes, 16
Tropidophorus grayi, 274
Trypanosomatidae, 167

470 SMITHSONIAN MISCELLANEOUS COLLECTIONS

Tulip tree, 53
Turkey, 278
Turtle, desert, 24, 26

painted, 272
Typhloblatta caeca, 14, 23
Typhloblattodes madecassus, 14, 23
Tyrophagus lintneri, 218

noxius, 218

Urinympha, 102
talea, 174, 175
Urodacus novaehollandiae, 213
Uropoda sp., 217
Uropodidae, 217

Vaccinium meridionale, 43, 160
Vanda, 151, 163
Veillonella parvula, 109
Vejovidae, 213
Verbenaceae, 161
V eromessor andrei, 316
Vertebrata, 268-290
Vertebrates, 3, 268
in biological control, 353-354
Vespa maculata, 318
Vespidae, 314
Vespula maculata, 61, 318
Vibrio comma, 105
Deneke’s, 105
metschnikovit, 105
sp., 106
types I and II, 106
tyrogenus, 105
Vinca minor, 161
Vireo latimeri, 281
Vireonidae, 281
Viruses, 103-104
Coxsackie, 103-104
encephalomyelitis, mouse, 104
poliomyelitis, 3, 103
Brunhilde type, 103
Columbia SK, 103

VOL. I4I

Viruses—C ontinued.
poliomyelitis—Continued.
Lansing strain, 103
unspecified strains, 103
yellow-fever, 104
Viverridae, 289
Vulpes sp., 288

Wasps, 91, 94, 314, 318, 348
cockroach-egg parasites, 234-254,
350, 352
cockroach-hunting, 255-266, 352
Water, balance, 72
drinking, 72
need for, 71
Water bug, 349
Wattle, 49
Webworm, 316
W einmannia sp., 155
Whipscorpion, 211
William Kieth, Steamship, 84
Willow, 47, 63, 152
Woodpeckers, 279-280
Wren, 281

Xerophiles, 25
Xestoblatta festae, 14, 87
immaculata, 14, 23
Xylosma suaveolens, 1590

Xysticus, 214

Yeasts, 3, 127, 120, 132
Yellow-fever virus, 104
Yucca elata, 54, 146

Zanthoxylum caribaeum, 66, 157

Zea mays, 142, 165

Zelotes, 314

Zeuxevamia splendidula, 93, 243, 249,
350

Zingiberaceae, 151

Zootermopsis, 103

Zygophyllaceae, 156

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Yd!I1 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S°

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> = > = Gs Ss

z w ae Ww . ae Ww)

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