SMITHSONIAN

MISCELLANEOUS COLLECTIONS

VOL. 141

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

(Publication 4470)

CITY OF WASHINGTON
PUBLISHED BY THE SMITHSONIAN INSTITUTION
1961

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

[ADVERTISEMENT]

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

Leonard Carmichael,
Secretary, Smithsonian Institution.

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 141 (WHOLE VOLUME)

THE BIOTIC ASSOCIATIONS OF
COCKROACHES

(With 37 Plates)

LOUIS M. ROTH

AND

EDWIN R. WILLIS

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

(Publication 4422)

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

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

SMITHSONIAN MISCELLANEOUS COLLECTIONS

VOLUME 141 (WHOLE VOLUME)

THE BIOTIC ASSOCIATIONS OF
COCKROACHES

(With 37 Plates)

LOUIS M. ROTH

AND

EDWIN R. WILLIS

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

(Publication 4422)

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

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

[FOREWORD]

People having only casual interest in insects usually express amazement when they learn how much is known about this most numerous group of animals. However, while entomologists have good reason to take pride in the accomplishments of their contemporaries and predecessors, they are more likely to be appalled by how much remains to be learned. We are indeed ignorant of even the identity of fully half and probably much more than half the total number of insect species. Of those that have been described, we have reasonably complete information about the behavior and basic environmental relationships for only a comparative few. The great majority of the remainder are known only from specimens found in museum collections. Such information as we have about these species usually amounts to no more than date and locality of collection.

This is true of the cockroaches, which now include approximately 3,500 described species. Conservative estimates based on partially studied museum collections and the percent of new species found in recent acquisitions, particularly from tropical and subtropical countries, indicate that at least 4,000 species remain unnamed. Although the group is well known in general terms to nearly all entomologists, there is an almost complete void of information about all except the few domestic species and, to a progressively diminishing degree, some 400 others. Many details about the lives of even those that share man's habitations are not fully understood. This then is a rough measure of how little is known about cockroaches.

With the exception of mosquitoes and a few other comparatively small groups of insects on which work has been concentrated, it is doubtful if any other comparable segment of the world's insect fauna is better known. Already an estimated 800,000 kinds of insects have been described, and since this figure is generally regarded as less than half the actual total, think what this means in terms of knowledge yet to be assembled. No wonder entomology is a growing science with a promising future, but the magnitude of the task also presents a serious obstacle to progress. Progress can continue only if the scattered literature resulting from the diversified labors of hundreds of contributors is brought together and summarized in thorough and well-organized compilations that can serve as a solid basis for future research.

The present work is such a compilation, for it assembles what has been gleaned from approximately 1,700 sources, including correspondence with a large number of other workers. Original observations during some eight years of concentrated effort in U. S. Army Quartermaster research laboratories are a valuable supplement to what others have done, and with this background of experience the authors are especially well qualified to appraise previous work. Seldom has a compilation been done so thoroughly or a single large group of insects been the subject of such uninterrupted effort.

The contents gives the categories of subject matter treated and the introduction discusses the value of this assembled information and offers suggestions for future study. No longer are cockroaches regarded only as disagreeable pests; many species appear to be important, actually or potentially, as carriers of disease. Recognition of this importance has grown considerably, even in the period since World War II. Consequently, anything that increases our knowledge of the basic bionomics of cockroaches will be consulted widely for factual information and for clues to new approaches.

In spite of this extensive compilation, the limitations of present information about cockroach bionomics must be kept in mind. The cited observations of many writers were fragmentary, or their conclusions disagreed. But it is fundamental to scientific inquiry that we should know and attempt to evaluate the results of previous study, and that is what Drs. Roth and Willis have done. Fortunately, their review is readily available. Sometimes, a piece of work fails to be of maximum value because the results are not generally accessible to later students. For this reason I am especially glad that the Smithsonian Institution, by disseminating the results of the authors' labors, has this opportunity to exercise one of its traditional functions—that of diffusing knowledge.

Throughout the period of research by Drs. Roth and Willis at Natick, I was in frequent correspondence with them, and I admire their many accomplishments. Our warmest commendations should go not only to them personally but also to those in administration who encouraged their fundamental research and who aided in the financial support of this publication.

Ashley B. Gurney
Entomology Research Division
United States Department of Agriculture

[CONTENTS]

						Page
Foreword						[iii]
List of Plates						[vii]
List of Illustrations						[viii]
	I.	Introduction				[1]
			Historical			[2]
			Methods			[4]
			Future work			[5]
			Illustrations			[7]
	II.	Species of cockroaches				[7]
	III.	Ecological relationships				[14]
			Cave habitats			[16]
				Cavernicolous cockroaches		[17]
				Cockroaches from burrows		[23]
			Desert habitats			[25]
				Desert cockroaches		[27]
			Aquatic habitats			[30]
				Amphibious cockroaches		[31]
			Outdoor habitats			[33]
				Cockroaches from outdoor habitats		[35]
			Structural habitats			[70]
				Land-based structures		[73]
					Cockroaches associated with land-based structures	[74]
				Ships		[82]
					Cockroaches associated with ships	[85]
				Aircraft		[87]
					Cockroaches associated with aircraft	[88]
	IV.	Classification of the associations				[91]
	V.	Mutualism				[96]
			Bacteroids			[96]
				Cockroaches in which bacteroids have been found		[99]
			Bacteria			[100]
			Protozoa			[101]
	VI.	Viruses associated with cockroaches				[103]
	VII.	Bacteria associated with cockroaches				[104]
	VIII.	Fungi and yeasts				[127]
			Fungi associated with cockroaches			[129]
	IX.	Higher plants associated with cockroaches				[139]
			Damage to plants by cockroaches			[162]
	X.	Protozoa associated with cockroaches				[166]
	XI.	Helminths associated with cockroaches				[190]
			Helminths for which cockroaches serve as primary hosts			[192]
			Helminths for which cockroaches serve as intermediate hosts			[203]
			Helminths whose eggs have been carried by cockroaches			[208]
	XII.	Arthropoda associated with cockroaches				[210]
			Arachnida			[211]
			Chilopoda			[222]
			Insecta			[224]
			Hymenoptera			[234]
			Predators and parasites of cockroach eggs			[234]
			Host selection by egg parasites			[254]
			Cockroach-hunting wasps			[255]
			Ants predaceous on cockroaches			[266]
	XIII.	Vertebrata associated with cockroaches				[268]
			Pisces			[268]
			Amphibia			[269]
			Reptilia			[272]
			Aves			[276]
			Mammalia			[283]
	XIV.	Checklist of cockroaches and symbiotic associates				[290]
	XV.	Cockroaches as commensals				[310]
			Hosts of commensal cockroaches			[310]
			Checklist of commensal cockroaches with their hosts			[315]
			Obscure associations			[316]
	XVI.	Cockroaches as predators				[319]
			Interspecies predation			[319]
			Intraspecies predation			[322]
	XVII.	Associations among cockroaches				[324]
			Familial associations			[325]
			Gregariousness			[330]
			Intraspecies fighting			[336]
			Interspecies compatibility			[337]
			Interspecies antagonism			[341]
	XVIII.	Defense of cockroaches against predators				[343]
	XIX.	The biological control of cockroaches				[348]
			Invertebrates			[349]
			Vertebrates			[353]
Acknowledgments						[354]
References						[356]
Index						[441]

[LIST OF PLATES]

Plate		Page
1	Blaberus craniifer, c. X 2. 1. (Photograph by Jack Salmon, Philadelphia Quartermaster Depot.).	[a]
2	Blaberus craniifer, nymph. (Photograph by Jack Salmon.)	[A-3]
3	Blaberus giganteus, c. X 2.2. (Photograph by Jack Salmon.)	[A-4]
4	Blatta orientalis, c. X 3.8. A, Male. B, Female. (Photographs by Jack Salmon.)	[A-5]
5	A-B, Blattella germanica, c. X 5.2. A, Male. B, Female. C-D, Blattella vaga, c. X 5.2. C, Male. D, Female with oötheca.	[A-6]
6	Byrsotria fumigata, c. X 2. A, Brachypterous male. B, Macropterous male. C, Female.	[A-7]
7	A and B, Cariblatta lutea minima, X 10. A, Male. B, Female with partly formed oötheca. C, Ectobius pallidus, female with completely formed oötheca, X 8. (C, From Roth and Willis [1957].)	[A-8]
8	A, Cryptocercus punctulatus, c. X 4.6. (Photograph by Jack Salmon.) B, Panesthia australis, X 2.8.	[A-9]
9	Cutilia sp. near sedilloti, c. X 5. A, Male. B, Female.	[A-10]
10	Diploptera punctata, c. X 5. A, Male. B, Female.	[A-11]
11	Eurycotis floridana, c. X 2.8. A, Male. B, Female. (Photographs by Jack Salmon.)	[A-12]
12	A-B, Gromphadorhina portentosa, c. X 1.5. A, Male nymph. B, Adult female. C, Coleolaelaps (?) sp., a mite from G. portentosa, c. X 32. (Glycerine jelly preparation and photograph of C by Dr. Barbara Stay.)	[A-13]
12A	Ischnoptera deropeltiformis, c. X 5.3. A, Male. B, Female.	[A-14]
13	Leucophaea maderae, c. X 2.2. A, Male. B, Female. (Photographs by Jack Salmon.)	[A-15]
14	Nauphoeta cinerea, c. X 3.4. A, Male. B, Female.	[A-16]
15	Neostylopyga rhombifolia, c. X 3.4. A, Male. B, Female with partially formed oötheca.	[A-17]
16	Panchlora nivea, X 4.5. A, Dead individual showing normal, pale green coloration. B, Dead individual showing the bright red coloration (very dark areas) characteristic of infection with Serratia marcescens. C, Living male. D, Living female.	[A-18]
17	A, Parcoblatta pensylvanica, female with completely formed oötheca, X 4. B, Parcoblatta virginica, female with partly formed oötheca, X 7.3.	[A-19]
18	Parcoblatta uhleriana, c. X 5.5. A, Male. B, Female with oötheca.	[A-20]
19	Periplaneta americana, c. X 3. A, Male. B, Female. (Photographs by Jack Salmon.)	[A-21]
20	Periplaneta australasiae, c. X 3.2. A, Male. B, Female. (Photographs by Jack Salmon.)	[A-22]
21	Periplaneta brunnea, c. X 2.9. A, Male. B, Female.	[A-23]
22	Periplaneta fuliginosa, c. X 2.9. A. Male. B. Female.	[A-24]
23	Platyzosteria novae seelandiae, c. X 2.9. A. Male. B, Female with oötheca.	[A-25]
24	Pycnoscelus surinamensis, c. X 3.7. A, Male from Hawaii. B, Macropterous parthenogenetic female from Florida. C, Brachypterous nonparthenogenetic female from Hawaii. D, Late instar nymph. (Photograph of nymph D, by Jack Salmon.)	[A-26]
25	Supella supellectilium, c. X 6.3. A, Male. B, Female.	[A-27]
26	Bacteroids from Blattella germanica. A, Part of abdomen showing mycetocytes in fat body, X 225. B, Lobe of fat body showing 3 mycetocytes, X 750. C, Single mycetocyte; bacteroids appear hollow as result of fixation in Carnoy's fluid, X 1725. D, Smear of fat body showing bacteroids in various stages, X 1800. (All preparations and photographs through the courtesy of Dr. Marion A. Brooks.)	[A-28]
27	Fungi parasitic on cockroaches. A, Herpomyces arietinus growing on antennae, legs, body, and cerci of a nymph of Parcoblatta virginica, X 7. B, Herpomyces stylopygae on antenna of Blatta orientalis, X 35. (Reproduced from Richards and Smith [1955, 1956].) C, Herpomyces sp. [probably H. stylopygae] on antenna of B. orientalis, X 132. (Photographs B and C through the courtesy of Dr. A. G. Richards.)	[A-29]
28	A-B, Gregarines (Diplocystis sp.?) from Blaberus craniifer. A, Organisms removed from intestine, X 50. B, Organisms removed from hemocoele, X 32. C, Gregarine cysts in feces of Leucophaea maderae, X 12.	[A-30]
29	A, Undetermined mermithid that parasitizes Ectobius pallidus. X 9. The worm has partly emerged from the neck region of the cockroach. (Reproduced from Roth and Willis [1957].) B, Undetermined gordian worm that parasitized Eurycotis floridana shown beside its host, X 1.8. (Specimen courtesy of Dr. T. Eisner.)	[A-31]
30	A, Heteropoda venatoria, a cockroach-hunting spider, slightly less than natural size, on bananas. (Reproduced from a Kodachrome transparency through the courtesy of Dr. B. J. Kaston.) B to E, Lycosa sp. (avida?) capturing and feeding on a nymph of Supella supellectilium in the laboratory, X 1.4.	[A-32]
31	The centipede Scutigera coleoptrata capturing and feeding on cockroaches in the laboratory. A to E, Pursuit, capture, and eating of a nymph of Supella supellectilium, c. X 1.2. F, Centipede feeding on adult of Blattella germanica, X 1.8.	[A-33]
32	The mantid Hierodula tenuidentata (?) devouring a nymph of Periplaneta australasiae, c. X 1.5.	[A-34]
33	A, Prosevania punctata ♂ beside an oötheca of Periplaneta americana, X 5. B, Hyptia harpyoides with oötheca of Parcoblatta uhleriana from which it had emerged, X 5. C, Larva of a lampyrid beetle feeding on Parcoblatta virginica in the laboratory, X 4.	[A-35]
34	Chalcid parasites of cockroach eggs. A, Anastatus floridanus ovipositing into an oötheca which is still being carried by Eurycotis floridana, c. X 4. B, Comperia merceti ovipositing into an oötheca of Supella supellectilium, c. X 13. C, Tetrastichus hagenowii ovipositing into an oötheca of Periplaneta americana, c. X 10. (C from Roth and Willis [1954b].)	[A-36]
35	A, The wasp finds a cockroach. B, She stings the 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.)	[A-37]
36	Chemical defense of Diploptera punctata against predators; the spray pattern is displayed on KI-starch indicator paper. A, Spray pattern after right mesothoracic leg was pinched. B, Cumulative spray pattern after left mesothoracic leg of the same insect was pinched. C, The defensive glands of the cockroach on the left had been excised, and it is under persistent attack by ants from a laboratory colony of Pogonomyrmex badius (Latreille). The intact cockroach on the right was also attacked by the ants, but it discharged a spray of quinones and repelled the attackers. (From Eisner [1958], through the courtesy of Dr. T. Eisner.)	[A-38]

[LIST OF FIGURES]

Fig.		Page
1	Diagram illustrating the relationship between a mature plant of Herpomyces stylopygae and the integument of Blatta orientalis. (Reproduced from Richards and Smith [1956], through the courtesy of Dr. A. G. Richards.)	[128]
2	Representative Protozoa associated with cockroaches. A, Monocercomonoides melolonthae, X 3094 (after Grassé). B, Coelosporidium periplanetae, X 1310 (after Sprague); trophozoite with spores and chromatoid bodies. C, Endamoeba blattae, X 273 (after Kudo); trophozoite. D, Lophomonas striata, X 330 (after Kudo). E, Lophomonas blattarum, X 660 (after Kudo). F, Retortamonas blattae, X 3094 (after Wenrich). G, Nyctotherus ovalis, X 175 (after Kudo). H, Gregarina rhyparobiae, c. X 52: mature trophozoite attached to intestinal wall of Leucophaea maderae. (Redrawn from J. M. Watson [1945].) I, Diplocystis schneideri, c. X 14.4 (after Kunstler). J, Gregarina blattarum, c. X 57 (after Kudo). K, Protomagalhaesia serpentula, X 36 (after Pinto). L, Gamocystis tenax, magnification not known (after Schneider). (All figures except H redrawn from Kudo [1954] after sources indicated.)	[168]
3	Protozoa from the gut of the wood-feeding cockroach Cryptocercus punctulatus. A, Eucomonympha imla, female above, male below, c. X 375. (From Cleveland [1950c].) B, Barbulanympha sp. (From Cleveland [1953].) C, Urinympha talea, c. X 712. (From Cleveland [1951a].) D, Rhynchonympha tarda, c. X 450. (From Cleveland [1952].) E, Trichonympha okolona or T. algoa, c. X 390. (From Cleveland [1949].) (All drawings reproduced through the courtesy of Dr. L. R. Cleveland.)	[175]
4	The cockroach mite, Pimeliaphilus podapolipophagus. (From Baker et al. [1956]; reproduced through the courtesy of Dr. E. W. Baker and the National Pest Control Association.)	[221]
5	Evania appendigaster. Left, dorsal view, X 8. Right, side view, X 5. (Reproduced with permission. British Museum [Natural History], 1951, figs. 1A and 1B.)	[238]
6	Ampulex canaliculata attacking Parcoblatta virginica. A, Female wasp stinging her prey, c. X 4.8. B, Wasp's egg attached to the coxa of the mesothoracic leg of the cockroach. C, Larva of A. canaliculata (about three-quarters grown) feeding on the internal organs of the host from the exterior, c. X 4. (Reproduced from F. X. Williams [1929], through the courtesy of Dr. F. X. Williams and F. A. Bianchi, Hawaiian Sugar Planters' Association.)	[258]
7	Cockroach-hunting wasps. A, Dolichurus stantoni leading a nymph of Blattella lituricollis to her nest, c. X 4. (Reproduced from F. N. Williams [1919].) B, Podium haematogastrum attaching her egg to an Epilampra sp. while on the side of a termite mound that contains the wasp's nest, c. X 1.6. C, Epilampra sp. parasitized by P. haematogastrum showing the wasp's egg attached to the right fore coxa, c. X 3.2. (B and C reproduced from Williams [1928], through the courtesy of Dr. F. X. Williams and F. A. Bianchi.)	[263]

THE BIOTIC ASSOCIATIONS OF COCKROACHES^[1]

By Louis M. Roth and Edwin R. Willis^[2]

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

(With 37 Plates)

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

Hanitsch (1928)

[I. INTRODUCTION]

Recently we brought together much of the literature linking cockroaches with the transmission of certain organisms that cause disease in man and other vertebrates. In that paper (1957a) we concluded that cockroaches, being potential vectors of pathogenic agents, should not be regarded simply as minor annoyances. Obviously the associations of cockroaches with agents of vertebrate diseases are of more immediate importance than their relations with pathogens of lower animals or with nonpathogens. On the other hand, cockroaches are of general economic as well as medical importance, and their control is sought by many who are unaware of their medical significance. That the control of domiciliary cockroaches is far from satisfactory may be inferred from current entomological and pest-control journals in which new insecticides are continually advocated to replace others found to be inadequate. Possibly new approaches to the control of cockroaches are needed. Whether these lie in the direction of increased use of parasites and predators for the biological control of these insects remains to be seen. In any event, the more we know about any insect, especially its ecology, the greater the likelihood of achieving satisfactory control. In order to advance knowledge in any field of science, new research should proceed from the results of prior investigations when these exist. We hope that the observations and experiments cited herein may suggest areas for future research and exploitation.

To the best of our knowledge no previous publication has brought together the vast literature on the parasites, predators, commensals, and other symbiotic associates of the Blattaria. For this reason, we have tried to assemble observations on all such known associations. Undoubtedly we have overlooked some records, as, for example, those buried in papers dealing with other phases of cockroach biology. We hope that such inadvertent omissions will not seriously impair the usefulness of this compilation. Whatever its defects, this review should be a unified source of information for all who are interested in the biotic associates of cockroaches.

In addition to previously published information, this monograph also contains original records and observations on the associations of cockroaches that are reported here for the first time. Although some of the observations were made by us, others were made by colleagues who have graciously made their knowledge available to us in private communications.

HISTORICAL

Chopard (1938) in his book La Biologie des Orthoptères reviewed much of the literature on cockroaches, but of the many biotic associations that exist he discussed only the commensal cockroaches, gregariousness, and familial associations. Asano (1937), who reviewed the natural enemies of cockroaches, mentioned about 10 groups of animals that attack cockroaches. Thompson (1951) in his Parasite Host Catalogue, which was based mainly on papers abstracted or noted in the Review of Applied Entomology, listed only 19 insect parasites of cockroaches. Eighteen of these were Hymenoptera which attack only cockroach eggs; the single dipteron listed (Sarcophaga lambens Wiedemann, supposedly parasitic on Pycnoscelus surinamensis) is not a parasite in this case, but deposits its eggs on the dead insects (see p. 229). Cameron (1955) listed as parasites and predators of the cockroach 24 species of hymenopterous egg parasites, 7 species of Ampulex which hunt nymphs and adults, 17 Protozoa, 13 nematodes, 5 bacteria, 2 mites, and a few other miscellaneous predators. In his classified list of the protozoan parasites of the Orthoptera of the world, Semans (1943) listed about 26 species from cockroaches. Linstow (1878, 1889) recorded 14 species of helminths from cockroaches. Van Zwaluwenburg (1928) listed 33 names of roundworms which are commensals or secondary parasites of cockroaches, but some of these names are synonyms. La Rivers (1949) extended this list with 13 additional species. Chitwood (1932) recognized 24 species of nematodes which are primary parasites (probably commensals) of blattids. Steinhaus (1946) gave many instances of biological relationships between cockroaches and bacteria, fungi, and yeasts, but the cockroaches were not discussed as an entity and the information is scattered throughout the book.

In surveying the literature on this subject we have collected a far more extensive list of animals and plants associated with cockroaches than one might have expected from an examination of any one of the previous papers on this subject. In our review of the medically important organisms associated with the Blattaria, we pointed out that in addition to many experimental associations cockroaches have been found to harbor, naturally, 4 strains of poliomyelitis virus, about 40 species of pathogenic bacteria, the eggs of 7 species of pathogenic helminths, and to serve as intermediate hosts of 12 other species of helminths pathogenic for vertebrates; cockroaches have also been found to carry, on occasion, 3 species of Protozoa that are pathogenic to man and 2 species of fungi which are sometimes found associated with pathological conditions.

In addition to the above organisms of medical importance, we have compiled records of other organisms, nonpathogenic to vertebrates, which are naturally associated in some way with cockroaches. None of the following numbers can be considered absolute because some names may be synonyms. However, we believe that these figures are very close to the actual numbers of species that have been isolated because we have attempted to refer all obvious synonyms to the currently accepted name for each organism. On this basis there are about 45 species of bacteria, 40 fungi, 6 yeasts, 90 Protozoa, and 45 helminths that have been found associated naturally with cockroaches. Of the arthropods there are about 2 species of scorpions, 4 spiders, 15 mites, 4 centipedes, and 90 insects. Of vertebrates there are 4 species of fish, 16 amphibians, 12 reptiles, 20 birds, and 27 mammals. Besides these there are many records of experimental associations that have been contrived in the laboratory.

Some idea of the increase in our knowledge of the biotic associations of cockroaches, during the last 70 years, may be gathered from a comparison of the above figures with those of Miall and Denny (1886) who presented " ... a long list of parasites which infest the Cockroach." This list included 2 bacteria, 6 Protozoa (some of the names are synonyms), 7 nematodes (some of these names are also synonyms), 1 mite, 1 wasp, and 1 beetle. In addition, they mentioned as other foes of the cockroach: monkeys, hedgehogs, polecats, cats, rats, birds, chameleons, and frogs.

METHODS

We have listed the organisms known to be associated with cockroaches systematically by phylum, class, order, and family. Within each family the organisms are listed alphabetically by genus and species. Under each organism the associated cockroaches are listed as natural or experimental hosts, vectors, or prey. Identified cockroaches are listed by the currently accepted name. Unidentified cockroaches are indicated by the word "Cockroaches." The name of each cockroach is followed by the country in which the observation was made, the authority for the record, and with a few exceptions [3] pertinent biological information, where this is known. Question marks following the names of organisms or countries indicate tentative or questionable identifications.

Records of predators capturing and feeding on cockroaches in zoos and on shipboard we consider natural, even though it is very likely that these particular predators would not normally have access to this prey in nature.

Experimental prey are cockroaches that were fed to predators in the laboratory. Although these predators may have little, if any, access to these cockroaches in nature, we have included such records to indicate the relative acceptability of cockroaches as food by a wide variety of animals.

Records of presumed or known cockroach associates that give no information about an associated cockroach are not included in this review, even though certain of these (e.g., species of Ampulex, Evania, Podium) probably prey upon or parasitize cockroaches exclusively.

The validity of a host-parasite or predator-prey record is dependent upon the accuracy and knowledge of the observer. In assembling these records we have had to accept, in most instances, the identifications of species made by the original authors. However, as a result of our studies on the biology of various species of cockroaches, including some work on their hymenopterous parasites, we have questioned certain records in the literature. Other dubious records which have been perpetuated from one publication to the next, but which apparently were not based on fact, have also been questioned or have been clarified with the aid of specialists in particular groups.

Because the records cover a period of many years, the names of many of the organisms as well as the names of some of the cockroaches have been changed. Although it would have been comparatively simple to list the names as they appeared in the original references, this would have resulted in misleading redundancy with the same organism being catalogued under several synonyms. We have attempted to list each organism by its currently accepted name. However, no attempt was made to prepare complete taxonomic synonymies; the only synonyms given are those that identify the organisms by the names used by the authors of the papers cited. The synonyms under which the cockroaches may have been cited originally are listed in section II. The synonyms of associated organisms are listed with each organism. Although authorities for the name changes of the cockroaches are given, these workers are not necessarily those who were initially responsible for the synonymies. Various sections have been checked by specialists in the particular groups. Although we have accepted name changes suggested by these reviewers, we assume full responsibility for the names.

FUTURE WORK

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

Our understanding of most predator-prey and parasite-host relationships has barely progressed beyond the taxonomic stage. The total effect of predators and parasites in limiting natural populations of cockroaches remains to be determined. It is still not known how, for example, predatory or parasitic wasps select specific cockroaches from among all other insects. Secretions produced by certain cockroaches (e.g., 2-hexenal by Eurycotis floridana) will ward off certain predators. The identities and biological activities of most cockroach secretions are unknown, but the use of protective chemicals against predators may be widespread among cockroaches. If so, how effective are these repellents in protecting the individual or the species? It is not known whether cockroaches are protected by apparent mimetic resemblances to other arthropods. There is no experimental proof that insect parasites can successfully attack the eggs of cockroaches that incubate their eggs while they are being carried by the female.

It is conceivable that biological control of cockroaches might be achieved in limited areas such as man-made structures or sewers, but this possibility has not been thoroughly explored. It would be informative to know what effects, if any, organisms such as bacteria, Protozoa (e.g., gregarines), intestinal nematodes, or other helminths have on cockroaches. Possibly pathogenic microorganisms can be used for biological control of cockroaches; this approach seems to have been little explored.

Associations of colonial cockroaches (e.g., Cryptocercus spp.) may be truly familial or they may merely result from gregariousness. Newly hatched nymphs of species that carry their oöthecae until the eggs hatch cluster near the mother. This may be a response to the mother as such, a search for shelter beneath the nearest object (thigmotaxis? or negative phototaxis?), or there may well be yet another explanation. Tepper in 1893 stated that the native cockroaches of Australia are almost wholly carnivorous; little supporting evidence for this claim has been brought forward since that time. The apparent supersedure of one species of domiciliary cockroach by another may result from antagonism between different species, or it may result from more rapid breeding and more effective utilization of available food and space; but which? Several species of cockroaches are frequently found associated with certain plants (e.g., bromeliads and bananas); the ecological relations in these associations remain to be determined. Many of the obscure associations between cockroaches and other insects, spiders, birds, and burrowing animals have never been adequately defined. The factors influencing cannibalism have never been thoroughly investigated experimentally. These are only a few ideas for future work that have occurred to us during the preparation of this review. We hope that these suggestions as well as other questions that may occur to readers will stimulate further research in areas where it is obviously needed.

ILLUSTRATIONS

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

[II. SPECIES OF COCKROACHES]

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

Agis orientalis Chopard

Aglaopteryx absimilis Gurney

diaphana (Fabricius) [Ceratinoptera diaphana Fabricius; Rehn and Hebard (1927)]

facies (Walker) [Aglaopteryx devia Rehn; Princis (1929). A. diaphana (Fabricius) in records from Puerto Rico only; Rehn (1932b); Gurney (1937)]

gemma Hebard [In Florida records = Ceratinoptera diaphana R. and H.; Hebard (1917)]

vegeta Rehn

ypsilon Princis

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

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

Amazonina emarginata Princis

Anaplecta asema Hebard

azteca Saussure

decipiens Saussure and Zehntner

fallax Saussure

hemiscotia Hebard

lateralis Burmeister

mexicana Saussure

Aneurina tahuata Hebard

viridis Hebard

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

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

Apteroblatta perplexa Shelford

Archiblatta hoevenii Vollenhoven

Archimandrita marmorata (Stoll)

tessellata Rehn

Arenivaga apacha (Saussure)

bolliana (Saussure)

erratica (Rehn)

floridensis Caudell

grata Hebard

roseni (Brancsik) [Heterogamodes roseni; Bei-Bienko (1950). "Polygamia" roseni is undoubtedly an erroneous citation of Polyphaga roseni, as there is no genus Polygamia (Gurney, personal communication, 1957)].

tonkawa Hebard

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

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

cavernicola J. W. H. Rehn

deplanatus (Saussure)

Attaphila aptera Bolívar

bergi Bolívar

flava Gurney

fungicola Wheeler

schuppi Bolívar

sexdentis Bolívar

Atticola mortoni Bolívar

Audreia bromeliadarum Caudell

jamaicana Rehn and Hebard

Balta godeffroyi (Shelford)

patula (Walker)

platysoma (Walker) [Temnopteryx platysoma (Walker); Hebard (1943)]

quadricaudata Hebard

scripta (Shelford)

torresiana Hebard

verticalis Hebard

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

Blaberus atropos (Stoll) [Blabera fusca Brunner; Hebard (1917)]

boliviensis Princis

craniifer Burmeister (pls. [1], [2])

discoidalis Serville [Blaberus cubensis Saussure; Hebard (1916)]

giganteus (Linnaeus) (pl. [3])

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

Blatta orientalis Linnaeus (pl. [4]) [Periplaneta orientalis; Hebard (1917)]

(Shelfordella) lateralis (Walker) [Shelfordella tartara (Saussure); Princis (1957). Periplaneta tartara Saussure; Bei-Bienko (1950)]

Blattella germanica (Linnaeus) (pls. [5], A, B; [31], F) [Blatella germanica; Gurney (1952). Phyllodromia germanica; Hebard (1917). Ectobius germanicus; Gurney (personal communication, 1957)]

humbertiana (Saussure) [Blatta humbertiana; Phyllodromia humbertiana; Hebard (1929)]

lituricollis (Walker) (fig. [7], A) [Blattella bisignata (Brunner); Bei-Bienko (1950)]

schubotzi Shelford

vaga Hebard (pl. [5], C, D)

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

Byrsotria cabrerae Rehn and Hebard

fumigata (Guérin) (pl. [6])

Cahita borero Rehn

nahua (Saussure)

Capucinella delicatula Hebard

Cariblatta antiguensis (Saussure and Zehntner)

cuprea Hebard

delicatula (Guérin) [Blattella delicatula Guérin; Cariblatta punctulata (Beauvois); Rehn and Hebard (1927)]

hylaea Rehn

imitans Hebard

insularis (Walker)

landalei Rehn and Hebard

lutea lutea (Saussure and Zehntner)

lutea minima Hebard (pl. [7], A, B)

nebulicola Rehn and Hebard

orestera Rehn and Hebard

punctipennis Hebard

reticulosa (Walker)

stenophrys Rehn and Hebard

Cariblattoides instigator Rehn and Hebard

suave Rehn and Hebard

Ceratinoptera picta Brunner

Chorisoneura barbadensis Rehn and Hebard

flavipennis Saussure and Zehntner

formosella Rehn and Hebard

parishi Rehn

specilliger Hebard

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

translucida (Saussure)

Choristima sp.

Choristimodes sp.

Chromatonotus infuscatus (Brunner)

notatus (Brunner)

Compsodes schwarzi (Caudell)

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

Cosmozosteria lateralis (Walker)

Cryptocercus punctulatus Scudder (pl. [8], A)

relictus Bei-Bienko

Cutilia nitida (Brunner)

soror (Brunner)

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

Cyrtotria capucina (Gerstaecker)

Dendroblatta sobrina Rehn

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

lampyrina Gerstaecker

porcellio Gerstaecker

Deropeltis autraniana Saussure

erythropeza Adelung

melanophila (Walker)

nigrita Saussure

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

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

Ectobius africanus Saussure

albicinctus (Brunner)

duskei Adelung

lapponicus (Linnaeus) [Ectobius perspicillaris Herbst, as used by Lucas (1920); Blair (1934)]

lucidus Hgb.

nicaeensis (Brisout)

pallidus (Olivier) (pls. [7], C; [29], A) [Ectobius lividus (Fabricius); Ectobius livens (Turton); Kevan (1952); Princis (in Roth and Willis, 1957)]

panzeri Stephens [Ectobius ericetorum (Wesmaël); Ramme (1923)]

panzeri var. nigripes Stephens

semenovi Bei-Bienko

sylvester (Poda) [Ectobius sylvestris (Poda); Ramme (1951)]

tadzihicus Bei-Bienko

vittiventer (Costa) [Ectobius vittiventris (Costa); Ramme (1951)]

Ellipsidion Saussure [Apolyta Brunner; Hebard (1943)]

affine Hebard

australe Saussure [Ellipsidion pellucidum (Brunner); Hebard (1943)]

bicolor (Tepper)

simulans Hebard

variegatum (Fabricius) [Ellipsidion aurantium Saussure; Hebard (1943)]

Epilampra abdomen-nigrum (De Geer)

annandalei Shelford

azteca Saussure

conferta Walker

conspersa Burmeister

grisea (De Geer)

maya Rehn

mexicana Saussure

mona Rehn and Hebard

notabilis Walker

sodalis Walker

tainana Rehn and Hebard

wheeleri Rehn

sp. (fig. 7, B, C)

Eremoblatta subdiaphana (Scudder)

Ergaula capensis (Saussure) [Dyscologamia capensis Saussure; Dyscologamia wollastoni Kirby; Princis (1957)]

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

Escala sp.

Euandroblatta palpalis Chopard

Eublaberus posticus (Erichson)

Eudromiella bicolorata Hebard

calcarata Bei-Bienko

Euphyllodromia angustata (Latreille)

liturifera [Euphyllodromia decastigmata Hebard; Princis (1959)]

Eurycotis bananae Bei-Bienko

biolleyi Rehn [Eurycotis carbonaria Biolley; Rehn (1918)]

caraibea (Bolívar)

decipiens (Kirby)

dimidiata (Bolívar)

ferrum-equinum Rehn and Hebard

floridana (Walker) (pl. II) [Platyzosteria ingens Scudder; Platyzosteria sabalianus Scudder and hence, by inference, Eurycotis sabalianus (Scudder); Hebard (1917)]

galeoides Rehn and Hebard

improcera Rehn

kevani Princis

lixa Rehn

manni Rehn

opaca (Brunner)

Euthlastoblatta abortiva (Caudell)

Euthyrrhapha nigra Chopard

pacifica Coquebert

Geoscapheus robustus Tepper

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

Gromphadorhina laevigata S. and Z.

portentosa (Schaum) (pl. [12], A, B)

Gyna kazungulana Giglio-Tos

maculipennis (Schaum) [Gyna vetula Brunner; Shelford (1909b)]

tristis Hanitsch

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

Hemiblabera brunneri (Saussure)

Henicotyle antillarum (Brunner)

Heterogamodes krügeri (Salfi)

rugosa (Schulthess)

Holocompsa azteca (Saussure)

cyanea (Burmeister)

fulva (Burmeister)

metallica Rehn and Hebard

nitidula (Fabricius)

zapoteca Saussure

Hololampra bivittata (Brullé)

chavesi (Bolívar)

maculata (Schreber) [Aphlebia maculata Schreber; Harz (1957); Gurney (personal communication, 1959)]

marginata (Schreber)

punctata (Charpentier) [Aphlebia punctata Charpentier; Ramme (1951)]

Hololeptoblatta sp.

Homalopteryx laminata Brunner

Hoplosphoropyga babaulti Chopard

Hormetica apolinari Hebard

laevigata Burmeister

ventralis Burmeister

Ignabolivaria bilobata Chopard

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

panamae Hebard

podoces Rehn and Hebard

rufa occidentalis Saussure

rufa rufa (De Geer)

schenklingi Karney

Karnyia discoidalis (Brunner)

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

remota Hebard

Lamproblatta albipalpus Hebard

meridionalis (Brunner)

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

lucifrons Hebard

rehni Hebard

vitrea (Brunner)

zapoteca (Saussure)

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

Leurolestes pallidus (Brunner)

Litopeltis biolleyi (Saussure)

bispinosa (Saussure) [Audreia marginata Caudell; Hebard (1920)]

deianira Rehn

musarum Rehn

Lobolampra subaptera Rambur

Loboptera decipiens (Germar)

thaxteri Hebard

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

Lophoblatta arawaka Hebard

Macropanesthia rhinocerus Saussure

Mareta acutiventris Chopard

Maretina uahuka Hebard

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

Megamareta verticalis Hebard

Melanosilpha capensis Saussure and Zehntner

Methana canae Pope

curvigera (Walker)

marginalis (Saussure)

Moluchia (?) dahli Princis

Monastria biguttata (Thunberg)

Muzoa madida Rehn

Myrmeblattina longipes (Chopard)

Myrmecoblatta rehni Mann

wheeleri Hebard

Namablatta bitaeniata (Stål)

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

Nauphoeta cinerea (Olivier) (pl. [14]) [Nauphoeta bivittata Burmeister; Zimmerman (1948)]

flexivitta (Walker) [Nauphoeta brazzae (Bolívar); Rehn (1937)]

punctipennis Chopard

Nelipophygus ramsdeni Rehn and Hebard

Neoblattella brunneriana (Saussure) [Blattella brunneriana; Gurney (personal communication, 1959)]

carcinus Rehn and Hebard

celeripes Rehn and Hebard

detersa (Walker)

dryas Rehn and Hebard

eurydice Rehn and Hebard

fratercula Hebard

fraterna (Saussure and Zehntner)

grossbecki Rehn and Hebard

laodamia Rehn and Hebard

nahua (Saussure) [Blattella nahua Saussure and Zehntner of Caudell (1914); Hebard (1920)]

proserpina Rehn and Hebard

semota Rehn and Hebard

tridens Rehn and Hebard

vatia Rehn and Hebard

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

Nesomylacris cubensis Rehn and Hebard

relica Rehn and Hebard

Nocticola bolivari Chopard

caeca Bolívar

decaryi Chopard

simoni Bolívar

sinensis Silvestri

termitophila Silvestri

Nothoblatta wasmanni (Bolívar)

Notolampra antillarum Shelford

Nyctibora azteca Saussure and Zehntner

brunnea (Thunberg)

laevigata (Beauvois)

lutzi Rehn and Hebard

mexicana Saussure

noctivaga Rehn

obscura Saussure

sericea Burmeister

stygia Walker

tomentosa Serville [Nyctibora latipennis Burmeister; Hebard (1917, p. 263)]

Oniscosoma granicollis (Saussure)

Opisthoplatia maculata Shiraki

orientalis (Burmeister)

Oulopteryx meliponarum Hebard

Oxyhaloa buprestoides (Saussure)

deusta (Thunberg)

Panchlora antillarum Saussure

exoleta Burmeister

fraterna Saussure and Zehntner

nivea (Linnaeus) (pl. [16]) [Panchlora cubensis Saussure; Gurney (1955). Pycnosceloides aporus Hebard; Hebard (1921c)]

peruana Saussure

sagax Rehn and Hebard

virescens (Thunberg)

Panesthia angustipennis (Illiger) [Panesthia javanica Serville; Hebard (1929)]

australis Brunner (pl. [8], B)

laevicollis Saussure

lobipennis Brunner

spadica (Shiraki)

Parahormetica bilobata (Saussure)

Parcoblatta americana (Scudder)

bolliana (Saussure and Zehntner) [Kakerlac schaefferi Rehn; Hebard (1917)]

caudelli Hebard [♀♀ of Ischnoptera insolita R. and H.; Ischnoptera uhleriana fulvescens S. and Z. (in part); Hebard (1917)]

desertae (Rehn and Hebard) [♂♂ of Ischnoptera insolita R. and H.; Hebard (1917)]

divisa (Saussure and Zehntner) [Ischnoptera divisa S. and Z.; Hebard (1917)]

fulvescens (Saussure and Zehntner) [Ischnoptera uhleriana fulvescens S. and Z. (in part); Hebard (1917)]

lata (Brunner) [Ischnoptera couloniana R. and H. (not Saussure); Ischnoptera major R. and H. (not S. and Z.); Hebard (1917)]

notha Rehn and Hebard

pensylvanica (De Geer) (pl. [17], A) [Ischnoptera pennsylvanica Saussure; Hebard (1917)]

uhleriana (Saussure) (pl. [18]) [Ischnoptera uhleriana Saussure; Hebard (1917)]

virginica (Brunner) (pls. [17], B; [27], A; [33], C; fig. [6]) [Ischnoptera borealis Brunner; Hebard (1917)]

zebra Hebard

Pelmatosilpha coriacea Rehn

kevani Princis

marginalis Brunner

purpurascens (Kirby)

rotundata Scudder

vagabunda Princis

Periplaneta americana (Linnaeus) (pls. [19], [35]) [Stylopyga americana; Blatta americana L.; Hebard (1917)]

australasiae (Fabricius) (pls. [20], [32])

brunnea Burmeister (pl. [21])

cavernicola Chopard

fuliginosa (Serville) (pl. [22])

ignota Shaw

lata (Herbst)

Perisphaerus armadillo Serville

glomeriformis (Lucas)

Phaetalia pallida (Brunner)

Phidon (?) dubius Princis

Phlebonotus pallens (Serville)

Pholadoblatta inusitata (Rehn)

Phorticolea boliviae Caudell

testacea Bolívar

"Phyllodromia" treitliana Werner

Phyllodromica brevipennis (Fischer)

graeca (Brunner)

irinae (Bei-Bienko)

maculata (Schreber)

megerlei (Fieber)

polita (Krauss)

pygmaea (Bei-Bienko)

tartara (Saussure)

tartara nigrescens Bei-Bienko

Platyzosteria analis (Saussure) [Polyzosteria analis Saussure; Kirby (1904)]

armata Tepper

bifida (Saussure)

castanea (Brunner)

novae seelandiae (Brunner) (pl. [23]) [Periplaneta fortipes Walker; Shelford (1912); Platyzosteria novae-zealandiae]

scabra (Brunner)

Plectoptera dorsalis (Burmeister)

infulata (Rehn and Hebard)

lacerna Rehn and Hebard

perscita Rehn and Hebard

poeyi (Saussure) [Plectoptera floridana Hebard; Rehn and Hebard (1927)]

porcellana (Saussure)

pygmaea (Saussure)

rhabdota (Rehn and Hebard)

vermiculata Rehn and Hebard

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

indica Walker [Polyphaga pellucida (Redtenbacher); Princis (1957)]

saussurei (Dohrn)

Polyzosteria limbata Burmeister

melanaria (Erichson)

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

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

laticornis Perty

septentrionalis Hebard

Pseudophoraspis nebulosa (Burmeister)

Pycnoscelus niger (Brunner)

striatus (Kirby) [Leucophaea striata Kirby; Gurney (personal communication, 1957)]

surinamensis (Linnaeus) (pl. [24]) [Leucophaea surinamensis (L.); Hebard (1917). Blatta melanocephala Stoll; Kirby (1904)]

Rhicnoda natatrix Shelford

Rhytidometopum dissimile Princis

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

orientis Hebard

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

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

Salganea morio (Burmeister)

Sibylloblatta panesthoides (Walker)

Simblerastes jamaicanus Rehn and Hebard

Spelaeoblatta gestroi Bolívar

Sphecophila polybiarum Shelford

ravana Fernando

termitium Shelford

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

Stictolampra buqueti concinula (Walker)

Styphon bakeri Rehn

Supella hottentotta (Saussure)

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

Symploce breviramis (Hanitsch)

cavernicola (Shelford) [Ischnoptera cavernicola (Shelford); Phyllodromia nigrocincta Chopard; Hebard (1929)]

curta Hanitsch

flagellata Hebard

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

jamaicana (Rehn)

kevani Chopard

parenthesis (Gerstaecker) [Phyllodromia parenthesis Gerstaecker; Rehn (1932)]

remyi (Hanitsch) [Ischnoptera remyi Hanitsch; Chopard (1938)]

ruficollis (Fabricius) [Symploce bilabiata Rehn and Hebard; Princis (1949a)]

Tartaroblatta karatavica Bei-Bienko

Temnopteryx obliquetruncata Chopard

phalerata (Saussure)

Theganopteryx straminea Chopard

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

Tivia australica Princis

brunnea (Chopard)

fulva (Burmeister)

macracantha Chopard

obscura (Chopard)

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

Typhloblattodes madecassus Chopard

Xestoblatta festae (Griffini)

immaculata Hebard

[III. ECOLOGICAL RELATIONSHIPS]

The ecology of extinct cockroaches is necessarily a highly speculative subject. From the coexistence of fossil cockroaches and fossil plants in the same geological stratum, one might conclude that there had been intimate associations between them during prehistoric life. Heer (1864) and Goldenberg (1877) suggested that Carboniferous cockroaches fed on the plants with which they have been found as fossils. Scudder (1879) concurred with this hypothesis. However, Bolton (1911), remarking on the noticeable associations of blattoid wings with vegetable remains, suggested that the cockroaches may have been partly carnivorous, feeding on the snails Spirorbis pusillus, which were attached to the leaves of Cordaites. Yet the proximity of fossil insects and plants in the same geological formation is hardly proof of a similar association during life. In fact, Sellards (1903), Bolton (1921), and Laurentiaux (1951) have all pointed out that the cockroach remains, particularly the more resistant wings, may have been washed into streams by heavy rains and transported with drifting plant material to places where permanent deposits were accumulating.

Some species of fossil cockroaches have long, well-developed ovipositors, very unlike present-day cockroaches whose ovipositors are small and nonprotruding. Brongniart (1889) and Zalesskii (1939, 1953) have suggested that certain Permian and Carboniferous cockroaches with long ovipositors may have inserted their eggs singly into trees and other plants, rather than protecting the eggs with an oötheca. However, Laurentiaux (1951), although conceding the possibility of egg laying in vegetable material, suggested that oviposition into the earth is more probable because of the unbending nature of the ovipositor.

Although the ecological associations of modern cockroaches should be well known from direct observation, actually most species are still little more than names on museum specimens, and our knowledge of them is fragmentary. All too frequently ecological observations have been only incidental to taxonomic or faunistic studies; yet the biological information that is contained in such papers is all that we know of many species. For this reason we have cited these observations in some detail, especially when they were brief; longer accounts of cockroach bionomics, of necessity, have been abstracted.

Very few exclusively ecological studies of insects have included cockroaches. The native woodroaches (Parcoblatta pensylvanica, P. uhleriana, and P. virginica) of the northern United States were included in ecological studies of the Orthoptera by Hubbell (1922), Strohecker (1937), and Cantrall (1943). Fifteen species of cockroaches were included in an ecological study of the Orthoptera of northern Florida by Friauf (1953). The original papers should be consulted for detailed descriptions of the habitats and accounts of the associated plants and other Orthoptera.

In this chapter the cockroaches are grouped into those that have been found in man-made structures and those that occur in other habitats. Certain species may appear in several categories because they live both indoors and out. The structural pests are divided into cockroaches that occur in land-based structures, those on ships, and those in aircraft. The nonstructural cockroaches are divided into those that occur in quite specific habitats (caves, water, and deserts) and those that occur generally out of doors. Nests of various arthropods serve as microhabitats of commensal cockroaches; these latter associations are discussed on pages 310-318.

In this chapter our discussion is limited to the physical environment and specific habitats of cockroaches, and only very general references are made to associated organisms. The relationships of cockroaches to the biota are examined in detail in subsequent chapters. To show the full extent of the associations, the associates, from bacteroids to vertebrates, are arranged phyletically. These associate-centered classifications serve admirably to relate various species of cockroaches within common bounds, but fail to give an integrated account of the total biotic relationships in the ecology of each species. Although physically separated in this monograph, the many associates of each species of cockroach should all be considered in appraising the ecology of that species. To assist the reader to achieve this end, we have included a checklist (p. [290]) which serves as a convenient index to certain organisms associated with particular species of cockroaches.

CAVE HABITATS

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

Cavernicolous cockroaches show varying degrees of dependence on and adaptation to these specialized habitats. Some of the common domiciliary species (Blatta orientalis, Blattella germanica, and Periplaneta americana) may have accompanied man into caves and remained there after he left (Chopard, 1929a, 1936, 1938). Other species, from the paucity of records noting their occurrence in caves, are undoubtedly accidental inhabitants that may never become established. Besides these, however, many other species of cockroaches have established large breeding colonies in caves. Although some of the latter species show very pronounced morphological adaptations to a cave life, many others resemble their noncavernicolous relatives. The possible origin of cavernicolous Orthoptera has been discussed by Chopard (1938).

Cavernicolous cockroaches have been segregated into four groups according to their ability to adapt to their environment and the degree of their specialized evolution (Chopard, 1936, 1938): (1) Trogloxenes: Cockroaches that occur in caves in a sporadic fashion (the domiciliary cockroaches and accidentals such as Ectobius and Heterogamodes). (2) Troglophiles: Cockroaches found habitually in caves (Symploce, Periplaneta cavernicola). (3) Guanobies: Cockroaches that live in the guano of cavernicolous vertebrates (Gyna, Acanthogyna, Dyscologamia, Pycnoscelus). (4) Troglobies: Cockroaches that apparently cannot live outside of caves and which show very marked adaptive characters (Alluaudellina, Nocticola, Spelaeoblatta, Typhloblatta). For complete discussions of these groups including descriptions of the adaptive characters shown by certain genera, the original sources should be consulted.

Although we know very little of the ethology of most of the cavernicolous cockroaches, it is intriguing that three of the six known species of Nocticola are cave dwellers, two are inhabitants of termite nests (p. [315]), and one (N. bolivari) was found under stones and cement blocks (Chopard, 1950b). In the rather extensive list of cavernicolous cockroaches only two (Arenivaga grata and Parcoblatta sp.) were taken from caves in North America north of Yucatan. All other records are from Africa, Asia, Central America, Europe, West Indies, East Indies, and the Philippine Islands. This we find puzzling. Packard (1888) in his extensive study of the cave fauna of North America listed no cockroaches. Dearolf (1941) found only the above-mentioned Parcoblatta in one of 37 caves in Pennsylvania. Kohls and Jellison (1948) listed no cockroaches among the arthropods from six bat caves in Texas. We would expect Periplaneta americana to inhabit mines in North America, but we have found no such records. Have cockroaches been ignored in fauna collections from North American caves, or has our cave fauna been less extensively studied than that of other parts of the world?

The two species of cockroaches found in mines (Blattella germanica and Periplaneta americana) are also found in caves. For this reason we have included them in the list headed Cavernicolous Cockroaches. On the other hand, the cockroaches found in animal burrows are generally different species from those found in caves, so we have grouped these together in a second list.

CAVERNICOLOUS COCKROACHES

Alluaudellina cavernicola

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

East Africa.—Chopard (1936).

Apotrogia angolensis

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

Apteroblatta perplexa

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

Arenivaga grata

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

Aspiduchus borinquen

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

Aspiduchus cavernicola

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

Blaberus atropos

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

Blaberus craniifer

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

Blaberus giganteus

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

Blatta lateralis

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

Blatta orientalis

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

Blattella germanica

South Africa.—Numerous in a gold mine on the Witwatersrand (Porter, 1930).

Tonkin.—Chopard (1929a); Colani (1952).

Byrsotria fumigata

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

Deropeltis erythropeza

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

Ectobius pallidus

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

Ectobius vittiventer

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

Ectobius sp.

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

Ergaula scarabaeoides

Sumatra.—West coast (Hebard, 1929).

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

Euthyrrhapha nigra

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

Gyna kazungulana

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

Gyna maculipennis

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

Gyna tristis

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

Heterogamodes krügeri

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

Holocompsa zapoteca

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

Hoplosphoropyga babaulti

Stated to be a troglophile by Chopard (1938).

Nocticola caeca

Philippine Islands.—Bolívar (1892).

Nocticola decaryi

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

Nocticola simoni

Philippine Islands.—Bolívar (1892).

Parcoblatta sp.

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

Periplaneta americana

IN CAVES

East Africa.—Its presence in the cave at Shimoni was thought to indicate that man had sought refuge there and brought the cockroaches in with baggage or provisions (Chopard, 1936).

India.—Many present in cave at Vengurla, the floor of which was covered with bird guano (Abdulali, 1942).

Madagascar.—Thought to have been introduced into the cave entrance by man (Chopard, 1945, 1949a).

IN MINES

Great Britain.—In a coal mine at Pontewydd where they had been established for some years (Lucas, 1916). In the Pentre Pit mine where they were abundant (Lucas, 1918). Abundant in a Welch mine 2,166 feet below the surface (Lucas, 1925). This species was found quite commonly in a number of South Wales coal mines; in one deep mine a white-eyed mutant form comprised about 5 percent of the cockroach population for the preceding 11 years (Jefferson, 1958).

India, western Bengal.—Very numerous in coal mines where the sole food apparently was human faeces (Chandler, 1926).

South Africa.—Numerous in four deep-level gold mines on the Witwatersrand.

Sumatra.—Numerous males and females from Sawah Lunto "'from a coal mine where they lived in great numbers on the faeces of miners'" (Hanitsch, 1929).

Periplaneta australasiae

Sarawak.—Found swarming on walls of caves and in soft bird guano in company with Symploce cavernicola (Moulton, 1912).

Tonkin.—Chopard (1929a); Colani (1952).

Periplaneta cavernicola

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

Periplaneta lata

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

Periplaneta sp.

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

Perisphaerus sp.

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

Polyphaga aegyptiaca

Turkmen S.S.R.—Females found in front part of Bakharden bat cave on several occasions (Vlasov, 1929).

Turkey.—At Magharadjik and Arab Dede, found in caves with various other animals (Lindberg, 1954).

Polyphaga sp.

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

Pycnoscelus niger

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

Pycnoscelus striatus

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

Pycnoscelus surinamensis

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

South Celebes.—Hanitsch (1932).

Spelaeoblatta gestroi

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

Symploce breviramis

South Celebes.—Hanitsch (1932).

Symploce cavernicola

Sarawak, Borneo.—Swarming on walls of caves and in soft bird guano on the cave floor (Moulton, 1912). Hanitsch (1931) noted that this species was first recorded by Shelford from a cave in Sarawak and that there is a series from a cave in the Oxford University Museum, taken by Banks in 1928.

Malaya.—On the walls of the inner cavern of a cave at Biserat; the insects covered the walls in places (Chopard, 1919).

Sumatra.—From Baso cavern, on the west coast (Hebard, 1929).

Symploce curta

South Celebes.—Hanitsch (1932).

Symploce remyi

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

Tivia macracantha

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

Tivia sp.

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

Typhloblatta caeca

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

Typhloblattodes madecassus

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

Xestoblatta immaculata

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

Unidentified cockroaches

Malaya.—The walls of a cave were covered by dense groups of a species of "Blatta" (Annandale, 1900).

England.—"The chief insect pests of the mines are cockroaches, which often swarm in hot mines and those with pit pony stables...." (Hardy, 1941).

COCKROACHES FROM THE BURROWS OF VERTEBRATES

Arenivaga apacha

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

Arenivaga bolliana

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

Arenivaga erratica

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

Arenivaga floridensis

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

Arenivaga roseni

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

Arenivaga tonkawa

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

Cariblatta lutea

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

Euthlastoblatta abortiva

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

Parcoblatta fulvescens

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

Polyphaga aegyptiaca

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

Polyphaga indica

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

Polyphaga saussurei

Tadzhikistan.—Found in burrows of turtles and rodents (Zmeev, 1936).

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

Pycnoscelus surinamensis

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

DESERT HABITATS

There is relatively little ecological information about cockroaches that live in deserts, even though certain species, notably Polyphaga aegyptiaca, have long been known to inhabit arid zones. In fact, so little is known about the ecology of arid-zone insects in general that it is more a subject for research than for review (Pradhan, 1957). In their account of the cockroaches of Northern Kenya and Jubaland, Kevan and Chopard (1954) describe in some detail the vegetational areas of this arid desert or semidesert country, which averages only about 10 inches of rain per year. The other sources that are cited below contain very little more biological information than the abstracted material that is given under each species.

Nearly all the Polyphaginae are said to be marked xerophiles whose distribution coincides with that of the deserts (Bei-Bienko, 1950). With the exception of Arenivaga floridana, the species of Polyphaginae in the United States all occur in the Southwest, where they are (with a few exceptions) the only cockroaches that inhabit the desert regions proper (Hebard, 1917). The Polyphaginae reach their greatest diversity in the deserts of Northern Africa and Anterior and South-Central Asia (Bei-Bienko, 1950). Some of the desert-inhabiting species have also been found under nondesert conditions. This only further exemplifies the plasticity of cockroaches in adapting to different environments.

The ability of desert insects to live under what appear to be extremely unfavorable conditions has been abundantly illustrated by Pradhan (1957). Uvarov (1954) has pointed out that a desert "covers a great variety of landscapes, which provide desert animals with a wide range of habitats, some of them offering very favorable conditions for life." Pradhan (1957) stated that many desert animals avoid the extremes of desert climates by choosing suitable microclimates for diurnal resting places, that a permanent or temporary underground existence is very common among insects in arid zones, and that many nocturnal Orthoptera burrow into the soil or hide under stones where temperatures are lower. For example, the type of Parcoblatta desertae was found under a boulder on the bare desert (Rehn and Hebard, 1909).

Symbiosis with burrowing animals is another solution to the problem of existence in the desert; in fact, symbiosis is a mode of life adopted by nearly half of the desert cockroaches about which we have any information. Vlasov and Miram (1937) found Polyphaga indica, Polyphaga saussurei, and Arenivaga roseni in the burrows of rodents and desert turtles. In the desert regions of Arizona, females of Arenivaga erratica were found commonly in burrows of the kangaroo rat (Vorhies and Taylor, 1922) and in dens of wood rats and ground squirrels (Ball et al., 1942). Arenivaga apacha and Arenivaga bolliana have also been found inhabiting the nests of wood rats (Hebard, 1917; 1943a). Bei-Bienko (1950) has suggested that the adaptation of desert-inhabiting cockroaches to rodent burrows might enable these insects to survive in the severe climatic conditions of deserts in summer.

Under desert conditions in southern Arizona, the relative humidity outside of the burrows of the kangaroo rat is 1 to 15 percent during the day and 15 to 40 percent at night; but inside the burrows the relative humidity is 30 to 50 percent, and the temperature, even during the day, is below 30° C. (Schmidt-Nielsen, 1949). Thus by living in rodent burrows during the day and going outside at night, the desert cockroaches could avail themselves of the most favorable microclimates obtainable. Presumably whatever food these insects eat provides them with sufficient water to enable them to survive under desert conditions. Bodenheimer (1953) has suggested that the extent of utilization of dew, which is sometimes heavy in the desert, should be investigated; he stated that tenebrionid beetles have been seen in the early morning eating dry [dead?] herbs that were still wet with dew. It is obvious that there is a need for additional detailed information without which we can only guess about the ecology of desert cockroaches.

In the following list we have cited only those species that were stated to have been found under desert conditions. Undoubtedly, related species that have been taken in similar localities are also desert-inhabiting forms, as, for example, other species of Arenivaga that were collected in Texas by Hebard (1943a). In the absence of specific information linking such other species with deserts, we have arbitrarily relegated those forms to the section on outdoor habitats. In addition to the species listed below, desert cockroaches are said to be found in the following genera: Anisogamia, Mononychoblatta, and Nymphytria (Chopard, 1938).

DESERT COCKROACHES

Agis orientalis

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

Arenivaga apacha

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

Arenivaga bolliana

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

Arenivaga erratica

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

The Biotic Associations of Cockroaches

THE BIOTIC ASSOCIATIONS OFCOCKROACHES

THE BIOTIC ASSOCIATIONS OF COCKROACHES[1]

HISTORICAL

METHODS

FUTURE WORK

ILLUSTRATIONS

CAVE HABITATS

CAVERNICOLOUS COCKROACHES

Alluaudellina cavernicola

Apotrogia angolensis

Apteroblatta perplexa

Arenivaga grata

Aspiduchus borinquen

Aspiduchus cavernicola

Blaberus atropos

Blaberus craniifer

Blaberus giganteus

Blatta lateralis

Blatta orientalis

Blattella germanica

Byrsotria fumigata

Deropeltis erythropeza

Ectobius pallidus

Ectobius vittiventer

Ectobius sp.

Ergaula scarabaeoides

Euthyrrhapha nigra

Gyna kazungulana

Gyna maculipennis

Gyna tristis

Heterogamodes krügeri

Holocompsa zapoteca

Hoplosphoropyga babaulti

Nocticola caeca

Nocticola decaryi

Nocticola simoni

Parcoblatta sp.

Periplaneta americana

IN CAVES

IN MINES

Periplaneta australasiae

Periplaneta cavernicola

Periplaneta lata

Periplaneta sp.

Perisphaerus sp.

Polyphaga aegyptiaca

Polyphaga sp.

Pycnoscelus niger

Pycnoscelus striatus

Pycnoscelus surinamensis

Spelaeoblatta gestroi

Symploce breviramis

Symploce cavernicola

Symploce curta

Symploce remyi

Tivia macracantha

Tivia sp.

Typhloblatta caeca

Typhloblattodes madecassus

Xestoblatta immaculata

Unidentified cockroaches

COCKROACHES FROM THE BURROWS OF VERTEBRATES

Arenivaga apacha

Arenivaga bolliana

Arenivaga erratica

Arenivaga floridensis

Arenivaga roseni

Arenivaga tonkawa

Cariblatta lutea

Euthlastoblatta abortiva

Parcoblatta fulvescens

Polyphaga aegyptiaca

Polyphaga indica

Polyphaga saussurei

Pycnoscelus surinamensis

DESERT HABITATS

DESERT COCKROACHES

Agis orientalis

Arenivaga apacha

THE BIOTIC ASSOCIATIONS OF
COCKROACHES

THE BIOTIC ASSOCIATIONS OF COCKROACHES^[1]