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).

Arenivaga roseni

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

Blattella vaga

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

Compsodes schwarzi

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

Cyrtotria capucina

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

Derocalymma lampyrina

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

Derocalymma porcellio

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

Deropeltis autraniana

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

Deropeltis melanophila

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

Deropeltis nigrita

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

Eremoblatta subdiaphana

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

Euandroblatta palpalis

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

Heterogamodes rugosa

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

Namablatta bitaeniata

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

Nauphoeta punctipennis

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

Parcoblatta desertae

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

Polyphaga aegyptiaca

Caucasus.—The wingless female was found buried in sand and dust (Burr, 1913).

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

Polyphaga indica

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

Polyphaga saussurei

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

Supella hottentotta

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

Symploce kevani

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

Theganopteryx straminea

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

Tivia brunnea

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

Tivia fulva

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

Tivia obscura

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

AQUATIC HABITATS

The so-called aquatic or amphibious cockroaches are all members of the subfamily Epilamprinae (Chopard, 1938). These forms are not nearly as aquatic as water beetles or aquatic Hemiptera, but in their relations to water they behave differently from nonamphibious cockroaches, which tend to avoid water except for drinking. There are apparently no special morphological characteristics that distinguish amphibious cockroaches (Shelford, 1907, 1909a; Chopard, 1938), although Takahashi (1926) listed several characters that he considered made Opisthoplatia maculata adapted for an aquatic life: (1) Back of body easily wetted; (2) long hairs on underside of thorax trap air; (3) terminal abdominal spiracles open into tubes that extend rearward; (4) long hairs on ventral surfaces of cerci "protect" terminal abdominal spiracles. Annandale (1906) also suggested that the position of the posterior abdominal spiracles, at the base of tubes that project rearward from beneath the seventh tergite, are an adaptation to an aquatic life. However, as Shelford (1907) and Chopard (1938) have pointed out, this same feature may be observed in many terrestrial cockroaches. The legs of amphibious cockroaches are similar to those of nonaquatic species and are not modified for swimming (Shelford, 1909a; Takahashi, 1926).

Biological observations have been made on relatively few species, but representatives of at least six genera occur in quasi-aquatic habitats. Strictly speaking, these cockroaches live on land at the edges of streams or pools and spend relatively brief periods in the water. A few species are found in water-filled bromeliads. The behavior of the known amphibious species of cockroaches in relation to their habitats is discussed below.

AMPHIBIOUS COCKROACHES

Audreia bromeliadarum

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

Dryadoblatta scotti

Trinidad.—This species was taken from the leaf bases of Tillandsia sp. at 3,100 feet; water had collected between the leaves and the insect was presumed to be more or less amphibious (Scott, 1912). Subaquatic in the bromeliad Glomeropitcairnia erectiflora: "This large and handsome species [D. scotti] is very common in the larger, water-filled, epiphytic bromeliads of the rain forest. Within these plants it is usually to be found, often in considerable numbers, just above the surface of the water or partly immersed in it. The cockroaches will descend rapidly into the water when alarmed and probably obtain their nourishment from the accumulated organic matter in the water. Floating material is probably taken and it seems less likely that they feed below the surface. They appear to be ovoviviparous." (Princis and Kevan, 1955.)

Epilampra abdomen-nigrum

Puerto Rico.—Abundant in wet "malojillo" meadows. The nymphs swim easily and remain under water for long periods, as do the adults (Seín, 1923; Wolcott, 1950).

Panama.—A swimming nymph, captured in a dipper with mosquito larvae in a lagoon of the Rio Chilibre, was kept under observations in an aquarium. If disturbed, the insect dived into the water from floating vegetation and swam rapidly below the surface for a minute or two. Finally becoming quiescent, the cockroach would then cling to submerged roots; twice it remained still for 15 minutes before climbing to the surface, where it remained for five or more minutes before emerging completely (Crowell, 1946).

Epilampra annandalei

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

Epilampra sp.

Siamese Malay States.—Wingless females rested on floating logs from which they would dive into the water upon the least disturbance; they remained under water for several minutes, then surfaced beneath the shelter of the log. In the jungle all females were taken either in the water or among matted roots on the sides of the stream. Winged males were seen rising from the surface of the water (Annandale, 1900).

Sarawak.—All specimens were immature; they swam and dived well, but were soon drowned if prevented from rising to the surface to breathe. "When at rest the body of the cockroach is almost entirely submerged, the tip of the abdomen alone projecting above the surface of the water; the abdomen moves gently up and down and every 30-40 seconds a bubble of air issues from the prothoracic spiracle on each side." (Shelford, 1901, 1916.)

India.—A nymphal female, found in a jungle stream at Chota Nagpur, could swim with belly or back upward. When held under water it drowned in a few minutes. The tip of the abdomen was held out of water (Annandale, 1906).

Shelford (1907) has suggested that the immature stages of terrestrial species of Epilampra may well be amphibious. This is an area that could profit by more field observations.

Opisthoplatia maculata

Formosa.—Invariably found under or between rocks near mountain streams. The wingless adult and the nymph have similar habits. Normally the cockroach lives on land, and when it goes into the water it returns to land within a few minutes. This cockroach rarely swims, but when it does, it maintains its body in a horizontal position just below the surface of the water. Ordinarily, it walks on the river bottom or on water-covered rocks. This insect feeds on decayed leaves and, according to Shikano, it will eat human feces. (Takahashi, 1926.)

This species has a large number of long hydrophobic hairs on the ventral sides of the thorax and anterior abdominal segments. When the insect submerges, air is trapped in these hairs. The thoracic and one pair of abdominal spiracles open into the bubble of trapped air. However, the insect apparently does not use this plastron of air to replenish its tracheal air supply, but, like Rhicnoda natatrix (see below), it inspires air while at the surface through its posterior abdominal spiracles and expires air into the bubble under the thorax. While the insect is submerged, the air bubble increases in volume until part of it breaks away and floats to the surface. (Takahashi, 1926.)

Opisthoplatia orientalis

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

Rhicnoda natatrix

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

Stictolampra buqueti concinula

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

Unidentified epilamprines

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

OUTDOOR HABITATS

This category is a catchall for all cockroaches that are not limited to the more circumscribed habitats that have been previously considered. Some cockroaches in this section select specific microhabitats (e.g., Cryptocercus spp., which live exclusively in rotten logs; and Neoblattella dryas, N. eurydice, and N. grossbecki in bromeliads). Others are found in a wide variety of habitats (e.g., Ischnoptera deropeltiformis and Parcoblatta spp.). But some species are so little known that their actual habitats are barely suggested in the collection data.

Williams (1941) made an ecological study of the floor fauna of the Panama rain forest. He found Orthoptera (nearly all were unidentified nymphal cockroaches) in the litter of dead leaves, twigs, and other plant products in over 90 percent of the quadrats he examined. These insects represented about 0.25 percent of the total animal population.

Delamare Deboutteville (1948) made a quantitative study of the animal population in suspended soil that had accumulated between the roots of forest epiphytes of the lower Ivory Coast. He analyzed 2 dm.³ samples of soil from an epiphyte located 45 meters above ground on a main branch of Parinarium, with these results: Horizon A.—Superficial zone of large rootlets, 6 cm. deep: 2 cockroaches, 4 arachnids, and 4 beetles. Horizon B.—Zone of fine rootlets, 6 cm. deep: 6 cockroaches and numerous other arthropods. Horizon C.—Humid zone, 8 cm. deep: 7 cockroaches and numerous other arthropods. Plants, such as Palissota, were also living in this very original biotype.

The species of cockroaches listed below have been found in the following kinds of outdoor microhabitats: In jungle, forest, and woodlands they have been found in rotten wood; under bark of living, dead, and fallen trees; in decay cavities in trees; burrowing in living bark; on foliage of trees, shrubs, bushes, and low herbage; on vines and in bromeliads and epiphytic ferns; under signs on trees and stumps; in piles of logs and firewood; under dead leaves and debris; in and under decaying fruit on the ground. Cockroaches have been found between the leaves and under leaf sheaths of sugarcane, corn, and other grasses; under dry fibers and fronds of coconut trees; in hollow stems and bases of tree-fern fronds; under bracts of banana blossoms and in bunches of bananas (p. [146]). Cockroaches also inhabit abandoned cocoons and larval tents, wasp nests, ant nests, termite nests, bird nests, rat nests, and burrows of other rodents (pp. 23-25, 310-319). Cockroaches have been found in rock crevices and under rocks; under boards and other objects on ground; under seaweed, drift, and other debris on beaches; burrowing in soil and under clods of earth; in marshes and swamps; in dumps and rubbish heaps.

The above list does not exhaust the available outdoor microhabitats that cockroaches find suitable for their continued existence, but it is fairly representative. Although we have no measurements to substantiate this conclusion, we suggest that the microhabitats cited above have a more constant temperature and a relatively higher humidity than is provided by the surrounding macrohabitats. We would expect insects such as cockroaches, whose water balance is dependent on a continuous supply of fluid water or moist food, to seek moist environments or to avoid situations in which their transpiration might increase. Deviations, presumably brief, from this expected behavior must occur to account for the cockroaches that are found under relatively unfavorable environmental conditions. Despite the apparent preference for cryptic habitats, some cockroaches are found in hot sunlight (Ellipsidion spp.; Tepper, 1893); Rehn (1945) has stated that many kinds are diurnal rather than nocturnal. Movement of cockroaches between habitats may be assumed to occur; but movement from an unfavorable environment to a more favorable one, following a shift in water balance, has not been observed in nature; however, laboratory experiments suggest that the mechanism for mediating such behavior is present in some species of cockroaches (Gunn and Cosway, 1938; Roth and Willis, 1952a). Obviously, additional research is needed on the bionomics of all species. Further conclusions based on current limited knowledge can only be speculative and possibly misleading.

COCKROACHES FROM OUTDOOR HABITATS

(Except Amphibious, Desert, and Cavernicolous Forms)

Aglaopteryx absimilis

Puerto Rico.—Living in rotten, wooden fence; living between leaves of Samanea saman and in abandoned cocoons of Megalopyge krugii on bucare trees (Wolcott, 1950).

Leeward Islands.—On coconut tree (Princis and Kevan, 1955).

Aglaopteryx facies

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

Aglaopteryx diaphana

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

Aglaopteryx gemma

Florida.—On Long Key, under coquina boulder in heavy scrub; under loose, dry fibers near head of standing coconut palm (Rehn and Hebard, 1912). Climbing on roots of red mangrove, Rhizophora mangle, in swamp; under loose bark on trunk of Exothea paniculata in dense jungle; under limestone boulder in keys scrub; under signs on oaks, sweet gum, and longleaf pines in southeastern and southern States (Hebard, 1917). Infrequent in the shrub growth of the Sandhills habitat (Friauf, 1953).

Texas.—In undergrowth of pine forest; under sign on oak near river; in Tillandsia sp. (Hebard, 1917). Usually in hiding places on trees; only once found under a stone on ground (Hebard, 1943a).

Allacta similis

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

Amazonina emarginata

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

Anaplecta asema

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

Anaplecta decipiens

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

Anaplecta fallax

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

Anaplecta hemiscotia

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

Anaplecta lateralis

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

Arenivaga bolliana

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

Arenivaga floridensis

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

Arenivaga grata

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

Aristiger histrio

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

Aspiduchus boriquen

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

Audreia bromeliadarum

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

Audreia jamaicana

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

Balta godeffroyi

Australia.—Under bark (Hebard, 1943).

Balta quadricaudata

Australia.—From sugarcane (Hebard, 1943).

Balta scripta

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

Balta torresiana

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

Balta verticalis

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

Blaberus atropus

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

Blaberus discoidalis

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

Blaberus giganteus

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

Blaberus spp.

Venezuela.—Only taken in the forests of the Orinoco near the trunks of rotten trees at night (Doumerc in Blanchard, 1837).

Panama.—Among dead leaves and debris on floor of rain forest (E. C. Williams, Jr., 1941).

Blatta lateralis

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

Blatta orientalis

Great Britain.—One female nymph under bark of tree 10 feet above the ground (Burr, 1900). Swarming within a rubbish heap in February (Lucas, 1912). In refuse tip under old sacks and sheets of linoleum (Hallett in Lucas, 1922). Male under bark of oak far from houses (Donisthorpe, 1918). One adult female and nymph in prone dead elm 50 yards from house (Burr, 1937). An immature male at the roots of Ballota nigra (Buck in Gardner, 1954). Four additional records of this species outdoors away from houses (Lucas, 1920).

Southern Crimea.—Under stones, dead leaves, and detritus in small copses of Quercus pubescens, Carpinus orientalis, Cornus mas, Paliurus aculeatus, and Dictamnus fraxinella; 19 specimens, apparently breeding outdoors (Adelung, 1907).

North-central U.S.—Observations since 1950 indicate a marked increase in frequency and duration of infestations outdoors; observed in bare soil, vegetation, debris, alongside foundations in sodded areas, along sidewalks, and at edge of parking areas throughout the year; in some urban residential areas, the yards of whole blocks of houses were "alive" with this species on warm summer nights; in winter they have been found under stones, leaf debris, and soil near structures (Shuyler, 1956).

Blattella germanica

Algeria.—Under moist leaves in woods (Lucas, 1849).

California.—Under rubbish and on date palms (Herms, 1926).

Connecticut.—In city dump under loose material, very numerous (Walden, 1922). Additional infestations of dumps by this species have been reported in New York (Felt, 1926, 1928) and New Jersey (Hansens, 1949, 1950)

England.—Swarming within a rubbish heap in February (Lucas, 1912).

Formosa.—Lives among fallen leaves on the ground (Takahashi, 1924).

North-central U.S.—Reported living outdoors near buildings and in soil under basementless buildings from early summer to late fall (Shuyler, 1956).

Blattella humbertiana

India.—Common among decaying vegetation and on trees (Chopard and Chatterjee, 1937).

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

Blattella vaga

Arizona.—Typically an inhabitant of irrigated fields and yards, it is found in fewer numbers on the dry desert. It is found under stones, plant debris, and clumps of earth; found in greatest numbers around decaying dates on ground (Flock, 1941a).

Texas.—Beneath duff under athel trees; rather abundant in clumps of Rhodes grass (Riherd, 1953).

Byrsotria cabrerae

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

Byrsotria fumigata

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

Cahita borero

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

Cahita nahua

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

Cariblatta antiguensis

Virgin Islands, St. Croix.—Common under heaps of rubbish (Beatty, 1944).

Trinidad.—On herbage below bananas; all stages on Hibiscus at night; in grass at dusk; on low herbage under old coconut (Princis and Kevan, 1955).

Cariblatta cuprea

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

Cariblatta delicatula

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

Cariblatta hylaea

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

Cariblatta imitans

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

Cariblatta insularis

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

Cariblatta jamaicensis

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

Cariblatta landalei

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

Cariblatta lutea lutea

North Carolina.—Under pine straw on ground in woods (Brimley, 1908).

Southeastern U.S.—Under dead oak leaves; under dead needles in longleaf-pine woods; in wire grass; under refuse; beaten from undergrowth in pine and oak woods (Rehn and Hebard, 1916). In undergrowth of shortleaf-pine, longleaf-pine, and oak woods; in heavy scrub in damp spot of sand dune area; from high bushes, Ilex coriacea [= lucida] along inland swampy area (Hebard, 1916a). "The species is in large part terrestrial, being usually found among dead leaves and litter on the ground. Occasional specimens are, however, sometimes beaten from bushes. Individuals are decidedly active and are usually to be found in the greatest numbers in sandy situations" (Hebard, 1917).

Florida.—Throughout winter and spring they are frequent beneath leaves and other debris on ground, especially in dry, sandy locations (Blatchley, 1920). Friauf (1953) found this species under debris, fallen leaves, leaf mold, or decaying wood in these habitats: Dry, ruderal grassland (infrequent), scrub (frequent), sandhills (dominant), xeric hammock (infrequent), mesic hammock (dominant), pond margin (infrequent), longleaf-pine flatwoods (frequent), bayhead (occasional), low hammock (frequent), and alluvial hammock (occasional). In the shrub stratum in these habitats: Scrub (frequent), sandhills (dominant), and xeric hammock (infrequent). In herbaceous stratum in these habitats: Sandhills (dominant), mesic hammock (dominant), and black-pine flatwoods (infrequent). On bare soil or bare sand under vegetation in these habitats: Sandhills (dominant), pond margin (infrequent), longleaf-pine flatwoods (frequent), and slash-pine flatwoods (frequent) (Friauf, 1953).

Cariblatta lutea minima

Florida.—Series of specimens captured on Long Key under dead petioles of coconut palm on moist ground at edges of pools of brackish water. Specimens from Key West were in dry dead grass under boards (Rehn and Hebard, 1912). Nymphs frequent under bark on decaying pine logs in pine woods; occasional in leaf mold in heavy junglelike scrub (Rehn and Hebard, 1914). In water-soaked leaves in heavy red-mangrove swamp (Hebard, 1915). Under dead petioles of coconut palm on sandy soil in grapefruit grove (Hebard, 1916a). Numerous at bases of tufts of coarse grass growing just back of sea beach (Blatchley, 1920). Friauf (1953) found this species in leaf duff, leaf mold, debris, or decaying wood in these habitats: Dry, ruderal grassland (occasional), scrub (infrequent), sandhills (infrequent), mesic hammock (infrequent), pond margin (occasional), longleaf-pine flatwoods (occasional), and low hammock (infrequent). On bare soil or bare sand under vegetation in these habitats: Longleaf-pine flatwoods (occasional) and slash-pine flatwoods (occasional). Dominant in the spartina marsh habitat in the grass stratum and duff around clumps. Frequent in the saw-grass marsh habitat in the grass stratum and, during the dry season, in decaying vegetation on the marsh floor.

Cariblatta nebulicola

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

Cariblatta reticulosa

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

Cariblatta stenophrys

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

Cariblatta spp.

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

Cariblattoides instigator

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

Cariblattoides suave

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

Ceratinoptera picta

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

Chorisoneura flavipennis

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

Chorisoneura formosella

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

Chorisoneura parishi

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

Chorisoneura specilliger

Panama.—In grass (Hebard, 1920).

Chorisoneura texensis

Florida.—"The almost impenetrable jungle on Key Largo was examined, and in its depths the two specimens of this species were secured by beating the lower branches of gumbo limbo, other trees and the lower bushes and shrubs, among which latter are to be found such tropical forms as Ocotea catesbyana [= Nectandra coriacea] and Citharexylum villosum" (Rehn and Hebard, 1912). In nests of webworm and beaten from bushes of bayberry, Myrica cerifera, along edge of pine woods (Rehn and Hebard, 1916). Beneath dead leaves in oak woods and beaten from foliage of oak and bayberry (Blatchley, 1920). Infrequent in the tall shrub stratum of the xeric hammock habitat (Friauf, 1953).

Texas.—The great majority of specimens were beaten from foliage of bushes (Hebard, 1943a).

Southeastern and southern U.S.—In undergrowth in pine woods; beaten from shrubbery, from bayberry bushes, from lower branches of gumbo limbo and other trees, from lower bushes and shrubs in jungle, and from low oaks on hills. In Texas, beaten from tall weeds in opening in river-plain jungle scrub (Hebard, 1917).

Chorisoneura translucida

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

Chromatonotus infuscatus

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

Chromatonotus notatus

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

Comptolampra liturata

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

Cryptocercus punctulatus

North Carolina.—"They were never found except in parts of the logs [chestnut] where the decayed wood was soft, punky and wet" (Rehn and Hebard, 1910).

Oregon.—In fir logs where sap wood was soggy (Hebard, 1917).

Virginia.—In decaying chestnut and pine logs; taken six times in chestnut and once in pine (Hebard, 1917). In rotten logs in deep ravines of moist woods (Davis, 1926).

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

Cryptocercus relictus

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

Cutilia soror

Marquesas Islands.—Males under stones and dead log (Hebard, 1933a).

Hawaii.—In soil about roots of pineapple (Illingworth, 1927). Often found about roots of grasses and weeds and other debris (Williams et al., 1931). Under stones and pineapple mulching paper (Fullaway and Krauss, 1945).

Wake Island.—Numerous, some from rotten logs. Found in bunch grass on Ocean Island (Bryan, 1926).

Cutilia spp.

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

Dendroblatta sobrina

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

Diploptera punctata

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

Raiatea, Society Islands.—Beaten out of bracken (Cheesman, 1927).

Uahuka, Marquesas Islands.—Under bark (Hebard, 1933a).

Dryadoblatta scotti

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

Ectobius africanus

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

Ectobius albicinctus

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

Ectobius duskei

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

Ectobius lapponicus

Southeastern Europe.—Numerous under stones on Trebovic (Burr, 1898).

U.S.S.R.—Found in wooded communities and peat bogs (in northern part of its range); males occur predominantly on herbaceous plants and bushes, but females hide under fallen leaves, moss, etc. (Bei-Bienko, 1950). It populated about 25 percent of the aspen trees in an experimental plot, feeding in galleries in the bark of young branches; there were 25 or more individuals per tree (Stark in Bei-Bienko, 1950).

Germany.—Abundant in woods; in pine woods in company with Stenobothrus vagans and Tettix kraussi. Numerous in low aspen bushes in forest. Numerous in deciduous and coniferous forests on trees and underbrush; under fallen leaves and moss; on oaks (Zacher, 1917). In foliage of young oak on top of mountain (Ramme, 1923).

Great Britain.—Under moss and dry leaves, among woodland undergrowth, and, generally, on vegetation close to the ground; occasional on bushes and trees (Lucas, 1920). Nymphs in heather in February and later; adults among rushes fringing pond in July (Lucas, 1925). Nymphs and males on rushy vegetation; unusually abundant on low herbage in dried-up swamp (Lucas, 1930).

Ectobius nicaeensis

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

Ectobius pallidus

Algeria.—Under stones; in moist places that are shaded and covered with plants (Lucas, 1849).

England.—Very abundant on sand dunes and among bracken in July (Buxton, 1914).

Germany.—In deciduous and coniferous forests; at edge of forest, from bare woods and bushes; numerous under leaves in oak woods and under moss (Zacher, 1917). In forest well lighted by the sun (Ramme, 1923).

Massachusetts.—Under loose lichens and bark on oak trees; under boxes, baskets, paper, etc., near houses; on Swiss chard (Flint, 1951). On roofs of houses, in shrubbery (Gurney, 1953). We have collected this species for several summers in a fairly dense, wooded area near dwellings, among fallen leaves and climbing on the erect stems and undersides of the leaves of periwinkle. Oöthecae were found on the ground under leaves and debris.

Ectobius panzeri

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

Germany.—In beech woods and in pine woods (Zacher, 1917).

Ectobius semenovi

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

Ectobius sylvester

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

Ectobius tadzhicus

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

Ectobius vittiventer

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

Ellipsidion affine

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

Ellipsidion australe

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

Ellipsidion bicolor

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

Ellipsidion simulans

Australia.—From sugarcane (Hebard, 1943).

Ellipsidion spp.

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

Epilampra abdomen-nigrum

Trinidad.—In dried-up drain; among grass; in debris under old cacao tree; under old leaves (Princis and Kevan, 1955).

Puerto Rico.—Abundant in damp lowlands (Seín, 1923). Under dead leaves in wet malojillo meadow (Wolcott, 1936).

This species is amphibious (p. [31]). Shelford (1907) suggested that immature stages of other species of the genus may be aquatic, which would place them in moist situations on the shores of rivers and other bodies of water.

Epilampra azteca

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

Epilampra mona

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

Epilampra tainana

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

Epilampra wheeleri

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

Epilampra spp.

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

Ergaula capensis

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

Eudromiella bicolorata

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

Euphyllodromia liturifera

Colombia.—In brushwood (Princis, 1946).

Eurycotis biolleyi

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

Eurycotis decipiens

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

Eurycotis dimidiata

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

Eurycotis ferrum-equinum

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

Eurycotis floridana

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

Eurycotis galeoides

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

Eurycotis kevani

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

Eurycotis opaca

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

Euthlastoblatta abortiva

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

Graptoblatta notulata

Tahiti.—On foliage in sun or concealed among dead leaves that collect between the fronds of tree ferns (Cheesman, 1927).

Hawaii.—Quite active during the day, occurring on sugarcane, etc., in the wetter districts; it is also a household insect (Williams et al., 1931).

Hemiblabera brunneri

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

Henicotyle antillarum

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

Holocompsa metallica

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

Hololampra bivittata

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

Hololampra chavesi

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

Hololampra maculata

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

Hololampra marginata

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

Hololampra sp.

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

Hololeptoblatta sp.

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

Homalopteryx laminata

St. Vincent.—In decaying leaves in forest (Rehn and Hebard, 1927).

Trinidad.—In forest debris and debris under old cacao trees; it is not uncommon under dry leaves; it feigns death when disturbed (Princis and Kevan, 1955).

Hormetica laevigata

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

Ignabolivaria bilobata

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

Ischnoptera deropeltiformis

North Carolina.—Under pine straw on ground in woods (Brimley, 1908).

Georgia.—Under dead oak leaves; under debris in garden; running on ground in pine and oak woods (Rehn and Hebard, 1916).

Indiana.—It is "a ground-frequenting, forest-loving insect, hiding beneath cover or about the edges of deep woodland, more frequently in damp places, and rarely taken beneath bark, signs, or at lights" (Blatchley, 1920).

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

Texas.—It preferred damp, open woodlands (Hebard, 1943a).

Eastern and southeastern U.S.—Under stone in heavy deciduous forest; under damp, dead leaves on edges of forests; under bark of pine log; in wire grass and sphagnum bordering stream thicket; in leaf mold and rubbish about pothole in pine woods, Pinus caribaea; under debris and leaf mold in hammock; under dead oak leaves in heavy deciduous forest (Hebard, 1917).

Florida.—"This species is distinctly geophilous and appears to prefer damp surroundings" (Rehn and Hebard, 1912). Under boards on very wet ground in everglades; in debris and leaf mold in heavy, junglelike areas of trees, bushes, and vines (Rehn and Hebard, 1914). Adults and numerous nymphs beneath weeds, grass, and other debris washed up on beach of Lake Okeechobee (Blatchley, 1920). Friauf (1953) found this species in leaf duff, leaf mold, and/or decaying wood on ground in these habitats: Dry, ruderal grassland (infrequent), scrub (infrequent), sandhills (occasional), xeric hammock (frequent), mesic hammock (dominant), shrubby, longleaf-pine flatwoods (infrequent), bayhead (dominant), and low hammock (dominant). On open bare soil or bare sand under vegetation in these habitats: Dry, ruderal grassland (infrequent), mesic hammock (dominant), moist, ruderal grassland (infrequent), pond margin (occasional), longleaf-pine flatwoods (infrequent), slash-pine flatwoods (infrequent), and low hammock (dominant). Infrequent in the herbaceous stratum of these habitats: Dry, ruderal grassland, moist, ruderal grassland, and longleaf-pine flatwoods. Infrequent in the shrub stratum of the dry, ruderal grassland habitat. (Friauf, 1953.)

Tennessee.—Taken in traps baited with cantaloupe in a parklike stand of oak, gum, hickory, and tulip trees in a creek bottom, and in a stand of oak on a dry ridge (Walker, 1957).

Ischnoptera panamae

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

Ischnoptera podoces

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

Ischnoptera rufa rufa

Virgin Islands, St. Croix.—Common under rubbish and on shrubbery at night (Beatty, 1944).

Barbados.—Occasionally found in cane fields (Tucker, 1952).

West Indies.—In Puerto Rico, under stones in cultivated area, under debris on alkalie flat. In Jamaica, under dry petioles of coconut palm in grassy area; under logs, logwood on docks, and litter on limestone and near beach. In Panama, under drift on edge of coral-sand beach; under rubbish at edge of jungle (Hebard, 1916c).

Jamaica.—Under limbs and leaf litter in mangrove swamp (Rehn and Hebard, 1927).

Lamproblatta albipalpus

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

Lamproblatta meridionalis

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

Latiblattella chichimeca

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

Latiblattella lucifrons

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

Latiblattella rehni

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

Latiblattella zapoteca

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

Leucophaea maderae

Barbados.—In cane fields (Tucker, 1952).

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

Litopeltis biolleyi

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

Litopeltis bispinosa

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

Litopeltis deianira

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

Litopeltis musarum

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

Lobolampra subaptera

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

Loboptera decipiens

France.—All stages common beneath stones (Blair, 1922). Under stones and dead leaves (Chopard, 1947).

Maltese Islands.—Quite common in open country under stones (Valletta, 1955).

Dalmatia.—On seashores under rocks and seaweed cast up on shore (Bei-Bienko, 1950).

Loboptera thaxteri

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

Lobopterella dimidiatipes

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

Lophoblatta arawaka

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

Macropanesthia rhinocerus

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

Megaloblatta blaberoides

Panama.—Under bark on tree (Hebard, 1920).

Ecuador.—Under a dense pile of dead leaves around base of tree (Campos R., 1926).

Megamareta verticalis

Australia.—In sugarcane (Hebard, 1943).

Methana canae

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

Methana curvigera

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

Methana marginalis

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

Moluchia (?) dahli

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

Muzoa madida

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

Nauclidas nigra

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

Nelipophygus ramsdeni

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

Neoblattella detersa

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

Neoblattella dryas

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

Neoblattella eurydice

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

Neoblattella grossbecki

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

Neoblattella proserpina

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

Neoblattella semota

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

Nesomylacris cubensis

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

Nesomylacris relica

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

Nocticola bolivari

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

Nyctibora laevigata

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

Nyctibora lutzi

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

Nyctibora obscura

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

Nyctibora stygia

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

Oniscosoma spp.

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

Opisthoplatia orientalis

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

Panchlora antillarum

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

Panchlora nivea

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

Texas.—Lives in foliage and in the green sheaths of plants (Hebard, 1943a).

Cuba.—On cane leaves; according to Gundlach this genus lives under the loose bark of trees (Rehn and Hebard, 1927).

Puerto Rico.—In rotting trunks of coconut palms (Seín, 1923). Most specimens have been collected from the very rotten interior of coconut palms (Wolcott, 1950).

Trinidad.—On corn; under old log; flies readily to lights (Princis and Kevan, 1955).

Panchlora sagax

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

Panesthia australis

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

Panesthia laevicollis

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

Parcoblatta bolliana

North Carolina.—Under pine straw on ground in pine woods (Brimley, 1908).

Texas.—Under dry cow dung in pine woods (Hebard, 1917).

Nebraska.—Under pile of old boards (Hauke, 1949).

Parcoblatta caudelli

North Carolina.—From under the bark of dead trees (Rehn and Hebard, 1910).

Virginia.—At night on shrubbery. In South Carolina, under sign on tree (Hebard, 1917).

Tennessee.—In traps baited with cornmeal, cantaloupe, or fish in a stand of oak on dry ridge, and in abandoned rocky field on a south-facing slope (Walker, 1957).

Parcoblatta desertae

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

Parcoblatta divisa

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

Parcoblatta fulvescens

Eastern and southeastern U.S.—Trapped in molasses jars: in heavy, barrier-beach forest; in typical pine-barrens undergrowth; in pine barrens with heavy, grassy undergrowth; on border of pine barrens and on edge of swamp; in heavy deciduous forest; in heavy oak woods. Found under debris in dead, shortleaf-pine needles; under dead leaves on edge of oak and shortleaf-pine woods; under bark of pine log; among dead leaves under live oaks; under sign on Pinus caribaea (Hebard, 1917).

Georgia.—From under bark of pine log, among dead leaves under live oaks, and under leaves on edge of oak and shortleaf-pine woods (Rehn and Hebard, 1916).

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

Parcoblatta lata

Southeastern and southern U.S.—Under bark of pine logs and stumps; in sweet-gum logs and stumps; moderately numerous under bark of dead shortleaf pines; under bark of longleaf-pine stumps; under signs on red oak and longleaf pines; in dead oak. In Texas, under bark of pine stumps (Hebard, 1917).

North Carolina.—All stages under loose bark of dead pines, both prostrate and upright, and stumps. "It seems to prefer the space under the bark to be rather damp" (Brimley, 1908). Under bark of dead pine trees (Rehn and Hebard, 1910).

Florida.—Infrequent in leaf duff and decayed wood of low hammock habitat (Friauf, 1953).

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

Missouri.—In leaf stratum of oak-hickory forest (Dowdy, 1951). Earlier, Dowdy (1947) reported finding numerous immature Pseudomopinae [presumably Parcoblatta sp.] in soil and leaf strata of oak-hickory forest.

Texas.—Captured in molasses-baited traps in low, wet, oak woods and in dry woodlot on hillside (Hebard, 1943a).

Parcoblatta pensylvanica

Eastern and southeastern U.S.—Trapped in molasses-baited jars; in oak and in chestnut forests, and on knoll with high deciduous trees. Found in oak and pine woods, under bark of decaying chestnut log and dead chestnut stump, and under signs on trees including oaks (Hebard, 1917).

North Carolina.—In all stages under loose bark of upright, dead pines, when the space under the bark was dry (Brimley, 1908).

Virginia, North and South Carolina.—Under signs on trees (white and red oaks); under bark of dead shortleaf-pine and sweet-gum logs and stumps (Rehn and Hebard, 1916).

Indiana.—Beneath bark of logs and stumps; empty oöthecae common beneath loose bark of logs, especially shellbark hickory (Blatchley, 1920). Under loose bark on logs in January (Blatchley, 1895).

Illinois.—In pine forest associes, in black oak forest on sand, in oak-hickory forest on clay, and in climax forest; it evidently moved into the pine associes nightly, great numbers of oöthecae were found under bark of pine logs, where, in October and November, hibernating nymphs were found (Strohecker, 1937). In nests of Vespula maculata (Balduf, 1936; McClure, 1936).

Missouri.—Usually in hollow trees, under loose bark, in woodpiles, and in cracks in rural buildings (Rau, 1940).

Michigan.—Common in oak-dune and beech-maple forests, under loose bark on dead trees and fallen logs, and under debris on forest floor (Hubbell, 1922). "A characteristic inhabitant of the low shrub-terrestrial and probably the terrestrial-hypogeic stratum." It occurred throughout the upland forests; groups were found established in and under logs 100 to 200 feet from the nearest forest (Cantrall, 1943).

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

Parcoblatta uhleriana

North Carolina.—Under pine straw on ground in woods (Brimley, 1908). Under bark of dead trees; 92 males attracted to lights (Rehn and Hebard, 1910).

Virginia.—Resting on woods foliage; at night on road (Rehn and Hebard, 1916).

Eastern and southeastern U.S.—Trapped in molasses-baited jars: in oak and pine woods, in heavy barrier-beach forest, in both scant and typical undergrowth on pine barrens, in heavy grassy undergrowth on pine barrens, on border of pine barrens, on edge of swamp, in heavy deciduous forest, in heavy oak woods, in upland oak and chestnut forest, in chestnut forest, in forested ravine, and on ridge with heavy oak, chestnut, and maple forest. Found under damp leaves on edge of forest, under bark of decayed chestnut log, inside decaying chestnut log with Cryptocercus punctulatus, under palmetto roots, under bark of pine stump, and in dry leaves under live oaks (Hebard, 1917).

Tennessee.—In traps baited with cornmeal or cantaloupe in maple-gum-oak forest in a mesic valley, and in a stand of oak on a dry ridge (Walker, 1957).

Indiana.—Beneath cover on slopes of high wooded hills. "This is essentially a forest-loving species; usually occurring beneath leaves and other debris on or along the borders of heavy hardwood timber." (Blatchley, 1920.)

Illinois.—In oak-hickory forest on clay and in climax forest (Strohecker, 1937).

Michigan.—In oak-dune woods (Hubbell, 1922). Restricted to woodlands, where it inhabited piles of moist dead leaves and rotten logs in oak-hickory forest (Cantrall, 1943).

U.S.A.—This species, P. uhleriana, and P. virginica were attracted at night to honeydew secreted by aphids on Pyrus sp. (Davis, 1918).

Parcoblatta virginica

New England.—Females under loose stones, boards, and other debris on ground; beneath loose bark (Morse, 1920).

North Carolina.—Under debris in dead shortleaf-pine needles (Rehn and Hebard, 1916).

Florida.—Infrequent in the shrub stratum of the scrub habitat. This was the only habitat of 25 studied in which this species was found (Friauf, 1953).

Eastern and southeastern U.S.—Trapped in molasses-baited jars: in pine and oak woods, in pine barrens, in pine woods with heavy grass undergrowth, in oak forest, in heavily forested ravine, on rocky slope with few deciduous trees, on knoll with high deciduous trees, in lofty chestnut forest, and in heavy low chestnut and oak forest on high ridge; under bark of decaying chestnut log and stump; under stones in chestnut forest; under bark of pine stumps (Hebard, 1917).

Indiana.—Frequents borders of open woods and fields; under debris, loose bark, and half-buried logs (Blatchley, 1920).

Illinois.—In black-oak forest on sand, in oak-hickory forest on clay, and in climax forest (Strohecker, 1937).

Michigan.—Common in oak-dune and beech-maple forests; under loose bark on dead trees and fallen logs and under debris on forest floor (Hubbell, 1922). Restricted to woodlands, where it inhabited piles of moist dead leaves and rotten logs in oak-hickory forest (Cantrall, 1943).

Texas.—Captured in molasses traps in moist woods of maple, oak, and pine with much undergrowth and a heavy layer of duff; in open, rather dry woodlot of Spanish oak and other trees; and in low wet woods of willow and oak along creek (Hebard, 1943a).

Parcoblatta zebra

Indiana.—Beneath log in cypress swamp (Blatchley, 1920).

Louisiana and Mississippi.—In decay cavity in sweet gum; under sign on shortleaf pine (Hebard, 1917).

Parcoblatta spp.

Alabama.—In the dry wall of a sweet-gum stump together with serropalpid and tenebrionid beetles (Snow, 1958).

Ohio.—Oöthecae under loose bark of fallen trees, where as many as 184 oöthecae were found within a few feet of each other; others found under boards and in piles of firewood (Edmunds, 1952).

Pelmatosilpha coriacea

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

Pelmatosilpha kevani

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

Pelmatosilpha purpurascens

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

Periplaneta americana

Bermuda.—Among and under decaying debris, just above high-tide line (Verrill, 1902).

Johnson Island.—Nocturnal, coming out at night in great numbers about Tribulus blossoms. Under timbers on French Frigate Shoals (Bryan, 1926).

United States.—Alleyways and yards may be overrun during the summer; adults and hundreds of nymphs found in decaying maple trees along residential street (Gould and Deay, 1938, 1940). Around fumaroles where a railroad fill was burning internally (Davis, 1927). Common in palm trees along the gulf coast of Texas, where they often fly around street lights at night (Zimmern in Gould and Deay, 1940).

Periplaneta australasiae

Bermuda.—Very abundant under stones (Rehn, 1910).

Jamaica.—Under bark of dead tree and under bases of leaves of coconut palms (Rehn and Hebard, 1927).

Virgin Islands, St. Croix.—Common in sugarcane fields and in woodlands (Beatty, 1944).

Florida.—Juveniles under bark of dead logs of Pinus caribaea (Hebard, 1915). Frequently found under signs on trees near borders of towns; under bases of dead petioles of cabbage palmetto (Hebard, 1917). Beneath logs, burlap bags, and other cover in old orange orchards (Blatchley, 1920).

Marquesas Islands.—Under coconut fronds and grass (Hebard, 1935).

Nihoa Island.—Nymphs only, on Sida, Pritchardia, bunch grass, and about camp (Bryan, 1926).

Periplaneta brunnea

Georgia.—Under signs on oaks (Rehn and Hebard, 1916).

Florida.—Beneath bark of stump (Blatchley, 1920).

Periplaneta fuliginosa

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

Phidon (?) dubius

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

Phoraspis spp.

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

Phyllodromica brevipennis

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

Phyllodromica graeca

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

Phyllodromica irinae

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

Phyllodromica maculata

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

Phyllodromica meglerei

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

Phyllodromica polita

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

Phyllodromica pygmaea

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

Phyllodromica tartara

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

Phyllodromica tartara nigrescens

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

Platyzosteria castanea

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

Platyzosteria novae seelandiae

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

Plectoptera dominicae

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

Plectoptera dorsalis

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

Plectoptera floridana

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

Plectoptera infulata

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

Plectoptera lacerna

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

Plectoptera perscita

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

Plectoptera porcellana

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

Plectoptera pygmaea

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

Plectoptera rhabdota

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

Plectoptera vermiculata

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

Polyphaga aegyptiaca

Algeria.—Nymphal females under decaying leaves at the end of November (Lucas, 1849).

Transcaucasia.—In burrows in argillaceous cliffs along ravines. Females often covered by attached clay particles, an indication, according to Bei-Bienko, that this species is ecologically connected to compact clay soils or at least does not avoid them (Bei-Bienko, 1950).

See also the section on desert habitats (p. [29]).

Polyphaga saussurei

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

Polyzosteria limbata

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

Poroblatta spp.

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

Pseudomops septentrionalis

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

Pycnoscelus surinamensis

Florida.—Under stones and rubbish; very abundant under coquina boulders in woods at Key West (Rehn and Hebard, 1905). "This species is common under planks, stones, and other debris on the ground ... also found at Long Key in the dry fibres at the base of the petioles of a coconut palm" (Rehn and Hebard, 1912). At Musa Isle, found burrowing in sand (Hebard, 1915). In fallen leaves and decaying wood in xeric and mesic hammock habitats (Friauf, 1953).

Hawaii.—The soil swarmed with young of various stages during the summer (Illingworth, 1915). In soil about roots of pineapple under mulching paper; feeding on pineapple roots (Illingworth, 1927, 1929).

Fakarava, Tuamoto Archipelago.—Numerous among dead leaves in tree holes (Cheesman, 1927).

West Indies.—Under decayed stalks of sugarcane and in siftings from mangrove swamps, Cuba. Under manure, bases of leaves of coconut palm, litter, logs, and stones on coral rock and in bromeliads, Jamaica. Under wood, tiles, and boards in stable yards; immature individuals bored into the soil, Puerto Rico. (Rehn and Hebard, 1927.)

Barbados.—Frequents cane fields (Tucker, 1952).

Puerto Rico.—"Altho primarily a xerophytic species: collected among dry stones on Mona Island, under dry cow dung at Boquerón, and under boxes at Guánica, it is reasonably common in the more humid parts of Puerto Rico" (Wolcott, 1950).

Virgin Islands, St. Croix.—Common under rubbish; frequently seen feeding on chicken feces around chicken roosts (Beatty, 1944). By feeding on chicken feces it may become the vector of the chicken eyeworm, Oxyspirura mansoni, as described in the references cited on page 204.

Egypt.—Large numbers were found in moist soil at the site of a manure pile (Chakour, 1942).

Germany.—Under greenhouse conditions the depth to which P. surinamensis penetrated the soil was determined; 21 dug down to a depth of 8 to 10 cm., 3 dug down 10 to 12 cm., but only one dug 13 cm. below the surface. Often the tubes in the soil ended in a chamber which the cockroach might not leave for several days; nymphs molted in such chambers and females bore their young there (Roeser, 1940).

Rhytidometopum dissimile

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

Riatia orientis

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

Simblerastes jamaicanus

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

Styphon bakeri

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

Supella supellectilium

Virgin Islands, St. Croix.—Under rubbish heaps; in sugarcane straw (Beatty, 1944).

Africa.—"A cosmotropical species which occurs both out of doors and as a household pest in many warmer parts of the world. It is apparently endemic to non-forested areas in much of Africa north of the Equator." (Kevan and Chopard, 1954.)

Symploce flagellata

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

Symploce hospes

Hawaii.—Under stones and rubbish (Illingworth, 1915).

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

Symploce jamaicana

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

Symploce ruficollis

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

Puerto Rico.—In siftings from sea-grape thicket on sandy soil (Rehn and Hebard, 1927). Often living under leaf-sheaths of sugarcane (Wolcott, 1950).

Tartaroblatta karatavica

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

STRUCTURAL HABITATS

In this category we include all man-made structures, whether inhabited by man or not, that may become infested with cockroaches. A nonexhaustive list of such structures would include dwellings, restaurants, mess halls, barracks, groceries, markets, bakeries, dairies, drug stores, department stores, hotels, hospitals, warehouses, mills, factories, packing houses, animal houses, breweries, incinerators, privies, sewers, sewage treatment plants, ships, aircraft, etc. Although dwellings are only one of the many kinds of structures that are colonized by cockroaches, the several species that have adopted this mode of life are generally referred to as domiciliary cockroaches. This term is adequate only if we remember that these cockroaches are not restricted to domiciles but are pests in other structures as well.

Associations between man and certain species of cockroaches possibly started as casually as the short-lived association that Beebe (1953) observed when he discovered three cockroaches in the newly built couch of an orang-utan. Obviously, when man came down from the trees, his fellow travelers found his cave dwellings and other abodes particularly favorable habitats. From such primitive beginnings, domiciliary cockroaches have spread into every kind of structure that man has since devised. We predict that when man develops a suitable vehicle, cockroaches will someday accompany him into space. Yet despite the apparent predilection of certain species of cockroaches for man, man is only incidental to these associations. Only the shelter and food that man unwittingly provides for these unwelcome guests attract cockroaches to him; man's physical presence is unnecessary.

Most, if not all, of the common domiciliary cockroaches apparently originated in the Tropics or sub-Tropics from whence they have spread, through normal commercial channels, into most of the inhabited world. At least eight domiciliary cockroaches originated in Africa (Rehn, 1945): Blatta orientalis, Blattella germanica, Leucophaea maderae, Nauphoeta cinerea, Oxyhaloa buprestoides, Periplaneta americana, P. australasiae, and Supella supellectilium; and, perhaps, Periplaneta brunnea as well; Neostylopyga rhombifolia was probably of Indo-Malayan origin; Pycnoscelus surinamensis was of oriental origin; and Leurolestes pallidus was endemic in the West Indies (Rehn, 1945). Princis (1954a) rejected Africa as the original home of Blatta orientalis and advanced reasons for placing its origin in Central Asia.

Several domiciliary species have become well established in temperate zones and some even in the Arctic. Bei-Bienko (1950) listed the following 10 species as sinanthropes in the Palearctic zone: Blatta lateralis, B. orientalis, Blattella germanica, Leucophaea maderae, Periplaneta americana, P. australasiae, Polyphaga saussurei, Pycnoscelus surinamensis, and Supella supellectilium. In the warmer parts of the temperate regions, as in their native Tropics, certain domiciliary species breed outdoors as well as indoors. In the less temperate extensions of their ranges most domiciliary species are nearly always found indoors. In regions with low winter temperatures these cockroaches do not survive in unheated structures; but in heated buildings Blattella germanica, for example, has been able to withstand the rigorous climate of Alaska, where it has caused severe infestations (Chamberlin, 1949).

The limiting factors that determine whether man-made structures will provide suitable habitats for cockroaches are favorable temperature and availability of water and food. The range of temperatures that man provides for his own comfort and protection fosters the rapid increase of cockroach populations indoors. Gunn (1934, 1935) has demonstrated that the preferred temperature range (zone of indifference) of Blatta orientalis is 20-29° C. The upper limit of the preferred temperature of Blattella germanica and Periplaneta americana is 33° C. (Gunn, 1935). The lower limits of temperature tolerance were not sharply defined in Gunn's work. However, less than optimum temperatures, if they last for only short periods, are not necessarily lethal. The 24-hour mortality for P. americana that had been held for one hour at 0° C. was only 2±2 percent (Knipling and Sullivan, 1957). Gunn (1934) observed that Blatta orientalis would not settle at temperatures above 33° C. and would react violently against higher temperatures (e.g., 39° C.) by running away; thus the thermotactic behavior of cockroaches might be presumed to bring them into favorable environments within structures. Thermal death points have been determined for the above three species by Gunn and Notley (1936).

It is common knowledge among those who rear cockroaches experimentally that, unless the water content of the food is high, fluid water is essential in the insects' dietary. Ten species of domiciliary cockroaches have been shown to be unable to survive as long on dry food alone as they could on food and water at 36-40 percent relative humidity (Willis and Lewis, 1957). Blatta orientalis, when in a state of normal water balance, usually spent more time in the drier part of a humidity gradient; but desiccated insects tended to become hygropositive (Gunn and Cosway, 1938). We presume that other domiciliary species behave similarly. If water is available nearby, it may be presumed that partially desiccated cockroaches could locate a source through the mediation of a humidity sense. Hygroreceptors have been demonstrated on the antennae of Blattella germanica (Roth and Willis, 1952a) and suggested for Blatta orientalis (Gunn and Cosway, 1938).

Drinking water is available to cockroaches in the traps of sinks, wash basins, tubs, and toilet bowls; in flush tanks; as condensation on cold pipes, flush tanks, and windows; around leaking pipes and faucets; as spillage; in miscellaneous water-filled containers, such as pet drinking dishes, aquaria, vases; empty beverage bottles; and drainage from ice boxes. Soft, juicy fruits and vegetables can provide both moisture and food. There seems to be a tendency for certain species (Blatta orientalis and Blattella germanica) to become established in the more humid parts of structures, such as basements, around sinks, and in bathrooms. Whether this is a reaction to a preferred humidity or merely a fortuitous aggregation near sources of drinking water and food has never been clearly demonstrated. The rather widespread dissemination of these species into zones of low as well as high humidity suggests that detailed studies of the microclimatic conditions of structural microhabitats will be needed before meaningful conclusions can be drawn about the stratification of cockroaches within structures according to species.

In nearly all structures infested by cockroaches, food of some kind is available, either in the structure itself or nearby. This may be the food stored by man for his own use or the use of kept animals; it may be crumbs, food spillage, garbage, or excreta; glues and pastes on cartons, boxes, stamps, envelopes, labels, and wall paper; sizing on cloth and book covers; various dried animal and plant products; dead insects; living plants; etc. In fact, it is almost impossible, despite good housekeeping, to keep any structure used by man free of all food suitable for cockroaches.

That the requisite temperature, water, and food are provided, more or less adequately, by a variety of structures is attested by the innumerable infestations of cockroaches that develop when control measures are relaxed. Within structures the accessibility of certain harborages to cockroaches probably depends on the habits of the species and to some extent on their size. Similar types of harborages in different structures may be used by the same species, although there seems to be some overlapping by different species into the same kinds of daytime shelters. The comparative ecology of domiciliary cockroaches has not been thoroughly investigated, so any interpretation of observational data is necessarily speculative and inconclusive at this time. Our discussion on pages 324 to 343 is also pertinent to this section.

LAND-BASED STRUCTURES

Dwellings provide a variety of microhabitats that are acceptable to cockroaches. It has been stated that old houses, or houses that have many cracks and crevices, or have basement kitchens that are not kept clean and in good repair are particularly liable to invasion by cockroaches (Laing, 1946; British Museum [Nat. Hist.], 1951). Although this statement is undoubtedly true, it has been our personal experience, as well as the experience of others, that new, clean, and well-planned houses and apartments are also easily and sometimes quickly invaded by cockroaches. Mallis (1954) has cited the following places that are frequently infested by cockroaches in homes. In the kitchen, cockroaches are found in and around sinks, in cupboards above and below sinks, under tables and chairs, in stoves, around breadboards, in utility cabinets, in kitchen closets, under linoleum, behind, under, and inside refrigerators and iceboxes. In living rooms cockroaches are found in furniture, studio couches, sewing machines, closets, and bookshelves; behind picture frames, pennants, calendars, and other wall ornaments. In bathrooms cockroaches are found in and behind utility cabinets and toilets; they may be found in wicker clothes hampers, in brooms and mops, and in door hinges. Ordinarily, cockroaches are not found in bedrooms unless they are abundant elsewhere in the dwelling. Additional harborages are cited under specific cockroaches in the list below.

In markets DeLong (1948) found the German cockroach in bags of potatoes and onions, in crates of citrus fruits, in pads and shredded papers in banana boxes, and in cases of bottled beverages. The insects were attracted by coffee and crawled into the folds of coffee bags. They were found in cartons of canned goods; in bread and baked goods; in cartons of packaged cookies, cakes, and crackers. Packaged cereals were attractive, and cockroaches were sometimes found in packages of cigarettes. The insects occurred in scales (by the hundreds) and in cash registers. Rather heavy infestations were found under stainless-steel cappings that covered wooden arms on the fish cleaning stand. The insects were numerous in display cases where they were warm and sheltered. They were also found behind mirrors above produce racks, in electrical switch boxes and conduits, and in telephone boxes, as well as generally in cracks and behind loose moldings or loose wall boards. Enclosed boxlike tables were frequently heavily infested.

In restaurants cockroaches may be found in the following places: Crevices in wood, plaster, concrete, and metal; in the bar; in the kitchen and in the associated equipment; in cupboards, lavatories, and garbage storage areas; and on the undersides of chairs and tables (Mallis, 1954).

In drug stores Frings (1948) found cockroaches behind the mirror and between the sink and the cooler. Thousands were found in hollow ornamental shelf edging. The hollow bases of malted-milk dispensers and drink mixers were cockroach havens.

In a hospital Frings (1948) found cockroaches in decorative trim around doorways, by the thousands in wicker laundry baskets, and in incubators for premature babies. In military hospitals we have seen cockroaches (Blattella germanica) in kitchens and dining halls in the usual hiding places mentioned above and on the undersides of stainless-steel serving tables.

In department stores cockroaches have been found in food departments, beauty salons, rest rooms, dressing rooms, linen departments, and stationery departments (Anonymous, 1952). The infestation in the linen department was traced to clean towels which, when returned from the laundry, contained at least 500 cockroaches per bundle. The insects were carried into the rest rooms and beauty salon when the towels were distributed.

The microhabitats of cockroaches in privies and sewers have not been studied. These habitats are particularly important in view of the demonstrated migrations of cockroaches from sewers into dwellings and the possible dissemination of pathogenic microorganisms from feces to food. The reader is referred to our 1957(a) paper for a summary of the known information on cockroach dispersal from sewers.

COCKROACHES ASSOCIATED WITH LAND-BASED STRUCTURES

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

Aglaopteryx ypsilon

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

Allacta similis

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

Blaberus craniifer

Cuba.—Household pest (Deschapelles, 1939). Particularly abundant in houses in Santiago and Havana (Rehn and Hebard, 1927).

Florida.—Under boards in woodshed (Rehn and Hebard, 1912, 1914).

Blaberus discoidalis

Ecuador.—In eating places (Campos R., 1926).

Hispaniola.—In houses (Rehn and Hebard, 1927).

New Jersey.—In greenhouse (Weiss, 1917).

Puerto Rico.—In homes (Seín, 1923). In fruit stores (Wolcott, 1950).

Blatta lateralis

Central Asia.—Household pest, often found in homes with clay floors (Bei-Bienko, 1950).

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

Arenivaga bolliana

THE BIOTIC ASSOCIATIONS OF
COCKROACHES

THE BIOTIC ASSOCIATIONS OF COCKROACHES^[1]