This dissertation has been 69-15,943 microfilmed exactly as received

MILLER, Henry Keith, 1934- COMPARATIVE MORPHOLOGICAL STUDIES OF PROV ENTRICULI IN THE ORDER DICTYOPTERA: SUBORDER BLATTARIA.

The Ohio State University, PiuD., 1969 Entomology

University Microfilms, Inc., Ann Arbor, Michigan COMPAR ATIVE MORPHOLOGICAL STUDIES OF

PROVENTRICULI IN THE ORDER

DICTYOPTERA: SUBORDER

BLATTARIA

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By

Henry Keith Miller, B. S. in Educ. , M. S. Ent.

s{s sje sjc $ 9jc #

The Ohio State University 1969

Approved by

A d v ise r Faculty of Entomology ACKNOWLEDGMENTS

My sincere thanks go to Dr. Frank W. Fisk for his in­ valuable advice and encouragement during the course of this investigation, as well as the generous contribution of specimens from his personal collection.

I would also like to express my deep appreciation for the numerous specimens contributed by the following persons:

Dr. Louis M. Roth, Pioneering Research Laboratory, U. S. Army

Natick Laboratories, Natick, Massachusetts; Dr. M. J. Mackerras,

Division of Entomology, Australian Commonwealth Scientific and

Industrial Research Organization; Dr. Ashley B. Gurney, National

Museum, Washington, D. C. ; and Dr. Charles A. Triplehorn, Curator of Insects, The Ohio State University.

Photographic illustrations herein were made possible by the generous cooperation of Dr. Paul A. Colinvaux, Faculty of Popula­ tion and Environmental Biology, The Ohio State University, who permitted the use of sophisticated photomicrography equipment.

Technical advice was generously contributed by Dr. Colinvaux,

Dr. Eileen K. Schofield, The Ohio State University, as well as

Harold "Bo" Hagen, regional representative of Leitz, Inc. Thanks are also extended to Dr. S. S. Y. Young and Dr. D. E.

Johnston of The Ohio State University Genetics and Entomology

Faculties respectively, for their suggestions on handling portions of the data obtained from this study. VITA

1934 ...... Born February 9th, Patton, Pennsylvania

1956 ...... B. S. in Educ. , Capital University, Columbus, Ohio

1956-1959 . . . Lt. , U. S. Air Force Intelligence

1959 ...... Teacher, Gahanna Lincoln H. S. , Gahanna, Ohio

1960 ...... Graduate Teaching Assistant in Zoology and Entomology, The Ohio State University

196 1 . Research Assistant, Columbus Psychiatric Hospital, Columbus, Ohio

1962-1966 . . . Instructor and Assistant Professor, Biology Department, Capital University, Columbus, Ohio

1965-1969 Summers and Part-time . . . Entomologist, Encephalitis Project, Ohio Department of H ealth

1968 Summer . Osburn Fund Fellow, Faculty of Entomology, The Ohio State University

PUBLICATION

"A Laboratory Investigation of Maturation in the Cuban Burrowing Cockroach, Byrsotria fumigata (Guerin). " R. W. Fleischmann, H. K. M iller, and R. V. Skavaril, Ohio Journal of Science, Vol. 68: 11-17, January 1968. FIELDS OF GRADUATE STUDY

Major Field: Entomology

Medical Entomology. Dr. C. E. Venard, The Ohio State U n iv e rsity

Insect Physiology, Dr. F. W. Fisk, The Ohio State U n iv e rsity

v TABLE OF CONTENTS

P ag e ACKNOWLEDGMENTS . ii

VITA ...... iv

LIST OF ILLUSTRATIONS...... ix

INTRODUCTION ...... 1

HISTORICAL REVIEW ...... 2

METHODS...... ; ...... 5

CLASSIFICATION ...... 8

GENERAL DESCRIPTION OF THE BLATTARIAN PROVENTRICULUS...... 17

TERMINOLOGY USED IN KEY ...... 18

PROVENTRICULUS DIAGRAM ...... 22

TENTATIVE KEY TO SUBFAMILIES OF BL ATT ARIA BASED ON PROVENTRICULI...... 23

PHOTOMICROGRAPHS...... 30

DESCRIPTIONS AND DISCUSSION...... 57

Family Cryptocercidae...... 57

Subfamily Cryptocercinae...... 57

vi Page Family Blattidae...... 58

Subfamily Lamproblattinae...... 59

Subfamily Blattinae ...... 60

Subfamily Polyzosterinae...... 61

Family Polyphagidae...... 63

Subfamily Polyphaginae...... 63

Subfamily Holocompsinae...... 65

Family Blattellidae...... 66

Subfamily Anaplectinae...... 66

Subfamily Ectobiinae...... 67

Subfamily Nyctiborinae...... 69

Subfamily Blattellinae...... 70

Subfamily Plectopterinae...... 72

Family Blaberidae...... 74

Subfamily Blaberinae...... 74

Subfamily Zetoborinae...... 75

Subfamily Epilam prinae...... 76

Subfamily Diplopterinae...... 79

Subfamily Panchlorinae...... 79

Subfamily Panesthiinae...... 81

Subfamily Perisphaerinae...... 83

v ii P age Subfamily Pycnoscelinae...... 84

Subfamily Oxyhaloinae...... 85

SUMMARY...... 86

BIBLIOGRAPHY ...... 88

v iii LIST OF ILLUSTRATIONS

P age Plate I. Diagram of Proventriculus...... 22

Fig. 1. Cryptocercus punctulatus Scudder...... 30

Fig. 2. Lamproblatta albipalpus Hebard ...... 30

Fig. 3. Lamproblatta meridionalis (Bruner)...... 30

Fig. 4. Lamproblatta romanii Rehn ...... 30

Fig. 5. Blatta orientalis Linnaeus ...... 30

Fig. 6. Celatoblatta subcorticaria Johns ...... ’ 30

Fig. 7. Celatoblatta vulgaris Johns ...... 31

Fig. 8. Deropeltis erythrocephala (Fabricius)...... 31

Fig. 9. Hebardina sp ...... 31

Fig. 10. Henicotyle antillarum (Brunner)...... 31

Fig. 11. Henicotyle n. sp ...... 31

Fig. 12. Neostylopyga rhombifolia (Stoll) ...... 31

Fig. 13. Periplaneta americana (Linnaeus) ...... 32

Fig. 14. Peripaneta australasiae (Fabricius) ...... 32

Fig. 15. Periplaneta brunnea Burm eister ...... 32

Fig. 16. Periplaneta fuliginosa (Serville) ...... 32

Fig. 17. Periplaneta japonica Karny ...... 32

Fig. 18. Cosmozosteria trifasciata (Tepper) ...... 32 ix P ag e Fig. 19. Drymaplaneta semivitta (Walker)...... 33

Fig. 20. Eurycotis biolleyi Rehn...... 33

Fig. 21. Eurycotis decipiens (Kirby)...... 33

Fig. 22. Eurycotis floridana (Walker)...... 33

Fig. 23. Eurycotis n. sp...... 33

Fig. 24. Methana convexa (Walker) ...... 33

Fig. 25. Pelmatosilpha coriacea Rehn ...... 34

Fig. 26. Platyzosteria castanea (Brunner)...... 34

Fig. 27. Platyzosteria melanaria (Erichson) ...... 34

Fig. 28. Polyzosteria limbata (Burmeister)...... 34

Fig. 29. Polyzosteria viridissima Shelford...... 34

Fig. 30. Temnelytra truncata (Brunner)...... 34

Fig. 31. Arenivaga cervarae (Bolivar)...... 35

Fig. 32. Compsodes delicatulis (Saussure and Zehntner) . . . 35

Fig. 33. Homoeogamia mexicana (Burm eister) ...... 35

Fig. 34. Latindia dohrniana (Saussure and Zehntner) ...... 35

Fig. 35. Paralatindia azteca (Saussure)...... 35

Fig. 36. Polyphaga aegyptiaca (Linnaeus) ...... 35

Fig. 37. Holocompsa sp ...... 36

Fig. 38. Zetha vestita (Brulle)...... 36

Fig. 39. Anaplecta fallax (Saussure)...... 36

x P age Fig. 40. Anaplecta mexicana Saussure...... 36

Fig. 41. Anaplecta sp ...... 36

Fig. 42. Anaplecta sp . 36

Fig. 43. Ectobius lapponicus (Linnaeus) ...... 37

Fig. 44.. Ectobius pallidus (Olivier) ...... 37

Fig. 45. Ectobius sylvestris (Poda) ...... 37

Fig. 46. Parellipsidion conjunctum (W alker) ...... 37

Fig. 47. Megaloblatta blaberoides (W alker) ...... 37

Fig. 48. Nyctibora obscura Saussure...... ' 37

Fig. 49. Nyctibora sp ...... 38

Fig. 50. Blattella germanica (Linnaeus) ...... 38

Fig. 51. Chromatonotus heterus (Hebard) ...... 38

Fig. 52. Gislenia australica (Brunner)...... 38

Fig. 53. Ischnoptera deropeltiformis (Brunner) ...... 38

Fig. 54. Ischnoptera galibi H ebard ...... 38

Fig. 55. Ischnoptera panamae Hebard ...... 39

Fig. 56. Ischnoptera rufa rufa (De G eer) ...... 39

Fig. 57. Loboptera decipiens (Germar) ...... 39

Fig. 58. Nelipophygus n. sp ...... 39

Fig. 59. Nesomylacris sp ...... 39

Fig. 60. Pulvillar microtrichiae from Nesomylacris n. sp. Approximately 450 X ...... 39

xi P age Fig. 61. Parcoblatta fulvescens (Saussure and Zehntner) . . 40

Fig. 62. Parcoblatta pensylvanica (De G eer)...... 40

Fig. 63. Parcoblatta virginica (Brunner)...... 40

Fig. 64. Pseudomops septentrionalis (Hebard) ...... 40

Fig. 65. Shawella couloniana (Saussure) ...... 40

Fig. 66. Symploce hospes (Perkins) ...... 40

Fig. 67. Symploce ruficollis (Fabricius)...... 41

Fig. 68. Symploce sp ...... 41

Fig. 69. Xestoblatta buscki (Gurney)...... 41

Fig. 70. Xestoblatta festae (Griffini) ...... 41

Fig. 71. Xestoblatta immaculata Hebard ...... 41

Fig. 72. Xestoblatta cantralli Fisk and Gurney...... 41

Fig. 73. Aglaopteryx facies (W alker) ...... 42

Fig. 74. Aglaopteryx sp ...... 42

Fig. 75. Aglaopteryx sp ...... 42

Fig. 76. Agmoblatta thaxteri (Hebard) ...... 42

Fig. 77. Amazonina conspersa (Brunner) ...... 42

Fig. 78. Amazonina platystylata (Hebard)...... 42

Fig. 79. Amazonina n. sp ...... 43

F ig. 80. C ahita nahua ( S a u s s u r e ) ...... 43

Fig. 81. Caloblatta sp ...... 43

xii P age Fig. 82. Cariblatta craticula Hebard...... 43

Fig. 83. Cariblatta fossicauda-Hebard...... 43

Fig. 84. Cariblatta imitans H ebard ...... 43

Fig. 85. Cariblatta lutea minima Hebard ...... 44

Fig. 86. Cariblatta plagia Hebard ...... 44

Fig. 87. Cariblatta sp ...... 44

Fig. 88. Cariblattoides sp...... 44

Fig. 89. Ceratinoptera n. sp ...... 44

Fig. 90. Ceratinoptera sp ...... 44

Fig. 91. Chorisoneura texensis Saussure and Zehntner.... 45

Fig. 92. Chorisoneura sp...... 45

Fig. 93. Dendroblatta cnephaia Hebard ...... 45

Fig. 94. Dendroblatta sobrina Rehn ...... 45

Fig. 95. Ellipsidion sp ...... 45

F ig. 96. E uphyllodrom ia an g u stata ( L a tr e ille ) ...... 45

Fig. 97. Euphyllodromia decastigmata Hebard ...... 46

Fig. 98. Euthlastoblatta n. sp ...... 46

Fig. 99. Imblattella fratercula H ebard...... 46

Fig. 100. Imblattella impar (Hebard) ...... 46

O Fig. 101. Latiblattella angustifrons H ebard ...... 46

Fig. 102. Latiblattella sp ...... 46

x iii Page Fig. 103. Lophoblatta arlei Albuquerque...... 47

Fig. 104. Lophoblatta brevis Rehn ...... 47

Fig. 105. Lophoblatta n. sp ...... 47

Fig. 106. Nahublattella nahua (Saussure)...... 47

Fig. 107. Nahublattella n. sp ...... 47

Fig. 108. Neoblattella borinquen Rehn and H ebard ...... 47

Fig. 109. Neoblattella vomer Rehn and Hebard ...... 48

Fig. 110. Onychostylus notulatus (Stal)...... 48

Fig. 111. Rhytidometopum sp...... 48

Fig. 112. Riatia fulgida (Saussure)...... 48

Fig. 113. Supella supellectum (Serville)...... 48

Fig. 114. Archimandrita tessellata Rehn ...... 48

Fig. 115. Aspiduchus borinquen Rehn...... 49

Fig. 116. Aspiduchus cavernicola Rehn...... 49

Fig. 117. Blaberus atropos (Stoll) ...... 49

Fig. 118. Blaberus craniifer Burm eister...... 49

Fig. 119. Blaberus discoidalis Serville...... 49

Fig. 120. Blaberus giganteus (Linnaeus)...... 49

Fig. 121. Blaberus parabolicus (W alker)...... 50

Fig. 122. Byrsotria fumigata (Guerin)...... 50

Fig. 123. Eublaberus distanti (Kirby)...... 50

xiv P age Fig. 124. Eublaberus posticus (Erichson)...... 50

Fig. 125. Galiblatta n. sp ...... 50

Fig. 126. Hemiblabera brunneri Rehn and Hebard ...... 50

Fig. 127. Petasodes moufeti (Kirby)...... 51

Fig. 128. Lauxoblatta sp...... 51

Fig. 130. Type II microspines from valvular plicae of Phortioeca phoraspoides. Approximately 450 X . . . 51

Fig. 131. Type III microspines from dental areas of Phortioeca phoraspoides. Approximately 450 X . . . 51

Fig. 132. Zetabora signaticollis Burm eister ...... 51

Fig. 133. Ataxigamia tatei Tepper ...... 52

Fig. 134. Audreia carinulata (Saussure)...... 52

Fig. 135. Epilampra abdomen-nigrum (De G eer) ...... 52

Fig. 136. Epilampra azteca Saussure...... 52

Fig. 137. Epilampra grisea (De G eer) ...... 52

Fig. 138. Epilampra mexicana Saussure ...... 52

Fig. 139. Epilampra wheeleri Rehn and Hebard ...... 53

Fig. 140. Laxta granicollis (Saussure)...... 53

Fig. 141. Litopeltis biolleyi (Saussure)...... 53

Fig. 142. Litopeltis bispinosa (Saussure) ...... 53

Fig. 143. Diploptera punctata (Eschscholtz) ...... 53

Fig. 144. Type I microspines from Diploptera punctata Approximately 450 X ...... 53

xv P age Fig. 145. Achroblatta luteola (Blanchard) ...... 54

Fig. 146. Capucina patula (W alker) ...... 54

Fig. 147. Panchlora nivea (Linnaeus)...... 54

Fig. 148. Proscratea complanata (Perty) ...... 54

Fig. 149. Geoscapheus robustus Tepper ...... 54

Fig. 150. Macropanesthia rhinocerus Saussure ...... 54

Fig. 151. Panesthia laevicollis Saussure...... 55

Fig. 152. Panesthia sp ...... 55

Fig. 153. Panesthia sp ...... 55

Fig. 154. Gyna capucina Gerstaecker ...... 55

Fig. 155. Pycnoscelus indicus (Fabricius)...... 55

Fig. 156. Pycnoscelus surimanensis (Linnaeus)...... 55

Fig. 157. Gromphadorhina portentosa (Schaum) ...... 56

Fig. 158. Henschoutedenia flexivitta (W alker)...... 56

Fig. 159. Leucophaea maderae (Fabricius) ...... 56

Fig. 160. Nauphoeta cineria (Olivier)...... 56

xvi INTRODUCTION

Considering the total body of taxonomic knowledge which currently exists regarding cockroaches, one finds a greater dif­ ference of opinion regarding their placement in higher taxonomic categories (i. e. superfamily and subfamily) than one finds at the genus or species level. Some clarification of relationships at these higher levels seems to be suggested.

The principal objectives of this study are two-fold: One, to compare the proventricular morphology of as broad a cross section of Blattarian species as possible; and two, using only proventricular characters, construct a key to the subfamilies of cockroaches. Hopefully, this key will show how well or how poorly proventricular characters alone can be used to construct a system­ atic arrangement which will m irror modern classification systems for the suborder Blattaria.

1 HISTORICAL REVIEW

Prior to the work of McKittrick (1964) the proventricular morphology of only a scattered few species of cockroaches was described. Wilde (1877) investigated the "gizzards" of various

Orthopterans, and Ramme (1913) discussed significances of proven- triculi in selected Coleoptera and Orthoptera. Sanford (1918) dealt with the proventriculus indirectly during his work on the physiology of digestion in Blattidae, and Eidmann (1924) investigated the physiology of the "gizzard" of Periplaneta orientalis. In 1934,

Cleveland discussed the proventriculus in his monumental work on the wood-feeding cockroach Cryptocercus, a study mainly devoted to cellulose-digesting symbionts present in the gut. Judd (1948) de­ scribed a number of different proventricular types in orthopteriod insects, but said nothing about relationships between Blattarian types.

M ackerras (1965) included a system for numbering proven­ tricular teeth in the genus Polyzosteria.

For additional references to proventricular studies in other insects, the reader is referred to the discussion in McKittrick1 s treatise (1964). 3

Cockroach classification originated with Linnaeus (1758) who described nine species in the genus Blatta under the Order

Coleoptera. Latreille (1810) initiated use of the term "Blattaria, " but referred to it as a family in the Order Orthoptera. Burmeister

(1838) and Serville (1839) both considered roaches as belonging to a

single family (Blattina and Blattaires respectively) with relatively few genera listed.

- Brunner (1865) in his "Nouveau Systeme des Blattaires"

greatly expanded cockroach classification using eleven tribes and fifty-five genera under the superfamily Blattariae, Order Orthoptera.

Walker (1868), utilized many of the present-day family names, under the division Blattariae, Order Dermaptera.

Kirby (1904) in his Cataloque of Orthoptera, Shelford (1906a) in his contributions to "Genera Insectorum, " and Hebard (1917,

1919) all assigned cockroaches to a single family, Blattidae.

Of the more recent taxonomists, Imms (1957), and

McKittrick (1964) place cockroaches as the Suborder Blattaria under the Order Dictyoptera, while Rehn (1951) considers them a super­ family under the Suborder Dictyoptera, and Princis (I960), and

Brues and Melander (1932) accord them Order (Blattariae) status.

Generally, the trend in cockroach taxonomy from the past to the present has been one of a gradual expansion to a current maximum illustrated by Princis (1962-1969) in his contributions to

M. Beier's "Orthopterorum Catalogus" in which he divides the world species of the Order Blattaria into four suborders, twenty-eight families, twenty-one subfamilies and 429 genera.

The current view toward classification sesms to recommend the application of as many characters as possible in placing a speci­ men. Shelford (1906b) was one of the first to recognize this. The extensive monograph by Roth and Willis (I960) strongly suggested that reproductive habits may have strong phyletic implications, though they did not attempt to alter classification schemes in their study.

McKittrick (1964) made the first genuine effort at cockroach classification based on many characters, both external and internal; a comparative study that emphasized male and female genitalia with associated musculature, and also included oviposition behavior and morphology of proventriculi. METHODS

The techniques used to remove, treat, and mount proven­ triculi were initially quite sim ilar to those of McKittrick (1964).

Gross dissection were; made of pinned (relaxed), wet preserved, or live cockroaches. They were then soaked in a warm (60°C) solution of 10 per cent aqueous potassium hydroxide for one to two hours.

This treatment served to remove attached muscles and other cellu­ lar matter. After removing the proventriculi from the potassium hydroxide, they were rinsed well in distilled water, and transferred to 70 per cent alcohol, where any remaining tracheal tubes or cellular matter were removed.

Difficulty was encountered using this technique with pinned museum specimens which had been dried out for several years, in that any fat present in the thorax and abdomen had hardened to a plaster-like consistency. Efforts to chip out the proventriculus prior to its exposure to potassium hydroxide often resulted in its destruction, especially in those species having delicate membranous proventriculi.

It was found that this problem could usually be overcome by

soaking the entire cockroach in potassium hydroxide prior to dissecting out the proventriculus. Since this treatment resulted in a much more transparent cuticle, a careful dissection, which usually- involved entrance to the body cavity between the first and second ventral abdominal sclerites, would allow removal of the structure wanted without damaging genitalia or external morphological features. The specimen could then be repinned, dried, and returned to the collection.

Following a wash in 70 per cent alcohol, McKittrick stained the preparation with Chlorazol black E, then counterstained in acid fuchsin. It was found that elimination of the counterstain did not noticeably detract from the value of the finished preparation, so it was omitted.

Additional problems arose in connection with the specimens' exposure to 100 per cent alcohol and xylene as suggested by

McKittrick. M aterial prepared under these conditions tended to be extremely fragile, frequently tearing while being cleaned of parti­ culate m atter or while being flattened under a coverslip.

To eliminate this fragility, a new set of procedures was devised, utilizing Methyl Cellosolve (Ethylene Glycol Monomethyl

Ether). Small (5 ml) beakers were used as carriers throughout the procedure, and following a distilled water wash, specimens were transferred to Cellosolve, then stained (17 per cent Cholrazol black E in Cellosolve) for two minutes, destained for ten to twenty seconds in fresh Cellosolve and transferred to cedar oil for clearing. While in cedar oil, specimens were slit longitudinally and cleared of parti­ culate material. Mounting was in Permount, directly from the cedar oil, with excess oil being blotted off before the mounting medium was applied.

The use of Cellosolve eliminated fragility problems, as well as shortening considerably the time required to process each slide.

To insure accuracy in relating the correct proventriculus with its donor, proventricular preparations were done individually with each occupying a slide previously etched with the appropriate scientific name and identity number.

A total of 162 species were examined, in most instances in­ volving only one slide per species, as proventricular characters did not vary with sex or age, but only in size.

All slides were examined under a Spencer AO phase- contrast microscope with magnifications from 100-430X. CLASSIFICATION OF SPECIMENS UTILIZED IN STUDY

Order: Dictyoptera

Suborder: Blattaria

Superfamily: Blattoidea

Family: Cryptocercidae

Subfamily: Cryptocercinae

Cryptocercus punctulatus Scudder. (Fig. 1)

Family: Blattidae

Subfamily: Lamproblattinae

Lamproblatta albipalpus Hebard. (Fig. 2)

Lamproblatta meridionalis (Bruner). (Fig. 3)

Lamproblatta romanii Rehn. (Fig. 4)

Subfamily: Blattinae

Blatta orientalis Linnaeus. (Fig. 5)

Celatoblatta subcorticaria Johns. (Fig. 6)

Celatoblatta vulgaris Johns. (Fig. 7)

Deropeltis erythroeephala (Fabricius). (Fig. 8)

Hebardina sp. (Fig. 9)

Henicotyle antillarum (Brunner). (Fig. 10)

8 Henicotyle n. sp. (Fig. 11)

Neostylopyga rhombifolia (Stoll). Fig. 12)

Periplaneta americana (Linnaeus). (Fig. 13)

Periplaneta australasiae (Fabricius). (Fig. 14)

Periplaneta brunnea Burmeister. (Fig. 15)

Periplaneta fuliginosa (Serville). (Fig. 16)

Periplaneta japonica Karny. (Fig. 17)

Subfamily Polyzosterinae

Cosmozosteria trifasciata (Tepper). (Fig. 18)

Drymaplaneta semivitta (Walker). (Fig. 19)

Eurycotis biolleyi Rehn. (Fig. 20)

Eurycotis decipiens (Kirby). (Fig. 21)

Eurycotis floridana (Walker). (Fig. 22)

Eurycotis n. sp. (Fig. 23)

M ethana convexa (W alker). (Fig. 24)

Pelmatosilpha coriacea Rehn. (Fig. 25)

Platyzosteria castanea (Brunner). (Fig. 26)

Platyzosteria melanaria (Erichson). (Fig. 27)

Polyzosteria limbata (Burmeister). (Fig. 28)

Polyzosteria viridissima Shelford. (Fig. 29)

Temnelytra truncata (Brunner). (Fig. 30) 10

Superfamily: Blaberoidea

Family: Polyphagidae

Subfamily: Polyphaginae

Arenivaga cervarae (Bolivar). (Fig. 31)

Arenivaga tonkawa Hebard. (No Fig. )

Compsodes delicatulus (Saussure and Zehntner). (Fig. 32)

Homoeogamia mexicana (Burmeister). (Fig. 33)

Latindia dohrniana (Saussure and Zehntner). (Fig, 34)

Paralatindia azteca (Saussure). (Fig. 35)

Polyphaga aegyptiaca (Linnaeus). (Fig. 36)

Subfamily: Holocompsinae

Holocompsa sp. (Fig. 37)

Zetha vestita (Brulle). (Fig. 38)

Family: Blattellidae

Subfamily: Anaplectinae

Anaplecta fallax (Saussure). (Fig. 39)

Anaplecta mexicana Saussure. (Fig. 40)

Anaplecta sp. (Fig. 41)

Anaplecta sp. (Fig. 42)

Subfamily: Ectobiinae

Ectobius lapponicus (Linnaeus). (Fig. 43)

Ectobius pallidus (Olivier). (Fig. 44) 11

Ectobius sylvestris (Poda). (Fig. 45)

Parellipsidion conjunctum (Walker). (Fig. 46)

Subfamily: Nyctiborinae

Megaloblatta blaberoides (Walker). (Fig. 47)

Nyctibora obscura Saussure. (Fig. 48)

Nyctibora sp. (Fig. 49)

Subfamily: Blattellinae

Blattella germanica (Linnaeus). (Fig. 50)

Chromatonotus heterus (Hebard). (Fig. 51)

Gislenia australica (Brunner). (Fig. 52)

Ischnoptera deropeltiformis (Brunner). (Fig. 53)

Ischnoptera galibi Hebard. (Fig. 54)

Ischnoptera panamae Hebard. (Fig. 55)

Ischnoptera rufa rufa (De Geer). (Fig. 56)

Loboptera decipiens (Germar). (Fig. 57)

N elipophygus n. sp. (Fig. 58)

Nesomylacris sp. (Figs. 59. 60)

Parcoblatta fulvescens (Saussure and Zehntner). (Fig. 61)

Parcoblatta pensylvanica (De Geer). (Fig. 62)

Parcoblatta virginica (Brunner). (Fig. 63)

Pseudomops septentrionalis (Hebard). (Fig. 64)

Shawella couloniana (Saussure). (Fig. 65) Symploce hospes (Perkins). (Fig. 66)

Symploce ruficollis (Fabricius). (Fig. 67)

Sym ploce sp. (Fig. 68)

Xestoblatta buscki (Gurney). (Fig. 69)

Xestoblatta festae (Griffini). (Fig. 70)

Xestoblatta immaculata Hebard. (Fig. 71)

Xestoblatta cantralli Fisk and Gurney.' (Fig.

Subfamily: Plectopterinae

Aglaopteryx facies (Walker). (Fig. 7 3)

Aglaopteryx sp. (Fig. 74)

Aglaopteryx sp. (Fig. 75)

Agmoblatta thaxteri (Hebard). (Fig. 76)

Amazonina conspersa (Brunner). (Fig. 77)

Amazonina platystylata (Hebard). (Fig. 78)

Amazonina n. sp. (Fig. 79)

Cahita nahua (Saussure). .(Fig. 80)

Caloblatta sp. (Fig. 81)

Cariblatta craticula Hebard. (Fig. 82)

Cariblatta fossicauda Hebard. (Fig. 83)

Cariblatta imitans Hebard. (Fig. 84)

Cariblatta lutea minima Hebard. (Fig. 85)

Cariblatta plagia Hebard. (Fig. 86) Cariblatta sp. (Fig. 87)

Cariblattoides sp. (Fig. 88)

Ceratinoptera n. sp. (Fig. 89)

C e ra tin o p te ra sp. (Fig. 90)

Chorisoneura texensis Saussure and Zehntner. (Fig. 91)

Chorisoneura sp. (Fig. 92)

Dendroblatta cnephaia Hebard. (Fig. 93)

Dendroblatta sobrina Rehn. (Fig. 94)

Ellipsidion sp. (Fig. 95)

Euphyllodromia angustata (Latreille). (Fig. 96)

Euphyllodrqmia decastigmata Hebard. (Fig. 97)

Euthlastoblatta n. sp. (Fig. 98)

Imblattella fratercula Hebard. (Fig. 99)

Imblattella impar (Hebard). (Fig. 100)

Latiblattella angustifrons Hebard. (Fig. 101)

Latiblattella sp. (Fig. 102)

Lophoblatta arlei Albuquerque. (Fig. 103)

Lophoblatta brevis Rehn. (Fig. 104)

L ophoblatta n. sp. (Fig. 105)

Nahublattella nahua (Saussure). (Fig. 106)

Nahublattella n. sp. (Fig. 107)

Neoblattella borinquen Rehn and Hebard. (Fig. 108) Neoblattella vomer Rehn and Hebard. (Fig. 109) 0 _ Onychostylus notulatus (Stal). (Fig. 110)

Plectoptera sp. (No Fig.)

Rhytidometopum sp. (Fig. Ill)

Riatia fulgida (Saussure). (Fig. 112)

Supella supellectum (Serville). (Fig. 113)

Family: Blaberidae

Subfamily: Blaberinae

Archimandrita tessellata Rehn. (Fig. 114)

Aspiduchus borinquen Rehn. (Fig. 115)

Aspiduchus cavernicola Rehn. (Fig. 116)

Blaberus atropos (Stoll). (Fig. 117)

Blaberus craniifer Burmeister. (Fig. 118)

Blaberus discoidalis Serville. (Fig. 119)

Blaberus giganteus (Linnaeus). (Fig. 120)

Blaberus parabolicus (Walker). (Fig. 121)

Byrsotria fumigata (Guerin). (Fig. 122)

Eublaberus distanti (Kirby). (Fig. 123)

JSublaberus posticus (Erichson). (Fig. 124)

Galiblattan. sp. (Fig. 125)

Hemiblabera brunneri Rehn and Hebard. (Fig. 126)

Petasodes moufeti (Kirby). (Fig. 127) 15

Subfamily: Zetoborinae

Lauxoblatta sp. (Fig. 128)

Phortioeca phoraspoides (Walker). (Figs. 129, 130, 131)

Zetabora signaticollis Burmeister. (Fig. 132)

Subfamily: Epilamprinae

Ataxigamia tatei Tepper. (Fig. 133)

Audreia carinulata (Saussure). (Fig. 134)

Audreia gatunae (Hebard). (No Fig.)

Epilampra abdomen-nigrum (De Geer). (Fig. 135)

Epilampra azteca Saussure. (Fig. 136)

Epilampra grisea (De Geer). (Fig. 137)

Epilampra mexicana Saussure. (Fig. 138)

Epilampra wheeleri Rehn and Hebard. (Fig. 139)

Hyporhicnoda reflexa (Saussure and Zehntner). (No Fig.)

Laxta g rani colli s (Saussure). (Fig. 140)

Litopeltis biolleyi (Saussure). (Fig. 141)

Litopeltis bispinosa (Saussure). (Fig. 142)

Subfamily: Diplopterinae

Diploptera punctata (Eschscholtz). (Figs. 143, 144)

Subfamily: Panchlorinae

Achroblatta luteola (Blanchard). (Fig. 145)

Capucina patula (Walker). (Fig. 146) Panchlora nivea (Linnaeus). (Fig. 147)

Proscratea complanata (Perty). (Fig. 148)

Subfamily: Panesthiinae

Geoscapheus robustus Tepper. (Fig. 149)

Macropanesthia rhinocerus Saussure. (Fig. 150)

Panesthia laevicollis Saussure. (Fig. 151)

P a n e sth ia sp. (Fig. 152)

Panesthia sp. (Fig. 153)

Subfamily: Perisphaerinae

Gyna capucina Gerstaecker. (Fig. 154)

Subfamily: Pycnoscelinae

Pycnoscelus indicus (Fabricius). (Fig. 155)

Pycnoscelus surimanensis (Linnaeus). (Fig. 156)

Subfamily: Oxyhaloinae

Gromphadorhina portentosa (Schaum). (Fig. 157)

Henschoutedenia flexivitta (Walker). (Fig. 158)

Leucophaea maderae (Fabricius). (Fig. 159)

Nauphoeta cinerea (Olivier). (Fig. 160) GENERAL DESCRIPTION OF THE BLATTARIAN PROVENTRICULUS

The cuticle-lined foregut of cockroaches, as in all insects, terminates in a narrow stomodeal valve which appears to regulate the passage of food from the crop to the midgut. In Blattaria a more sophisticated structure, the armarium (McKittrick, 1964), is structurally and functionally allied with the stomodeal valve, and together they comprise the proventriculus.

Basic proventricular organization is usually quite consistent throughout the Blattaria, and involves six major panels or plicae which extend the total length of the organ. Each plica is in turn equipped with an anterior sclerotized tooth, tubercle, or dental area; a median invagination of variable size and shape, referred to as the pulvillus or pulvillar area; and a posterior valvular area comprising the stomodeal valve (Plate I).

Few basic differences exist in the valvular plicae of cock­ roaches, but the dental and pulvillar belts as well as the interdental areas vary greatly, and form the basis for differentiating between various taxonomic groupings.

17 DEFINITIONS OF SOME TERMS USED IN KEY

Antedental m icrotrichial tuft (admt) - - A group or collection of m icro-

trichiae commonly found just anterior to each tooth and

secondary sclerotization. Most common in subfamily

Plectopterinae, but also in some members of the subfamily

Blattellinae (Plate I, Fig. 89).

Capped tubercles--Tubercles in which the tips are more heavily

sclerotized than the bases (as in the subfamily

Pycnoscelinae). The sclerotized portions of each tubercle

extend back to a distinct line which crosses the midline of

the tubercle, resulting in a capped appearance.

Dental area--That portion of the dental belt which bears the teeth or

tubercles, regardless of whether or not either is present.

Dental tubercle (tu)--A lightly sclerotized protuberance of the dental

belt, much smaller than a tooth, and commonly found in

the anterior quarter of the dental belt (Fig. 157). Usually

total six in number, but vary from 1 to 24 in different

species. Any projections which are smaller than those

pictured in Figs. 118-120, are referred to as tubercles.

18 19

Intercalates (ic)--Sclerotized rays flanking both secondary

sclerotizations and interdentaries in interdental areas.

May or may not be present when secondary sclerotizations

and/or interdentaries are present (Plate I, Fig. 61).

Interdental area (ia)--That area between two teeth or tubercle

areas (Plate I).

Interdentaries (id)--Sclerotized rays which occur in pairs, one each

flanking the secondary sclerotization of each interdental

area. Never present when secondary sclerotizations are

absent, and do not occur in proventriculi without teeth

(Plate I, Fig. 71).

Microspines (cf. Peterson, 1948, p. 284)--Scalelike, or spinelike

cuticular projections (spinules) found in most proventriculi.

Usually discernible only under high power magnifications,

and grouped in the following categories: Type I (Fig. 144);

Type II (Fig. 130); and Type III (Fig. 131).

Microtrichiae (Fig. 60) (cf. Richards, p. 268)--Hairlike or setalike

cuticular projections found in most proventriculi ("setules"

of some authors). Larger than most microspin.es and

generally broader, they are easily discernible under low

power magnifications. When present in heavy concentra­

tions the area involved may be referred to as setulose. 20

Obvious micro spines--M icro spines which are easily seen when viewed

under low-power microscope objectives.

Prim ary valvular plicae (pvp) - -Main panels in valvular portion of

proventriculus, usually six in number (Plate I, Fig. 160).

Tooth #l--Also referred to as "constricted tooth. " The central tooth

in subfamilies Blattinae and Polyzosterinae which is_con­

stricted mid-dorsoventrally or mid-laterally, and serves as

the starting point for assignment of tooth numbers 1-6

(Figs. 7, 23).

Tooth (to)--Large distinctly sclerotized projection of the dental belt

(Plate I). Usually, but not always, totaling six in number,

each one being at least one-half the dental belt or greater in

length. A proventriculus is referred to as toothed if at least

one dental sclerotization is present and is as large or larger

than those in Figs. 118-120.

Secondary sclerotization (ss)--The one sclerotized panel bisecting

the interdental area between each pair of teeth (Plate I,

Fig. 99). Not present in proventriculi without teeth, except

for five members of the subfamily Polyphaginae.

Undifferentiated proventricular panel (upv) - -A characteristic of the

subfamily Panesthiinae, in which one quarter of the proven­

triculus is composed of a wide, totally membranous panel extending the total length of the organ. Not to be confused with plica I (McKittrick, 1964), which is not as wide and possesses an anterior tubercle and pulvillus (Figs. 151-153). 22

admt ss

PL A T E I

Diagram of generalized cockroach proventriculus, showing parts (proventriculus slit longitudinally and laid open) admt--antedental microtrichial tuft; db--dental belt; ia--interdental area; ic-- intercalary; id--interdentary; pb-^pulvillar belt; p--pulvillus; pvp--prim ary valvular plicae; to--tooth; ss--secondary sclero­ tization; sv--stomodeal valve. TENTATIVE KEY TO SUBFAMILIES OF BLATTARIA BASED ON PROVENTRICULI

I Non-toothed, tubercles may be present or lacking

(Figs. 37, 38, 44, 155) ...... 2

II Toothed (Figs. 2, 12, 25) ...... 17

2(1) Tubercles (at least one) present ...... 8

2' Tubercles absent...... 3

3(2') Tubercle and interdental areas lightly sclerotized

as in Figs. 11, 111, or as in Figs. 32-35 . . Polyphaginae

3' Tubercle and interdental areas not as above...... 4

4(3') Totally membranous, no recognizable dental belt,

pulvilli, or valvular plicae; microtrichia few and

widely scattered (Figs. 37, 38)...... Holocompsinae

4' Not totally membranous, with some character­

istic organization of the dental belt, pulvilli, and

p l i c a e ...... 5

23 24

5(4') Wide undifferentiated panel present

(Fig. 1 5 0 ) ...... Panesthiinae (part)

5' Wide undifferentiated panel not present ...... 6

6(5') Scalelike type III microspines present on

anterior tooth areas (Figs. 128, 129) . . Zetoborinae (part)

6' Scalelike type III micro spines not present on

anterior tooth areas ...... 7

7(6') Scalelike type I microspines present on

dental and interdental areas (Fig. 140) . Epilamprinae (part)

7' No concentrations of any type microspines on

dental areas (Figs. 84, 145, 147) . . . Panchlorinae (part)

8(2) Wide undifferentiated panel always present,

usually with 7 tubercles (Figs. 149, 151-153) . Panesthiinae

8' Wide undifferentiated panel present or lacking,

tubercles number less than 7 or greater than 7 .... 9

9(8') Wide undifferentiated present;

9 tubercles (Fig. 154) ...... Perisphaerinae

9' No undifferentiated panel present, tubercles

usually less than 9; rarely more than ...... 9 10 Tubercles appear capped, smooth, evenly- tapered toward point; interdental area quite broad and flat, except for rectangular pouch

(Figs. 155, 156) ...... Pycnoscelinae

Tubercles and interdental areas not as above 11

Tubercle and tubercle edges smooth, not

scaly, often with 4 sclerotized rays in

valvular plicae; tubercle number usually 6

but varies from 1 (Fig. 134) to 24 (Fig. 141)

(Figs. 133, 135-140, 142) ...... Epilamprinae

Tubercle and tubercle edges rough and/or

scaly, no sclerotized rays in valvular plicae 12

Pulvilli bases rectangular (Figs. 146, 148),

square (Fig. 147), or rarely indistinct

(Fig. 1 4 5 ) ...... Panchlorinae

Pulvilli bases not rectangular, or square,

commonly indistinct in shape ...... 26

13(12') Tubercle areas triangular in shape

(Fig. 132), or as in Figs. 128, 129 ...... Zetoborinae

13' Tubercle areas not triangular in

shape or as in Fig. 128, 129 ...... 14

14(13') Tubercles not scaly, tubercle edges rough,

plica I noticeably broad (Fig. 143) ...... Diplopterinae

14' Tubercles scaly...... 15

15(14') Tubercles scaly and with roughened

edges (Figs. 9, 100, 142, 158) ...... Oxyhaloinae

15' Tubercles not as above...... 16

16(15') Tubercles at least four times as long as

wide, beset with microspines (Figs. 43, 44). Ectobiinae (part)

16' Tubercles variable in shape, not four times

as long as wide, no obvious microspines,

tubercles not scaly, edges usually smooth;

all plicae quite broad (Figs. 115, 116,

122, 125-127) ...... Blaberinae (part) 27

17(1) Teeth club-shaped, sculptured,

heavily sclerotized and all six

similar in appearance (Figs. 2-4) .... Lamproblattinae

17' Teeth not club-shaped...... 18

18(17') Teeth variable in shape, commonly with

2 m irror pairs, bladelike, with tooth #1

constructed along leading edge

(Figs. 5-17) ...... B lattin ae

18' Teeth not usually blade-like, with

tooth #1 constricted laterally ...... 19

19(18') Teeth extremely variable in shape, very

heavily sclerotized, seldom bladelike,

2 m irror pairs present with tooth #1

equally constricted laterally and often

Y-shaped (Figs. 18-30) ...... Polyzosterinae

19" Teeth moderately sclerotized, more

similar in shape, no m irror pairs or

tooth #1 p r e s e n t ...... 20 28

20(19') Secondary sclerotizations present in

interdental areas ...... 22

20' Secondary sclerotizations absent in

interdental areas ...... 21

21(20') Teeth very lightly sclerotized, quite

narrow, each with stout microtrichia

as apex; narrow folds of interdental

areas extend from crop to mid-pulvillar

line (Figs. 45, 46) ...... Ectobiinae

21' Teeth generally reduced in size, mod­

erately sclerotized, usually quite similar

in shape with blunt tips and an overall

triangular appearance (Figs. 114, 117-121,

124)...... B lab erin ae

22(21') Pulvilli and anterior valvular plicae each

with two sclerotized tubercles (Fig. 1) . . . Cryptocercinae

22' Pulvilli and anterior valvular plicae

without sclerotized tubercles...... 23 29

23(22') Teeth moderately sclerotized, highly

sculptured, rectangular (lateral view),

usually no single point or projection

on each tooth (Figs. 39-42) ...... Anaplectinae

23' Teeth not highly sculptured...... 24

24(23') Teeth almost identical, each usually with

a single long blunt tip arising from pos­

terior end of tooth and curving posteriorly

(Figs. 48, 49). or with anterior teeth tips

rounded, blunt, and hooked posteriorly

(Fig. 4 7 ) ...... Nyctiborinae

24' Teeth not rounded, blunt, or hooked

posteriorly ...... 25

25(24') Antedental microtrichial tufts usually

present along crop-dental belt junction;

apices of teeth usually with one or more

sharp microtrichiae; microspines on mid­

blade usually scattered (Figs. 73-113). . . Plectopterinae

25' Antedental microtrichial tufts usually

absent at crop-dental belt junction;

apices of teeth without m icrotrichiae;

microspines on midblade often with bases touching (Figs. 50-72) ...... Blattellinae Fig. 1. Cryptocercus punctulatus Scudder. Cuticular proventriculus.

Fig. 2. Lamproblatta albipalpus Hebard. The same.

Fig. 3. Lamproblatta meridionalis (Bruner). The same.

Fig. 4. Lamproblatta romanii Rehn. The same. _ .

Fig. 5. Blatta orientalis Linnaeus. The same.

Fig. 6. Celatoblatta subcorticaria Johns. The same. Fig. 7. Celatoblatta vulgaris Johns. Cuticular proventriculus. to #1--constricted tooth.

Fig. 8. Deropeltis erythrocephala (Fabricius). The same.

Fig. 9. Hebardina sp. The same.

Fig. 10. Henicotyle antillarum (Brunner). The same.

Fig. 11. Henicotyle n. sp. The same.

Fig. 12. Neostylopyga rhombifolia (Stoll). The same. Fig. 13. Periplaneta americana (Linnaeus). Cuticular proventriculus.

Fig. 14. Periplaneta australasiae (Fabricius). The same.

Fig. 15. Periplaneta brunnea Burmeister. The same.

Fig. 16. Periplaneta fuliginosa (Serville). The same.

Fig. 17. Periplaneta japonica Karny. The same.

Fig. 18. Cosmozosteria trifasciata (Tepper). The same. Fig. 19. Drymaplaneta semivitta (Walker). Cuticular proventriculus.

Fig. 20. Eurycotis biolleyi Rehn. The same.

Fig. 21. Eurycotis decipiens (Kirby). The same.

Fig. 22. Eurycotis floridana (Walker). The same.

Fig. 23. Eurycotis n. sp. The same, to #1 --constricted tooth.

Fig. 24. Methana convexa (Walker). The same. Fig. 25. Pelmatosilpha coriacea Rehn. Cuticular proventriculus.

Fig. 26. Platyzosteria castanea (Brunner). The same.

Fig. 27. Platyzosteria melanaria (Erichson). The same.

Fig. 28. Polyzosteria limbata (Burmeister). The same.

Fig. 29. Polyzosteria viridissima Shelford. The same.

Fig. 30. Temnelytra truncata (Brunner). The same. Fig. 31. Arenivaga cervarae (Boliver). Cuticular proventriculus.

Fig. 32. Compsodes delicatulis (Saussure and Zehntner). The same.

Fig. 33. Homoeogamia mexicana (Burmeister). The same.

Fig. 34. Latindia dohrniana (Saussure and Zehntner). The same.

Fig. 35. Paralatindia azteca (Saussure). The same.

Fig. 36._ Polyphaga aegyptiaca (Linnaeus). The same. O-lmm

Fig. 37. Holocompsa sp. Cuticular proventriculus.

Fig. 38. Zetha vestita (Brulle). The same.

Fig. 39. Anaplecta fallax (Saussure). The same.

Fig. 40. Anaplecta mexicana Saussure. The same.

Fig. 41. Anaplecta sp. The same.

Fig. 42. Anaplecta sp. The same. Fig. 43. Ectobius lapponicus (Linnaeus). Cuticularproventriculus.

Fig. 44. Ectobius pallidus (Olivier). The same.

Fig. 45. Ectobius sylvestris (Poda). The same.

Fig. 46. Parellipsidion conjunctum (Walker). The same.

Fig. 47. Megaloblatta blaberoides (Walker). The same.

Fig. 48. Nyctibora obscura Saussure. The same. ^.O Y n m

Q.*5 v n m Qovrtm

Fig. 49. Nyctibora sp. Cuticular proventriculus.

Fig. 50. Blattella germanica (Linnaeus). The same.

Fig. 51. Chromatonotus heterus (Hebard). The same.

Fig. 52. Gislenia australica (Brunner). The same.

Fig. 53. Ischnoptera deropeltiformis (Brunner). The same.

Fig. 54. Ischnoptera galibi Hebard. The same. 0-3vmVn

Q .sm tn

Fig. 55. Ischnoptera panamae Hebard. Cuticular proventriculus.

Fig. 56. Ischnoptera rufa rufa (De Geer). The same.

Fig. 57. Loboptera decipiens (Germar). The same.

Fig. 58. Nelipophygus n. sp. The same.

Fig. 59. Nesomylacris sp. The same.

Fig. 60. Pulvillar microtrichiae from Nesomylacris n. sp. Approximately 450 X. Fig. 61. Parcoblatta fulvescens (Saussure and Zehntner). Cuticular proventri cuius, ic--intercalary.

Fig. 62. Parcoblatta pensylvanica (De Geer). The same.

Fig. 63. Parcoblatta virginica (Brunner). The same.

Fig. 64. Pseudomops septentrionalis (Hebard). The same.

Fig. 65. Shawella couloniana (Saussure). The same.

Fig. 66. Symploce hospes (Perkins). The same. Fig. 67. Symploce ruficollis (Fabricius. ) Cuticular proventriculus.

Fig. 68. Symploce sp. The same.

Fig. 69. Xestoblatta buscki (Gurney). The same.

Fig. 70. Xestoblatta festae (Griffini). The same.

Fig. 71. Xestoblatta immaculata Hebard. The same. id--interdentary.

Fig. 72. Xestoblatta cantralli Fisk and Gurney. The same. Fig. 73. Aglaopteryx facies (Walker). Cuticular proventriculus.

Fig. 74. Aglaopteryx sp. .The same.

Fig. 7 5. Aglaopteryx sp. The same.

Fig. 76. Agmoblatta thaxteri (Hebard). The same.

Fig. 77. Amazonina conspersa (Brunner). The same.

Fig. 78. Amazonina platystylata (Hebard). The same. Fig. 79. Amazonina n. sp. Cuticular proventriculus.

Fig. 80. Cahita nahua (Saussure). The same.

F ig. 81. Caloblatta sp. The same.

F ig. 82. Cariblatta c rati evil a Hebard. The same.

F ig. 83. Cariblatta fossicauda Hebard. The same.

F ig. 84. Cariblatta imitans Hebard. The same. Fig. 85. Cariblatta lutea minima Hebard. Cuticular proventriculus.

Fig. 86. Cariblatta plagia Hebard. The same.

Fig. 87. Cariblatta sp. The same.

Fig. 88. Cariblattoides sp. The same.

Fie. 89. Ceratinoptera n. sp. The same. admt. --antedental m icrotrichial tuft.

Fig. 90. Ceratinoptera sp. The same. J0.lvw>n i ^

Fig. 91. Chorisoneura texensis Saussure and Zehntner. Cuticular proventriculus.

Fig. 92. Chorisoneura sp. The same.

Fig. 93. Dendroblatta cnephaia Hebard. The same.

Fig. 94. Dendroblatta sobrina Rehn. The same.

Fig. 95. Ellipsidion sp. The same.

Fig. 96. Euphyllodromia angustata (Latreille). The same. • " ‘ .V 'v K '

0 3 rr»tr»

v yifx

O-'vnYn

Fig. 97. Euphyllodromia decastigmata Hebard. Cuticular proventriculus.

Fig. 98. Euthlastoblatta n. sp. The same.

Fig. 99. Imblattella fratercula Hebard. The same. ss--secondary sclerotization. Fig. 100. Imblattella impar (Hebard). The same.

Fig. 101. Latiblattella angustifrons Hebard. The same. Fig. 102. Latiblattella sp. The same. Fig. 103. Lophoblatta arlei Albuquerque. Cuticular proventriculus.

Fig. 104. Lophoblatta brevis Rehn. The same.

Fig. 105. Lophoblatta n. sp. The same.

Fig. 106. Nahublattella nahua (Saussure). The same.

Fig. 107. Nahublattella n. sp. The same.

Fig. 108. Neoblattella borinquen Rehn and Hebard. The same. Fig. 109. Neoblattella vomer Rehn and Hebard. Cuticular proventriculus.

Fig. 110. Onychostylus notulatus (Stal). The same.

Fig. 111. Rhytidometopum sp. The same.

Fig. 112. Riatia fulgida (Saussure). The same.

Fig. 113. Supella supellectum (Serville). The same.

Fig. 114. Archimandrita tessellata 'Rehn. The same. I.OVnVn

Q.'SvnVn

Fig. 115. Aspiduchus borinquen Rehn. Cuticular proventriculus.

Fig. 116. Aspiduchus cavernicola Rehn. The same.

Fig. 117. Blaberus atropos (Stoll). The same.

Fig. 118. Blaberus craniifer Burmeister. The same.

Fig. 119. Blaberus discoidalis Serville. The same.

Fig. 120. Blaberus giganteus (Linnaeus). The same. Fig. 121. Blaberus parabolicus (Walker). Cuticular proventriculus.

Fig. 122. Byrsotria fumigata (Guerin). The same.

Fig. 123. Eublaberus distanti (Kirby). The same.

Fig. 124. Eublaberus posticus (Erichson). The same.

Fig. 125. Galiblatta n. sp. The same.

Fig. 126. Hemiblabera brunneri Rehn and Hebard. The same. Fig. 127. Petasodes moufeti (Kirby). Cuticular proventriculus.

Fig. 128. Lauxoblatta sp. The same.

Fig. 129. Phortioeca phoraspoides (Walker). The same.

Fig. 130. Type II microspines from valvular plica of Phortioeca phoraspoides. Approximately 450 X. Fig. 131. Type III microspines from dental area of Phortioeca phoraspoides. Approximately 450 X.

Fig. 132. Zetabora signaticollis Burmeister. Cuticular proventriculus. Fig. 133. Ataxigamia tatei Tepper. Cuticular proventriculus.

Fig. 134. Audreia carinulata (Saussure). The same.

Fig. 135. Epilampra abdomen-nigrum (DeGeer). The same.

Fig. 136. Epilampra azteca Saussure. The same.

Fig. 137. Epilampra grisea (De Geer). The same.

Fig. 138. Epilampra mexicana Saussure. The same. Fig. 139. Epilampra wheeleri Rehn and Hebard. Cuticular proventriculus.

Fig. 140. Laxta granicollis (Saussure). The same.

Fig. 141. Litopeltis biolleyi (Saussure). The same.

Fig. 142. Litopeltis bispinosa (Saussure). The same.

Fig. 143. Diploptera punctata (Eschscholtz). The same.

Fig. 144. Type I microspines from Diploptera punctata. Approximately 450 X. 54

Fig. 145. Achroblatta luteola (Blanchard). Cuticular proventriculus.

F ig. 146. Capucina patula (Walker). The same.

F ig. 147. Panchlora nivea (Linnaeus). The same.

F ig. 148. Proscratea complanata (Perty). The same.

F ig. 149. Geoscapheus robustus Tepper. The same.

F ig. 150. Macropanesthia rhinocerus Saussure. The same. 55

Fig. 151. Panesthia laevicollis Saussure. Cuticular proventriculus. upv--undifferentiated proventricular panel. Fig. 152. Panesthia sp. The same. upv--undifferentiated proventricular panel. Fig. 153. Panesthia sp. The same. upv - -undifferentiated proventricular panel.

Fig. 154. Gyna capucina Gerstaecker. The same.

Fig. 155. Pycnoscelus indicus (Fabricius). The same.

Fig. 156. Pycnoscelus surimanensis (Linnaeus). The same. Fig. 157. Gromphadorhina portentosa (Schaum). Cuticular proventriculus. tu--tubercle.

Fig.. 158. Henschoutedenia flexivitta (Walker). The same.

Fig. 159. Leucophaea maderae (Fabricius). The same.

Fig. 160. Nauphoeta cineria (Olivier). The same, pvp--prim ary valvular plica. DESCRIPTIONS AND DISCUSSION

Family Cryptocercidae

This family of wood-eating cockroaches is considered

(McKittrick and M ackarras, 1965) to be the most primitive of all

Blattarians, and as such will be the first group discussed.

Subfamily Cryptocercinae (Fig. 1)

The proventriculus of Cryptocercus punctulatus is charac­ terized by teeth having greatly reduced and flattened edges or blades, and sclerotized tubercles on all pulvilli and valvular plicae.

The teeth are quite flat, being as long (anterior to posterior) as those in Blattidae, but lack the massive sculpturing and sclerotiza- tion found in Blattidae. Five of the six teeth lack points, and single moderately sclerotized ridges are the only projections on the other­ wise flattened bases.

The development of interdental sclerotizations is not as extensive as that found in Blattidae, each interdental area having one secondary sclerotization and two interdentaries. Intercalates are completely absent. Both the secondary sclerotization and the interdentaries are fairly broad, being knobbed anteriorly, with a

57 58 moderately wide membranous area between them.

The pulvilli are oval in shape and not obviously supplied with microspines or microtrichiae, but possess two unevenly-shaped tu b e rc le s.

The valvular plicae also have no obvious microspines or microtrichiae, but possess a single tubercle each, located just posterior to the pulvilli.

The presence of sclerotized tubercles on the pulvilli and valvular plicae seems to be a definitive characteristic, as this condi­ tion was not observed in any other single specimen or group in the suborder Blattaria.

Family Blattidae

Members of this family are characterized by having large thick proventriculi equipped with six large, heavily-sculptured teeth and extensive development of secondary sclerotizations, interden­ taries, and intercalaries. In the subfamilies Blattinae and

Polyzosterinae, striking differences between teeth in the same proventriculus lend themselves to a numbering system (1-6) which aids in subfamily assignments. Interdental sclerotizations are pres­ ent in all subfamilies but vary in shape from subfamily to subfamily. 59

Subfamily Lamproblattinae (Fig. 2-4)

The teeth in this subfamily are quite different from those in

either the Blattinae or Polyzosterinae. They can be characterized by their overall club-shaped appearance and the sim ilar shape and position of all six teeth in the same proventriculus. This similarity

of adjacent teeth precludes the designation of tooth #1 (the con­

stricted tooth), and subsequent numbering of the remaining teeth.

Other than in the overall club-shaped appearance, tooth morphology does not lend itself to definitive characterization in this subfamily.

Interdental sclerotization in Lamproblattinae also differs from that in the other two subfamilies of Blattidae. Although second­

ary sclerotizations, interdentaries, and intercalaries are all present,

they are all much wider, with wider membranous areas between them.

In Lamproblatta romanii (Fig. 4) the posterior ends of the secondary

sclerotizations and interdentaries are heavily sclerotized.

Pulvilli are rectangular and shorter than the teeth, and ap­

pear dark, due to a dense covering of microtrichiae. Type II m icro-

spines were seen in only one of the specimens examined.

Valvular plicae follow the Polyzosterinae pattern very

closely, being dark anteriorly and not so dark posteriorly, due to

their extensive complement of microtrichiae. 60

Subfamily Blattinae (Fig. 5-17)

The teeth in this subfamily tend to be more generally blade -

like than in either Polyzosterinae or Lamproblattinae. All represent­

atives of Blattinae seem to possess one tooth which is notched or

constricted. Following the terminology of McKittrick (1964) and

M ackerras (1967) the author has labeled this tooth the constricted

tooth and arbitrarily assigned it the number "1". In only one speci­

men (Fig. 7) is tooth #1 difficult to assign. The remaining five teeth were assigned the numbers 2-6, proceeding clockwise from tooth #1.

Although in no specimens are all six teeth similar or identi­

cal (radial symmetry) m irror pairs do exist. In all specimens

observed, teeth two and six and teeth three and five form two pairs

of m irror opposites.

Although the presence of m irror pairs is also a character­

istic of the subfamily Polyzosterinae and to a minimal extent sub­

family Lamproblattinae, this condition does not exist in other

cockroaches, and therefore seems a reliable character.

Interdental sclerotizations in the subfamily Blattinae are

highly developed, with secondary sclerotizations, interdentaries,

and intercalaries being present in all specimens.

Several exceptions exist, but the predominant shape of

interdental sclerotizations is a very narrow one, in fact, a series 61 of narrow plates with little membranous area between them

(Figs. 9. 14-16). Two exceptions to this pattern occur in Henicotyle antillarum (Fig. 10) and Henicotyle n. sp. (Fig. 11), where the secondary sclerotizations and interdentaries are more spadelike and bulbous respectively.

Proventricular pulvilli in this subfamily are quite obvious and well delineated, being rectangular in shape and considerably shorter than the teeth. In addition, they are always obviously covered with a dense mat of microtrichiae, in many instances creating a dark, setulose appearance.

The valvular plicae also appear quite hairy, with micro- trichia present on the anterior one-third and type I microspines on the remainder.

Subfamily Polyzosterinae (Figs. 18-30)

Teeth in this subfamily vary to a greater extent within an individual proventriculus and from specimen to specimen than in the subfamily Blattinae.

In five of the Australian Polyzosterinae (Figs. 18, 26-29), tooth #1 is very conspicuously Y-shaped, while tooth #4 is distinctly oval or globular in shape.

Unfortunately, the remaining members of this subfamily are not as easily separated from those in Blattinae, but some less pronounced differences are apparent to the careful observer.

The notched bladelike tooth #1 that is regularly found in

Blattinae, was found in only one specimen of Polyzosterinae (Fig. 21)

The Polyzosterinae type of constricted tooth is more commonly con­ stricted laterally (Figs. 19, 23, 24) rather than from the leading edg

Tooth #1 in the remaining specimens was selected by eliminating the two m irror pairs and picking the most obviously constricted tooth from the remaining two (Figs. 20, 22, 30).

Teeth in this subfamily are generally less bladelike than those in Blattinae, not projecting nearly as far into the arm arial lumen, and generally being more rectangular (anterior to posterior) in shape.

As is the case in Blattinae, the interdental sclerotizations are highly developed, being spadelike (Figs. 18, 26-29) in the

Australian specimens and very narrow in the remaining members of the subfamily. Intercalates are more numerous in most members of this subfamily than in any other group, often lying in such close proximity to each other that counting them is quite difficult.

As in Blattinae, the pulvilli are quite well delineated, being rectangular in shape and much shorter than the teeth or the second­ ary sclerotizations. Microtrichiae are very numerous, marking this subfamily as having the darkest and most obviously setulose pulvilli in the suborder Blattaria. 63

The valvular plicae in Polyzosterinae are generally darker and more obviously setulose than in Blattinae. Often the anterior one-third of all plicae are as densely equipped with microtrichiae as the pulvilli, which is not the case in Blattinae. In addition, the intraplical folds are also densely supplied with microtrichiae, par­ ticularly the anterior one-third. M icrotrichiae on the posterior two- thirds of the plicae are obvious and uniform, but not dark.

Microspines in this region are commonly type II.

Family Polyphagidae

A single characteristic proventriculus which would be generally representative of all members of this family does not ex­ ist in the small sample available for this study.

Generally, but certainly not in all specimens, the proven- triculi found in this group can be typified as being membranous with little if any sclerotization. In several species, a total lack of tradi­ tional proventricular organization exists, with no j.nterdental sclerotizations, no delineated dental belt, pulvilli, or plicae.

Subfamily Polyphaginae (Figs. 31-36)

The teeth and interdental areas of Arenivaga cervarae and

Polyphaga aegyptica are quite sim ilar, being equipped with varying numbers of very large, blunt, and moderately sclerotized 64 microspines which are grouped at the anterior end of each plica and interdental area (Figs. 31, 36). No secondary sclerotizations, interdentaries or intercalaries are present.

The pulvilli of Arenivaga cervarae are more readily recognized than those of Polyphaga aegyptica and both possess a pul- villar and plical covering of scalelike microspines. Type I m icro­ spines are found in Arenivaga cervarae, and these are replaced by very fine microtrichiae in Polyphaga aegyptica.

Compsodes delicatulus (Fig. 32), Paralatindia azteca

(Fig. 35), Latindia dohrniana (Fig. 34), and Homoeogamia mexicana

(Fig. 33) were all placed in this subfamily by the author, a move motivated more by traditional taxonomic considerations rather than by proventricular similarities. They were placed in the closely related families Latindiidae and Homoegamiidae by Princis (1963) and not considered by McKittrick (1964). Realistically, they are suf­ ficiently different in proventricular appearance to warrant placing each of them in other groups. The first three exhibit tooth and inter­ dental areas that are very lightly sclerotized, quite flat, and beset with fine blunt microspines which are sim ilar to, but much smaller than those found in the dental areas of Arenivaga cervarae and

Polyphaga aegyptica. In addition, each tooth area is equipped with several long curved microtrichiae at its most anterior end. 65

With the exception of Paralatindia azteca and Latindia dohrniana, pulvilli and valvular plicae are not sharply delineated.

Pulvillar microtrichiae of Latindia dohrniana can just be classed as obvious, while the type II microspines of Paralatindia azteca are definitely not in the "obvious" category.

Homoeogamia mexicana has less organization than any other members of this subfamily, having a scattering of obvious type II microspines on the dental areas. The pulvilli are recognizable but not distinct, and their uniform covering of hairlike microtrichiae is not at all obvious.

In regard to interdental sclerotizations, all members of this subfamily save Homoeogamia mexicana possess broad, flat, and very lightly sclerotized secondary sclerotizations and interdentaries, though they are not sharply defined.

Subfamily Holocompsinae (FigB. 37, 38)

Both representatives of this subfamily are characterized by their total lack of organization. No recognizable dental belt, pul­ villar belt, or valvular plicae can be seen, as they are totally membranous with several scattered patches of microtrichiae throughout both. Family Blattellidae

This family is represented by the largest number of speci­ mens in the study (75), and as might be expected in terms of biological diversity, the many examples are quite sim ilar in some morphological aspects and quite dissim ilar in others.

Proventricular teeth are quite sim ilar within subfamilies, but vary between subfamilies.

Interdental sclerotizations (secondary sclerotizations, interdentaries, and intercalates) are present in four of the five sub­ families, and pulvillar morphology is quite similar throughout the fam ily.

Subfamily Anaplectinae (Figs. 39-42)

Teeth in this subfamily are quite sim ilar in overall shape, being rather square in form, as viewed from the side. They must be classed as sculptured, though they are not as heavily sclerotized, nor do they assume the characteristic club-shaped appearance found in the subfamily Lamproblattinae (Family Blattidae), the only group with which they could be confused. The tooth edges are uneven and

each tooth possesses several irregular, more heavily sclerotized

areas along the blades. In addition, three of the four specimens ex­

hibit antedental m icrotrichial tufts which overhang the anterior ends

of most teeth and secondary sclerotizations. 67

Within the same proventriculus, individual teeth are very sim ilar with no unique character on any_one tooth, and because of this do not lend themselves to numbering or pairing.

Interdental sclerotizations in Anaplectinae are broad and very distinct, with a secondary sclerotization plus a pair of inter­ dentaries being found in each interdental area. No intercalaries were present in the four specimens examined.

Pulvilli in this subfamily are fairly distinct, being some­ what broader than those in Blattidae, but still considerably shorter than the teeth. M icrotrichiae are usually quite obvious but not so dense as in Blattidae.

Very fine microtrichiae are present on the anterior one- third of all valvular plicae, with type I microspines rather evenly scattered over the more posterior portions.

Subfamily Ectobiinae (Figs. 43-46)

Teeth in this subfamily are so small, lightly sclerotized, and so poorly developed, that two of the four specimens (Figs. 43,

44) are classified as having tubercles rather than teeth. Ectobius sylvestris (Fig. 45) and Parellipsidion conjunctum (Fig. 46) ex­ hibit the most distinctive tooth form, with 6 very sim ilar narrow teeth whose tips are single, stout, curved microtrichiae. In both instances, however, the teeth extend over most of the dental belt 68

(anterior to posterior), a condition that does not exist in the two specimens possessing tubercles. Ectobius lapponicus (Fig. 43) and Ectobius pallidus (Fig. 44) show little dental sim ilarity within the same proventriculus, and their tubercles may be characterized as being little more than irregular collections of pointed micro - spines which give the tubercle edges a serrated appearance. In addition, the tubercles are considerably shorter than the interdental areas that flank them.

Interdental sclerotizations are completely absent in this sub­ family, but the membranous tissue in the interdental areas is organized into a regular pattern (Fig. 44, 46) common to proven- triculi with interdental sclerotizations present. In Parellipsidion conjunctum these patterned folds have their inner margins edged with tiny blunt microspines, and the folds are much longer than in other subfamilies, extending back from the crop to the posterior ends of the pulvilli.

The pulvilli of Ectobiinae are not sharply delineated but can be characterized by their long sac-like shape. Although the limits of the dental belt are rather obscure, it seems that pulvillar length closely approximates the length of the dental belt. Only Ectobius pallidus (Fig. 44) exhibits pulvillar microtrichiae, and even here they are not at all obvious. 69

Valvular plicae are obviously beset with microtrichiae in

Ectobius pallidus and Ectobius sylvestris but bear type II m icro­ spines in Parellipsidion conjunctum. M icrotrichiae and microspines are absent in Ectobius lapponicus.

Subfamily Nyctiborinae (Figs. 47-49)

The teeth of Nyctibora obscura (Fig. 48) and Nyctibora sp. (Fig. 49) are quite sim ilar in that both exhibit rather long posteriorly-curved, blunt-tipped shapes. Those of Megaloblatta blaberoides (Fig. 47) lack the long tapering development of the for­ mer two, but still curve posteriorly and exhibit blunt tips.

Within each proventriculus the teeth are quite similar, varying slightly only in size.

Interdental sclerotizations are present in all three speci­ mens, but are difficult to resolve in Nyctibora obscura and Nyctibora

sp. In the former, secondary sclerotizations, interdentaries and intercalaries are long, extremely narrow, and beset with large numbers of fine microtrichiae which contribute to a setulose ap­ pearance. In the latter, they are broader, but exhibit the same

setulose appearance. Megaloblatta blaberoides (Fig. 47) on the other

hand, possesses more heavily sclerotized interdental plates, which

are very obvious. The presence of microtrichiae is not nearly as

noticeable in this specimen as in the other two. 70

It is somewhat difficult to delineate the pulvillar limits in the two members of the genus Nyctibora. Dental belt and pulvillar belt seem to merge with no sharp distinction between them. The pulvilli appear to arise at the base of the teeth, and like those in the subfamily Ectobiinae at least equal the length of the teeth.

In Megaloblatta blaberoides the pulvilli are more readily seen, being somewhat shorter than those in the genus Nyctibora but still quite large and equal to the teeth in length.

Pulvillar microtrichiae are obvious in all representatives of this subfamily, although much more dense in Megaloblatta blaberoides.

Obvious microtrichiae are present on the valvular plicae of all but Nyctibora obscura, which is supplied with a covering of type II microspines.

Subfamily Blattellinae (Figs. 50-72)

The teeth in this subfamily are moderately sclerotized and very obvious. Tooth sim ilarities between specimens and within an individual proventriculus are very evident.

Intraproventricular comparisons show a high degree of

sim ilarity between all six teeth, often to the point where groupings

of two or four teeth might be called identical. However, the desig­

nation of tooth #1 would be an arbitrary maneuver since no 71 constriction (as referred to in Blattinae and Polyzosterinae) is pres­ ent. The absence of this central starting point (tooth #1) and the fact that in many instances the identical teeth are side by side, precludes numbering them.

Large microspines are frequently present on the fore and midblades of the teeth, varying in shape but in most cases so closely spaced that their bases are in contact with each other.

With only two exceptions, Xestoblatta buscki and Xestoblatta festae (Figs. 69, 70), the crop-dental belt line is not marked by the presence of antedental m icrotrichial tufts, a characteristic frequently found in members of the subfamily Plectopterinae.

Interdental sclerotizations in the subfamily Blattellinae vary from broad to narrow, but with secondary sclerotizations, inter­ dentaries, and intercalaries being found in all specimens.

Pulvilli in this subfamily are much shorter than the teeth, being somewhat rounded and generally very obvious. As might be expected in a large group of diverse specimens, pulvillar micro- trichiation varies throughout the subfamily, being obvious and dark in some instances and much less obvious in others.

As with the pulvilli, microtrichiae found on valvular plicae also vary in size and degree of sclerotization, being quite obvious in

some instances and barely detectable in others. Both microtrichiae 72 and type I microspines are found on the valvular plicae with the usual condition being characterized by more obvious investment on the anterior one-third.

In term s of overall proventricular morphology, no one character seems foolproof in separating members of Blattellinae from those of Plectopterinae. It is, rather, a combination of tooth shape and microspination, plus the absence of antedental micro- trichial tufts, that are most valuable.

Subfamily Plectopterinae (Figs. 73-113)

The 41 specimens representing this subfamily present con­ siderable diversity in proventricular form, particularly in regard to teeth. No one basic tooth shape typifies this large group. Tooth shapes can be loosely grouped into the following categories: (1) rec­ tangular or disc-shaped (Figs. 77, 78, 89, 97, 98, 103-105, 107-

109); (2) d ro p -sh a p e d (F ig s. 80, 96, 99); (3) sp in y -ed g ed (Fig. 113);

(4) clam-shaped (Fig. 106); and (5) hooked (Fig. 111). All specimens in the five groups above exhibit a high degree of sim ilarity between the six teeth. This is not true, however, in group six (Figs. 73-76,

79, 81-88, 91-95, 100-102, 110, 112) whose 22 specimens possess teeth that vary not only from specimen to specimen but also within a single proventriculus. 73

Generally, teeth in this subfamily are equipped with large,

blunt or pointed microspines on the mid and foreblades. Unlike the

relationship found in Blattellinae, where microspine bases were often

touching, microspines on the teeth of Plectopterinae are usually fur­

ther apart and appear more randomly distributed.

Overall, the teeth of Plectopterinae are generally much more

rectangular than those occurring in Blattellinae but the two sub­

families are not easily separated on the basis of tooth differences

alone.

The presence of antedental microtrichial tufts and secondary

sclerotizations is a characteristic shown by a majority of the speci­

mens in this subfamily (e. g. Figs. 12, 84, 88, 92), but a few (Fig.

91, 94, 95, 112) do not exhibit this condition.

Interdental sclerotizations in Plectopterinae are in most

cases secondary sclerotizations, interdentaries, and intercalaries,

though intercalaries are absent in a number of specimens. The

— -secondary sclerotizations and interdentaries are universally quite

broad, whereas the intercalaries, when present, are often quite

narrow. Only one specimen, Cahita nahua (Fig. 23) possessed no

interdental sclerotizations.

The pulvilli of Plectopterinae vary but slightly in length

and width, but are universally round and shorter than the teeth. 74

Pulvilli are commonly supplied with obvious microtrichiae, although in some they are completely absent. Type I and type II microspines are also present in some instances.

Microspination of the valvular plicae also varies from very obvious to non-existent. Where present, type I or type II predominate.

Family Blaberidae

The nine subfamilies making up the family Blaberidae repre­ sent a tremendous diversity in proventricular form. Teeth are pres­ ent in relatively few specimens, and both teeth and dental tubercles are absent in the subfamily Panchlorinae. Dental tubercles commonly occur elsewhere in Blaberidae, but very greatly in form and number.

Interdental sclerotizations are typically absent, and several very atypical pulvilli and plical arrangements are noted.

Subfamily Blaberinae (Figs. 114-127)

Sclerotizations of the dental areas in members of this sub­ family range from the six rather heavily sclerotized short, smooth teeth which characterize members of the genus Blaberus (Figs. 117 —

120) to the uneven, lightly sclerotized tubercles of Petasodes moufeti (Fig. 127). Galiblatta n. sp. could not be located in the lit­

erature, and was placed in the subfamily Blaberinae on the basis of 75 its proventricular sim ilarities with other members of the subfamily.

The teeth exhibited by members of the genus Blab.erus

(Figs. 117-120) are utilized as an arbitrary dividing line between teeth and tubercles. Any dental sclerotizations smaller than those in the genus Blaberus are classified as dental tubercles.

Most specimens of Blaberinae studied were consistent in having six teeth or six tubercles each, but within the same proven­ triculus the teeth or tubercles were frequently dissim ilar in shape.

Interdental sclerotizations are noticeably absent, the inter­ dental areas usually being composed of a single fold, and beset with fine microspines of type I or type II, which are not at all obvious.

The pulvilli of Blaberinae are characteristically obvious and noticeably longer than any of the teeth or tubercles anterior to them. M icrotrichiation is heavy and obvious in the genus Blaberus

(Figs. 117-120) but far less distinguishable in other specimens, where type I and type II micro spines predominate.

Valvular plicae are usually well-defined, with microspines of type II predominating, but with type I also represented in several in sta n c e s.

Subfamily Zetoborinae (Figs. 128, 129, 132)

Both teeth and tubercles are absent in the proventriculi of

Phortioeca phoraspoides (Fig. 129) an

Limits of the dental belt are obscure, but tubercle areas as well as the interdental areas exhibit a scaly covering of type III micro spine s.

Zetabora signaticollis (Fig. 132) possesses six small, rough-edged tubercles which have either blunt or multipointed ends.

Tubercle areas are roughly triangular in shape, and interdental areas are quite broad, flat, and marked by a small anterior- posterior pouch.

Pulvilli in this subfamily are universally longer than the tubercles or tubercle areas, though they vary in length. Those of

Zetabora signaticollis are the shortest and least conspicuous of the three. Those of Lauxoblatta sp. are noticeably longer than the for­ mer and more sack-shaped. The pulvilli of Phortioeca phoraspoides are the longest of the group, exceeding the valvular plicae in length.

Valvular plicae of Zetoborinae are conventionally organized and tend to be moderately broad.

Microspination in this subfamily is uniform but not obvious in any of the three specimens. With the exception of tubercle areas, type II microspines are the only ones found throughout the remain­ ing portions of the proventriculi.

Subfamily Epilamprinae (Figs. 133-142)

Teeth as they appear in Blattoidea do not exist in this sub­ family. Anteriorly, the tooth areas usually exhibit greatly reduced, 77 lightly-sclerotized tubercles, though even these tubercles are lacking in some specimens. When present, the tubercles characteristically appear as irregular multipointed projections, usually 6 in number, but 24 in Litopeltis biolleyi (Fig. 141), 5 in Litopeltis bispinosa

(Fig. 142) and only one in Audreia carinulata (Fig. 134).

Generally, dental tubercles in this subfamily have serrated rather than smooth edges, and are far less prominent than those ex­ hibited by tubercle-bearing members of Blaberinae.

Special mention must be made regarding the dental tuber­ cles of Litopeltis biolleyi (Fig. 141), since their number and a r­ rangement show no similarity to any other specimen or group of specimens. Each of the six dental areas is anteriorly modified into two separate membranous flaps and each bears a pair of small, lightly-sclerotized, sometimes multipointed tubercles. The 24 tubercles in this species label it as unique, since no more than nine are found elsewhere in any one proventriculus.

On the opposite end of the scale is Laxta granicollis

(Fig. 140) which lacks dental tubercles and therefore bears little

proventricular similarity to the other members of Epilamprinae. It

is also somewhat distinctive among the cockroaches examined in that

its dental areas are beset with concentrations of type III microspines

(Fig. 131), one of only three specimens so equipped. Princis 78

(1962-69 Pars 4) designates Laxtinae and Epilamprinae as quite separate subfamilies, while McKittrick (1964) lumps them as one.

Limits of the dental belt in Epilamprinae are difficult to define, but the areas surrounding the tubercle bases are covered with a scalelike layer of type I microspines.

Interdental sclerotizations are absent in this subfamily, the area usually assuming a rounded pouch-like appearance flanking each tubercle. These pouches, like the tubercle bases, are beset with a fine scalelike covering of type I microspines. Litopeltis bispinosa exhibits a greater interdental organization than is found throughout the remaining representatives of the subfamily in having three rounded pouches between each pair of tubercles and three greatly expanded interdental areas.

Pulvilli are also, in most instances, difficult to delineate.

They extend posteriorly from the tubercles to the more conventionally organized valvular plicae, and are considerably longer than the tuber­ cles themselves. They are often covered with a uniform scattering of type I or type II microspines.

Five members of the genus Epilampra (Figs. 135-139) plus

Audreia gatunae (not illustrated) possess four sclerotized rays in the folds between the valvular plicae. These rays are unique for

Epilampra and Audreia at least, and seem to be a much more reliable characteristic than tubercle number or morphology.

The remaining members of the subfamily Epilamprinae un­ fortunately exhibit nothing unusual in the organization of their valvular

plicae. Individual panels can easily be discerned and they exhibit a

rather even dispersal of non-obvious type I or type II microspines, with type II in the majority of specimens.

Subfamily Diplopterinae (Fig. 143)

Proventricular teeth as such are absent in the one specimen

representing this subfamily. Dental tubercles are present, having

the same jagged, serrated edges, but being somewhat broader than

those found in Epilamprinae.

Interdental sclerotizations are absent, with interdental areas

being broad and not noticeably pouch-shaped.

As in Epilamprinae, pulvilli are not well defined, but are

definitely longer than the tubercles. Microspination is just obvious

and is chiefly of type II microspines.

Valvular plicae are well defined and quite conventional save

for the presence of one exceptionally wide plica.

Subfamily Panchlorinae (Figs. 145-148)

With one exception, proventriculi of this subfamily have

neither teeth nor dental tubercles, and bear little if any resemblance 80 to each other. The proventriculi of Achroblatta luteola (Fig. 145),

Papucina patula (Fig. 84), and Panchlora nivea (Fig. 147), are totally membranous in composition, while the proventriculus of

Proscratea complanata (Fig. 148) has dental tubercles sim ilar to those found in the subfamily Diplopterinae.

Limits of the dental belt are quite obscure in all repre­ sentatives, but both the dental and interdental areas are well supplied with microspines of type I or II. M icrotrichiae are also present.

Microspination is not obvious except in Achroblatta luteola (Fig. 145) where it appears moderately dense.

Pulvilli in this subfamily are also not well delineated, but are somewhat rectangular in Panchlora nivea (Fig. 147), but much longer (anterior to posterior) in the three remaining specimens. In all cases, were tubercles present, the pulvilli would be considerably lo n g er.

Pulvillar microtrichiation is obvious only in Achroblatta luteola (Fig. 145). When present, type II microspines are most com­ mon, but two of the specimens exhibit no microspination whatsoever.

Valvular plicae are fairly well defined in all specimens ex­ cept Achroblatta luteola (Fig. 145). Microspines generally are absent except for widely scattered type II in Capucina patula. 81

It must be admitted that members of this subfamily evidence little proventricular morphology in common. Placement was in­ fluenced to a greater degree by current taxonomic literature than by proventricular similarities.

Subfamily Panesthiinae (Figs. 149-152)

Teeth are absent in the proventriculi of this subfamily, in­ stead, tiny sclerotized tubercles which vary in form and number from specimen to specimen, are found. Limits of the dental belt are ob­ scure and individual tubercles appear scaly under high magnifications.

Members of the genus Panesthia (Figs. 151-153) show a great deal of similarity in their dental areas, as all three possess seven definite, usually rough and scaly-edged tubercles, a situation which is unique to the subfamily Panesthiinae.

The remaining specimens, Geoscapheus robustus (Fig, 149) and Macropanesthia rhinocerus (Fig. 150), seem to be somewhat dif­ ferent from those in Panesthia in that tubercles are fewer in number and are generally less obvious. Geoscapheus robustus (Fig. 149) possesses only three tubercles which are definite projections; the remaining sclerotizations exist only as scaly patches. Macropanes- thia rhinocerus is less completely developed than the former. No projecting tubercles are evident, and only two scaly sclerotized patches exist over the entire dental area. The scaly appearance of 82 tubercles, tubercle bases, and some interdental areas is caused by- concentrations of type I and III microspines.

A proventricular character which is apparently unique to both Panesthiinae and Perisphaerinae, is the existence of a wide un­

organized membranous panel or plica extending the total length of

the proventriculus. McKittrick (1964) labels plica number one in

some of her figures, but this is predominately a feature of the valvu­

lar portion of the proventriculus.

In Panesthiinae, this wide panel accounts for approximately

one quarter of the total diameter of the proventriculus, does not bear

tubercles or other sclerotization anteriorly, and is not equipped with

pulvilli. The function of this panel is not known at this time, but it

remains unique to the two subfamilies mentioned above.

Only one member of the subfamily, Geoscapheus robustus,

possesses sclerotization of any kind in the interdental areas, where

fine microspines appear in several interdental areas.

Limits of the pulvillar and dental belts in Panesthiinae can

not be defined anteriorly or posteriorly, appearing as continuous

panels from the crop, to and continuous with the valvular plicae.

The pulvillar area of Macropanesthia rhinocerus (Fig. 150) is less

organized than that in the other three specimens, having a rather

variable (in width) series of folds with a general anterior to posterior

organization. 83

Microspination of the pulvillar and valvular plical areas

is a non-obvious type I in the genus Panesthia, and type II in the two

remaining species.

Subfamily Perisphaerinae (Fig. 151)

As in all but one subfamily of the family Blaberidae, the

single representatave of Perisphaerinae examined lacks teeth. It is

unique, however, in its possession of nine tubercles, the second

highest number observed in the entire study.

Four of the nine tubercles are quite small, very lightly

sclerotized, and appear to be little more than concentrations of

type I and II microspines. The five remaining tubercles exhibit more

form, being somewhat triangular in shape.

As in Panesthiinae, Gyna capucina (Fig. 154) also possesses

a wide unorganized panel extending the length of the proventriculus.

In this case the panel does exhibit obvious microspination (primarily

type II) which the panels in Panesthiinae do not.

The limits of the dental belt and interdental areas are ob­

scure, but interdental areas are liberally supplied with the same

type of microspines which invest the dental areas.

Pulvilli in Gyna capucina are considerably longer than the

dental tubercles, being rather bulbous posteriorly. M icrotrichia-

tion is just apparent, but not heavy, and consists of a uniform

O 84 scattering of type I and II microspines.

The valvular plicae are not well delineated, and their in­ vestment with scattered type II microspines is much less obvious than on the pulvilli.

Subfamily Pycnoscelinae (Figs. 116, 117)

Teeth are absent in both representatives of this subfamily, with six very small, smooth-tipped tubercles in each proventriculus.

Unlike other tubercles in the family Blaberidae, their tips are quite pointed, and are more heavily sclerotized than their bases, creating the impression that most of the tubercles are capped (see definition, p. 18).

The interdental areas appear primarily as rather wide pouches extending posteriorly from the crop to a point midway along the length of the pulvilli.

The pulvilli can be readily seen and are much longer than the tubercles, though not as bulbous posteriorly.

Valvular plicae are conventionally organized, though they appear wider than in other subfamilies of Blaberidae.

Microspination throughout the proventriculus is uniform but not obvious, being entirely of type II microspines on dental areas, pulvilli, and valvular plicae. Subfamily Oxyhaloinae (Figs. 157-160)

Teeth are absent in this subfamily, though the tooth area is equipped with very small unevenly shaped tubercles. Unlike those found in Diplopterinae and Epilamprinae, the individual tubercles ap­ pear to be composed of concentrations of microspines and therefore have uneven edges and a scaly appearance under high magnifications.

In three of the four specimens, type I microspines make up the dental tubercles, while type II are found in the fourth. Tubercle number is consistently six in all specimens examined.

Interdental sclerotizations are absent in this subfamily, with a short rectangular pouch present in two instances and no pouches or organization present in the other specimens.

In Nauphoeta cinerea (Fig. 160) and Leucophaea maderae

(Fig. 159) microtrichiae are present in the interdental areas, but are clustered and scalelike rather than uniformly distributed as in

Gromphadorhina portentosa (Fig. 157), and Henschoutedenia flexivitta (Fig. 158).

Pulvilli are fairly well delineated in this subfamily, and are all longer than the dental tubercles. Microspination is just obvious in two specimens and not at all obvious in the other two. Type II predominate, in all cases. Valvular plicae are reasonably well delineated, and m icro­ spination is identical with the pulvilli. SUMMARY

The construction of a key to the subfamilies of cockroaches on the basis of proventricular characters alone, presented a problem common to any key-construction based on a single set of characters.

That problem involved the considerable morphological diversity pres­ ent between the genera possessing tubercles or no sclerotizations of the dental belt, and resulted in less definitive placement of several examples into subfamilies, than the author desired. It was felt, however, that these differences were more a function of inherent biological diversity between genera, than the number of specimens involved.

On the one hand, of the 162 species examined in this study, approximately 108 separate easily and distinctly into nine of the sub­ families (Cryptocercinae, Blattinae, Lamproblattinae, Polyzoster- inae, Blattellinae, Plectopterinae, Anaplectinae, Ectobiinae, and

Nyctiborinae) listed by McKittrick (1964). These 108 specimens have in common the possession of distinctive tooth morphology and this plus the presence or absence of antedental microtrichial tufts combine to facilitate classification.

86 87

On the other hand, the morphological diversity exhibited by the remaining 54 species (11 subfamilies; Polyphaginae, Holocomp- sinae, Zetoborinae, Blaberinae, Panesthiinae, Pycnoscelinae,

Diplopterinae, Panchlorinae, Oxyhaloinae, Epilamprinae,

Perisphaerinae) often defied the most dedicated efforts to group them into subfamilies. Tubercle number and morphology, microspine type, undifferentiated panels, and sclerotized rays in the valvular plicae provide excellent key characters, but were not always present in all members of a particular subfamily.

Based strictly on proventricular morphology, the species in this study generally seemed to group more naturally under the con­ cise taxonomic system of McKittrick (1964) than under the more ex­ tensive listings of Princis (1963), a system which involved 28 families as compared with the 5 of McKittrick. Little correlation, however, was found between proventricular morphology and the "complexes"

(a category lying between family and subfamily, Blaberoid,

Panchloroid, Epilamproid) proposed by McKittrick.

Overall, it is felt that the use of proventricular morphology

can be of significant value to the cockroach taxonomist (particularly

since identification is not dependent upon the age or sex of the cock­

roach) when utilized independently, or more effectively still, when

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