The Male Genitalia of Blattaria. IV. Blaberidae Blaberinae

The Male Genitalia of Blattaria. IV. Blaberidae Blaberinae to In this *Manuscript received by the editor Zlpril 8, 1970. 197o] Roth Blattaria to the subfamily Blaberinae. The genera Hyporhicnoda, Phoetalia, and Oxycercus, which were placed in subfamilies other than the Blaberinae by McKittrick and Princis, have male genitalia characteristics of members of Blaberinae and I include them in this subfamily. MATERIALS AND METHODS The technique for preparing the male genitalia is given in Roth (969). Unless otherwise indicated in the explanation of figures, the : phallomeres L2d (Fig. 41) and L (Fig. 43) were mounted dorsal side up and the genital hook Ra (Fig. 42) is oriented ventral side uppermost. The source of each of the specimens illustrated is given using the following abbreviations: (AMNH) American Museum of Natural History, New York; (ANSP) Academy of Natural Sciences, Philadelphia; (BMNH) British Museum (Natural History), London (CUZM) Copenhagen University, Zoological Museum, Denmark; (L) Zoological Institute, Lund, Sweden; (LEM) Lyman Entomological Museum, Macdonald College, Quebec, Canada; (N) U. S. Army N atick Labs., N atick, Mass.; (USNM) United States National Museum, Washington, D. C. Geographical collection data and the names of specialists who identified the specimens., if known, :ollow these abbreviations. The number preceding the abbreviations refers to the number assigned the specimen and its corresponding genitalia (on a slide) which are deposited in their respective museums. RESULTS AND DISCUSSION The male phallomeres which have proved useful in studies of Blaberidae are a retractable hook on the right side (R:z) (Fig. 42) a median sclerite (L:vm) which may or may not be solidly attached to a sclerite L:zd (L2 dorsal); and lying below or partly surrounding L:d is a membrane, the prepuce, which almost always bears spines of various sizes and shapes (Fig. 4I). The phallomere (LI) on the left side, has a heavily sclerotized cleft (Fig. 43) and a well defined membranous lobe which overlaps the cleft area; because of its transparency this membrane is difficult to .see in mo3t of the photographs. Rehn and Hebard (I9:Z7, pp. :57-:58), in discussing the Blaberinae, concluded that there are "... at least three lines of development or phyla, which are as follows: I. Archimandrita, Blaberus, Eublaberus 3 IO Psyche [September Blaltica, Byrsotria, Hemiblabera, dsliduchus, Monachoda 3. Monastria, Petasodes The first group is well defined, although relationship with Byrsotria is evident The second group is well defined rom the first, but whether Monachoda should be considered the maximum differentiation o the second group, or a less highly modified member of the third one, is a matter o opinion." The above linear arrangement was the one preferred by Rehn and Hebard. Princis’ (1963) arrangement o some o the genera in the Blaberinae follows Rehn and Hebard, but several additional genera were included. Brachycolinae" Brachycola, Hormetica, Parahormetica, Sibylloblatta, Bionoblatta Blaberinae: ,/lrchimandrita, Blaberus, Mesoblaberus, Eublaberus, Blaptica, Byrsotria, Hemiblabera, ./lsliduchus, Monachoda, Monastria, Minablatta, Petasodes, Cacoblatta, ,Hiereoblatta, Stylhon McKittrick (1964) studied genera o Blaberinae and "or convenience" arranged them in the ollowing tribes: I. Blaberini: glrchimandrita, Blaberus, Blatica 2. Brachycolini: Brachycola, Hormetica, Parahormetica, Petasogte$ 3. Byrsotriini: Byrsotria, Eublaberus, Hemiblabera 4- 1Vionastriini" Monastria Male genitalic characters support the separation o( genera Blaberinae into 3 (Blaberini, Monastriini, and Brachycolini) rather than 4 tribes; and the genta.lia o Petasodes indicate that it does not belong in the Brachycolini as suggested by .-cKittrick. The present study tends to support Rehn and Hebard’s arrangement, especially with regard to the placement o Petasodes. I. Blaberini: ,4rchimandrita, ts#iduchus, Blaberus, Bla#tica, Byrsotria, Eublaberus, Hemiblabera, HyIorhicnoda, Minablatt I-I 8ty[hon (Figs. 17). Figs. 1-9. 1. (11 ANSP)../lrchimandrita marmorata (Stoll). San Mareos, Niearagua. 2. (61 USNM). ./lrch’imandrita tessellata Rehn. San Jos6, Costa Riea (det. Rehn). 3. (18 USNM). Blaberus #iyanteus (L.). St. Jean, French Guiana (det. Roth.). 4. (5 USNM). Blaberus craniifer Burmeister. Key West, Florida (det. Roth). 5. (N). Blaberus discoidalis Serville. Panama (det. Roth). 6. (516 L). Blaberus boli,iensis Prineis. Guayaquil, Ecuador (det. Princis). 7. (514 L). Blaberus anisitsi Branesik (det. Prineis). 8. (N). Blaberus atrolos (Stoll). Trinidad (det. Roth). 9. (N). Blaberus lmrabolicus Walker. Puraquequara, Rio Negro, Amazonas, Brazil (det. Roth). (stale 10 ram). 1970] Roth Blattaria 3II Psyche [September (19 ANSP). Blaberus brasilianus Saussure. Natal, Brazil. (23 ANSP). Blaberus scutatus Saussure (paratype of B. scutata var. obscura S. and Z.). 12. (8 ANSP). Blaberus minor Saussure. Argentina (det. Hebard). 13. (14 ANSP). Blaberus fusiformis Walker. Santa Cruz de la Sierra, Bolivia (det. Hebard). 14. (32 ANSP). Blaberus colosseus (Illiger). Fyzabad, Trinidad (det. Roth). 15. (N). Eublaberus posticus (Eriehson). Panama (det. Roth). 16. (N). Eublaberus distanti (Kirby). Trinidad (det. Roth). (scale 10 mm). Figs. 10-16. To a considerable extent the male genitalia of the genera placed in the Blaberini, show distinct trends (as do the female genitalia and proventricular characters used by McKittrick). It is difficult to separate them into the 2 tribes Blaberini and Byrsotriini which McKittrick used and I have combined them into the Blaberini. However, certain of the genera show closer relationships to one another than to others and therefore I have arranged them in generic groups. In the Blaberini, L2d is solidly joined to L2vm, and may or 1.970] Roth Blattaria may not be hookshaped. The prepuce is a flexible membrane bearing characteristic spines. When flattened, the margins of the preputial membrane often extend beyond the G.utline of Lad and the preputial spines are readily visible. The genera may be arranged as follows: Generic Group I. A rchimandrita (Figs.. I, 2), Blaberus (Figs. 4-I4), Blaptica (Figs. 26, 27, 29). I have reported on the genus Blaber.us in a recent paper (Roth, 969) and for this reason have illustrated R2 (Fig. 42) and L (Fig. 43) for only one of the species. However, for the sake of completeness I have included the L2d for the known species (Figs. 4, 44-55); these structures allow the separation of 3 species groups of Blaberus Giganteus Group" L2d is recurved dorsally and slightly to the right, often forming a hooklike structure. Extending dorsally and laterally on the left, about where L2vm and L2d are joined, is a tumorlike outgrowth. Spines on the preputial membrane are relatively small. (Blaberus giganteus, Figs. 3, 41-43; B. craniifer, Figs. 4, 44)Atropos Group: L2d is not hookshaped and the tumorlike outgrowth on the left side is absent. Truncate or rounded elevations are usually present only on the left side and are generally much larger and more robust than spines on the right. The preputial spines are often arranged in a single row on the left, and a single or sometimes double or partially double row on the right; spines on the right usually more numerous than on the left. (Blaberus discoidalis, Figs. 5, 47; B. boliviensis, Figs. 6, 49; B. anisitsi, Figs. 7, 48; B. atropos, Figs. 8, 45; B. parabolicus, Figs. 9, 46). Brasilianus Group" L2d not hookshaped and tumorlike outgrowth absent as in the Atropos Group. Anterior elevations not greatly enlarged on the left side and about the same size as those on the right. Preputial spines numerous, usually present on the left and right .sides and often occur in more than a single row. (Blaberus brasilian.us, Figs. O, 50; B. scutatus, Figs. I, 52; B. minor, Figs. :, 5 B. [usi[ormis, Figs. I3, 53; B. colosseus, Figs. I4, 54-55). There is considerable intraspecific variation in the male genitalia of Blaberus spp. often making specific determination of certain-species difficult. However, the species group separations are reasonably clear cut. In Zlrchimandrita spp. (Figs. 3:, 35, 38), L2d is not hookshaped, but otherwise this sclerite and the associated preputial fringe of spines are closest to the Gig’anteus Group of Blaherus. Psyche [September In the genus Blaltica the preputial spines are small and occur in multiple rows in obscura (Fig. 56) and dubia (Fig. 6:) but are reduced in number and arranged in a single row in interior (Fig. 59). According to Hepper (I965), Blaptica l)ereyrai Hepper is near B. dubia, but Hepper noted that the preputial spines of lereyrai are fewer in number than in dubia. The number of spines shown in Hepper’s (I965) figure I closely approximate those found in B. interior. The number of preputial spines in species of Blaberus is variable (Roth, I969) and this is probably also true for the genus Blatica. Rehn (I93O, p. 6:) stated that Styhon "... is a distinctive genus occupying an isolated position and not closely related to any of the other previously known genera.." The genus is found in the Dutch Leeward Islands and Rehn (I93O, p. 67) further stated "It is most unexpected to find on these semiarid islands of-[ the north coast of South America an entirely unrecognized genus which has no known close relative on the adjacent.mainland. The genus Parahormetca, which is in some respects the nearest relative, is an inha.bitant of regions widely removed in South America." The L:d and prepuce of Styl)hon (Fig. 64) indicate a closer relationship to Blaptica (Fig. 59) than to Parahormetica (Figs. I99, :oz, :o3) the latter is a member of the B rachycolini (see below). Generic Group :. dspiduchus (Figs. 21-23), Byrsotria (Figs. :4, :5), Eublaberus (Figs. I5-I6), Hemiblabera (Figs. 17-2o). In this group Ld is strongly hookshaped. The genera dsl)iduchus (Figs. 97, IOO, IO3, IO6), Byrsotria (Figs. 75, 78, 81), and Hemiblabera (Figs. 84, 87, 9o, 93, 96) all have very similar L:d’s and the preputial spines are somewhat elongated and arranged principally in a single row. Bonfils’ (1969, p. I32) drawing of the L:d and Figs. 17-25. 17. (24 ANSP). He’mibIabera granulata Saussure. Antigua, West Indies. 18. (50 USNM). Hemiblabera sp. probably tenebricosa Rehn and Hebaid. Rojo Jeeo, Mr. Samana, San Domingo (det. Gurney). 19. (26 ANSP). Herniblabera brunneri (Saussure). Puerto Rico (det. Rehn). 20. (16 MCZ). Hemiblabera labulabor Rehn and Hebard. Andros Island, Bahamas (det. Gurney). 21. (16 ANSP). Asliduchus caerncola J. W. H. Rehn. Paratype. Barrio Monte, Cabo Ro.jo, Puerto Rieo. 22. (15 ANSP). A sliduchus borinquen J. \V. H. Rehn. Paratype. Corozal, Puerto Rieo. 23. (29 AMNH). Aspiduchus borinquen. Corozal, Puerto Rieo. (det. by J. A. G. Rehn as ,4. deplanatus; J. W. H. Rehn (1951) synonimized Rehn and Hebard’s deldanatus with borinquen. 24. (25, ANSP). Byrsotria cabrerM Rehn and Hebard. Matanzas Provlnee, Cuba (det. Rehn). 25. (N.) Byrsotria ]umig.ata (Gu6rin). (det. Roth). (stale ,--" 5 mm) Roth Blattaria Psyche [September Figs. 26-31. 26. (20 ANSP). Blaptica interior Hebard. Formos.a, Argentina. (det. Rehn). 27. (22 ANSP). Blaptica obscura Saussure and Zehntner. So Leopoldo, Rio Grande do Sul, Brazil (det. Rehn.) 28. (64 USNM). Minablatta bipustulata (Thunberg). Vi(.osa, Minas Geraes, Brazil (det. Rehn). 29. (21 ANSP). Blaptica dubia (Serville). Buenos Aires, Argentina (det. Hebard). 30. (N). Hyporhicnoda litomorpha Hebard. Panama. (det. Gurney). 31. (6 ANSP). Styphon bakeri Rehn. Paratype, Curaqao, Dutch West Indies. (scale mm). 1970] Roth Blattaria Figs. 32-40. 32-34. (62 USNM). Xrc.himandrita tessellata. Canal Zone, Panama (det. Hebard). 35-37. (61 USNM). X. tessellata. (from specimen shown in Fig. 2), 38-40. (11 ANSP). drchimandrita rnarmorata. (from specimen shown in Fig. 1). (scale 0.3 ram). prepuce of Hemiblabera tristis Bontils shows these structures similar to other members of the genus. to, be Rehn and Hebard (I927, p. 268) stated that Hemiblabera a position intermediate between Byrsotria and dspiduchus The species brunne’ri has a distinct tendency toward Byrsotria, xvhile conversely H. pabulator and granulata exhibit more of an approach toward dspiduchus.’" The male genitalia are basically too similar to support Rehn and Hebard’s ’... clearly occupies Psyche [-September Figs. 41-49. 41-43. Blaberus giganteus. 41. (1 AMNH). Caripito, Venezuela. (L2d dorsal left phallomere; L2vm ventromedial left sclerite; P prepuce.) 42-43. (27 ANSP). Muzo, Colombia. (det. Roth). (SI Subapical incision of hooked right phallomere R2; C--- Cleft of left phallomere L1). 44. (46 USNM). Blaberus craniifer. Cordoba, Mexico. (det. Roth). 45. (34 USNM). Blaberus atropos. Charleston Quarantine (? Colombia) (det. Roth). 46. (6 AMNH). Blaberus parabolicus. San Martin, Peru (det. Roth). 47. (N). Blaberus discoidalis. Panama (det. Roth). 48. (514 L). Blaberus anisitsi. (from specimen shown in Fig. 7). 49. (516 L). Blaberus bolivicnsis. (from specimen shown in Fig. 6). (scale mm). 1970] Roth Blattaria Figs. 50-55. 50. (19 ANSP). Blaberus brasilianus. (from specimen shown in Fig. 10). 51. (62 ANSP). Blaberus minor. Paraguay. 52. (23 ANSP). Blaberus scutatus. (from specimen shown in Fig. 11). 53. (56 ANSP). Blaberus ]usiformis. Provineia Sara, Dept. Vera Cruz, Bolivia. 54-55. Blaberus colosseus. 54. (1 USNM). Trinidad (det. Roth). 55. (30 ANSP). Caparo, Trinidad (det. Hebard). (scale 0.3 ram). Psyche [September Figs. 56-65. 56-58. (22 ANSP). Blaptica obscura. (from specimen shown in Fig. 27). 59-61. (20 ANSP). Bl’aptica interior. (from specimen shown in Fig. 26). 62-63. (21 ANSP). BlaItica dubia. (from specimen shown in Fig. 29). 64-65. (6 ANSP). Styphon bakeri. (from specimen shown in Fig. 31). (Fig. 65 is a ventral view.) (scale-- 0.3 mm). 1970] Roth Blattaria Figs. 66-74. 66-68. (19 MCZ). Eublaberus losticus. (Type 16074 of Blabera ]emorata Seudder). Napo or Marafi6n. 69-71. (27 AMNH). Eublaberus distanti. Moyobamba, San Martin, Peru. (det. Roth). 72. (N). Eublaberus distanti. Trinidad. (det. Roth). 73. (N.) Eublaberus losticus. Panama. (det. Roth). 74. (4 BMNH). Eublaberus sp. Para, Brazil. (det. 0,3 mm). Roth). (scale Psyche [September Figs. 75-83. 75-77. (N). Byrsotria fumi.qata (det. Roth). 78-80. (25 ANSP). Byrsotria cabrerai. (from specimen shown in Fig. 24). 81-83. (15 MCZ). B. cal, rerai. Coast below Pico Turquino, Cuba. (det. Gurney). 0.3 mm). (scale 1970] Roth Blattaria Figs. 84-92. 84-86. (13 ANSP). Hemiblabera pabulator. Paratype. Nassau, Bahamas. 87-89. (16 MCZ). H. pabulator. (from specimen shown in Fig. 20). 90-92. (24 ANSP). Hemiblabera granulator. (from specimen shown in Fig. 17). (scale-- 0.3 mm). Psyche [September Figs. 93-102. 93-95. (26 .A_NSP). Hemiblabera brunneri. (rom specimen shown in Fig. 19). 96. (50 USNM). Hemiblabera sp. probably tenebricosa. (from specimen shown in Fig. 18). 97-99. (15 ANSP). lsliduchus borinquen. (from specimen shown in Fig. 22). 100-102. (16 ANSP). lsliduchus ca,ernicola. (from specimen shown in Fig. 21). (scale ram). 1970] Roth Blattaria Figs. I03-111. 10:t-108. 4stiduchus borinquen. 103-105. (30 AMNH). Corozal Cave, Puerto Rico. (misidentified by Rehn and Hebard as detlanatus). 106-108. (29 AMNH). (from specimen shown in Fig. 23). 109-111. (64 USNM). AIinablatta bibustulata. (from specimen shown in Fig. 28). (scale 0.3 ram). Psyche [September Figs. 112-117. 112-114. (177 USNM). Hyorhicnoda reflexa (Saussure and Zehntner). Turrialba, Costa Rica (det. Gurney). 115-117. Hyorh!cnoda litomorlha Hebard. Panama (det. Gurney). (scale 0.3 mm). 1970] Roth Blattaria suggestion about the relationship of species of Hemiblabera to Byrsotria and Aspiduchus. Rehn and Hebard I927, p. 278) in erecting the genus Aspiduchus stated that it "... is apparently related on one hand to Hemiblabera Saussure, and on the other to Monachoda Burmeister." The male genitalia of Aspiduchus (Figs. 21-23) indicate a close relationship to Hemiblabera (Figs. 17-2o) but less so to Monachoda (Figs. I42, 144, 145) which I place in the Monastriini (see below). Rehn’s (i95I p. 7) drawings of the Lad’s of Aspiduch.us cavernicola (Fig. io’o) and A. borinquen (Figs. 97, o3, IO6) show relatively small differences. It is doubtful that the male genitalia can be used to distinguish between these 2 species. In Eublaberus spp. the preputial spines occur in more than a single row (Figs. 66, 69, 72-74). These spines are more slender and less sclerotized in E. distanti (Figs. 69, 72) than in E. posticus (Figs. 66, 73), as noted by Princis (I950). Hebard (1920 p. 116) states that Eublaberus is closely related to Blaberus. However L2d is more distinctly hookshaped in Eublaberus than it is in most species of Blaberus and for this reason I place it in the second generic group. Generic Group 3. Hyporhicnoda (Fig. 3o), Minablatta (Fig. 8). In this group Ld is not hookshaped. Rehn (94o) stated that Minablatta is related to Blaptica and Eublaberus but i.s probably more closely related to the former. The absence of a hookshaped Ld in Minablatta (Fig. o9) places the genus closer to Blatica. However, the elongated preputial spines, occurring in a single row in Minablatta (Fig. o9) show a similarity to the spines found in .dspiduckus (Fig. o6), Byrsotria (Fig. 75), and Hemiblabera (Fig. 96). Hyporkicnoda has been placed in the Epilamprinae by Hebard (9o), Princis (96o), and McKittrick (964). However, the prepuce of the male has a fringe of spines (Figs. 5) which is characteristic of the Blaberinae. Spines are not found on the prepuce of any genera belonging to the Epilamprinae; nor have I seen these spines in males other than Blaberinae, representing more than 90 genera of Blaberidae. In H.porkicnoda the apex of Lvm is more darkly pigmented than Led so that the junction of the 2 sclerites is readily discernible. Ionastriini: Hiereoblatta (Fig. 6), _84onackoda (Figs. I23-I2_), Monastria (Fi:s. I2I-I22). Petasodes (Figs. II8-I20). Psyche [September In this tribe the apex of L2vm is solidly joined to a relatively broad L2d. In Monastria (Figs. 127-138), the prepuce (Figs. 127, I3O, 133, 36, 138) is almost entirely covered by rows of small, closely packed spines and extends we’ll below the posterior margin of L2d. In Monachoda (Figs. 39-I 5 O), Hiereoblatta (Figs. 15 I53), and Petasodes (Figs. 154-162) the prepuce extends little or not at all beyond the margin of Lad and many of the preputial spines (considerably fewer in number than in Mon.astria) are usually hidden under L2d. In Petas’odes reflexa (Fig. 16o) there is a marked reduction in number of preputial spines and these are very lightly sclerotized. McKittrick (1964) placed Petasodes in the Brachycolini, but all the members of this tribe have L2d clearly separated from L2vm by a membrane (see below). Rehn’s (1937, p. 24) statement that Hiereoblatta is related to Monastria is justified by the similarities in male genitalia. However, the shape of L2d and the preputial spines hidden under it in Hiereoblatta (Figs. 148 151 indicate a closer relationship to Monachoda (Figs. 42, 44, 45) than to Monastria (Figs. 127, 13o, 133, 136, 138). Brachycolini" Bionoblatta, Brachycola, Hormetica, Oxycercus, Parahormetica, Phoetalia, Sibylloblatta (Figs. I63-II). This tribe is clearly differentiated t?rom the Blaberini and Monastriini in that L2d is not solidly attached to L2vm, but is separated from it clearly by a membrane. The spines on the prepuce vary in size from relatively large in Phoetalia (Figs.. 175, 178) to small in Parahormetica (Figs. 199, 202, 2o3), Oxycercrus (Fig. 2o.6), Bionoblatta (Fig. 2o,9), Sibylloblatta (Figs. 95, I98), and Brachycola (Fig. I93). Princis (I96O) placed Phoetalia in the Epilamprinae and McKittrick (I964) assigned it (as Leurolestes) to the Diplopterinae. The male genitalia of Phoetalia are not characteristic of either of these Figs. 118-126. 118. (17 ANSP). Pe:tasodes dominicana (Burmeist.er). Rio de Janeiro, Brazil (det. Hebard). 119. (7 ANSP). Petasodes mou[eti (Kirby). So Paulo, Brazil. (det. Hebard). 120. (18 ANSP). Petasodes reflexa (Thunberg). 46 m. south of Bahia, Brazil. (det. Rehn). 121. (10 ANSP). Monastria biguttata (Thunberg). Rio de Janeiro, Brazil (det. H.ebard). 122. (963 L). lllonastria similis (Serville). Misiones, Argentina (det. Princis). 123. (42 CUZM). Monachoda latissima Brunner. Bahia, Brazil. 124. (6 MCZ). Monachoda burmeisteri Saussure. 125. (21 BMNH). JlConachoda 7rossa (Thunberg). 126. (964 L). Hiereoblatta cassidea 10 mm). (Eschscholtz). Lages, Brazil (det. Princis). (scale Roth Blattaria 33o Psyche [September Figs. 127-135. 127-129. (963 L). Monaslria similis (from specimen shown in Fig. 122). 130-132. (52 USNM). Monastria sp. possibly similis. So Paulo, Brazil (det. Gurney). 133-135. (10 ANSP). Monastria biguttara. (from specimen shown in Fig. 121). (scale 0.3 mm). 1970] Roth Blattaria 136-138. (12 MCZ). Monastria biguttata. Brazil. 139MCZ). Monachoda burmeisteri. (from specimen shown in Fig. 124). The ventral surface of L2d (Fig. 139) was cut away and pulled downward to show the preputial spines. 142-144. (2 LEM). Monachoda grossa. 143-144. (21 BMNH). [onachoda grossa. (from specimen shown in Fig. 0.3 ram). 125). (scale Figs. 136-144. 141. (6 Psyche [September Figs. 145-153. 145-147. (42 CUZM). ,Monachoda latissima. (from specimen shown in Fig. 123). 148-150. (964 L). Hiereoblatta cassidea. (from specimen shown in Fig. 126). 151-153. (120 ANSP). Hiereoblatta cassidea. Paratype of Monastria semialata Saussure. Capivary, Santa Catharina, Brazil (from specimen Fig. 23 in Rehn, 1937). A piece of L2d (arrow, Fig. 151) is torn away from the main body of the selerite. (scale ram). 1970] Roth Blattaria Figs. 154-162. 154-156. men shown in Fig. 118. (17 ANSP). Petasodes dominicana. (from speci157-159. (7 ANSP). Petasodes moufeti. (from specimen shown in Fig. 119). 160-162. (18 ANSP). Petasodes reflexa. (from specimen shown in Fig. 120). The prepuce has been pulled down so that the lightly selerotized spines are visible. (scale- 0.3 mm). Psyche [September Figs. 163-168. 163. (1 ANSP). Hormetica laeigata Burmeister. So Paulo, Brazil (det. Rehn). 164. (2 ANSP). Horrnetica apolinari Hebard. Susumuco, Colombia (det. Hebard). 165. (14 MCZ). Hormetica apolinari. Villavicencio, Colombia (det. Roth). 166. (4 ANSP). Hormetica subclncta (Walker). Villavicencio, Colombia (det. Hebard). 167. (44 CUZM). Hormetlca qaerrucosa Brunner. Colombia. 168. (5 ANSP). Brachycola sexnotata (Thunberg). 92 Km from Rio de Janelro, Brazil (det. Rehn). (scale mm). Roth Blattaria Figs. 169-174. 169. (5 MCZ). SibyllobIatta lanesthoides (Walker). Type 1621 of Hormetica adena Seudder. Belmont, Massachusetts (adventive). 170. (961 L). Parahormetica bilobata (Saussure). Lages, Brazil (det. Prineis). 171. (3 ANSP). Parahormetica bilobata. Minas Geraes, Brazil (det. by Hebard as P. tumulosa Brunner a synonym of bilobata). 172. (962 L). Parahormetica cicatricosa Saussure. Brazil (det. Prineis). 173. (972 L). Oxycercus leruaaianus Bolivar. Carpapata, Peru (det. Prineis). 174. (49 USNM). Bionoblatta itatiayae (Miranda Ribeiro). Serra do Itatiaya, Brazil (det. Gurney). (scale mm). Psyche [September Figs. 175-180. 175-177. (57 MCZ). Phoetalia allida (Brunner). Point Antonio, Jamaica. 178-180. (38 BMNH). Phoetalia circumvagans Burmeister. (seale 0.3 ram). 1970] Roth Blattaria Figs. 181-190. 181-183. (14 MCZ). Hormetica apolinari. (from specimen shown in Fig. 165). 184. (2 ANSP). Hormetica alolinari. (from specimen shown in Fig. 164). 185-187. (44 CUZM). Hormetica .errucosa. (from specimen shown in Fig. 167). 188-190. (4 ANSP). Hormetica subcincta. (from specimen shown in Fig. 166). (scale 0.3 mm). -- Psyche [September Figs. 191-202. 191-192. (1 ANSP). (from specimen shown in Fig. 163). 193. (5 ANSP). Brachycola sexnotata. (from specimen shown in Fig. 168). 194. (20 MCZ). Brachycola sexnotata. 195-197. (1 LEM). Sibylloblatta tanesthoides. England (adventive) (det. McE. Kevan). 198. (5 MCZ). 8ibylloblatta [anesthoides. (from specimen shown in Fig. 169). 199-201. (961 L). Parahormetica b;,lobata, (from specimen shown in Fig. 170). 202. (3 ANSP). Parahormetica bilobata. (from specimen shown in Fig. 0.3 mm). 171). (scale Roth Blattaria Figs. 203-211. 203-205. (962 L). Parahormetica cicatricosa. (from specimen shown in Fig. 172). 206-208. (972 L). Oxycercus lberuianus. (from specimen shown in Fig. 173). 209-211. (49 USNM). Bionoblatta itatiayae. (from specimen shown in Fig. 174). (scale 0.3 mm. Psy c he S eptembe two subfamilies and the fringe of preputial spines (Figs. 75, 178) place it in the Blaberinae. Hebard (97) who placed Leurolestes ( Phoetalia) in the Epilamprinae noted that the male’s subgenital plate had "... features comparable to those found in Blaberus [Blaberinae]." McKittrick (964, p. 34) concluded that "The Zetoborinae are closely related to the Brachycolini..." This is supported by the similarity in shape of L2d of certain species of Brachycolini (especially Phoetalia, Figs. 175, 178) with those found in many males of Zetoborinae (Roth, 97ob) however, there are no preputial spines in the Zetobo.rinae. Hebard (I919, p. I28) grouped Hormetica alolinari and H. verrucosa together based on the black markings on the tegmina. The preputial spines of aolinari arise from a heavily sclerotized region (Figs. 18 I, I84) and though this region and the spines are reduced in verrucosa (Fig. I85) the shapes of their Lad’s indicate a close relationship. The L2d and preputial spines are much reduced in Hormetica subcincta (Fig. I88), and show a resemblance to these structures in Brachycola (Fig. I93). Hormetica laevigata (the genotype) is unique in lacking preputial spines (Fig. I9I) and is the only member of the Blaberinae I have seen in which these spines are completely absent (except for rare aberrant specimens of Blaberus spp., Roth, I969). The fringelike indentations of the preputial membrane of H. laevigata are reminiscent o.f this membrane in Phortoeca horaskoides (Walker) (Roth, I97o’b), though the indentations are broader in the fo.rmer species. Hebard’s (I92I, p. S I) conclusion that Hormetica and Parahormetica are closely allied as indicated by Brunner is supported by the present study. Rehn (I937, p. 248) .concluded that the genus Sibylloblatta was a member of endemic neotropical Brachycolae (Hormetica, Parahormetica, Brachycola, Styhon, and Bion [-- Bionoblatta] ). The male genitalia of 8tyhon (Fig. 64) are clearly tho.se of Blaberini, but the other genera grouped together by Rehn are Brachycolini. Rehn (I937, p. 252) placed Bionoblatta (Figs. 2o9-:,.II) between Hormetica and Parahormetica but closer to Parahormetica (Figs. 199-2oI ), and much less closely related to Brachycola (Figs. I93I94) than to either of the other 2 genera. The marked reduction in preputial spines does indicate a close relationship to Parahormetica (e.g.P. bilobata, Figs. I99, 2o2), but also to Oxycercus (Fig. 2o6). The hook R2 of Bionoblatta (Fig. 2IO) differs somewhat in shape from this phallomere in other species of the tribe and the subapical incision extends beyond the middle of the hook; in the other genera 1970] Roth Blattaria 34I the subapical incision is to.und closer to the tip of the hook (e.g., Fig. 2o7). Princis (I96o) placed Oxycercus in the Laxtinae, a subamily which McKittrick did not recognize and which Princis consider.s provisional (Roth, I9o7b). The L2d and prepuce of Oxycercus (Fig. 2o6) are clearly those o.f Brachycolini and near Parahormetica (Fig. 2o3). Caudell’s record of Oxycercus peruvianus trorn Peru is an error (Proc. U. S. Nat. Museum, 44: P- 35o: 19I 3). I examined his .specimen at the USNM and found it to be a species of Parahormetica. SUMMARY Based on male genitalia, 3 tribes as follows: 2I genera of Blaberinae are assigned to Blaberini" ./lrchimandrita, /tspiduchus, Blaberus, Blaptica, Byrsotria, E.ublaberus, Hemiblabera, Hyporhicnoda, Minablatta, Styphon. Monastriini" Hiereoblatta, Monachoda, Monastria, Petasodes. Brachycolini: Bionoblatta, Brachycola, Hormetica, Oxycercus, Parahormetica, Phoetalia, Sibylloblatta. ACKNOWLEDGEMENTS I thank the following for the loan of Museum ma.teria’l: Drs. M. G. Emsley and N. D. Jago, Academy ot Natural Sciences, Philadelphia; Dr. Ashley Gurney, U. S. National Museum; Dr. D. R. Ragge, British Museum (Natural History); Dr. S. L. Tuxen, Zoological Museum, Copenhagen; Dr. V. R. Vickery, Lyman Entomological Museum; Dr. Jerome G. Rozen, Jr., American Museum of Natural History; Dr. Karl Princis, Zoological Institute, Lund University. I am grateful to Mr. Samuel Cohen for taking the photographs. http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png

The Male Genitalia of Blattaria. IV. Blaberidae Blaberinae

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Abstract

to In this *Manuscript received by the editor Zlpril 8, 1970. 197o] Roth Blattaria to the subfamily Blaberinae. The genera Hyporhicnoda, Phoetalia, and Oxycercus, which were placed in subfamilies other than the Blaberinae by McKittrick and Princis, have male genitalia characteristics of members of Blaberinae and I include them in this subfamily. MATERIALS AND METHODS The technique for preparing the male genitalia is given in Roth (969). Unless otherwise indicated in the explanation of figures, the : phallomeres L2d (Fig. 41) and L (Fig. 43) were mounted dorsal side up and the genital hook Ra (Fig. 42) is oriented ventral side uppermost. The source of each of the specimens illustrated is given using the following abbreviations: (AMNH) American Museum of Natural History, New York; (ANSP) Academy of Natural Sciences, Philadelphia; (BMNH) British Museum (Natural History), London (CUZM) Copenhagen University, Zoological Museum, Denmark; (L) Zoological Institute, Lund, Sweden; (LEM) Lyman Entomological Museum, Macdonald College, Quebec, Canada; (N) U. S. Army N atick Labs., N atick, Mass.; (USNM) United States National Museum, Washington, D. C. Geographical collection data and the names of specialists who identified the specimens., if known, :ollow these abbreviations. The number preceding the abbreviations refers to the number assigned the specimen and its corresponding genitalia (on a slide) which are deposited in their respective museums. RESULTS AND DISCUSSION The male phallomeres which have proved useful in studies of Blaberidae are a retractable hook on the right side (R:z) (Fig. 42) a median sclerite (L:vm) which may or may not be solidly attached to a sclerite L:zd (L2 dorsal); and lying below or partly surrounding L:d is a membrane, the prepuce, which almost always bears spines of various sizes and shapes (Fig. 4I). The phallomere (LI) on the left side, has a heavily sclerotized cleft (Fig. 43) and a well defined membranous lobe which overlaps the cleft area; because of its transparency this membrane is difficult to .see in mo3t of the photographs. Rehn and Hebard (I9:Z7, pp. :57-:58), in discussing the Blaberinae, concluded that there are "... at least three lines of development or phyla, which are as follows: I. Archimandrita, Blaberus, Eublaberus 3 IO Psyche [September Blaltica, Byrsotria, Hemiblabera, dsliduchus, Monachoda 3. Monastria, Petasodes The first group is well defined, although relationship with Byrsotria is evident The second group is well defined rom the first, but whether Monachoda should be considered the maximum differentiation o the second group, or a less highly modified member of the third one, is a matter o opinion." The above linear arrangement was the one preferred by Rehn and Hebard. Princis’ (1963) arrangement o some o the genera in the Blaberinae follows Rehn and Hebard, but several additional genera were included. Brachycolinae" Brachycola, Hormetica, Parahormetica, Sibylloblatta, Bionoblatta Blaberinae: ,/lrchimandrita, Blaberus, Mesoblaberus, Eublaberus, Blaptica, Byrsotria, Hemiblabera, ./lsliduchus, Monachoda, Monastria, Minablatta, Petasodes, Cacoblatta, ,Hiereoblatta, Stylhon McKittrick (1964) studied genera o Blaberinae and "or convenience" arranged them in the ollowing tribes: I. Blaberini: glrchimandrita, Blaberus, Blatica 2. Brachycolini: Brachycola, Hormetica, Parahormetica, Petasogte$ 3. Byrsotriini: Byrsotria, Eublaberus, Hemiblabera 4- 1Vionastriini" Monastria Male genitalic characters support the separation o( genera Blaberinae into 3 (Blaberini, Monastriini, and Brachycolini) rather than 4 tribes; and the genta.lia o Petasodes indicate that it does not belong in the Brachycolini as suggested by .-cKittrick. The present study tends to support Rehn and Hebard’s arrangement, especially with regard to the placement o Petasodes. I. Blaberini: ,4rchimandrita, ts#iduchus, Blaberus, Bla#tica, Byrsotria, Eublaberus, Hemiblabera, HyIorhicnoda, Minablatt I-I 8ty[hon (Figs. 17). Figs. 1-9. 1. (11 ANSP)../lrchimandrita marmorata (Stoll). San Mareos, Niearagua. 2. (61 USNM). ./lrch’imandrita tessellata Rehn. San Jos6, Costa Riea (det. Rehn). 3. (18 USNM). Blaberus #iyanteus (L.). St. Jean, French Guiana (det. Roth.). 4. (5 USNM). Blaberus craniifer Burmeister. Key West, Florida (det. Roth). 5. (N). Blaberus discoidalis Serville. Panama (det. Roth). 6. (516 L). Blaberus boli,iensis Prineis. Guayaquil, Ecuador (det. Princis). 7. (514 L). Blaberus anisitsi Branesik (det. Prineis). 8. (N). Blaberus atrolos (Stoll). Trinidad (det. Roth). 9. (N). Blaberus lmrabolicus Walker. Puraquequara, Rio Negro, Amazonas, Brazil (det. Roth). (stale 10 ram). 1970] Roth Blattaria 3II Psyche [September (19 ANSP). Blaberus brasilianus Saussure. Natal, Brazil. (23 ANSP). Blaberus scutatus Saussure (paratype of B. scutata var. obscura S. and Z.). 12. (8 ANSP). Blaberus minor Saussure. Argentina (det. Hebard). 13. (14 ANSP). Blaberus fusiformis Walker. Santa Cruz de la Sierra, Bolivia (det. Hebard). 14. (32 ANSP). Blaberus colosseus (Illiger). Fyzabad, Trinidad (det. Roth). 15. (N). Eublaberus posticus (Eriehson). Panama (det. Roth). 16. (N). Eublaberus distanti (Kirby). Trinidad (det. Roth). (scale 10 mm). Figs. 10-16. To a considerable extent the male genitalia of the genera placed in the Blaberini, show distinct trends (as do the female genitalia and proventricular characters used by McKittrick). It is difficult to separate them into the 2 tribes Blaberini and Byrsotriini which McKittrick used and I have combined them into the Blaberini. However, certain of the genera show closer relationships to one another than to others and therefore I have arranged them in generic groups. In the Blaberini, L2d is solidly joined to L2vm, and may or 1.970] Roth Blattaria may not be hookshaped. The prepuce is a flexible membrane bearing characteristic spines. When flattened, the margins of the preputial membrane often extend beyond the G.utline of Lad and the preputial spines are readily visible. The genera may be arranged as follows: Generic Group I. A rchimandrita (Figs.. I, 2), Blaberus (Figs. 4-I4), Blaptica (Figs. 26, 27, 29). I have reported on the genus Blaber.us in a recent paper (Roth, 969) and for this reason have illustrated R2 (Fig. 42) and L (Fig. 43) for only one of the species. However, for the sake of completeness I have included the L2d for the known species (Figs. 4, 44-55); these structures allow the separation of 3 species groups of Blaberus Giganteus Group" L2d is recurved dorsally and slightly to the right, often forming a hooklike structure. Extending dorsally and laterally on the left, about where L2vm and L2d are joined, is a tumorlike outgrowth. Spines on the preputial membrane are relatively small. (Blaberus giganteus, Figs. 3, 41-43; B. craniifer, Figs. 4, 44)Atropos Group: L2d is not hookshaped and the tumorlike outgrowth on the left side is absent. Truncate or rounded elevations are usually present only on the left side and are generally much larger and more robust than spines on the right. The preputial spines are often arranged in a single row on the left, and a single or sometimes double or partially double row on the right; spines on the right usually more numerous than on the left. (Blaberus discoidalis, Figs. 5, 47; B. boliviensis, Figs. 6, 49; B. anisitsi, Figs. 7, 48; B. atropos, Figs. 8, 45; B. parabolicus, Figs. 9, 46). Brasilianus Group" L2d not hookshaped and tumorlike outgrowth absent as in the Atropos Group. Anterior elevations not greatly enlarged on the left side and about the same size as those on the right. Preputial spines numerous, usually present on the left and right .sides and often occur in more than a single row. (Blaberus brasilian.us, Figs. O, 50; B. scutatus, Figs. I, 52; B. minor, Figs. :, 5 B. [usi[ormis, Figs. I3, 53; B. colosseus, Figs. I4, 54-55). There is considerable intraspecific variation in the male genitalia of Blaberus spp. often making specific determination of certain-species difficult. However, the species group separations are reasonably clear cut. In Zlrchimandrita spp. (Figs. 3:, 35, 38), L2d is not hookshaped, but otherwise this sclerite and the associated preputial fringe of spines are closest to the Gig’anteus Group of Blaherus. Psyche [September In the genus Blaltica the preputial spines are small and occur in multiple rows in obscura (Fig. 56) and dubia (Fig. 6:) but are reduced in number and arranged in a single row in interior (Fig. 59). According to Hepper (I965), Blaptica l)ereyrai Hepper is near B. dubia, but Hepper noted that the preputial spines of lereyrai are fewer in number than in dubia. The number of spines shown in Hepper’s (I965) figure I closely approximate those found in B. interior. The number of preputial spines in species of Blaberus is variable (Roth, I969) and this is probably also true for the genus Blatica. Rehn (I93O, p. 6:) stated that Styhon "... is a distinctive genus occupying an isolated position and not closely related to any of the other previously known genera.." The genus is found in the Dutch Leeward Islands and Rehn (I93O, p. 67) further stated "It is most unexpected to find on these semiarid islands of-[ the north coast of South America an entirely unrecognized genus which has no known close relative on the adjacent.mainland. The genus Parahormetca, which is in some respects the nearest relative, is an inha.bitant of regions widely removed in South America." The L:d and prepuce of Styl)hon (Fig. 64) indicate a closer relationship to Blaptica (Fig. 59) than to Parahormetica (Figs. I99, :oz, :o3) the latter is a member of the B rachycolini (see below). Generic Group :. dspiduchus (Figs. 21-23), Byrsotria (Figs. :4, :5), Eublaberus (Figs. I5-I6), Hemiblabera (Figs. 17-2o). In this group Ld is strongly hookshaped. The genera dsl)iduchus (Figs. 97, IOO, IO3, IO6), Byrsotria (Figs. 75, 78, 81), and Hemiblabera (Figs. 84, 87, 9o, 93, 96) all have very similar L:d’s and the preputial spines are somewhat elongated and arranged principally in a single row. Bonfils’ (1969, p. I32) drawing of the L:d and Figs. 17-25. 17. (24 ANSP). He’mibIabera granulata Saussure. Antigua, West Indies. 18. (50 USNM). Hemiblabera sp. probably tenebricosa Rehn and Hebaid. Rojo Jeeo, Mr. Samana, San Domingo (det. Gurney). 19. (26 ANSP). Herniblabera brunneri (Saussure). Puerto Rico (det. Rehn). 20. (16 MCZ). Hemiblabera labulabor Rehn and Hebard. Andros Island, Bahamas (det. Gurney). 21. (16 ANSP). Asliduchus caerncola J. W. H. Rehn. Paratype. Barrio Monte, Cabo Ro.jo, Puerto Rieo. 22. (15 ANSP). A sliduchus borinquen J. \V. H. Rehn. Paratype. Corozal, Puerto Rieo. 23. (29 AMNH). Aspiduchus borinquen. Corozal, Puerto Rieo. (det. by J. A. G. Rehn as ,4. deplanatus; J. W. H. Rehn (1951) synonimized Rehn and Hebard’s deldanatus with borinquen. 24. (25, ANSP). Byrsotria cabrerM Rehn and Hebard. Matanzas Provlnee, Cuba (det. Rehn). 25. (N.) Byrsotria ]umig.ata (Gu6rin). (det. Roth). (stale ,--" 5 mm) Roth Blattaria Psyche [September Figs. 26-31. 26. (20 ANSP). Blaptica interior Hebard. Formos.a, Argentina. (det. Rehn). 27. (22 ANSP). Blaptica obscura Saussure and Zehntner. So Leopoldo, Rio Grande do Sul, Brazil (det. Rehn.) 28. (64 USNM). Minablatta bipustulata (Thunberg). Vi(.osa, Minas Geraes, Brazil (det. Rehn). 29. (21 ANSP). Blaptica dubia (Serville). Buenos Aires, Argentina (det. Hebard). 30. (N). Hyporhicnoda litomorpha Hebard. Panama. (det. Gurney). 31. (6 ANSP). Styphon bakeri Rehn. Paratype, Curaqao, Dutch West Indies. (scale mm). 1970] Roth Blattaria Figs. 32-40. 32-34. (62 USNM). Xrc.himandrita tessellata. Canal Zone, Panama (det. Hebard). 35-37. (61 USNM). X. tessellata. (from specimen shown in Fig. 2), 38-40. (11 ANSP). drchimandrita rnarmorata. (from specimen shown in Fig. 1). (scale 0.3 ram). prepuce of Hemiblabera tristis Bontils shows these structures similar to other members of the genus. to, be Rehn and Hebard (I927, p. 268) stated that Hemiblabera a position intermediate between Byrsotria and dspiduchus The species brunne’ri has a distinct tendency toward Byrsotria, xvhile conversely H. pabulator and granulata exhibit more of an approach toward dspiduchus.’" The male genitalia are basically too similar to support Rehn and Hebard’s ’... clearly occupies Psyche [-September Figs. 41-49. 41-43. Blaberus giganteus. 41. (1 AMNH). Caripito, Venezuela. (L2d dorsal left phallomere; L2vm ventromedial left sclerite; P prepuce.) 42-43. (27 ANSP). Muzo, Colombia. (det. Roth). (SI Subapical incision of hooked right phallomere R2; C--- Cleft of left phallomere L1). 44. (46 USNM). Blaberus craniifer. Cordoba, Mexico. (det. Roth). 45. (34 USNM). Blaberus atropos. Charleston Quarantine (? Colombia) (det. Roth). 46. (6 AMNH). Blaberus parabolicus. San Martin, Peru (det. Roth). 47. (N). Blaberus discoidalis. Panama (det. Roth). 48. (514 L). Blaberus anisitsi. (from specimen shown in Fig. 7). 49. (516 L). Blaberus bolivicnsis. (from specimen shown in Fig. 6). (scale mm). 1970] Roth Blattaria Figs. 50-55. 50. (19 ANSP). Blaberus brasilianus. (from specimen shown in Fig. 10). 51. (62 ANSP). Blaberus minor. Paraguay. 52. (23 ANSP). Blaberus scutatus. (from specimen shown in Fig. 11). 53. (56 ANSP). Blaberus ]usiformis. Provineia Sara, Dept. Vera Cruz, Bolivia. 54-55. Blaberus colosseus. 54. (1 USNM). Trinidad (det. Roth). 55. (30 ANSP). Caparo, Trinidad (det. Hebard). (scale 0.3 ram). Psyche [September Figs. 56-65. 56-58. (22 ANSP). Blaptica obscura. (from specimen shown in Fig. 27). 59-61. (20 ANSP). Bl’aptica interior. (from specimen shown in Fig. 26). 62-63. (21 ANSP). BlaItica dubia. (from specimen shown in Fig. 29). 64-65. (6 ANSP). Styphon bakeri. (from specimen shown in Fig. 31). (Fig. 65 is a ventral view.) (scale-- 0.3 mm). 1970] Roth Blattaria Figs. 66-74. 66-68. (19 MCZ). Eublaberus losticus. (Type 16074 of Blabera ]emorata Seudder). Napo or Marafi6n. 69-71. (27 AMNH). Eublaberus distanti. Moyobamba, San Martin, Peru. (det. Roth). 72. (N). Eublaberus distanti. Trinidad. (det. Roth). 73. (N.) Eublaberus losticus. Panama. (det. Roth). 74. (4 BMNH). Eublaberus sp. Para, Brazil. (det. 0,3 mm). Roth). (scale Psyche [September Figs. 75-83. 75-77. (N). Byrsotria fumi.qata (det. Roth). 78-80. (25 ANSP). Byrsotria cabrerai. (from specimen shown in Fig. 24). 81-83. (15 MCZ). B. cal, rerai. Coast below Pico Turquino, Cuba. (det. Gurney). 0.3 mm). (scale 1970] Roth Blattaria Figs. 84-92. 84-86. (13 ANSP). Hemiblabera pabulator. Paratype. Nassau, Bahamas. 87-89. (16 MCZ). H. pabulator. (from specimen shown in Fig. 20). 90-92. (24 ANSP). Hemiblabera granulator. (from specimen shown in Fig. 17). (scale-- 0.3 mm). Psyche [September Figs. 93-102. 93-95. (26 .A_NSP). Hemiblabera brunneri. (rom specimen shown in Fig. 19). 96. (50 USNM). Hemiblabera sp. probably tenebricosa. (from specimen shown in Fig. 18). 97-99. (15 ANSP). lsliduchus borinquen. (from specimen shown in Fig. 22). 100-102. (16 ANSP). lsliduchus ca,ernicola. (from specimen shown in Fig. 21). (scale ram). 1970] Roth Blattaria Figs. I03-111. 10:t-108. 4stiduchus borinquen. 103-105. (30 AMNH). Corozal Cave, Puerto Rico. (misidentified by Rehn and Hebard as detlanatus). 106-108. (29 AMNH). (from specimen shown in Fig. 23). 109-111. (64 USNM). AIinablatta bibustulata. (from specimen shown in Fig. 28). (scale 0.3 ram). Psyche [September Figs. 112-117. 112-114. (177 USNM). Hyorhicnoda reflexa (Saussure and Zehntner). Turrialba, Costa Rica (det. Gurney). 115-117. Hyorh!cnoda litomorlha Hebard. Panama (det. Gurney). (scale 0.3 mm). 1970] Roth Blattaria suggestion about the relationship of species of Hemiblabera to Byrsotria and Aspiduchus. Rehn and Hebard I927, p. 278) in erecting the genus Aspiduchus stated that it "... is apparently related on one hand to Hemiblabera Saussure, and on the other to Monachoda Burmeister." The male genitalia of Aspiduchus (Figs. 21-23) indicate a close relationship to Hemiblabera (Figs. 17-2o) but less so to Monachoda (Figs. I42, 144, 145) which I place in the Monastriini (see below). Rehn’s (i95I p. 7) drawings of the Lad’s of Aspiduch.us cavernicola (Fig. io’o) and A. borinquen (Figs. 97, o3, IO6) show relatively small differences. It is doubtful that the male genitalia can be used to distinguish between these 2 species. In Eublaberus spp. the preputial spines occur in more than a single row (Figs. 66, 69, 72-74). These spines are more slender and less sclerotized in E. distanti (Figs. 69, 72) than in E. posticus (Figs. 66, 73), as noted by Princis (I950). Hebard (1920 p. 116) states that Eublaberus is closely related to Blaberus. However L2d is more distinctly hookshaped in Eublaberus than it is in most species of Blaberus and for this reason I place it in the second generic group. Generic Group 3. Hyporhicnoda (Fig. 3o), Minablatta (Fig. 8). In this group Ld is not hookshaped. Rehn (94o) stated that Minablatta is related to Blaptica and Eublaberus but i.s probably more closely related to the former. The absence of a hookshaped Ld in Minablatta (Fig. o9) places the genus closer to Blatica. However, the elongated preputial spines, occurring in a single row in Minablatta (Fig. o9) show a similarity to the spines found in .dspiduckus (Fig. o6), Byrsotria (Fig. 75), and Hemiblabera (Fig. 96). Hyporkicnoda has been placed in the Epilamprinae by Hebard (9o), Princis (96o), and McKittrick (964). However, the prepuce of the male has a fringe of spines (Figs. 5) which is characteristic of the Blaberinae. Spines are not found on the prepuce of any genera belonging to the Epilamprinae; nor have I seen these spines in males other than Blaberinae, representing more than 90 genera of Blaberidae. In H.porkicnoda the apex of Lvm is more darkly pigmented than Led so that the junction of the 2 sclerites is readily discernible. Ionastriini: Hiereoblatta (Fig. 6), _84onackoda (Figs. I23-I2_), Monastria (Fi:s. I2I-I22). Petasodes (Figs. II8-I20). Psyche [September In this tribe the apex of L2vm is solidly joined to a relatively broad L2d. In Monastria (Figs. 127-138), the prepuce (Figs. 127, I3O, 133, 36, 138) is almost entirely covered by rows of small, closely packed spines and extends we’ll below the posterior margin of L2d. In Monachoda (Figs. 39-I 5 O), Hiereoblatta (Figs. 15 I53), and Petasodes (Figs. 154-162) the prepuce extends little or not at all beyond the margin of Lad and many of the preputial spines (considerably fewer in number than in Mon.astria) are usually hidden under L2d. In Petas’odes reflexa (Fig. 16o) there is a marked reduction in number of preputial spines and these are very lightly sclerotized. McKittrick (1964) placed Petasodes in the Brachycolini, but all the members of this tribe have L2d clearly separated from L2vm by a membrane (see below). Rehn’s (1937, p. 24) statement that Hiereoblatta is related to Monastria is justified by the similarities in male genitalia. However, the shape of L2d and the preputial spines hidden under it in Hiereoblatta (Figs. 148 151 indicate a closer relationship to Monachoda (Figs. 42, 44, 45) than to Monastria (Figs. 127, 13o, 133, 136, 138). Brachycolini" Bionoblatta, Brachycola, Hormetica, Oxycercus, Parahormetica, Phoetalia, Sibylloblatta (Figs. I63-II). This tribe is clearly differentiated t?rom the Blaberini and Monastriini in that L2d is not solidly attached to L2vm, but is separated from it clearly by a membrane. The spines on the prepuce vary in size from relatively large in Phoetalia (Figs.. 175, 178) to small in Parahormetica (Figs. 199, 202, 2o3), Oxycercrus (Fig. 2o.6), Bionoblatta (Fig. 2o,9), Sibylloblatta (Figs. 95, I98), and Brachycola (Fig. I93). Princis (I96O) placed Phoetalia in the Epilamprinae and McKittrick (I964) assigned it (as Leurolestes) to the Diplopterinae. The male genitalia of Phoetalia are not characteristic of either of these Figs. 118-126. 118. (17 ANSP). Pe:tasodes dominicana (Burmeist.er). Rio de Janeiro, Brazil (det. Hebard). 119. (7 ANSP). Petasodes mou[eti (Kirby). So Paulo, Brazil. (det. Hebard). 120. (18 ANSP). Petasodes reflexa (Thunberg). 46 m. south of Bahia, Brazil. (det. Rehn). 121. (10 ANSP). Monastria biguttata (Thunberg). Rio de Janeiro, Brazil (det. H.ebard). 122. (963 L). lllonastria similis (Serville). Misiones, Argentina (det. Princis). 123. (42 CUZM). Monachoda latissima Brunner. Bahia, Brazil. 124. (6 MCZ). Monachoda burmeisteri Saussure. 125. (21 BMNH). JlConachoda 7rossa (Thunberg). 126. (964 L). Hiereoblatta cassidea 10 mm). (Eschscholtz). Lages, Brazil (det. Princis). (scale Roth Blattaria 33o Psyche [September Figs. 127-135. 127-129. (963 L). Monaslria similis (from specimen shown in Fig. 122). 130-132. (52 USNM). Monastria sp. possibly similis. So Paulo, Brazil (det. Gurney). 133-135. (10 ANSP). Monastria biguttara. (from specimen shown in Fig. 121). (scale 0.3 mm). 1970] Roth Blattaria 136-138. (12 MCZ). Monastria biguttata. Brazil. 139MCZ). Monachoda burmeisteri. (from specimen shown in Fig. 124). The ventral surface of L2d (Fig. 139) was cut away and pulled downward to show the preputial spines. 142-144. (2 LEM). Monachoda grossa. 143-144. (21 BMNH). [onachoda grossa. (from specimen shown in Fig. 0.3 ram). 125). (scale Figs. 136-144. 141. (6 Psyche [September Figs. 145-153. 145-147. (42 CUZM). ,Monachoda latissima. (from specimen shown in Fig. 123). 148-150. (964 L). Hiereoblatta cassidea. (from specimen shown in Fig. 126). 151-153. (120 ANSP). Hiereoblatta cassidea. Paratype of Monastria semialata Saussure. Capivary, Santa Catharina, Brazil (from specimen Fig. 23 in Rehn, 1937). A piece of L2d (arrow, Fig. 151) is torn away from the main body of the selerite. (scale ram). 1970] Roth Blattaria Figs. 154-162. 154-156. men shown in Fig. 118. (17 ANSP). Petasodes dominicana. (from speci157-159. (7 ANSP). Petasodes moufeti. (from specimen shown in Fig. 119). 160-162. (18 ANSP). Petasodes reflexa. (from specimen shown in Fig. 120). The prepuce has been pulled down so that the lightly selerotized spines are visible. (scale- 0.3 mm). Psyche [September Figs. 163-168. 163. (1 ANSP). Hormetica laeigata Burmeister. So Paulo, Brazil (det. Rehn). 164. (2 ANSP). Horrnetica apolinari Hebard. Susumuco, Colombia (det. Hebard). 165. (14 MCZ). Hormetica apolinari. Villavicencio, Colombia (det. Roth). 166. (4 ANSP). Hormetica subclncta (Walker). Villavicencio, Colombia (det. Hebard). 167. (44 CUZM). Hormetlca qaerrucosa Brunner. Colombia. 168. (5 ANSP). Brachycola sexnotata (Thunberg). 92 Km from Rio de Janelro, Brazil (det. Rehn). (scale mm). Roth Blattaria Figs. 169-174. 169. (5 MCZ). SibyllobIatta lanesthoides (Walker). Type 1621 of Hormetica adena Seudder. Belmont, Massachusetts (adventive). 170. (961 L). Parahormetica bilobata (Saussure). Lages, Brazil (det. Prineis). 171. (3 ANSP). Parahormetica bilobata. Minas Geraes, Brazil (det. by Hebard as P. tumulosa Brunner a synonym of bilobata). 172. (962 L). Parahormetica cicatricosa Saussure. Brazil (det. Prineis). 173. (972 L). Oxycercus leruaaianus Bolivar. Carpapata, Peru (det. Prineis). 174. (49 USNM). Bionoblatta itatiayae (Miranda Ribeiro). Serra do Itatiaya, Brazil (det. Gurney). (scale mm). Psyche [September Figs. 175-180. 175-177. (57 MCZ). Phoetalia allida (Brunner). Point Antonio, Jamaica. 178-180. (38 BMNH). Phoetalia circumvagans Burmeister. (seale 0.3 ram). 1970] Roth Blattaria Figs. 181-190. 181-183. (14 MCZ). Hormetica apolinari. (from specimen shown in Fig. 165). 184. (2 ANSP). Hormetica alolinari. (from specimen shown in Fig. 164). 185-187. (44 CUZM). Hormetica .errucosa. (from specimen shown in Fig. 167). 188-190. (4 ANSP). Hormetica subcincta. (from specimen shown in Fig. 166). (scale 0.3 mm). -- Psyche [September Figs. 191-202. 191-192. (1 ANSP). (from specimen shown in Fig. 163). 193. (5 ANSP). Brachycola sexnotata. (from specimen shown in Fig. 168). 194. (20 MCZ). Brachycola sexnotata. 195-197. (1 LEM). Sibylloblatta tanesthoides. England (adventive) (det. McE. Kevan). 198. (5 MCZ). 8ibylloblatta [anesthoides. (from specimen shown in Fig. 169). 199-201. (961 L). Parahormetica b;,lobata, (from specimen shown in Fig. 170). 202. (3 ANSP). Parahormetica bilobata. (from specimen shown in Fig. 0.3 mm). 171). (scale Roth Blattaria Figs. 203-211. 203-205. (962 L). Parahormetica cicatricosa. (from specimen shown in Fig. 172). 206-208. (972 L). Oxycercus lberuianus. (from specimen shown in Fig. 173). 209-211. (49 USNM). Bionoblatta itatiayae. (from specimen shown in Fig. 174). (scale 0.3 mm. Psy c he S eptembe two subfamilies and the fringe of preputial spines (Figs. 75, 178) place it in the Blaberinae. Hebard (97) who placed Leurolestes ( Phoetalia) in the Epilamprinae noted that the male’s subgenital plate had "... features comparable to those found in Blaberus [Blaberinae]." McKittrick (964, p. 34) concluded that "The Zetoborinae are closely related to the Brachycolini..." This is supported by the similarity in shape of L2d of certain species of Brachycolini (especially Phoetalia, Figs. 175, 178) with those found in many males of Zetoborinae (Roth, 97ob) however, there are no preputial spines in the Zetobo.rinae. Hebard (I919, p. I28) grouped Hormetica alolinari and H. verrucosa together based on the black markings on the tegmina. The preputial spines of aolinari arise from a heavily sclerotized region (Figs. 18 I, I84) and though this region and the spines are reduced in verrucosa (Fig. I85) the shapes of their Lad’s indicate a close relationship. The L2d and preputial spines are much reduced in Hormetica subcincta (Fig. I88), and show a resemblance to these structures in Brachycola (Fig. I93). Hormetica laevigata (the genotype) is unique in lacking preputial spines (Fig. I9I) and is the only member of the Blaberinae I have seen in which these spines are completely absent (except for rare aberrant specimens of Blaberus spp., Roth, I969). The fringelike indentations of the preputial membrane of H. laevigata are reminiscent o.f this membrane in Phortoeca horaskoides (Walker) (Roth, I97o’b), though the indentations are broader in the fo.rmer species. Hebard’s (I92I, p. S I) conclusion that Hormetica and Parahormetica are closely allied as indicated by Brunner is supported by the present study. Rehn (I937, p. 248) .concluded that the genus Sibylloblatta was a member of endemic neotropical Brachycolae (Hormetica, Parahormetica, Brachycola, Styhon, and Bion [-- Bionoblatta] ). The male genitalia of 8tyhon (Fig. 64) are clearly tho.se of Blaberini, but the other genera grouped together by Rehn are Brachycolini. Rehn (I937, p. 252) placed Bionoblatta (Figs. 2o9-:,.II) between Hormetica and Parahormetica but closer to Parahormetica (Figs. 199-2oI ), and much less closely related to Brachycola (Figs. I93I94) than to either of the other 2 genera. The marked reduction in preputial spines does indicate a close relationship to Parahormetica (e.g.P. bilobata, Figs. I99, 2o2), but also to Oxycercus (Fig. 2o6). The hook R2 of Bionoblatta (Fig. 2IO) differs somewhat in shape from this phallomere in other species of the tribe and the subapical incision extends beyond the middle of the hook; in the other genera 1970] Roth Blattaria 34I the subapical incision is to.und closer to the tip of the hook (e.g., Fig. 2o7). Princis (I96o) placed Oxycercus in the Laxtinae, a subamily which McKittrick did not recognize and which Princis consider.s provisional (Roth, I9o7b). The L2d and prepuce of Oxycercus (Fig. 2o6) are clearly those o.f Brachycolini and near Parahormetica (Fig. 2o3). Caudell’s record of Oxycercus peruvianus trorn Peru is an error (Proc. U. S. Nat. Museum, 44: P- 35o: 19I 3). I examined his .specimen at the USNM and found it to be a species of Parahormetica. SUMMARY Based on male genitalia, 3 tribes as follows: 2I genera of Blaberinae are assigned to Blaberini" ./lrchimandrita, /tspiduchus, Blaberus, Blaptica, Byrsotria, E.ublaberus, Hemiblabera, Hyporhicnoda, Minablatta, Styphon. Monastriini" Hiereoblatta, Monachoda, Monastria, Petasodes. Brachycolini: Bionoblatta, Brachycola, Hormetica, Oxycercus, Parahormetica, Phoetalia, Sibylloblatta. ACKNOWLEDGEMENTS I thank the following for the loan of Museum ma.teria’l: Drs. M. G. Emsley and N. D. Jago, Academy ot Natural Sciences, Philadelphia; Dr. Ashley Gurney, U. S. National Museum; Dr. D. R. Ragge, British Museum (Natural History); Dr. S. L. Tuxen, Zoological Museum, Copenhagen; Dr. V. R. Vickery, Lyman Entomological Museum; Dr. Jerome G. Rozen, Jr., American Museum of Natural History; Dr. Karl Princis, Zoological Institute, Lund University. I am grateful to Mr. Samuel Cohen for taking the photographs.

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