Louis M. Roth.
The Male Genitalia of Blattaria. IV. Blaberidae: Blaberinae.
Psyche 77:308-342, 1970.

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THE MALE GENITALIA OF BLATTARIA, IV
BLABERIDAE : BLABERINAE*
BY Louis M. ROTH
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01 760
In this paper I shall illustrate and use male genitalic structures to show the probable relationships of the following 21 genera of Bionoblatta Rehn, Blaberus Serville, Blaptica Stil, Brachycola Ser- ville, Byrsotria Stil, Eublaberus Hebard, Hemiblabera Saussure, Hiereoblatta Rehn, Hormetica Burmeister, ffyporhicnoda Hebard, Minablatta Rehn, Monachoda Burmeister, Monastria Saussure, Oxycercus Bolivar, Parahormetica Brunner, Petasodes Saussure, Phoetaliu Shelford, Sibylloblatta Rehn, and Styphon Rehn. Princis (1960) placed most of the above genera in 2 subfamilies (Blaberinae and Brachycolinae) of Blaberidae. He also included Cacoblatta Saussure and Mesoblaberus Princis in the Blaberinae but I have not seen any males of species belonging to these genera. Anchoblatta Shelford, which Princis (1960) included in the Brachy- colinae with a (?) and listed under this subfamily in his 1963 Catalogus, has male genitalia characteristic of the Panchlorinae and I have assigned it to this subfamily (Roth, 1971 ). McKittrick (1964, p. 34) stated "The Blaberinae are the largest subfamily in this complex [Blaberoid Complex], and it forms a peculiarly diverse group made up entirely of tropical and subtropical New World forms. It includes here both the Blaberinae and Brachycolinae of Rehn (1951) and Princis (1960). Because of overlapping characteristics, any subdivision at this level would neces- sarily be arbitrary. As may be seen in tables VI [I? genitalia] and VII [proventriculus], there are no clear divisions within the group, only distinct trends." Previous studies (Roth, 1969, 1970a, 1970b) have indicated that the male genitalia of species of Blaberidae pos- sess excellent characters for showing subfamily, tribal, generic, and species group characteristics, and in the present study this was found to be true for the Blaberinae.
I shall follow McKittrick in assigning all of the genera she studied *Manuscript received by the editor April 8, 1970. 308




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19701 Roth - Blattaria 309
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 char- acteristics of members of Blaberinae and I include them in this subfamily.
The technique for preparing the male genitalia is given in Roth (1969). Unless otherwise indicated in the explanation of figures, the 2 phallomeres L2d (Fig. 41 ) and LI (Fig. 43 ) were mounted dorsal side up and the genital hook R2 (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 Natick Labs., Natick, Mass.; (USNM) = United States National Museum, Washington, D. C. Geographical collection data and the names of specialists who identified the specimens, if known, follow 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.
The male phallonieres which have proved useful in studies of Blaberidae are a retractable hook on the right side (R2) (Fig. 42) ; a median sclerite {Lzvm) which may or may not be solidly attached to a sclerite L2d (L2 dorsal) ; and lying below or partly surround- ing Lad is a membrane, the prepuce, which almost always bears spines of various sizes and shapes (Fig. 41). 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 ( 1927, pp. 257-258), in discussing the Blabe- rinae, concluded that there are ". . . at least three lines of develop- ment or phyla, which are as follows:
I. Archinzandrita, Blaberus, Enblaberus




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3 10 Psyche [September
2. Blaptica, Byrsotria, Hemiblabera, Aspiduchus, Monachoda 3. Monastria, Petasodes
The first group is well defined, although relationship with Byrsotria is evident . . . The second group is well defined from the first, but whether Monachoda should be considered the maximum differentiation of the second group, or a less highly modified member of the third one, is a matter of opinion." The above linear arrange- ment was the one preferred by Rehn and Hebard. Princis'
(1963) arrangement of some of the genera in the Bla- berinae follows Rehn and Hebard, but several additional genera were included.
Brachycolinae: Brachycola, Hormetica, Parahormetica, Sibyllo- biatta, Bionoblatta
Blaberinae : Archimandrita, Blaberus Mesoblaberus, Eublaberus, Blaptica, Byrsotria, Hemiblabera, hpiduchus, Monachoda, Mon- astria, Minablatta, Petasodes, Cacoblatta, Hiereoblatta, Styphon McKittrick (1964) studied 11 genera of Blaberinae and "for convenience" arranged them in the following tribes: I. Blaberini : Archimandrita, Blaberus Blaptica 2. Brachycolini: Brachycola, Hormetica, Parohornzeticn, Peta- sodes
3. Byrsotriini: Byrsotria, Eublaberus, He//7ih/ahcra 4. Monastriini : Monastria
Male genitalic characters support the separation of genera of Blaberinae into 3 (Blaberini, Monastriini, and Brachycolini) rather than 4 tribes; and the gentalia of Petasodes indicate that it does not belong in the Brachycolini as suggested by McKittrick. The present study tends to support Rehn and Hebard's arrangement, especially with regard to the placement of Petasodes. I. Blaberini : Archimandrita, Aspiduchus, Blaberus, Blaptica, Byrsotria, Eublaberus, Hemiblabera, Hyporhicnoda, Mina/,Zat/a Styphon (Figs. 1-117).
Figs. 1-9. 1. (11 ANSP). Archimandrita marmorata (Stoll). San Marcos, Nicaragua. 2. (61 USNM) . Archimandrita tessellata Rehn. San Jose, Costa Rica (det. Rehn). 3. (18 USNM). Blaberus giganteus (L.). St. Jean, French Guiana (det. Roth.). 4. (5 USNM). Blaberus craniifer Burrneister. Key West, Florida (det. Roth) . 5. (N) . Blaberus discoidalis Serville. Panama (det. Roth) . 6. (516 L) . Blaberus boliviensis Princis. Guayaquil, Ecuador (det. Princis). 7. (514 L). Blaberus anisitsi Brancsik (det. Princis). 8. (N). Rlaberus atropos (Stoll). Trinidad' (det. Roth); 9. (N). Blaberus farabolicus Walker.
Puraquequara, Rio Negro, Amazonas,
Brazil (det. Roth). (scale ==. 10 mm),




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Psyche [September
Fig's. 10-1 6. ( 19 ANSP). Bhberus brasilianus Saiissiii e. Natal, Brazil. 11. (23 AN?#). Btalieru~ scvtatus Saussure (paratype of B. sculda var. obscura S. and 2.). 12. (8 ANSP). RSaterus minor Saussure. Argentina (det. Hebard). 13. (14 ANSP). Blaberus fusiformis Walker. Santa Cruz de la Sierra, Bolivia {det. Hebard). 14. (32 ANSP). Blabervs colosseui (llliger). Fyzabad, Trinidad (det. Roth), IS. (N). EuWaberus poiiiciis (Erichson). Panama (det. Roth). 16. (N). Eublaberus distanti (Kirby). Trinidad (det. Roth), (scale = 10 mm).
To a considerable extent the male genitalia of the genera placed in the BIaberini, show distinct trends (as do the female genitalia and proventricular characters used by McKittricIc). It is difficult to separate them into the 2 tribes Blaberini and Byrsotriini which Mc- Kittrick used and
I have combined them into the BlaberinL How- ever, certain of the genera show closer relationships to. one another than to others and therefore I have arranged them in generic groups. In the BIaberini, L2d is solidly joined to Lavm, and may or



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19701 Roth - Blattaria 313
may not be hookshaped. The prepuce is a flexible membrane bearing characteristic spines. JVhen flattened, the ~nargins of the ~l-e~utial membrane often extend beyond the cutline of L2d and the preputial spines are readily visible.
The genera may be arranged as follows:
Generic Group I. A rchimandrita ( Figs. I, 2 ) , BZaberus (Figs. 4-14), BZaptica (Figs. 26, 27, 29).
1 have reported on the genus Blabems in a recent paper (Ruth, 1969)
and for this reason have illustrated R2
(Fig. 42) and LI
(Fig. 43)
for only one of the species. However) for the sake of completeness I have included the L2d for the known species (Figs. 41, 44-55) ; these structures allow the separation of 3 species groups of BZaberus:
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 Lzvm and L2d are joined, is a. tumorlike outgrowth. Spines on the preputial membrane are rela- tively small. (BZaberus giganteus, Figs. 3, 41 -43 ; B. craniifer, Figs. 4, 44).
Atropos Group: L2d is not hookshaped and the tumorlike out- growth 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. (BZaberus discoidaZis, Figs. 5, 47; B. boliviensis, Figs. 6, 49; B. anisitsi, Figs. 7, 48; B. atropos, Figs. 8, 45; B. paraboZicus, Figs. 9, 46). Brailianus Grou$: L2d not hookshaped and tumorlike out- growth 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 brasiZianlus, Figs. 10, 50 ; B. scutatz~s, Figs. I I ) 52 ; B. minor, Figs. 12, 5 I ; B. fusiformis, Figs. 13, 53 ; B. coZosseus, Figs. 14, 54-55). There is considerable intraspecific variation in the male genitalia of BZaberus spp. often making specific determination of certain-species difficult. However, the species gro~~p separations are reasonably clear cut.
In Archimondrita spp. (Figs. 32. 35, 381, L2d is not hookshaped, but otherwise this sclerite and the msociatecl preputial fringe of spines are closest to the Gigmteus Group of Bl~berus.



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314 Psyche [September
In the genus BZaptica the preputial spines are small and occur in multiple rows in obscura (Fig. 56) and dubia (Fig. 62) but are reduced in number and arranged in a single row in interior (Fig. 59). According to Hepper (1965)) BZaptica pereyrai Hepper is near B. dubia, but Hepper noted that the preputial spines of pereyrai are fewer in number than in dabia. The number of spines shown in Hepper's
(1965) figure I I closely approximate those found in I?. interior. The number of preputial spines in species of BZaberus is variable (Roth, 1969) and this is probably also true for the genus Blaptica.
Rehn
(1930) p. 62) stated that Styphon ((. . . is a distinctive genus occupying an isolated position and nat closely related to any of the other previously known genera." The genus is found in the Dutch Leeward Islands and Rehn (1930, p. 67) further stated "It is most unexpected to find on these semiarid islands off the north coast of South America an entirely unrecognized genus which has no known close relative on the adjacent mainland. The genus Para- hormeticd, which is in some respects the nearest relative) is an inhabitant of regions widely removed in South America." The L2d and prepuce of Styphon (Fig. 64) indicate a closer relationship to Blapticu (Fig. 59) than to Parahormetica (Figs. 199, 202, 203) ; the latter is a member of the Brachycolini (see below). Generic Group 2. Aspiduchus (Figs. 21-23), Byrsotria (Figs. 24, 25) ) Eublaberus (Figs. I 5-16), Hemiblabera (Figs. I 7-20). In this group L2d is strongly hookshaped. The genera Aspiduchus (Figs. 97, IOQ) 103, 106), Byrsotria (Figs. 75) 78, 81)) and Hemi- blabera (Figs. 84) 87, 90, 93, 96) all have very similar L2d's and the preputial spines are somewhat elongated and arranged principally in a single row. Bonfils' (1969, p. 132) drawing of the L2d and Figs, 17-25. 17. (24 ANSP). Hefmiblabera granulata Saussure. Antigua, West Indies. 18. (SO USNM). Hemiblabera sp. probably tenebricosa Rehn and Hebard. Rojo Jeco, Mt. Samana, San Domingo (det. Gurney). 19. (26 ANSP). Hemiblabera brunneri (Saussure) . Puerto Rico (det. Rehn) . 20. (16 MCZ). Hemiblabera $ahdabor Rehn and Hebard. Andros Island, Bahamas (det. Gurney).
21, (16 ANSP). As~iduchus cawmzicola J. W. H. Rehn. Paratype. Barrio Monte, Cab0 Rojo, Puerto Rico. 22. (15 ANSP). As$iduchus borinpen J. W. H. Rehn. Paratype. Corozal, Puerto Rico. 23. (29 AMNH). Aspiduc~us borinquen. Corozal, Puerto Rico. (det. by J. A. G. Rehn as A. deplana~us; J. W. H. Rehn (1951) synonimized' Rehn and Hebard's deplanatus with borinquen. 24. (25 ANSP) . Byrsotria cabrerai Rehn and Hebard. Matanzas Province, Cuba (det. Rehn). 25. (N.) Byrsotria fumigata (Gu6rin). (det. Roth). (scale = 5 mm)



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psyche [September
Figs. 26-31. 26. (20 ANSP). Blapticu interior Hebard. Formosa, Argen- tina. (det. Rehn). 27. (22 AMSP). Bla$iica oh~xrfi Saussure and Zehnt- ner- S3o Leopoldo, Rio Grande do Sul, Brazil (det. Rehri.) 28. (64 USKM). Mintzblatta h;pu~~tihtu (Thunherg), Vi<osa, Minas Geraes, Brazil (det. Rehn) . 29. (21 ANSP) . Bla$ticu didk ( ServiUe]. Buenos Aires, Argentina (det, Hebard). 30. (3 1, Hy$orhicnodu lifomorphu Hebard. Panama. (det. Gurney). 31. (6 *WSP). &y$hn hkeri Rehn. Paratype, Curapo, Dutch West Indies. (scale = 5 mrn j.



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Figs. 32-40. 32-34, (62 USNM). Archiinandri~a itssclluta. Canal Zone, Panama (det. Hebard). 35-37, (61 USNM). A, t~~dlatu, (from specimen shown in Fig. 2). 38-40. (11 ANSP). Archimrmdrita murmomta. (from specimen shown in Fig. 1). (scale = 0.3 mrn), prepuce of Hiv~iibhbero iristis Bod& sl~ows these structures to be similar to other members of the genus.
Rehn and Hebard (1927, p. 268) stzted that HemMabmz I'. . . clearly occupies a position intti-mediate between Byrsotria and Asfiiduchux, . , . The speck &runnwi has a distinct tendency towsrd Byrsotria, while convemly H. +abulator and granuZda exhibit more of an approach toward Aspiduchus." The male pitaiia are basicalIy too similar to support Rthn and Hebard's



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Psyche
[September
Figs. 41-49. 41-43. Bhberus giganteus. 41. (1 AMNH). Caripito, Vene- zuela, (L2d = dorsal left phallomere; L2vm = ventromedial left sclerite; P = prepuce.)
42-43. (27 ANSP). Mum, Colombia. (det. Roth). (SI = Subapical incision of hooked right phallomere R2; C = Cleft of left phallomere LI). ++. (46 USNM). Blabems crartiifcr, Cordoba, Mexico. (det. Roth). 45. (34 USNM). Blabcrus airapos. Charleston Quarantine ( ? Colombia) (det. Roth). 46 (6 AMNH). Riaberus parabal.cm. San Martin, Peru (det. Roth). 47. (N). Blaberus discoidah. Panama (det. Roth). 48. (514 L). Biahervs anishi. (from specimen shown in Pig, 7). 49. (516 L). Blaberus botivirmis. (from specimen shown in Fig. 6). (scale = 0.3 mm),




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19701 Roth - Blaitaria 319
Figs. 50-55. 50. (19 ANSP). Blaberus brasilianus. (from specimen shown in Fig. 10). 51. (62 ANSP). Blabems minor. Paraguay. 52. (23 ANSP). Biallerus stuiatus.
(from specimen shown in Fig. 11).
53. (56 ANSP).
Blaberus fusiformis. Provincia Sara, Dept. Vera Cruz, Bolivia. 54-55. SJaberus colotseus. $4. (1 USNM). Trinidad (det. Roth). 55. (30 ANSP). Caparo, Trinidad (det. Hebard).
(scale = 0.3 mm).




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Psyche [September
Figs. 56-65. 56-58. (22 ANSP), Bluptica obscura. {from specimen shown in Fig. 27). 59-61. (20 ANSP). Blaptica interior. (from specimen shown in Fig. 26). 62-63. (21 ANSP). Blapfica dubia. (from specimen shown in Fig. 29). 64-65. (6 ANSP). Slyphon bakeri. (from specimen shown in
Fig. 31).
(Fig. 65 is a ventral view.) (scale = 0.3 mm).



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Roth - Bialtaria
Figs. 66-74. 66-68. (19 MCZ). Eubla&cras posticus. (Type 16074 of Blabera jcmorata Scudder). Nap or MaramSn. 69-71. (27 AMNH). EubIaIf erui di~tmti. Moyobamba, San Martin, Peru. (det. Roth) . 72. (N 1. Euhlaberus distant'!. Trinidad. (det. T'inth). 73. /N) Eubluherus poslica~. Panama. (dct. Roth), 74. (+ BMNH). Eublaberits sp. Para, Brazil. (det. Roth). (scale = 0.3 mm).




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Psyche [September
Figs. 75-83. 75-77. (N). Byrsotria fumlgata (det. Roth). 78-80. (25 ANSP). Bynotria mbrrrai. (from specimen shown in Fig. 24). 81-83. (IS
MCZ). B. cahrerai. Coast below Pico Turquino, Cuba. (det. Gurney). (scale := 0.3 mm).




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Figs. E4-92. 8446. (13 ANSP) . Hemibiabera pabitlator, Paratype. Naa- sau, Bahamas. 87-89. (16 MCZ). H. falidator. (from specimen shown in Fig. 20). 90-92. (24 ANSP). Henitbhbera gramlatar. (from specimen shown in Fig. 17).
(scale = 0.3 mrn).




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324 Psyche [September
Figs. 93-102. 93-95. (26 ANSP) . Hemibiabera brunneri. (from specimen shown in Fig. 19). 96. (SO USNM), Hemiblabera sp. probably trne- bricosa. (from specimen shown in Fig, 18). 97-99. (IS ANSP). Aspi- duchus barinyuen. (from specimen shown in Fig. 22). 100-102. (16 ANSP). ^spiduchus: cavwnicola. (from specimen shown in Fig. 21). (scale == 0.3 mm).




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Figs. 103-111, 103-108. Aspidwhus barinquen, 103-105. (30 AMNH). Corozal Cave, Puerto Rico.
(misidentified by Rehn and Hebard as de- planatus). 106-108. (29 AMNH). (from specimen shown in Fig. 23). 109-111. (64 USNM). Afinablatta bipwtulata. (from specimen shown ID Fig. 28). (scale = 0.3 mrn).




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Psyche [September
Figs. 112- 117. 112-114. (177 USNM) , Hyporhimuda refiexu (Sausuure and Zehntner). Turrialba, Costa Rica (det. Gurney). 115-117. HypÌörkc
soda lhomof'fha Hebard. Panama (det. Gurney), (acale = 0.3 tam).



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19701 Roth - Blattaria 327
suggestion about the relationship of species of Hemiblabera to Byrsotria and Aspiduchus.
Rehn and Hebard (1927, 13. 278) in erecting the genus Aspiduchus stated that it ". . . is apparently related on one hand to Hemi- blabera Saussure, and on the other to Monachoda Burmeister." The male genitalia of Aspiduchus (Figs. 21-23) indicate a close rela- tionship to hiblabera (Figs. 17-20) but less so to Monachoda (Figs. 142, 144, 145) which I place in the Monastriini (see below). Rehn's ( 195 I, p. 7) drawings of the L2d's of Asp'iduchus cawed- cola (Fig. 100) and A. borinquen (Figs. 97, 103, 106) show rela- tively 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. distant! (Figs. 69, 72) than in E. posticus (Figs. 66, 73), as noted by Princis ( I 950). Hebard ( 1920, p. I I 6) 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), Adinablatta (Fig. 28).
In this group Lad is not hooksha~ed. Rehn (1940) stated that Minablatta is related to Blaptica and Eublaberus but is rob ably more closely related to the former. The absence of a hookshaped L2d in Minahlatta (Fig. 109) places the genus closer to Blaptica. However, the elongated preputial spines, occurring in a single row in Minablatta (Fig. 109) show a similarity to the spines found in &iduchus (Fig. 106), Byrsotria (Fig. 75), and Hemiblabera (Fig. 96).
~Hyporhicnoda has been placed in the Epilamprinae by Hebard ( 1920)) Princis ( I 960), and McKittrick ( 1964). However, the prepuce of the male has a fringe of spines (Figs. I 12, 1 I 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 go genera of Blaberidae. In Hyporhicnoda the apex of Lsvm is more darkly pigmented than L2d so that the junction of the 2 sclerites is readily discernible.
2. Monastriini : Hiereoblatta (Fig. 126), Monachoda (Figs, 123-1 25 ) , Monastria (Figs. 12 1-1 22 ) . Peinsodes (Figs. I I 8- I 20).



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