Louis M. Roth.
The Male Genitalia of Blattaria. I. Blaberus Spp. (Blaberidae: Blaberinae).
Psyche 76:217-250, 1969.

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PSYCHE
Vol. 76 September, I 969 No. 3
THE MALE GENITALIA OF BLATTARIA.
I. BLABERUS SPP.
(BLABERIDAE : BLABERINAE)
BY Lours M. ROTH
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01 760
"Like many other genera the forms of which are variable and the specific features hard to ascertain and express, the genus Blaberus has been a despair to the systematist." (Rehn and Hebard, 1927). The use of male genitalia, specifically the aedeagus and preputial spines, has helped to alleviate some of the taxonomic difficulties of several species of Blaberus. Burmeister ( 1838 ; in Princis, 1946) first mentioned the preputial spines in Blaberus trapezoideus Bur- meister and Hebard ( I 91 7) described them in Blaberus craniifer Burmeister and B. atropos (Stoll). Princis (1946) illustrated the aedeagus and prepuce of the following species of Blaberus: giganteus (Linn.), trapezoideus, craniifer atropos, discoidalis Serville, para- bolicus Walker, anisitsi Brancsik, and bolh~iensis Princis. Lefeuvre ( 1960) illustrated the genitalia of craniifer, Quiaoit ( 1961) de- scribed them for craniifer and giganteus, and McKittrick (1964) illustrated discoidalis.
With the exception of Princis (1946) and Lefeuvre (1960), in- traspecific variations were not mentioned by the above workers. I have found considerable more variation in Blaberus genitalia than was indicated by Princis and Lefeuvre. In this paper I shall illus- trate the male genitalia of 12 species of Blaberus, describe group and specific differences, including intraspecific variations, and discuss the probable evolution of the aedeagus and prepuce in this genus. MATERIALS AND METHODS
The following 5 species of Blaberus were available in cultures:



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218 Psyche [September
craniifer, giganteus, paraholicus, atropos, and discoidalis. In addition I have examined the genitalia of museum specimens of these species as well as those of B. co/osseus (Illiger), B. brasilianus Saussure, B. minor Saussure, B. fusiformis Walker, B. scutatus Saussure and Zehntner, B. anisitsi Brancsik, and B. boliviensis Princis. Of the 14
Blahus listed by Princis ( I 963) assellus (Thunb.) and latissimus (Herbst) were described from nymphs and are questionable species. For reasons given below, I consider Blaberus colosseus, which Hebard ( 1921 ) synonymized with B. giganteus, a valid species and B. trape- zoideus a synonym of B. craniifer.
As suggested by Princis (1946) the tips of the abdomens of dried specimens were dipped in hot water for about a minute, or the speci- mens were placed in a relaxing chamber. Once softened, the ab- domen was slit along the lateral membranes and the genitalia were removed usually without serious damage to the subgenital or supra anal plates. All specimens were treated with 10% KOH, cleared, and mounted in Permount. The hooked right phallomeres were mounted ventral side up and phallomeres LI and L2d were mounted dorsal side uppermost. The preparations of the prepuce were spread and flattened to show the spines. This should be taken into account when examining the illustrations. Normally the prepuce partly en- velopes L2d (see Fig. 121 in McKittrick's 1964 monograph which illustrates the folding of the prepuce in B. discoidalis). Although the principal genitalic characters used are L2d and the prepuce, I have also included photographs of R2 and LI for compara- tive purposes. Although these 2 phallomeres are very similar or have minor differences in all species of Blaberus (Figs. 1-24) they show family or subfamily differences and their inclusion should be useful in future studies of the genitalia of Blaberidae. Wherever known I have given locality data for the illustrated specimens, and the identity of the entomologist who determined the species. The abbreviations for the sources of this material are as follows (original geographical source, if known, follows the abbre- viations in the explanation of figures) : (N) = Natick culture ; (ANSP) = Academy Natural Sciences, Philadelphia; (IVICZ) =
Museum of Comparative Zoology, Harvard University; (L) = Zoological Institute, Lund, Sweden; (AMNH) = American Mu- seum of Natural History; (USNM) = United States National Museum ; ( BMNH) = British Museum (Natural History). Slides of genitalia are deposited with their respective males in the above museums.




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19691 Roth - Genitalia of Blattaria 219
RESULTS AND DISCUSSION
The male genitalia of Blaberidae consist of 3 main structures ( McKittrick7s, I 964 terminology) . The right phallomere (R2) (Figs. 1-12) is a retractable hook, and all the species have a sub- apical incision (Fig. 2, arrow). The median sclerite (Lavm) is solidly attached (in Blaberus spp.) to L2d (L2 dorsal = the virga, penis, or aedeagus) (Fig. 52). The prepuce (mantle of Hebard's, 1917 terminology) is a soft, flexible membrane (Fig. 52) bearing characteristic spines, or truncate or rounded sclerotized elevations. The left side of the prepuce is solidly attached by sclerotization to the side of the L2 phallomere, whereas the right side is usually connected by a flexible membrane which permits it to fold partly around the virga.
One of the sclerites (LI) (Figs. 13-24) of the left phallomere in all the species of Blaberus studied are more or less similar and have a heavily sclerotized cleft, noted by McKittrick (1964).
Based on body size, color, and shape of the pronoturn, Hebard ( I 93 I ) placed fuxif ormis, brasilianus, anisitsi, and scutatzis in the Bra~ilian~us Group of the genus. Princis ( 1946) divided 8 species into the Giganteus and Atropos Groups, basing his divisions on the shape of L2d and the spines present on the prepuce. He did not examine the genitalia of minor, brasilianus, fusiformis, and scutatus, but suggested that the Brasilianus Group, established by Hebard, probably should be included in the Atropos Group. As a result of my examination of the 4 species not investigated by Princis, I believe Hebard was correct in erecting the Brasilianus Group and I place the above 4 species in this group.
Species of Blaberus can be readily b laced in their respective Groups, by the shape of the virga and preputial spines. However, variation is such that specific determination is often difficult. The 3 Blaberus Groups may be distinguished in the following key: I. L2d recurved dorsally and slightly to the right, usually forming a hooklike structure (Fig. 52). Extending dorsally and later- ally on the left, about where Lavm and Lad are solidly joined, is a sclerotized tumorlike outgrowth (Fig, 52, T). There are no large truncate elevations and all of the spines on the prepu- tial membrane are relatively small (Figs. 28 -40, 47-57, 62-74, 215 Giganteus Group.
L2d not hookshaped. Tumorlike outgrowth on the left side ab- sent. Prepuce with anterior truncate or rounded elevations almost always present on the left, and sometimes on the right side of the preputial membrane
2.




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220 Psyche [September
Figs. 1-12. Right hooked phallamere (R2) of Blaberus spp. 1-2. GigaÌöteu Group. 1. B. crani~fer. (ANSP), Juxtiahuaca Cave, Colotlipa, Mexico (Jet. as B. irapezoidrv.~ by Rehn). 2. B. gigastev-s. (ANSP), Muzo, Colom- bia (arrow indicates the subapical incision). 3-6. bras ilia nu^ Group. 3. B.
scututux.
(ANSF), Ceara Mirim, Rio Grande do Norte, Brazil. 4. B. fu~i- formis. (ANSP), Provincia Sara, Dept Vera Cruz, Bolivia. 5. B, braat- lianus. (ANSI'), Natal, Brazil. 6. 5. minor. (ANSP) , Miiisian Tacaagk, Formosa, Argentina (det. Hebard). 7-10. Atropes Group. 7. B. atropos, (MCZ), Mexico. 8. 3. fafahvl'icus. (N), Ecuador. 9. B. discoidaff;. {N), Panama. 10. R. boSw'cn~i~. {L), Guayaquil, Ecuador (det. Princis). 11-12. Brua'ilianus Group. 11. B. cotosseus. (ANSP), Fyzabad, Trinidad. 12. 5. sp. D (probably ~olossfits). (ANSP), St. Laurent du Maroni, French Guiana (from specimen shown in Fig. 208). (all to scale shown in Fig. 12).



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19691 Roth - Genitalia of Biattwia 22 I
Figs. 13-24, Left phallomere (Ll) of Maim spp. 13-14. Gtgantfvs Group, 13. 3. g'tganietts, (ANSP), Muzo, Colombia. 14. S. (raniijer (ANSP), Juxtlahuaca Cave, ColotSipa, Mexico (det. as B. trapemideus by Rehn). 15-18, 22. Brasitiunw Group. IS. B. brus'tl'iantis. (ANSP), Natal, Brazil. 16. B. scufahs. (ANSP), Ceara Mirim, Rio Grande do Norte, Brazil, 17. R. minor, (ANSP), Villa Ana F.C.S.F., Argentine Republic (det. Hebard). 18. S, fiiiformis. fANSP), Santa Cruz de 5a Sierra, Bolivia (det. Hebard). 19-21, 23, 24. Atropus Group. 19. B. faraffolicus. (N), Ecuador. 20. 3. discoidalis (N), Panama. 21. h'. atropus. (N), Trini- dad. 22. B. caiossw, (ANSP), Caparo, Trinidad (det. Hebard). 23, B. a~isitsi. (L), (det. Princis) . 24. B. bo/i&ensis. (L), Guayaquil, Ecuador (det. Princis). (all to scale shown in Fig. 18).



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222 Psyche [September
Fig. 35. Upper. Tcgmen of B. craniifrr showing the distribution of the long setae on the marginal and acaptilar fie!& Lower. Anterior view of part of the tegmen showing the characteristic hairlike setae. 2. Anterior elevations usually rounded, present on the left and often on the right sides. Differences in size between anterior eleva- tions on the right and left sides not great. Preputial spines numerous, usually on the left and right sides and often occur in more than a single row (Figs. 76, 77, 79-82, 84-91, 93-1 I I, 21 1-214). In colossm the left preputial spines usually occur in a single row (Figs. 1 16-129) .. ,. Brasilianus Group. Truncate or rounded elevations usually present only on the left side and generally much larger and more robust than spines on the right. Preputial spines usually less numerous than in the Brasilianus Group, and are often arranged (when present) in a single row on the left, and single or sometimes double or ~artial double row on the right:. Spines on the right side usually more numerous than on the left (Figs. 133-153, 155-170, 174- 198, 200, 201, 203, 204, 210) .. . Atropos Group. Giganteus Group. - Two species of Blaberus hieanteus. Figs. 26, 27; craniifer, Figs. 41-46, 58-61) belong to this group. A useful tegminal character which Princis (1946) described can be used to distinguish B. craniifer from all other species of the genus. In
craniifer the marginal field and scapular field of the tegmina have diffuse projecting hairs (Fig, 25). According to Pincis, no other



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19691 Roth - Genitalia of Blattaria 223
Figs. 25-44. 5. giganitus (Giganteus Gioup). 26. (N). 27. (ITSNM), St. Jean, French Guiana (det. as B. coloiseus by Hebard) (scale =z 10 mm). 28-40. L2d and prepuce (all to scale shown in Fig. 34). 28. (CSNM)
(from specimen shown in Fig. 27). 29. (USNM), Cablma, Panama. 30.
(USNM), Ft. Clayton, Canal Zone. 31. (AMNH), Barro Cobrado Island, Canal Zone. 32. (USNM), Chilibrillo Cave, Buenos Aires, Canal Zone. 33. (USNM), Puerto Berrio, Colombia, 34-35. (ANSP), Muzo, Colombia. 36. (USNM) , Atlantico, Colombia. 37. (AMNH) , Colombia. 38. ( AMNH) , Caripito, Venezuela. 39-40. (N) .




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Figs. 41-57. B. rraniifa- (Gigan~us Group). 43-46. Adult males (scale 10 mm). 41. (USNM), Cordoba. Mexico (det, as B, trafmoidnus by Rehn). 42. (N). 43, (USNM), Rancho Qernado, Re. $5, Mexico. 44. (USNM), Teapa, Tabasco, Mexico. 45. (USNM), Key West, Florida. 46. (N), 47-57. L2d and prepuce (all to scale shown in Fig. 51). 47. (USNM) , Mexico (from specimen shown in Fig. 41). 48. (USMM), Mexico



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19691 Roth - Genitalia of Blattaria 225
Blaberus has this character, although I have seen a few very minute hairs in some specimens of B. giganteus. In craniifer the tumorlike sclei-otized outgrowth on the left side of L2 extends caudally for a short distance and usually merges gradually into the border of the prepuce where small spines begin and form a fringe around the membrane. The sclerotized extension of the lateral outgrowth varies somewhat in length but is generally distinct (Figs. 47-51, 53-57, 62-64, 66-74). Exceptions are seen in Fig. 52 and 65. In B. giganteus there is usually little or no sclerotized extension from the tumorlike outgrowth into the preputial membrane, so' that the preputial spines begin more abruptly at the outgrowth (Figs. 28-35, 38-40). Exceptions to this are shown in Fig. 36 and 37. In both species the preputial spines may occur in more than a single row, and in some individuals there may be a reduction in the number of preputial spines (Figs. 37, 65, 70, 74). If Princis' (1946) tegminal character is valid for B. craniifer then I have not seen any correctly determined specimens of B. trapezoideus. All the specimens determined by Hebard or Rehn as tra?ezoide,us (Figs. 41, 58-61 ) have hairy tegmina and I therefore consider them to be light forms of B. craniifer. The genitalia of these "trapezoideus" (e.g., Figs. 47, 63, 64, 66) are indistinguishable from craniifer. Two specimens identified as B. trapezoideus, received from the University Zoological Museum, Copenhagen, Denmark were actually B. craniif er (Costa Rica) and B, parabolicus (Peru). According to Princis (1946) the preputial spines of trapezoideus are similar to giganteus but are smaller and more numerous, though always clearly separated from each other. Considering the variation in size, number, and spacing of preputial spines I doubt if this character can be used to distinguish trapezoideus from giganteus. Princis also (1958) states that the pronotum of trupezoideus is laterally truncated with approxi- mately parallel sides. Some individuals of our light-phased form of craniifer (in culture) (Fig. 42) also have the pronotum laterally truncate. It is possible that trapezoideus and craniifer are simply variants of the same species. The type localities of the 2 spp. of thme (from specimen shown in Fig. 43). 49. (USNM), Vera Cruz, Mexico. 50. (USNM), Tuxtepec, Oaxaca, Mexico (labelled trapezoideus) . 51. (USNM), Mexico (from specimen shown in Fig. 44). 52. (MCZ), Chichen Itza, Yuca- tan, Mexico. 53. (USNM), El Salvador. 54. (USNM), San Salvador, El Salvador. 55. (USNM), Benque Viejo, British Honduras. 56. (MCZ), Colon, Panama. (Abbreviations for Fig. 52. L2d dorsal left phallomere; L2vm = ventromedial left sclerite (fused to L2d) ; P = prepuce; T = tumorlike outgrowth.)




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226 Psyche [September
FIRS. 58-74. B. cmnnfrr [Giganteus Group). 58-61. Adult malcq (scale = 10 mm). 5s. (ANSI'), Guatemala. 59. (ANSP), Vera Cruz, Mexico. 60. (ANSP). 61, (ANSP), Juxtlahuaca Cave, Colotlipa, Mexico. (These 4 males were determined as B. trupzoidcus by Rehn, or Hebard.) 62. (MCZ), Havana, Cuba (det. Rehn). 63. (USNM), Santiago-Vegas, Cuba. 64. (ANSP) (from specimen shown in Fig. 60). 65. (AMNH), Turrialba, Costa Rica. 66. (ANSP) (from specimen shown in Fig. 61). 67. (MCZ), Havana, Cuba. 68-69. (MCZ), Col6n, Panama. 70. (N). 71, (MCZ). 72. {ANSP), San Miguel, Vera Paz, Guatemala (det. as B. cola~si-us by Hebard). 73. (USNM), Florida (from specimen shown in Fig. 45). 74. (3).




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19691 Roth - Genitalia of Blattaria 227
Gzganteus Group are : giganteus - "America" ; craniifer - Cuba. (see addendum regarding trafiezoideus) . Although color markings are variable in several species of Blaberus, they are especially so in B. craniifer. This species is represented by light (Figs. 41-43) and dark (Figs. 45, 46) forms with intermediates (Fig. 44) connecting the extremes. Markings of light phased individ- uals resemble B. giganteus (cf. Figs. 26, 27). Lefeuvre ( 1960) has described some color varieties which occurred in his laboratory culture of craniifer and we have cultures of both light and dark forms which crossed successfully. Lefeuvre claims that rearing craniifer for a number of years favored the formation of an "artificial subspecies'' which differed from the original in I) coloration of the pronoturn, 2) general coloration, in particular the male, and 3) the morphology of the penis and prepuce.
Lefeuvre suggested that the original B.
craniifer may have hybridized with a cIoseIy related species. The variations in craniifer which Lefeuvre described can be seen in museum specimens from different geog;-aphic localities. I have never seen any dark forms of B. giganteus comparable to dark craniifer. Brasilianus Group. - The 5 species in this Group, namely, scutatus (Fig. 75) ) brailianus (Fig. 781, fusiformis (Fig. 83), minor (Fig. 92), and colosseus (Figs. 112-1 15), show some genitalic differences in L2d and preputial spines but variation is so great within 4 of these species (Figs. 76, 77, 79-82, 84-91, 93-1 I I ) , that specific determina- tions, using genitalia alone, are often imposible. In some individuals of fusiformis (Figs, 85-87, 89, 91 ) and minor (Figs. 94-95> 100, 106) there is a marked reduction or loss of preputial spines and they may occur in a single row, usually on the left side. Characteristic of this group is the anterior elevations which are generally fused on the right side, and are about the same size as those on the left.
Rarely are the anterior spines on the right larger than those on the left (e.g., Figs. 84, 85, 88). The preputial spines de- crease only slightly in size from the anterior to posterior position. When the spines are numerous and occur in more than a single row they are often closely spaced and form a more or less dense uniform fringe around the preputial membrane (e.g.) Figs. 76, 77, 79-82, 88, 90) 93, 97, 99, 101) 105)
Hebard ( 1921, pa 148) stated) "From a study of the material in the Philadelphia collections, as well as specimens recently received from the Guianas, we are finally convinced that B. colosseus (Illiger) was based on a mere individual variation of giganteus, unworthy of nominal recognition." Princis ( I 963) followed Hebard and listed



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Figs. 75-82. Bm~ilium~ Group. 75-77. B, scutatux, 75-76, ( ANSP), Pernambo, Brazil (paratype of 3. ~mtuha var. obscura S. and 2.). 77. (ANSP), Ceara Mirim, Rio Grande do Norte, Brazil. 78-82, B. .husi~ianus. 78-79. (ANSP), Natal, Brazil. SO. (ANSP), Independencia, Parahybq Brazil. gl. (MCZ), Brazil. 82. (USNM), Eatal, Brazil. (Scale for addm =z 10 mm; all genitalia to scale shown in Fig. 79.)



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Figs. 83-91.
E. ju~ijurmi~ (Bradiunus Group). 83. Adult male (scale = 10 mm). (ANSP), Santa Cruz de la Sierra, Bolivia (det. Hebard). 84-91.
L2d and prepuce (all to scale shown in Fig. 85). 84. (ANSP), Carurnbo, Matto Grosso, Brazil (a portion of the prepuce on the right side is missing). 8.5. (ANSF), San Francisco, Argentina. 86. (ANSP), Provincia Sara, Dept. Vera
Cruz, Bolivia (det. Hebard).
87. (ANSP),
Bolivia (from specimen shown in Fig, 83). 88. (ANSP), Provincia Sara, Dept. Vera Cruz, Bolivia. g9. (ANSP), Jundiahy, Brazil. 90. (USNM), Wtiariti KO, Matto Grosso, Brazil. 91. (ANSP).



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Psyche
[September
Figs. 92-107.
B. minor (Bradi~nus Group). 92. Adult male (scale = I0 mm). (ANSF), Argentina (det, Hebard). 93-107. L2d and prepuce (all to scale shown in Fig. 93). 93. (ANSPj, Paraguay. 94. (USNM), Natal, Brazil. 95. (USNM), Bmoklin, S5o Paula, Brazil, 96. (ANSP), Mission Tacaaglk, Formosa, Argentina. 97-98. (ANSF), Formosa, Argen- tina. 99. (ANSI'), Argentina (from specimen shown in Fig. 92). 100-102. (ANSP), Cham del Santiago del Ester0 Bords du Rio Selsdo Environs D'kaiio, Argentina. 103. { ANSP) , Formosa, Argentina. 104. (ANSF'). 105. (ANSP), Chaco de Santiago del Estero Rio Saiado, Argentina. 106-1 07. (ANSP) , Gran Chaco, Argentina.



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19691 Roth - Genitaih of Bhttm-iu 23 1
Figs. 108-111.
Brmiliunw Group. Right and left sides of the prepuce. 108. 3. $cutatus (from Fig. 76). 109. B. bmdian~ds (from Fig. 79). Ilk B. minor (from Fig. 99). 111. 3. fu~ijormis (from Fig. 87). (scale = 0.1 mm).




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232 Psyche [September
Figs. 112-122. B. colosseus (Brasilianuf Group). 112-1 15. Adult males (scale = 10 mm). 112. (ANSI'), Caparo, Trinidad. (This specimen is Fig. 4 in Hebard, 1916.) 113. (ANSP), Fyzabad, Trinidad. 114. (ANSP), Caparo, Trinidad, (This specimen is Fig. 5 in Hebard, 1916.) 115. {MCZ), Mexico. 116-122. L2d and prepuce (scale =: 0.2 mm). 116. (ANSP) (from specimen shown in Fig. 112). 117. (ANSP) (from specimen shown in Fig. 113). 118. (MCZ) (from specimen shown in Fig. 115). 31 19. (MCZ), Mexico. 120. (USNM), Trinidad, 121-122. (ANSP) (from specimen shown in Fig. 114; L2d and prepuce have been mounted separately).



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19691 Roth - Genitaiia of Blattaria 233
Figs. 123-129, B. calosstus (Brasilianw Group). 123. (AMNH), Trini- dad. 1%. (USNM), Trinidad (some of the anterior elevations on the left side are broken off). 125. (AMNH), Rancho Grande, near Maracay, Vene- zuela. 126-129. Right and left sides of the prepuce. 126. (MCZ) (from Fig. 119). 127 (ANSP) (from Fig, 116). 128. (ANSP) (from Fig. 117). 129. (MCZ) (from Fig. US) (scale = 0.2 mm).



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234 Psyche [September
Figs. 130-141. 3. atrapos (4fropss Group). 130-132. Adult males (scale == 10 mm). 130. (N), Trinidad. 131. (MCZ), Mexico. 132, (USNM), Colombia (taken in quarantine on bananas at Charleston, S.C.). 133-141. L2d and prepuce (all to scale shown in Fig, 140). 133, (MCZ) (from specimen shown in Fig. 131). 134. (USNM) (from specimen shown in Fig. 132). 135. (USNM), Georgetown, British Guiana, 136-138. (USNM), Trinidad. 139-141. (N), Trinidad.




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Figs, 142-159. Prepuce of B. alrofin.
From Narick culture which ori-
ginated in TrSridad (ail to scale ahown in Pig. 142). - *
sA
4.
I




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Psyche
-.
[September
Figs. IS+-165. 3. parubolicris (.'3iropos Group). 154. Adult male. (N), Puraquequara, Rio Negro, Amazonas, Brazil (scale = 10 mm). 155-165.