Louis M. Roth.
The Male Genitalia of Blattaria. X. Blaberidae. Pycnoscelus, Stilpnoblatta, Proscratea (Pycnoscelinae), and Diploptera (Diplopterinae).
Psyche 80:249-264, 1973.

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THE MALE GENITALIA OF BLATTARIA.
X. BLABERIDAE. PYCNOSCELUS, STILPNOBLATTA, PROSCRATEA (PYCNOSCELINAE), AND
DIPLOPTERA (DIPLOPTERINAE) .*
BY Louis M. ROTH
Pioneering Research Laboratory
U.S. Army Natick Laboratories
Natick, Massachusetts 01 760
McKittrick ( 1964) grouped the Pycnoscelinae, Diplopterinac, Panchlorinae, and Oxyhaloinae, in the Fanchloroid Complex of Blaberidae. The male genitalia of the latter two subfamilies have been described (Roth, 1971a, 1g71b). In this paper I shall illustrate the male genitalia of several species of Pycnoscelinae and two' species of Diplopterinae.
The genitalia were treated with 10% KOH and mounted in Permount. The source of each of the specimens illustrated is given using the following abbreviations: (ANSP) = Academy of Natural Sciences, Philadelphia; (BMNH) = British Museum (Natural History) ; (L) = Zoological Institute, Lund, Sweden; (MCZ) = Museum of Comparative Zoology, Harvard University; (VM) = Vienna Museum Natural History, Vienna, Austria. Geographical collection data and the names of specialists who identified the speci- mens, if known, follow these abbreviations. The number preceding the abbreviation refers to the number assigned to the specimen and its corresponding genitalia (on a slide) which are deposited in their respective museums.
RESULTS AND DISCUSSION
Pycnoscelinae
Pycnoscelus surinamensis (Linn.) is the type species but it is parthenogenetic and normally only exists as females. P. indicus (Fab.) is bisexual and apparently the parent stock from which surinamensis arose. Occasionally parthenogenetic males occur in cultures of surinamensis but thev are non-functional when mated to parthenogenetic females (Roth, 1967).
*Manuscript received by the editor August 27, 1973



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19731 Roth - Blattaria 25 1
The arrangement of the male genital phallomeres is shown in figure I. In Pycnloscelus indicus all 3 phallomeres are well devel- oped and sclerotized (Figs. 7-9). The L2d is separated from L2vm; the obliquely more or less truncate ends of these 2 sclerotized struc- tures (Figs. 7, ID, 21, 24, 31 ) have the appearance of having been broken off and separated from L2vm. The outer lower curved por- tion of L2d is spicular (Figs. I, 7, 10-13), and the underlying prepuce is densely "hairy" but otherwise not unusually shaped (Fig. 7, P). The curved genital hook (R2) lacks a subapical incision, is heavily sclerotized, somewhat truncate at the apex, the inner curved margin with (Figs. 8, I 4-1 8) or without (Figs. I 9-20) small projections. The LI is very well developed with the cleft turned upward and its margins heavily sclerotized (Figs. I, 9). The genitalia of Pycnoscelus surinamensis (Figs. 2 1-25), and Pyncoscdus nigra ( Brunner ) (Figs. 3 1-33) are indistinguishable from those of P. indicus. Habitus photographs of P. indicus and P. nigra are shown in figures 2 and 4.
The shapes of the L2d and R2 (Figs. 26, 27, 29, 30) of Pycnos- celus semivitreus Princis (Fig. 3) differ from those structures in P. indicus, P. surinamensis, and P. nigra; however, the shape of Li in all 4 of the above species is similar (cf. Figs. 9, 23, 28, 33 (dis- torted in preparation) ). Two of the genital phallomeres of Pycnos- celus striata (Kirby) are distinguishable from those of the other species of the genus. Its L2d (Fig. 34) differs in shape, lacks the spicular surface characteristic of the outer lower curved region and is only slightly separated from L2vm (cf. Fig. 31). The curved
portion of R2 (Fig. 35) of striata is more elongate and slender than in semiwitreus (Figs. 27, 29) and more uniform in width than in P. indicus (Figs. 14-20) or nigra (Fig. 32). Princis (1964) included Stilfinoblatta in the Pycnoscelidae and the genitalia of S. opaca (Walker) (Figs. 37-39) tends to support this conclusion, though I relegate his family to subfamily rank (McKittrick, 1964).
Especially notable is the marked similarity in appearance of the LI of Stilfinoblatta (Fig. 39) with those of- Pycnoscelus (Figs. 23, 36). The L2d (Fie. 37) of S. ~aca is. greatly reduced and irregular in outline and is widely separated from Fig. 1. Male genitalia (dorsal view). Top. (106 MCZ). Pycnoscelui indieus. Zamboanga, Philippine Islands, (det. Roth). Bottom. (70 BMNH) . Proseratea complanata. S2o Gabriel, Rio Negro, Brazil, 27.1X.1927, J. F. Zikan. (Ll = first sclerite of left phallomere ; L2d1 dorsal sclerite of L2 ;.
L2vm = ventromedial sclerite ; M saclike membrane above L2d ; R2 = hooked sclerite of right phallomere).




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Figs. 2-6. Males of Fytnoscelinae. 2. Pymnsteins ittdicits. Sakaerat District, Thailand. From a laboratory colony. 3. (100 MCZ). Fycnawtu~ fernwitreus, Manila, Philippine Islands. (det. Primis). This species was described (Princia, 1967, p. 148) from 2 8 $ torn Java, Depofe. 4. (25 BMNH). Pymoscelui niffra. Southwest China, Yunaao, Ho-an (leg. J. W. Gregory, 26.VJ932). 5. (1447 L). Pymo~celus $triufa. Kariorang, Borneo, (det. Princis). 6. (144 ANSP). Stitfinobiatia opaca. Butawa, Modera, S. P. Ceylon (det. by Hebard a; S, besgalmsis (Sauas) ). (scale = S mm).



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Roth - Blattaria 253
Figs. 7-20. Male genital phallomeres of Pymoscelw mdicus. 7-9. L2d, R2, and Ll from a specimen originating from Hawaii (laboratory culture). P = prepuce. 10-13. LZd's. 10. Sakaerat District, Thailand (laboratory culture). 11. (101 MCZ), Paaay, Philippine Island's. 12-13. Hawaii (lab- oratory culture). 14-20. R2's. 14. Sakaerat District, "Thailand (laboratory culture). 15-19. Hawaii (laboratory culture). 20. f 101 MCZ) , Pasay, Philippine Islands.
(scak = 0.2 mm; figures 10-13 magnified to scale shown in fig. 7 ; figures 14-20 magnified to scale &own in fig. 8).



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254 Psyche [September
Figs. 21-30. Male genital phallomeres of Pymowlus spp. 21-25. P. sari- nammsk. Two males which occurred in a culture originating front Ftaser Island, Australia. 26-30. P. tdvUreuf. 26-28, (102 MCZ). Manila, PhiIip- pine Islands (de~ Princis). 29-30. (100 MCZ). From 3 shown in figure 3. (scale = 0.2 mm; figures 25, 27, 29 magnified to scale shown in fig. 22).



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19733 Roth - Blattaria 355
Figs. 31-39. Male genital phailomeres of Pycnoscelinae, 31-33. (25 BMNH). Pycnosceltts nigra; LI (Fig. 33) was distorted in preparation, (from $ shown in figure 4). 34-36. (1447 I). Pycnoicdua Nrhta (from 8 shown in figure 5). 37-39. (144 ANSP). SUlfnoblatfa opaca (from 8 ihown in Sgure 6). (@cafe = 0.2 mm). ,




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256 Psyche
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Lavm. The R2 (Fig. 38) is broad, relatively short and, as in Py cnoscelus, lacks a subapical incision. Princis ( 1964) lists 4 valid species of Proscraiea: peruanu Sauss., inequalts (Walker), funebris Burmeister, and complanata (Perty). Bmnner ( 1865) synonymized P. peruana with P. wmpiamta with a ( ?) and Kirby ( 1904) listed them as synonyms. Hebard (1926) placed funebris as a synonym of cwnplanata, but Rehn (1932) felt that this was not warranted until additional information became available. Princis ( I 963 ) concluded that the above synonymies were incorrect, and showed differences in pronotal shapes and color mark- ings of the 4 species. I have examined the type of P. inequalu and find that its genitalia are so different from those of P. complanafa (type of genus) that it undoubtedly does not belong to this genus. I collected P. complunata (det. by Gurney) in Brazil and estab- lished a colony which was maintained for several years at the Natick Laboratories. Habitus figures of adults and a nymph (from the cul- ture) are shown in figures 40-44. The adult pronotal markings may vary (Figs. 40-42) (see Rehn, 1932, p. 71 ) and the pattern of the specimen shown in figure 42 resembles that shown by Princis ( 1963, p. 148) for funebrIS. The specimen provisionally determined by Rehn as peruana has pronotal markings (Fig. 45) similar to complanata. Princis ( 1964) placed Panchlora, Proscratea, and Phorfioecoides in the Panchloridae. Male genitalia clearly place Phortioecoides in die Zetoborinae { Blaberidae) (Roth, 19703). The male phallomeres of the Panchlorinae (5 genera) are notable for their marked reduc- tion or absence (Roth, 1971 b, Gurney and Roth, 1972). The geni- talia of Proscrai~u are not characteristic of the Panchlorinae. Hebard (1926) and Rehn (1932) believed that Proscratea be- longed to that section of the Perisphaeriinae which included Para- nauphoeta Brunner and its allies. I have examined the male genitalia of 6 species of Paranauphoeta and all 3 phallomeres differ markedly from those of Proscratea.
McKittrick (personal communication) placed Proscrutea in the Diplopterinae. However, the shape of the Lad and R2 in Proscratea are more like those of Pycnoscelus and I tentatively place Proscratea in the Pycnoscelinae. McKittrick ( I 964) shows Diplopiera, Leuro- lesfes, and Pycnoscelus, as arising from a common stock. Phoetalia (^ Leurolestes) which she placed with Diploptera in the Diplop- terinae belongs to the Blaberinae (Roth, ig70b). The shape of the spermatophore of Proscratea con~pZanata looks like a bowling pin and strongly resembles the spermatophore of Diploptera suggesting a relationship between these 2 genera. However, other genera in



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Figs. 40-45. Proferatea spp. 40-44. P. com$lanata. From laboratory colony which originated from Serra Tamendaui, Rio Negro, Amazonas. 4-0. Bra- chypterous mate. 41-42. Macropterous males. 43. Brachypteroua female. 44. Female nymph. 45. (129 ANSP). P. "peruund' $, Hacienda San Juan, Colonio Peren&, Peru, June 23, 1920; Come11 Univ. Exped. (det. Rehn). (scale = 5 mm; scale for figures 40-44 shown in fig. 43).



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Figs. 415-57. Male genital phallomeres of Proscratea spp. 46-48. (129 ANSP). P. "fwiana" (from 8 shown in feure 45). 49-57. P. comphata. 49-51. Tapurucuara, Rio Negro, Amawnaa. 53, 56'. Urucurutuba, Rio Madeira, Amamas. 54, 57. Serra Tamendaui, Xo Negro, Amazonai. (scale = 0.3 mm).




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19731 Roth - Blattaria
259
different subfamilies (e.g., Capucina [Zetoborinae,] Nauphoeta and Gromphadorhina [Oxyhaloinae] ) have spermatophores (Graves, 1969) somewhat similar in shape to those of Diploptera and Pros- crated.
The L2d of Proscratea is well developed, somewhat crescent- shaped (but variable) and widely separated from Lavm (Figs. 46, 49, 52-54). The membrane above Lad is modified to form a sac-like projection whose surface is covered with microspicules; this structure is not found in the other Pycnoscelinae. The R2 is short, stout, and lacks a subapical incision (Figs. 47, 50, 55-57). The shape of the
cleft in the LI of Proscratea does not curve upwards (Figs. 48, 5 I ) as it does in Pycnoscelus (Fig. 9) or Stilpnoblatta (Fig. 39). The genitalia of Proscratea oomplanata (Figs. 46-48) are indistinguishable from the specimen provisionally determined by Rehn ( 1932, pp. 71- 72) to be Proscratea peruana (Figs. 49-5 I ) . Because of the differences between the LI and the presence of the modified membrane over the L2d of Proscratea, I suggest 2 tribes in this subfamily :
I. Pycnoscelini : Py cnoscelus, Stilpno blatta 2. Proscrateini : Proscratea
Princis (1965) placed the genera Diploptera and Diplopterina in the Diplopteridae. The genitalia of Diplopterina are closer to cer- tain members of the Perisphaeriinae (unpublished observations) and I consider it to be a member of this subfamily. McKittrick ( 1964) included 2 genera, Diploptera and Phoetalia (= Leurolestes) in Diplopterinae. As indicated above, the genitalia of Phoetalia place it in the Blaberinae (Roth, 197ob). Diploptera punctata (Esch- scholtz) is the only viviparous cockroach known and, at present, I consider this genus to be the only member of the Diplopterinae. Other members of the Blaberidae, whose reproduction has been in- vestigated, are ovoviviparous. Princis (1965) lists 7 species of Diploptera and it would be of interest to determine if those, other than punctata, are viviparous also.
The arrangement of the phallomeres of Diploptera minor Brunner is shown in figure 78. The male genital phallomeres of D. punctata (type of genus) are shown in figures 60-62. The curved hook (R2) lacks a subapical incision, is more slender and elongate, and more strongly curved (Fig. 61) than in most of the species of Pycnos- celinae. The inner margin of the curved portion of the hook has



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260 Psyche [September
Figs. 58-67. Supra-anal and aubgenital plates, and genital phatlomeres of male Diploptera spp, 58-62, D. pmictata (from laboratory colony originating in Hawaii). 58. Supra-anal plate (dorsal). 59. Subgenital plate (ventral). 60-62. Genital phallomeres. 63-67. (67 BMNH). D. ap., Honolulu, Hawaii. 63. Supra-anal plate (dorsal), 64, Subgenital plate (ventral). 65-67. Genital phallomeres. (scale: supra-anal and subgenital plates "- 0.5 nun; phallo- mere9 = 0.2 mm).




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Figs. 63-77. Supra-anal and subgenital plates, and genital phallomeres of Diploptera app. 68-72. (68 BMNH). Samoa Island. 68. Supra-anal plate (dorsal). 69. Subgenita! plate (ventrai). 70-72. Genital phallomeres. 73- 77. (69 BMNH). Henderson Island, South Pacific. 73. Supra-anal plate (dorsal), 74. Subgenital plate (ventral). 75-77. Genital phatlomeres. (scale: supra-anal and subgenital plates = 0.5 mm; phhllomeres 0.2 mm).



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262 Psyche
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Figs. 78-79. (17 VM). Difloptera minor. Type 8, Philippines. 78. Genitalia and subgenital plate (SP) (ventral), E' = prepuce ; other abbe+ viations as in figure 1.
79. Supra-anal plate (dorsal). The terminal seg- ments of the cerci were missing. The paraprocts are visible beneath the supra-anal plate in the cleared specimen. (scale = 0.5 mm).




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19731 Roth - Blattaria
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minute undulating irregularities reminiscent of some specimens of P. indicus (cf. Fig. 61 with Figs. 14-1 5, 18). The L2d is apparently represented by a small sclerotized region of the prepuce widely sep- arated from the apex of L2vm (Fig. 60). The vertical downward curvature of the cleft and indentation on the outer margin of the lower lobe of LI
(Fig. 62) is unique for this phallomere in the Blaberidae ; the lower lobe of LI lacks setae. The supra-anal and subgenital plates of D. punctata are shown in figures 58-59. The genital phallomeres of 3 other undetermined specimens of Diploptera (Figs. 65-67, 70-72, 75-77) are indistinguish- able from D. punctata. The subgenital plates of these specimens (Figs. 64, 69, 74) are similar to D. punctata (Fig. 59)) but the shapes of the supra-anal plate of 2 of the specimens differ somewhat. In the specimen shown in figure 63, there is a deep invagination on the posterior margin; however because of the slight asymmetry of the indentation this may be an aberrant punctata. This specimen is from Hawaii and although D. punctata is widespread in the Pacific it is the only species of the genus recorded from Hawaii (Princis, 1965). Except for size, the phallomeres (fig. 78) of D. mintor are almost indistinguishable from D. punctata; the sclerotization (L2d) of the prepuce found in D. punctata is absent in minor. The supra-anal plate of D. minor is slightly more rounded (fig. 79) than that of punctata (fig. 58).
The writer thanks Dr. David Rentz, Academy of Natural Sciences, Philadelphia ; Dr. David Ragge, British Museum (Natural History) , London; Dr. Karl Princis, Zoological Institute, Lund, Sweden; Dr. Howard Evans, Museum of Comparative Zoology, Harvard Univer- sity; and Dr. A. Kaltenbach, Vienna Museum of Natural History, Austria, for the loan of museum specimens. I am grateful to Mr. Samuel Cohen for taking the photographs. I collected Proscratea complanata as a member of Phase C of the Alpha Helix expedition to the Amazon in 1967. I thank the National Science Foundation for support on the Amazon expedition under Grant NSF-GB-5916. BRUNNER, VON WATTENWYL, C.
1865. Noveau systkme des Blattaire. Vienna, 426 pp. GRAVES, P.
1969. Spermatophores of the Blattaria. Ann. Entomol. Soc. Amer., 62: 595-602.




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264 Psyche
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GURNEY, A. B. AND L. M. ROTH.
1972. A generic review of the cockroaches of the subfamily Panchlori- nae (Dictyoptera, Blattaria, Blaberidae) . Ann. Entomol. SOC. Amer., 65: 521-532.
HEBARD, M.
1926. The Blattidae of French Guiana, Proc. Acad. Nat. Sci. Phil.! 78: 135-244.
KIRBY, W. F.
1904. A synonymic catalogue of Orthoptera. Vol. I. London, pp. 61-205.
MCKITTRICK, F. A.
1964. Evolutionary studies of cockroaches. Cornell Univ. Agr. Exp. Sta. Mem. No. 389, 197 pp.
PRINCIS, K.
1963.
Kleine Beitrage zur Kenntnis der Blattarien und ihrer Verbrei- tung. VII. Opusc. Entomol., 28 : 147-155. 1964. Orthopterorum Catalogus. Edit. M. Beier. Pars. 6. Blattariae: Subordo Blaberoidea. 's - Gravenhage, pp. 174-281. 1965. Orthopterorum Catalogus. Edit, M. Beier, Pars. 7. Blattariae: Subordo Blaberoidea. 's - Gravenhage, pp. 284-400. 1967. Kleine Beitrage zur Kenntnis der Blattarien und ihrer Verbrei- tung. X. Opusc. Entomol., 32 : 141-151.
REHN, J. A. G.
1932. Wissenschaftliche Ergebnisse der schwedischen entomologischen Reisen des Herrn Dr. A. Roman in Amazonas 1914-1915 und 1923-192+. Arkiv for Zoologi, 2,4A: 1-71. ROTH, L. M.
1967. Sexual isolation in parthenogenetic Pyconoscelus surinamcnsis and application of the name Pycnoscelus indicus to its bisexual relative (Dict~optera : Blattaria: Blaberidae : Pycnoscelinae) . Ann. Entomol. SOC. Amer., 60: 774-779.
1970a. The male genitalia of Blattaria. 111. Blaberidae: Zetoborinae. Psyche, 77: 217-236.
1970b. The male genitalia of Blattaria. IV. Blaberidae: Blaberinae. Psyche, 77 : 308-342.
1971a. The male genitalia of Blattaria. V1. Blaberidae: Oxyhaloinae. Psyche, 78: 84-106.
1971b. The male genitalia of Blattaria. VIII. Panchlora, Anchoblatta, Biolleya, Pelloblatta, and Achroblatta (Blaberidae : Panchlorinae) . Psyche, 78 : 296-305.




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