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Louis M. Roth.
The Male Genitalia of Blattaria. III. Blaberidae: Zetoborinae.
Psyche 77:217-236, 1970.

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THE MALE GENITALIA OF BLATTARIA.
111. BLABERIDAE : ZETOBORINAE.
BY Louis M. ROTH
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01 760
In this paper I shall illustrate and characterize the male genitalia of 9 genera of blaberids which belong in the subfamily Zetoborinae. These genera are Capucina Saussure, Lanxoblatta Hebard, Phortioeca Saussure, Phortioecoides Rehn, Schistopeltis Rehn, Schizopilia Bur- meister, Tribonium Saussure, Zetobura Burmeister, and Zetoborella Hebard. Princis (1960) includes Tribonovlea Shelford in this sub- family but I have been unable to obtain a male of the one species in this genus.
The male genitalia of most species of Blaberidae are characterized by the presence of three phallomeres: a median sclerite, a sclerite on the left, and a retractable hook on the right side. This led McKittrick (1964, p. 72) to conclude that "The male genitalia are extraordi- narily uniform throughout the Blaberidae." She also stated (1964, p. 35) that "The male genitalia differ slightly in hook shape and the configuration of the posterior tip of the median sclerite within the Blaberidae, but otherwise show little variation within that group," and on p. IIO "The great uniformity of blaberid male genitalia pre- cludes drawing many conclusions regarding subfamily relationships, except in the case of the Blaberinae and Panesthiinae." "The great uniformity" of the male genitalia of Blaberidae is generally true only for the usual presence of three distinct phallomeres; in certain genera one 01- more of the phallomeres may be greatly reduced and even absent (e.g. R2 in Panchlora). However, the shapes of the phallo- meres, especially the retractable hook (Rz), prepuce, and L2d are often so distinctive that they may show subfamily, tribal, generic, species Group characteristics, and in some genera may be used for specific determination (Roth and Gurney, 1969; Rcth, 1969, 1970). The technique for preparing the genitalia has been described in pre- vious papers (Roth, 1969, 1970). The source of each of the specimens illustrated is given, using the following abbreviations: (ANSP) = Academy of Natural Sciences, Philadelphia; (BMNH) = British



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218 Psyche [June
Museum (Natural History), London ; ( CUZM) = Copenhagen University, Zoological Museum, Denmark ; ( MCZ) = Museum of Comparative Zoology, Harvard University ; ( N ) = Natick culture, U. S. Army Natick Labs., Massachusetts; (USNM) = United States National Museum, Washington, D.C. Geographical collection data, and the names of specialists who iden- tified the specimens, if known, follow these abbreviations. The num- ber preceding the abbreviations refers to the number assigned the specimen and its corresponding genitalia (on a slide) which are de- posited in the museums indicated.
The basic features of the 3 phallomeres of the male genitalia of Zetoborinae are as follows: R2 (hooked right phallomere) is well developed and usually has a subapical incision (Fig. 32). The pre- putial membrane has a heavily sclerotized area (L2d) which extends dorsally or dorsolaterally to the right of L2vm (Fig. 28) ; L2d is flattened in the preparations and it should be remerxbered that it usually extends dorsally. LI is very well developed and various areas are darkly sclerotized (Fig. 30).
Based on the degree of development of L2d and shape of Lzvm, the Zetoborinae may be divided into 3 Tribes as follows: I. Zetoborini (Z~tobora and Phortioecoides, Figs. 1-5, 22-39). L2d is least developed in this tribe and is a thin sclerotization of the preputial membrane which extends dorsally (Figs. 22, 25, 27, 28, 3 I, 34, 37). The subapical incision of R2 is clearly defined (Figs. 23, 29, 32, 35, 38). LI is very similar in all the species in this tribe (Figs. 24, 26, 30, 33, 36, 39).
Hebard ( 1921, p. 245) stated that "The differences shown by the forms iecognized by us as Tribonidium [= Zetobora] monasticurn, transversum, and amplum, may indicate full specific distinction, geo- graphic racial differentiation or, indeed, mere individual variation in a single exceptionally plastic species." The male genitalia of the above 3 species (Figs. 22-26, 28-36) are so similar that they are of little help in distinguishing the various species of Zetobora. Rehn ( 1937) placed Phortioecoides in the Panchlorinae. Princis (1961) followed Rehn and placed it in the Panchloridae. Princis' Panchloridae is based principally on the shape of the subgenital plate in the male. However, the subgenital plate of the type male of Phortioecoides guarani Rehn (the only species of the genus) is not at all like that of Panchlora. Also the genitalia of Panchlora are



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Figs. 1-5. 1. (105 ANSF). Zefohwa ~ignaticollis Burm. Curityba, Brazil (det. Hebard). 2. (6 BMNH). Zeiobora transversa (Brunnet). Rio de Janeiro, Brazil. 3. (106 ANSP). Xetobora ampla (Hebard). Paratype. Montagnes Des Orgues, Prov. de Rio Janeiro, near Tijuca, Brazil. 4, (7 BMNH). Zetoboru mamiticu Sauss. 5. (113 ANSI?). T'hrtioecoides guarani Rehn. Paratype. Horqueta, Paraguay - 40 Krn east of Rio (57'W, 2i0N), (scale ==: 5 mm.)




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230 Psyche
Figs. 6-9. 6. (5 ANSP), hxoblatta lata (Shelford). Muzo, Boyad, Colombia (det. Hebard). 7. (4 ANSP) . Lanxoblatta /rater Hebard. Paratype. Aotioquia, Colombia. 8. (2 ANSP}. Lfinxablatta emurflkata (Burm.). La Forestitre, Haut Maroni, French Guiana (det. Hebard}. 9. (97 ANSP) . Schtza#hilk jissicoli~ (Serv.) . La Foresti&re, Haut Maroni, French Guiana. (det. Hebard). (scale == 5 mm.)



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Roth - Blattaria
Figs. 10-13. 10. (N). Capucina pafula (Walker). Co-ita Rica (At. Gurney). 11. (102 ANSP). Zetohore//a gummido Hebard. Topetype. St. Jean du Maroni, French Guiana. 12. (N.) ~hortiofca phoras#eidrs



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222 Psyche
Figs. 14-17. 14. (101 ANSP). Phortioeca nimbata (Burm.). St. Jean du Maroni, French Guiana. IS. (100 ANSP). Phortiotca afolinari Hebard. Paratype. Villavicencio, Colombia. 16. (2 CUZM). Phwt'oeca werrucosu (Sauss.). Riacho det Fra. (det, Princia). 17. (5 BMNH). PLvr'.'atca



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Figs. 18-21, 18. (104 ANSP). Trtlionium ro~$crsv-m (Guerin). Nova Teutonia, Brazil. 19. (103 ANSP). Tribok coinwhirurn Hebard. Para- type. "New Grenada" (= Colombia). 20. (73 MCZ). Tdbonium sp. Brazil. 21. (US ANSP). Schisfopc~~is peculiaris Rehn. Hoiotype 5244. Porto Velho, Rio Madeira, Brazil. (scale = 5 rnm.)



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224 Psyche [June
greatly reduced and poorly developed or sclerotized. Most PanchZora la,ck an L2d and R2, and LI is usually represented by a weakly sclerotized cleft.
In erecting the genus Phortioecoides, Rehn (1937, p. 234) stated that it is "Related to Phortioeca Saussure, Capucinella Hebard, and Capucina Saussure, showing certain features of agreement with each, but differing from each one by combinations of characters." The genitalia of Capucinella (Lzd and R2) show none of the typical characters of Zetoborinae; the genus probably belongs in the Epi- lamprinae. The shape of L2cl of Phortioecoides (Fig. 37) more closely approaches that of Zetobora (Figs. 22, 27, 28) and I place Phortioecoides closer to this genus than to Capucina, 2. Phortioecini ( Capucina, Lanxoblatta, Phortioeca, Schizopilia, and Zetoborella, Figs. 6-17, 40-98). L2d (Figs. 40, 43, 45, 48, 50, 51, 54, 57, 60-62, 63, 66-p~, 72, 75, 78, 81, 84) is considerably more developed and robust than in the Zetoborini, and reaches its greatest development in certain species of Phortioeca (Figs. 87, 90, 93, 96-98). R2 has a well defined subapical incision (Figs. 41, 44, 46, 52, 55, 58, 64, 71) except for several species of Phortioeca (Figs. 73, 76, 79, 82, 85).
Hebard (1921, p. 210) compared Zetoborclla with Zetobora spp. The genitalia of Zetoborella (Figs. 40, 41) are closer to other mem- bers of the Phortioecini (e.g. Figs. 43-46) than to Zetobora. Capucina patula (Fig. 10) was placed in the Panchlorinae by McKittrick ( I 964)
and in the Laxtinae by Princis ( I 960). The Laxtinae of Princis is not recognized by McKittrick and she has assigned several of the genera in this grouping to other subfamilies. Princis (personal communcation) regards his subfamily Laxtinae ". . . as provisional. Further study will probably show that it is not at all a homogeneous group and must be sooner or later split up." McKittrick used proventricular and female genitalic characters in placing Capucina in the Panchlorinae. However many of the char- acter differences she used in comparing ovipositors and proventriculi seem to be very subtle, and I believe the male genitalia offer char- acters which show more clear cut relationships in the Blaberidae than do the proventriculus or ovipositor. Miller ( 1969) found that the morphological diversity of the proventsiculus in 9 subfamilies of Blaberidae ". . . often defied the most dedicated efforts to group them into subfamilies."' The male genitalia of Capucina are so similar basically to those of other members of the Phortioecini that I do not hesitate to assign this genus to the Zetoborinae.



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Figs. 22-33. 22-24. (179 USNM) . Zetobora sigmtiwllh. Santo Amaro, SSo Paula, Brazil. (det. Gurney). 25-26. (180 TJSNM). 2. sipaticollis. Santo Amaro, SZo Paulo, Brazil. 27. (181 USNM), Zetobora sp. Serra Caraqa, M. G. Brazil. 28-30. (6 BMNH), Zeiobora transvi-rsa. (from adult shown in Fig. 3). 31-33. (7 BMNH). Zetobora mwasfica. (from adult shown in Fig. 4). LI :=: first aclerite of left phallomere; C = cleft of LI; L2vm = median sderite (L2 ventromedial) ; L2d = dorsal sclerite of 1^; R2 = hooked aclerite of right phallomere; SI = subapical incision of R2. (scale = 0.2 mm.)




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226 Psyche
Figs. 34-44, 34-36. (106 ANSP). Zetabora ampla. (from adult shown in Fig, 3; Fig, 34 la a ventral view). 37-39. (113 ANSP). Phoriioecsidct
parad. (from adult shown in Fig. 5). 44-42. (102 ANSP). ZdaboreUa pmmicwla, (from adult shown in Fig. 11). 43-44 (97 ANSP), Schism fills fwtollis. (from adult shown In Fig. 9). (scale = 0.2 mm.)



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19701 Roth - Blatfaria
Figg. 45-53. 45-49, Lawcob/atia frater. 45-47. (3 ANSP). Paratype. Antioquia, CoIdia. 48-49. (4 ANSP). (from adult shown in Fig, 7). 50. (1 ANSI'). Lanxoblatta Ma. Mum, Boyd, Colombia. 51-53. (K). Lanxoblaita emarghafa. From laboratory colony originating in Florea, Manaun, Amazonas, Brazil. (scale = 0.3 mm.)



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228 Psyche
Fig& 54-62. Lanxoblatta emar9inaia. 54-36. (2 AMP). (from adult shown in Fig. 8). 57-59. (9 MCZ). Ecuador. 60. (10 MCZ). Probably Brazil 61. (6 ANSP).
St. John du Marod, French Guiana. 62. (7 ANN'). Antioquia, Colombia, (scale = 0.2 mm.)




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Figs. 63-71. Capucha paiula. (N). From a laboratory culture origi- nating in Costa Rica. (al! IO scale shown in Fig. 71).



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Figs. 7240. 72-77. Phrtioeca nimbata. 72-74. (101 AN SP) . (from adult shown in Fig. 14). 75-77. (107 ANSP). La Forestiire, Haut Marmi, French Guiana. 78-80. (5 BMNH). Phortiwa mm'imiliuni, (from adult ihown in Fig. 17). (scale = 0.2 mm.)




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Figs. 81-89, S 1-83. (100 ANSP) , Phartiocca apolinari. (from adult shown in Fig. IS). 84-86.
(2 CUZM). Phortiocca verrucom
(from adult
shown in Fig. 16). 87-89. (98 ANSP). Phortioeca peruana, (from adult
shown in Fig. 13). L = lacuna in sclerotized basal area of L2d. (scale = 0.2 mm.)




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Psyche
Figs. 90-9s. Phort 'oeru phoraipoides (Walker). 90-92. (99 ANSP). Mum, Colombia (det. Hebard). 93-98. (N). Laboratory culture originating from Panama. F = fringe around preputial membrane. (scale = 0.2 mm.)



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19701 Roth - Blattaria 233
In Phortioeca peruana (Fig. 87) and P. phoraspoides (Fig. 98), the sclerotization of the preputial membrane, which forms the base of L2d, may be incomplete leaving a lacuna or window. However, it is clear that this character is variable (Figs. 90, 93, 96-98) and cannot be used for specific determination. In P. phoraspoides the rounded margin of the preputial membrane which surrounds the sclerotized base of L2d has more or less uniformly spaced indentations (Fig. 93) which are absent in other species of Phortioeca examined. Based on the shape of R2, Phortioeca ninzbatu, maximiliani, apolinari, and verrucosa can be grouped together. In these species the curved hook portion of R2 is relatively narrow and its subapical incision is poorly developed or absent (Figs. 73, 76, 79, 82, 85). Phortioeca peruana and P. phoraspoides both have stouter and more robust R2's and the subapical incisions (Figs. 88, 94) are distinct. Rehn (1932) stated that the genera Zetobora, Lanxoblatta, Zeto- horella, and Sc'hizopilia ". . . are clearly derivatives of a single phylum," whereas Schistopeltis and Trihonium ". . . typifies a dis- tinct and clearly marked phylogenetic series." In general the male genitalia tend to support Rehn's hypothesis. However, I have sep- arated Zetobora from the other 3 genera of his grouping because of the relatively poorly developed L2d.
3.
Triboniini ( Tribonium and Schistopeltis, Figs. I 8-2 I, 99- I I 3 ) . In this tribe, Lavm has a well developed posterior arm or extension ( Fig. 105 ) which extends dorsally ( laterally in the photographs because of flattening) and L2d extends upward and curves towards this outgrowth (Figs. 99, 102, 105, 108, I 1 I ). The subapical in- cision of R2 is clearly defined and occurs at about the middle of the hook (Figs. ioo, 103, 106, 109, 112).
Based on the shape of L2vm and Lad of the male genitalia, 9 genera of Zetoborinae are divided into 3 tribes as follows: I. Zetoborini (Zetobora and Phortioecoides). 2. Phortioecini (Capucina, Lanxo blatta, Phortioeca, SchizopiZia, and Zetoborda).
3. Triboniini (Tribonium and Schistopeltis).



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234 Psyche
Figg. 99-107. 99+101. (103 ANSP). Triboniurn c o h b h . (from adult shown in Fig. 19). 102-104. (104 ANSP). Trihortium canspersum (from adult shown in Fig. 18). 105-107. (1 CUZM). Xribon'Mm sp. E = dorsal extension of L2vm. (scale = 0.2 mm.)



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19701 Roth - Blatta& 235
Figs. 10B-113. 108-110. Tribonium ip. 108. (73 MCZ). (from adult &own in Fig. 20). 109-110. (31 MCZ). 111-113. (115 ANSP). Schhto- peltis perulih.
(from adult shown in Fig. 21). (scale = 0.2 mm.)



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236 Psyche [June
I thank the following for the loan of Museum material: Dr. M. G. Emsley and Dr. N. D. Jago, Academy of Na,tional Sciences, Philadelphia; Dr. Ashley Gurney, U. S. National Museum, Wash- ington, D.C.; Dr. D. R. Ragge, British Museum (Natural History) London; Dr. S. L. Tuxen, Zoological Museum, Copenhagen. I col- lected living specimens of Lanxoblatta enzarginata during 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. I thank Mr. Samuel Cohen for taking the photographs.
HEBARD, M.
1921. South American Blattidae from the Museum National d'Histoire Naturelle, Paris, France. Proc. Acad. Nat. Sci. Phil., 73 (11) : 193-304.
MCKITTRICK, F. A.
1964. Evolutionary studies of cockroaches. Cornell Univ. Agr. Exp. Sta. Memoir 389, 197 pp.
MILLER, H. K.
1969. Comparative morphological studies of proventriculi in the Order Dictyoptera : suborder Blattaria. Ph.D. Thesis, The Ohio State University, Columbus, Ohio, 91 pp.
PRINCIS, K.
1960. Zur s~stematik der Blattarien. Eos, 36: 427-449. REHN, J. A. G.
1932. Wissenschaftliche Ergebnisse der Schwedischen entomologischen Reisen des Herrn Dr. A. Roman in Amazonas 1914-1915 and 1923-1924. Arkiv for Zoologi, 24A, 1-73. 1937. New or little known neotropical Blattidae (Orthoptera). Num- ber four. Trans. Amer. Entomol. Soc. 63: 207-258. ROTH, L. M.
1969. The male genitalia of Blattaria. I. Blaberus spp. (Blaberidae: Blaberinae). Psyche, 76: 217-50.
1970. The male genitalia of Blattaria. 11. Poeciloderrhis spp. (Bla- beridae: Epilamprinae). Psyche, 77 : 104-119. ROTH, L. M. AND A. B. GURNEY.
1969. Neotropical cockroaches of the Epilampra abdomennigrum com- plex; a clarification of their systematics (Dictyoptera, Blattaria). Ann. Entomol. Soc Amer. 62: 617-27.




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