Louis M. Roth.
The Male Genitalia of Blattaria. VII. Galiblatta, Dryadoblatta, Poroblatta, Colapteroblatta, Nauclidas, Notolampra, Litopeltis, and Cariacasia (Blaberidae: Epilamprinae).
Psyche 78(3):180-192, 1971.

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THE MALE GENITALIA OF BLATTARIA. VII.
GALIBLATTA, DRYADOBLATTA, POROBLATTA,
COLAPTEROBLATTA, NAUCLIDAS, NOTOLAMPRA,
LITOPELTIS, AND CARIACASIA.
(BLABERIDAE : EPILAMPRINAE) .
BY Louis M. ROTH
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01 760
The male genitalia of cockroaches have proved to be extremely use- ful in showing generic relationships (Roth, 1970a, 1970b). This study of 8 genera again shows the importance of wing internal male genital structures in grouping genera of Blattaria. The genitalia of species of the following genera are illustrated in this paper: Galiblatta Hebard, Dryudoblatta Rehn, Poroblatta Heb- ard, Nauclidns Rehn, Notolampra Saussure, Colapteroblatta Hebard, Lttopeltis Hebard, and Cariacasia Rehn. Princis ( 1960) placed Dryadoblatta and Notolampm in the Epilampridae (Epilamprinae and Phoraspinae respectively) and the other 6 genera in the Blabe- ridae, subfamily Laxtinae. McKittrick (1964) placed Laxta in the Epilamprinae and Princis (in Roth, 1970a) considered his subfamily Laxtinae provisional and predicted it probably would be split up. McKittrick (1964) placed Litopeltis, Poroblatta (with a query), and Galiblatta (in Roth, 1968) in the Epilamprinae. I follow Mc- Kittrick in placing all ovoviviparous cockroaches in Blaberidae and consider all the above genera as belonging to the Epilamprinae. Other genera of Epilamprinae will be treated in future publications. MATERIALS AND METHODS
The source of each of the museum specimens illustrated is given using the following abbreviations: (ANSP) = Academy of Natural Sciences, Philadelphia; ( BMNH ) = British Museum (Natural History), London; (L) == Zoological Institute, Lund, Sweden; (MCZ) Ì Museum of Comparative Zoology, Harvard University, Cambridge, 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. RESULTS AND DISCUSSION
McKittrick (1964, p. 37) stated that "the slight differences evi- dent in the character systems barely justify the designation of



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19711 Roth - Bla~aria I 8 I
tribes . . ." within the Epilamprinae. However, she tentatively divided the 13 genera of Epilamprinae which she studied into 5 tribes. She included Epilampra, Litopeltis, and Poroblatta (with a query) in the Epilamprini.
I have found that the male genitalia of many genera of Epi prinae may be used to make tribal designations. In the present s the male genitalia clearly fall into 3 groups based on distinct ferences in the L2d and prepuce.
I. Poroblattini (Poroblatta [Fig. I], Colapteroblatta [Fig. Dryadoblatta [Fig. 31, Galiblatta [Fig. 41, Nauclidas [Fig. 51' am-
udy
dif-
21 *
-
In this tribe the L2d is elongated, curved, 'aclerotized, tapers slightly toward the tip, and is separated from L2vm (Figs. 9, 12, 15, 18, 21. 24). Apparently there is no distinctive prepuce. The R2 has a sub- apical incision (Figs. 10, 13, 16, 19, 22, 25 ) and the shapes of LI are all basically similar (Figs. I I, 14, I 7, 20, 26). Hebard ( 1919) claimed that Poroblatta (Figs. 9-11) is related to Colapteroblatta (Figs. 12-14) but showed closer affinity to Acroporoblatta, and the nearest relative of Colapteroblatta was Poroblatta. I have not seen any males of Acroporoblatta, but the genitalia support Hebard's con- clusion regarding a close relationship between Poroblatta and Cola$- teroblatta. According to Hebard (1926, p. 236) Galiblatta is ap- parently nearest Colapteroblatta. The close relationship between these 2 genera is seen in their genitalia but I would place Galiblatta closer to Dryadoblatta (cf. Figs. 21-23 and Figs. 24-26) than to Colapteroblatta (Figs. I 2- I 4). The male genitalia of Galiblatta cribrosa differs from G. williamsi in the shape and microscopic sur- face of the tip of L2d (Figs. I 8, 2 I, in Roth, I 968). Rehn and Hebard
(1927, p. 319) not having access to males tentatively assigned the West Indian species Parasphaeria nigra Brunner to the genus Poroblatta. Later Rehn (1930) p. 58) erected the genus Nauclidas using P. nigra as the type genus; he stated that Nauclidas ". . . belongs to the assemblage which also comprises Colapteroblatta, Poroblatta, Acroporoblatta, and Galiblatta." Rehn placed Nauclidas nearer Galiblatta than to any of the other genera. The male genitalia of Nauclidas (Figs. 15-17) confirm this close relationship to members of the Poroblattini. Rehn (1930, p. 56-58) based the genus Dryadoblatta on Homalop- teryx scotti Shelford. He believed that Dryadoblatta was ". . . prob- ably as near to Pinaconota Saussure as to any other genus known at this writing . . . In the present incomplete state of our knowledge of the diagnostic features of the genera placed in the Epilamprinae, and in the absence of any phylogenetic concept of their classification, it seems best to compare Dryadoblatta with Pinaconota. Future



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I 82 Psyche [September
study may show the two genera are not closely related, but it is not possible at this writing to attempt an analytic treatment of the genera of the subfamily. It is certain, however, that Dryadoblatta is not closer in relationship to any of the other genera, and its agree- ment with Pinaconota in many features is marked.'' I have examined a male specimen which Rehn determined as Pinaconota sp. (Fig. 52), and have also seen the male type of Ischnoptera ( ?) sicca Walker which Kirby synonymized with Pina- conota bifasciata (Saussure) and which Princis (1958, p. 68) lists as a synonym of this species. The male shown in Fig. 51 is similar to the type of sicca, and I collected all stages of this species in the hanging nest of an oriole in the Amazon. Princis (personal com- munication) examined my specimens of sicca and concluded that Ischnoptera sicca Walker is not a Pinaconota. The male genitalia indicate clearly that neither Ischnoptera sicca (Figs. 53-55) nor Rehn's Pinaconota sp. (Figs. 56-58) are closely related to Dryado- blatta (Figs. 24-26), a genus obviously related to Galiblatta (Figs. 21-23). The genitalia of Pinaconota sp. and ('I." sicca are quite different and support Princis's conclusion that they are not congeneric. 2. Notolamprini (Notolampra [Figs. 6, 8a] ) . - Rehn and Hebard (1927, p. 202) noted that the 3 species of Notolampra have a markedly convex dorsal surface but are more elongate than Phora- spis, which is a genus whose species are also strongly convex and resemble cassidid Chrysomelidae. According to Rehn and Hebard, Notolmnpra ". . . marks a transition from the more normal epilam- prine type to that of the specialized phoraspid offshoot of the family." Princis (1960) placed Notolampra in the Phoraspinae; but the male genitalia of Phoraspis differ considerably from those of Noto- lampra and I have placed Phoraspis in the Phoraspini of the Epilam- prinae (Roth, 1972).
The genitalia of 2 species of Notolampra which I have seen differ markedly from each other. In N. gibba (Type genus) the L2d (Fig. 27) is much more robust than the L2d of members of Poroblattini, and does not taper toward the apex. RI (Fig. 28) is long and slender and has a subapical incision; LI (Fig. 29) differs in shape from the LI of Poroblattini (cf. Figs. I I, 14, I 6, 20, 23, 26). In N. antillarum, the shape of L2d (Figs. 30, 33) differs from that of N. gibba (Fig. 27) and is partially covered by minute spines. The l~hallomeres RI (Figs. 31, 34) and LI (Figs. 32, 35) are very similar to those of Poroblattini. Notolampra gibba is found in Brazil, and N. antillarum is West Indian.
3. Epilamprini (Litopeltis [Figs. 7, 7a], Cariacasia [Fig. 81. - The genitalia of Litopeltis and Cariacasia are sufficiently close to



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Roth - Blattaria d3
Figs. 1-4. Adult males of Epilamprinae (Foroblattini). 1. (118 ANSP). Parolilatia sp. Sierra San Lorenzo, Magdaiene, Colombia. 2. (116 ANSP). Col&pteroblatta comfsa Hebard. Type. San Lorenzo, Santa Marta, Colom- bia. 3. (17 MCZ) . Dryudo/~htta $cotti (Shelford) . Mount Tucuche, Trini- dad (type locality) (det. Darlington). 4. (USNM). Galibhita. wi///umsi Roth. Tarurna-Acu, about 15 Km. northeast of Manaus, Amazonas, Brazil (from Roth, 196S).
(scale = 5 mm).




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I 84 Psyche [September
Fig*;. 3-83. Ad~~ltri of Epilamprinae. 5, Poroblattiiii. Xaltrlida~ nigru {Brunner). (9 ; adventive obtained from the Iiriti'-.h rnuiieust)). 6, (1469L). Notolamprini. ffotola~tipra afif'illtlrum Shelford. Trinidad (det. Ptincisl. 7. (119 ANSP). Litopdtis areas Rehn, Paratype. Santa Maria de Dota Costa Rica. 7a. (172 ANSP). Lifopiiliii ffidlcyi (Saussure). Costa Rica (det. Rehn). 8. (114 ANSP). Cariacada cu#uck Rehn. Type 1123. Carilla, Costa Rica. Sa. (175 ANSP). h'otolampra g'bba (Thtinberg). Pernambuco, Brazil (det. by Hebard as h'otolamfra cassidra (Burm.), a s-ynonym).




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Figs, 9-17. Cockroach male genitalia of Epilamprinae ( Poroblattini) . 9-11. (I18 ANSP). Poroblatta sp. (from specimen shown in Fig. 1). 12-14. (116 ANSP). Cotafterobiaiia compsa (from specimen shown in Fig. 2). 15-17, Nauclidas nigra. (from adventive on bananas probably originating in the West Indies; specimen from a mail culture established at the British Museum). (LI ÌÔ first sderite of left phallomere; L2vn = median sclerite; L2d = dorsal sclerite of L2; R3 = hooked scierite of right phallo- mere; SI = subapical incision}. (scale = 0.3 mm).



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Figs. 18-26. Cockroach male genitalia of Epilarnprinae (Poroblattinij. IS-20. (ANSP). Gaiibhtta cribrosa Hebard. Type No. 1029. St. Jean du Maroni, French Guiana. 21-23. (USNM). Galiblalta williamsi. Tarunaa- Acu, about IS Km. northeast of Manaus, Amawnas, Brazil (det. Roth). 24-26. (17 M'CZ). Dryudobtatta scolti. (from specimen shown in Fig. 3). {Figa. 18-25 from Roth, 1968).
(scale = 0.3 mm).




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19711 Roth - Blattarh 187
Figs. 27-35. Cockroach male genitalia of EpiIamprinae (Notolamprini). 27-39. (175 ANSP). Notulamfra ybta (from specimen shown in Fig. 8a). 30-35. Natolampra anfillarvm. 30-32, (24 BMNH) . Trinidad. 33-35. (1469 I.). (from specimen shown in Fig. 6). (der. Princis). (scale = 0.3 mm).




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188 Psyche
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Figs. 36-44. Cockroach male genitalia of Epilamprinae (Epilamprini). 36-38. Litapdtis inspinosa (Saussure). Puntarenas Province, Costa Rica (det. Fisk). 39-41. (168 ANSP). L, bispinosa. Porto Bello, Panama (det. Rehn). 42-44, (172 ANSP), Litoprliis bmlteyi. (from specimen shown 111 Fig. 7a). (L2d = dorsal sclerite of L2; P = prepuce). (scale = 0.1 mm).



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Roth - Blatfaria
Figs. 45-50. Cockroach male genitalia of Epilamprinae (Epilamprini). 45-47. (119 ANSP). LitopeStis oreas. Paratype. {from specimen shown in Fig. 7). 48-50. (114 ANSP) . Cariacasia capucina, Type. (f corn specimen slicrivn in Fig. 8).
(scale = 0.3 mm).




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Psyche [September
Figs. 51-58. Adult males and male genitalia of Bhberidae. 51. "Isrh- noptera" sicca Waiker. Near Serra Tamendaui, Ria Ne~ro, Amazonai (det. Roth). 52. (189 USNM). Pinaronoia sp. liha daa Afcatraises, Sio Paulo, Brazil. (det. Rehn).
53-55. "Ischnoptera" i':rca (same data as speci- men shown in Fig. 51). 56-St. (189 USNM). Pinaconolia sp. (from å´ipeci men shown in Fig. 52). (scale for adults = S mm., for genitalia = 0.2 mm).



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19711 Roth - Blattaria 191
most Epilanzpra (Roth, 1971) to place them in Epilamprini ; L2d is a variably shaped dark sclerite separated from L2vm and the prepuce is usually a distinctively shaped lobe covered by microtrichia (Fig. 39). The RI's of Litopeltis (Figs. 37, 40, 43, 46), and Cariacasia (Fig. 49) have a subapical incision and the shapes of LI (Figs, 38, 41, 44, 47, 50) are similar. The differences in the genitalia of the 3 species of Litopeltis are so minor (Figs. 36-47) that it would be impossible to use them to distinguish species. When Hebard (1920, p. 140) described the genus Litopeltis he stated that it ". . . belongs to the second section of the Perisphaerinae, containing Stenopilma Sauss. p= Cyrtotria] and its allies. To this section also belong the American genera Colapteroblatta? Poroblatta and Acroporoblatta Hebard and Mioblatta . . . . nearest relationship with Colapteroblatta exists, this indicated by the general similarity of tegminal and wing form and venation and limb armament." The genitalia of Litopeltis (Figs. 36-47)
are sufficiently different from
those of Colapteroblatta (Figs. 12-14) to place them in different tribes.
Rehn (1928, p. 190) in discussing the genus Cariacasia placed it in the Perisphaeriinae and claimed it was related to Litopeltis and Mioblatta Saussure. However, he also stated that ". . . the male of Litopeltis superficially looks more like the epilamproid genus Leuro- lestes f - ~ Phoetalia]. The relationship of the two genera here treated is, however, more intimate than a casual glance, even at individuals of the same sex, would indicate." Phoetalia has male genitalia characteristic of Blaberinae and I recently assigned it to this subfamily (Roth, 197ob). Because of differences in tarsal arm- ariient, Rehn (1930, p. 59) removed Litopeltis and Cariacasia . . . from the vicinity of the Poroblatta complex, although their general appearance much suggests the latter assemblage." The geni- talia of Litopeltis (Figs. 36-47) and Cariacasia (Figs. 48-50) are very similar showing a close relationship, and differ from those of Poroblattini, thus supporting Rehn's conclusions. Based on male genitalia, 8 genera of Epilamprinae are placed into 3 tribes as follows:
I. Poroblattini. - Poroblatta, Nmclidas, Galihlatta, Dryado- blutta, and Colapteroblatta.
2. Notolamprini. - 'Notola?npra.
3. Epilamprini. - Litopeltis, Cariacasia.



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1 92 Psyche
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ACKNOWLEDGMENTS
I thank Dr. N. Jago, Academy of Natural Sciences, Philadelphia, Dr. Ashley Gurney, U. S. National Museum, Washington, D.'C.j Dr. David Ragge, British Museum (Natural History), London, Dr. Frank Fisk, Ohio State University, Columbus, and 'Dr. Karl Princis, Lund, Sweden, for the loan of specimens. I collected specimens of "Ischnoptera" sicca during Phase C of the Alpha Helix expedition to the Amazon in 196'7. I thank the National Science F'oundation for support on the Amazon expedition under Grant NSF-GB-5916. I am grateful to Mr. Samuel Cohen for taking the photo,graphs.
REFERENCES
HEBARD, M.
1919. Studies in the Dermaptera and Orthoptera of Colombia. First paper. Trans. Amer. Entomol. Soc. 45: 89-179. 1920. The Blattidae of Panama. Mem. Amer. Entomol. Soc. 4: 1-148. (1919).
1926.
The Blattidae of French Guiana.
Proc. Acad. Nat. Sci. Phil. 78:
135-244.
MCKITTRICK, F. A.
1964. Evolutionary studies of cockroaches. Cornell Univ. Agric. Exp. Sta. Memoir 389, 197 pp.
PRINCIS, K.
1958. Revision der Walkerschen und Kirbyschen Blattarientypen im British Museum of Natural History, London. 11. Opus. En- tomol. 23 : 59-75.
1960. Zur systematik der Blattarien. Eos 36: 427-449. REHN, J. A. G.
1928. New or little known Neotropical Blattidae (Orthoptera). Num- ber one. Trans. Amer. Entomol. Soc. 54: 125-194. 1930. New or little known Neotropical Blattidae (Orthoptera). Num- ber two. Trans. Amer. Entomol. Soc. 56: 19-71. REHN, J. A. G. AND M. HEBARD
1927. The Orthoptera of the West Indies. Number 1. Blattidae. Bull. Amer. Mus. Nat. Hist. 54: 1-320.
ROTH, L. M.
1968. A new species of Galiblatta from Brazil (Blattaria, Blaberidae). Psyche 75: 249-255.
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.
1971. The male genitalia of Blattaria. V. Epilampra spp. (Blaberidae: Epilamprinae). Psyche 77 : 436-486.
1972. The male genitalia of Blattaria. IX. Blaberidae. Gyna spp. (Perisphaeriinae), Phoraspis, Thorax, and Phlebonotus (Epi- lam~rinae). Trans. Amer. Entomol. Soc. (in press).



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