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Louis M. Roth.
The Male Genitalia of Blattaria. V. Epilampra spp. (Blaberidae: Epilamprinae).
Psyche 77:436-486, 1970.

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THE MALE GENITALIA OF BLATTARIA. V.
EPILAMPRA SPP.
(BLABERIDAE : EPILAMPRINAE) .
BY Louis M. ROTH
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01 760
'The genus Epilampra is one of those assemblages which have de- veloped within the tropics of both hemispheres a vast number of species, often quite distinct, again closely related and difficult to distinguish.
With a general type of coloration the fluctuations of which make definite and exact characterization difficult, if not at times virtually impossible, it combines a uniformity of development in numerous other features, that in general in the family are suffi- ciently varied to prove of value to the systematic student. To add to the uncertainty of a situation difficult at best, we find many of the published descriptions almost valueless to aid in the recognition of these forms. As a whole the genus is one of the most difficult, ob- scure and generally unsatisfactory to study in the entire Blattidae." (Rehn and Hebard, 1927, p. 209).
Princis (1967) lists 60 species of Epilampra all of which are found only in the New World. At least five of these species [atriventris (Saussure) , cribrosa ( Burrneister) , ferruginea ( Brunner) , proximo (Brunner) , and verticalis (Burmeister) 1 have males with tergal glands, and their genitalia are so distinctly different from the males which lack tergal glands that I
(1970) have placed them in the
genus Poeciloderrhis Stil. This study of about 30 of the remaining 55 species of Epilampra listed by Princis ( 1967) shows that the male genitalia are useful not only for specific determinations of many species, but they may also indicate species relationships. MATERIALS AND METHODS
The technique of preparing slides of genitalia has been described in earlier papers (Roth, 1969b, 1970).
The source of each of the specimens illustrated is given, using the following abbreviations: (AMNH) = American Museum of Nat- ural 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; (MCZ) =



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Fig. I, Male genitalia (dorsal view) of Epilampra abdomenninpurn from Tapurucuara, Rio Negro, Brazil. (C = cleft of Ll; LIZ first sclerite of left phallmnere; Lib =: setal brush of LI; L3vm ==: median sclerite L2 ventromedial) ; L2d = dorsal sclerim of 12; P = prepuce; R2 :^ hooked sderite of tight phallomere).
Museum of Comparative Zoology, Harvard University, Cambridge, Mass.; (N) = U. S. Army Labs., Natick, Mass.; (USNM) = United States National Museum, Washington, D.C. Geographical collection data, if known, follow these abbreviations. The number preceding the abbreviations refers to the number assigned the speci- men and its corresponding genitalia (on a slide) which were deposited in the museum indicated. These numbers are used in the text where the identifications of certain species are discussed. If known, the taxonomists who identified the species are given. In several cases these specialists of the Elattaria disagreed in their determinations, emphasizing the difficulty in identifying species of *!era from literature descriptions. Unfortunately male type tna- terial was not always available so that several questions still remain unresolved. In spite of this drawback the results point up the value of using male genitalia in the taxonomy of a difficult genus. The phallorneres characteristic of Ep2ampra male genitalia are shown in Figure I.
Prepuce - Usually distinctively shaped with a definite marginal outline and often covered by microtrichia (Fig. I, P) .



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438 Psyche [December
Lad - This sclerite (Fig. I, L2d) is always separated from L2vm (Fig. I) by a membrane, and may or may not be clearly sep- arated from and lie above the prepuce. In some species (e.g. Mexi- cans Group) most of Lad is a sclerotized and integral part of the prepuce.
R2 (retractable genital hook, Fig. I, R2) -A subapical incision is often found on the ventral surface (Fig. 18, SI) . LI -A deep lateral curved incision or cleft is present (Fig. I, C) and a setal brush (Fig. I, Lib) may or may not be found on the dorsal surface.
Rehn and Hebard ( 1927, p. 210) without specifying the characters used, tentatively erected 5 species Groups for Epilampra, primarily for West Indian species. These Groups and the species included were : I. Cubensis Group : - cubensis Bolivar.
2. Mexicana Group: - insularis Bolivar, tainana Rehn and Hebard.
3. Abdomennigrum Group : - abdomennigm (De Geer), mona Rehn and Hebard.
4. Burmeisteri Group : - gundlachi Rehn and Hebard, burmeis- teri (Guirin) , wheeleri Rehn, haitensis Rehn and Hebard, sabulosa Walker.
5. Grisea Group : - quisqueiana Rehn and Hebard. Rehn and Hebard (1927) indicated that other species would prob- ably fall into these groups. The male genitalia do not support the placement of a number of the above species in the groups erected by Rehn and Hebard. Based on genital characters I suggest the follow- ing species groups of Epilampra: Mexicana, A bdomennigrum, Bur- moisten, Sodalis, Shelfordi, Heusseriana, and Yersiniana. Mexicana Group
[Epilampra mexicana Saussure (Figs. 2-13) ; E. fallax Saussure and Zehntner (Figs. 14-23) ; E. conferta Walker (Figs. 24-43)]. This group includes species in which most of Lad is a flattened sclerotized plate which is an integral part of, and does not lie above, the prepuce (Fig. 5). In mexicana only a small part of L2d on the left side is separated from and lies above the prepuce, and on the right side the L2d tapers and extends upward toward the Lzvm (Figs. 2, 5, 8, 10, I I, 12). In fallax the lateral extension of the right side of Lad is quite long (Figs. 14, 17, 210, 21). In some con- ferta the L2d extends well beyond the left side of the prepuce (Figs. 24, 27, 29-32, 34) and the extension on the right may vary consid- erably in length. The prepuce of mexicana is deeply notched and is readily distinguished from the other members of the group.



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Roth - Blattaria
Figs. 2-1 3. Cockroach male genitalia. Epdampra m~xiruna. 2-4. (63 USNM). Guatemala. 5-7. (109 USNM). Turrialba, Coma Rica. 8-9. (106 USNM). Turrialba. Costa Rica- 10. (11 1 USNM). Chis. Soyalo, [Rt. 195, Km 241. Mexico. (2-10, det. Gurney). 11. (52 AMSP). Central Mexico. 12-13. (110 USNM). Ver. Rio TacoSopan, [Rt. 195, Krn 241, Mexico. (11-13 det. Roth). (L2d = dorsal acleritc of L2; P = preptice). (scale = 0.2 mm)




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440 Psyche [December
Figs. 14-23, Cockroach male genitalia, Epitampra jailax. 14-16. ( 51 ANSI'). Sapucay, Paraguay (Jet. Roth), 17-18. (66 USNM). Santa Cata- rina, Brazil (det. Albuquerque), (SI ^= Subapical Incision). 19-20. (73 WSNM).
Santa Catarina, Brazil. (det. Albuquerque). 23-23. (103 USNM). Rio Lujer, Buenos Aires, Argentina (det. Albuquerque). (in Fig. 21 the prepuce is collapsed), (scale "= 0.2 mm)



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Figs. 24-35, Cockroach male genitalia. Epiiumpra ronfcrtu. 24-Zb. ( 15+ US). El Valle, Panama (det. Princis). 27-28. (136 USNM). Barro Colorado Uland, Panama. (det. Princis). 29. (135 USNM). Barro Col~radO Island, Panama. (det. Roth). 30. (50 ANSP). ChirKpi, Panama. (labeled sligmosa in pencil), 31. (105 USNM). San Isidrodel, General Cattago, Costa Rica (det. Princis). 32-33. (HZ USNM). Napo, Santa Cecilia, Rio A , Ecuador. (det. Roth). 34-35, (133 USNM). Same locality as 32-33. (det. Roth). (scale == 0.2 mm)




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Psyche [December
i
s
, ,
t
b
1"
Figs. 36-46.
Cockroach male genitalia. 36-38. (4 CUZM). E. conferta. Callanga, Peru. (det. Princis) . 39-40. (121 USNM) . E. cozferta. Callanga, Dept. of Cudco, Peru. (dct Roth). 41-43. (67 USNM). Epilampra ~p. Rio Blanco or Ecuador. (det as E. mexicam by Albuquerqut). The sctal brush 5-1 Fig. 43 is very lightly pigmented and difficult to see in the photograph. 44-46. (168 USNM). Eftihmpra sp, Cundinamarca, Colombia. (wale = 0.3 mm)




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19701 Roth - Blattaria 443
In species of the Mexicana Group, R2 (Figs. 3, 6, 15,18, 22, 25, 28, 33, 37, 40, 42, 45) has a subapical incision, and a setal brush (Figs. 4, 7, 9, 13, 16, 19, 23, 26, 35, 38, 43, 46) occurs on LI. Princis ( I 958, p. 63) synonymized Epilampra stigmosa Giglio-TOS with Epilampra conferta Walker.
The specimens determined by
Princis (Figs. 24-28, 31) as conferta are similar to a specimen, in the Philadelphia Academy collection, which was labeled (in pencil) E. stignzosa (Fig. 30). However, the E. conferta identified byHebard, Albuquerque, and Rehn (Figs. 229-237) have entirely different geni- talia from Princis' conferta (Figs. 24-28, 31, 36-38). The type of Walker's conferta (Brazil) is a female, whereas Giglio-Tos' type of stigmosa is a male.
According to Gurney (personal communication) "E. stigmosa G.- T. was based on 4 males from Ecuador. Giglio-Tos also described E. josephi from 2 males from S. Jose, Ecuador (stigmosa was from the valley of Santiago, Ecuador) ; they apparently were similar to stigmosa for the most part. . . . No. 76 [conferta, det. Albuquerque] (Figs. 235-237), seems rather small to be conferta, judging from Walker's description (though type is a female), but size may be quite variable. I would be inclined to use the name stigmosa for No. 132 [Figs. 32-33], No. 134 [Figs. 24-26], and No. 136 [Figs. 27-28], and perhaps No. 105 [Fig. 311, but am more uncertain about No. 76 [Figs. 235-2371 being conferta. . . . No. 105 [Fig. 311 has the face darker than 132, et. al., the interocular space is wider, and the ventral surface of the abdomen is much darker; however, if the genitalia agree this may be just variation." It is apparent that Gurney is not convinced that stigmosa and con- ferta are the same species. However, for the present, I am following Princis' conclusions. It is highly probable that more than one spe- cies is involved here which are very similar in external appearance. The problem may be partly solved by examining the male genitalia of the type of stigmosa. Unfortunately the Type of conferta, as in- dicated above, is a female. The prepuce and L2d of conferta speci- mens from Ecuador (Figs. 32, 34) differ somewhat from these structures in specimens from Panama and Costa Rica (Figs. 24, 27, 29, 30, 3 I ) and from specimens from Peru (Figs. 36, 39). The R2
(Figs. 37, 40) of Peruvian males are noticeably stouter than the genital hooks (Figs. 25, 28) of Panamanian specimens. Epilam- pra conferta may well be a complex of sibling species. The genitalia of two undetermined or questionably determined species belonging to this group are shown in Figures 41-46. One
specimen from Ecuador, determined by Albuquerque as E. mexicana



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444 Psyche [December
is not this species, according to Princis (personal communication) and its genitalia (Figs. 41-43) are distinctly different from those of mexicana (cf. Figs. 2-13).
A bdomennigrum Group
[Epihzpra abdomennigrum (De Geer) (Figs. 50-55) ; E. maya Rehn (Figs. 47-49) ; E. sagit& Hebard (Figs. 59-67) ; E. taira Hebard (Figs. 56-58) ; E. grisea (De Geer) (Figs. 68-96) ; E. jorgenseni (Rehn)
(Figs. 97-1 13) ; E. berlandi Hebard (Figs. I 14- 117); E. guianae Hebard (Figs. 119-127)]. In this group L2d is a thick, black, variably shaped sclerite, which lies above the prepuce.
The presence of a setal brush on LI dis- tinguishes it from the following Burmeisteri Group. The size of the setal brush is inter- and intraspecifically variable and sometimes the area covered by the setae is small, or the setae are lightly sclerotized (Figs. 58, 99) and difficult to see. The hook (R2) has a subapical incision in all the species listed in this Group Epilampra maya (Figs. 47-49), until recently considered a synonym of E. abdomennigrum (Figs. 50-55), was shown to be a valid species by Roth and Gurney (1969). They illustrated the genitalia of a large number of individuals of both species to show the extent of variation which occurs in the phallomeres. Epilampra abdomenni- grum is found in South America, Trinidad, and the West Indian Islands of Grenada and St. Lucia, but whether it occurred in Puerto Rico was uncertain (Roth and Gurney, 1969). The Puerto Rican record reported here (Figs. 50-52) suggests that Rehn and Hebard (1927, p. 228)
were probably correct in regarding Sein's ( 1923) record of wheeleri in Puerto Rico as actually being abdomennigrum. Epilampra maya occurs in Central America and Mexico. The male
taken in Boston Quarantine
(Figs. 47-49) had Jamaica as the lo-
cality but it is possible that the specimen boarded ship in a Central American port.
Rehn (1902) stated that E. maya is closely related to E. conspersa and E. azteca and that it is separated from the latter by the shape of the supraanal plate. E. maya is very close to abdomennigrum with which it has been confused', and the genitalia of azteca (Figs. 247-249) are decidedly different and I have placed it in the Bur- rneisteri Group.
Hebard (1929, p. 366) stated that E. sagitta is near (by markings) E. colunzbiana and E. opaca. However, the shape of L2d and prepuce of s&ta (Figs. 59, 62, 65) appear to be closer to those of abdomen- nigfium (Figs. 50, 53-55) than to columbiana (Figs. 208-219) and



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Psyche [December
Figs. 56-67. Cockroach male genitalia. 56-58. (83 USNM) . Epilum$ra taira. Surinam (dm. Gurney) (arrow in 58 indicates setai brush). 59- 67, Efilampra sagitta. 59-61. (74 U-SNM). Amapii, Brazil (det. Albuqucr- que ; confirmed by Frincis), 62-64. (182 ANSP). Type 1135. Teffe, Ama- zonan, Brazil. 65-67. (N). Florea, Manaun, Brazil (det Hoth). (scale = 0.2 mm)




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19701 Roth - Blattaria
447
fig^. 68-76. Cockroach male genitalia. EpiSaiafra &#a. 68-70. (42 ANSP) . Bariica, British Guiana (diet. Hebard). 71-73. (34- AMNH) . Surinam (labeled E. lu(i/upa Rehn, a synonym of grisea). 74-76. (157 USNM). Rosario, Lake Rogagua, Bolivia (det. Roth). (scale = 0.2 mm)



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Psyche [December
Figs. 77-96. Cockroach male genitalia. Epil~mpra grisea. 77. (.W USNM). Trinidad, British West Indies. 78. (93 USNM). Lelydorp, Suma- trawcg, Surinam. 79. (94 USNM). Surinam. 80. (96 USNM). Popogai- mama Creek, Surinam. 81-82. (156 USNM) . Blancaflor, Beni, Bolivia. S3-93. (N). 83-85. Flores, Manaua, Brazil. 86. Adolpho Ducke Forestry Preserve, about 60 Km. from Manaua, Brazil. 87. Puraquequara, Rio Negro, Amazonan, Brazil. 88-89. Moura, Rio Negro, Amazonas, Brazil. 90-93. Tapurucuara, Rio Negro, Amazonas, Brazil. 94. (107 USNM). 95. (108 USNM). At Plant Quarantine, Miami, Florida; with some bromeliads, possibly from Peru. 96. (65 WSNM).
At Miami, in plane, probably from
South America. (all det. Roth). (scale = 0.2 rnm]



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19701 Roth - Blattarh 449
Figs. 97-108.
Cockroach male genitalia. EfUainfra jorgensmi. 97-105. Farawpea of Efilamfra dgmaiiphra Rehn (=: E. jorqenseni). Miaionea, Argentina. 97-99. (86 AMP).
(arrow in 100 indicates setal brush}. 100- 102. (96 ANSP). 103-105. (48 AMSF). (Fig. 103 is a ventral view; part of the prepuce (arrow) in this specimen is missing). 106-109. (164 USNM), Between Coronel Oveido and Asuncion, Paraguay (det. Roth). (scale = 0.2 mm)




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450 Psyche [December
Figs. 109-117. Cockroach male genitalia. 109-1 13. Epitampra jargmseni. 109-1 10. (94 ANSP). Paratype of Ep'itampra stigm.uii$hora Rdn (= E. ]or- genscd). Misiono, Argentina. 111. (95 ANSP). Misiones, Argentina (det. Rehn). 112-113. (81 ANSP). same data as figs. 109-110. 114-117. Epilanpra
bcrhdi. 114, (47 ANSP}. Provincia Sara, Bolivia (det. Hebard). 115- 117. (83 ANSP). Paratype. Icafio, Santiago del Estero, Argentina. (arrow in 117 indicates setal brush). (scale = 0.2 rnm)



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Roth - Btattaria
Figs. 118-126. Cockroach male genitalia. Ep lampra guianae, 118-120. (35 ANSP), Paratype, Rockstone, British Guiana. 121-323. (91 USNM). Surinam. 124. (90 USNM). 125. (89 USNM). Brokopondo, Surinam. 126. (92 USNM). Surinam, (rips of prepuce (arrows) in figureh 125 and 126 missing). ( 121-126, det. Roth). (scale å´== 0.2 mm)



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452 Psyche [December
Figs. 127-135. Cockroach male genitalia. 127-133. Epilampra burmei- attri}. 127-129. (2.5 MCZ). Cuba (labeled E. raraibea S. and Z., which ia a synonym of bttrmeisteri). 130-131. (24 MCZ). Yimque de Baracoa, Oriente Province, Cuba (Set. Gurney). 132-133. (39 ANSP). Cuba (dct. Rehn). 134-135. (21 MCZ). Epilamfra takuna. Mountains north of Imias, Orierrte Province, Cuba (det. Gurney). The LI of taintma wag lost in preparation of the slide and the species is tentatively placed in die Bw- mei$lerf Group; all other Cuban Spilumpra belong to this Group. (scale = 0.2 mm)




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Figs. 136-144.
Cockroach male genitalia, Epiiumpra qtkqueiana. Para- types. 136-158. (37 ANSP). 139-141. (84 ASSP). San Lorenzo, Province of Saman4, Dominican Republic, Hispaniola. 142-144. (82 ANSP). San Francisco Mountains, Province of Santo Domingo, Dominican Republic. (scale = 0.2 mm)




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454 Psyche [December
opaca (Figs. 178-195 ) which I place in the Burmeisteri Group be- cause of the absence of a setal brush on LI. The prepuce of E. taira (Fig. 56) is relatively broad and extends well beyond the margins of the L2d. These structures resemble those of E. opaca but the right side of the prepuce of opaca is more extended and pointed (Fig. I 78).
The prepuce of E. grisea is greatly elongated, tapers to a point posteriorly, and shows remarkably little variation (Figs. 68, 71, 74, 77-81, 83-92, 94-96) over a wide geographical range. Based on Princis' (1967) catalogue, new geographical records for this species are Bolivia (Figs. 74-76)', Trinidad (Fig. 77)) and possibly Peru (Fig. 95).
The L2d of E. jorgenseni is unusual in being very large and deeply indented. The left side of the L2d tends to follow the contour of the underlying prepuce (Figs. 97, 100, 103, 106, 109, 112). One of the specimens (det. Rehn) apparently has an aberrant shaped L2d ( Fig. I I I ) . The genitalia of E. berlandi (Figs. I I 5-1 I 7 ) are very similar to those of jorgenseni and support Hebard's ( 1921, p. 283) claim that the two species are closely related. The Paraguayan record for jorgenseni (Figs. 106-108) is new.
The prepuce of E. guianae is unique for this group, and i,s deeply notched (Figs. I I 8, 12 I, 124-126) ; in this respect the prepuce is reminiscent of that of mexicana (Figs. 2, 5, 8, 10-12)) though the general shape is different between these 2 species. The prepuce of one unusual specimen of guianae has 2 indentations in the prepuce (Figs. 125).
Burmeisteri Group
[Epilampra burmeisteri (Guirin) (Figs. I 27- I 33 ) ; E. tainana Rehn and Hebard (Figs. 134-135) ; E. quisqueiana Rehn (Figs. I 36-144) ; E. sabulosa Walker (Figs. 145-150) ; E. wheeler! Rehn (Figs. 15 I- 156) ; E. gundlachi Rehn and Hebard (Figs. 157-162) ; E. haitensis Rehn and Hebard (Figs. 163-165) ; E. hamiltoni (Rehn) (Figs. 166- 168) ; E. bromel'iadarum (Caudell) (Figs. 169-1 7 I ) ; E. exploratrix (Gurney)
(Figs. 359-361 ) ; E. gatunae (Hebard) (Figs. 172-174) ; E. fugax (Bonfils) (Figs. I 75-177) ; E. opaca (Walker) (Figs. 178- 195 ) ; E. substrigata Walker (Figs. 196-207) ; E. columbiana Saus- sure (Figs. 208-219) ; E. latifrons Saussure and Zehntner (Figs. 299- 301) ; E. histriga Walker (Figs. 220-228) ; E. thunbergi Princis (Figs. 238-243) ; E. castanea Brunner (Figs. 244-246) ; E. azteca Saussure (Figs. 247-280) ; E. crossea Saussure (Figs. 293-298) 1. This Group is essentially similar to the Abdomennigrum Group



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19701 Roth - Blattaria 45 5
but is separated from it by the absence of a setal brush on LI. R2
usually has a subapical incision but a few species lack this character. The prepuce is usually well defined but in a few species it is markedly reduced.
The Burmeisteri Group includes the largest number of species of Epilampra and may be further divided into subgroups based on the relative sizes, shapes, and extent of development of L2d and the prepuce. Although not all of the species will fit readily into the fol- lowing subgroups many of them do show a close relationship and I believe an attempt at sub-divisions is worthwhile. Subgroup A:
(burmeisteri, tainana, quisqueiana, sabulosa, wheel- eri, gundlachi, haitensis, hamiltoni, bromeliadarum, gatunae, f ugax) : In thi,s subgroup the area of Lad is relatively small in relation to, and covers only a small anterior part of the prepuce. In burmeisteri (Figs. 127, 130, 132), quisqueiana (Figs. 136, 139, 142), and sabulosa (Figs. 145, 148, 150) the L2d is roughly divided in 2 parts, the left half usually being larger and sometimes more darkly pigmented


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