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Louis M. Roth.
The Male Genitalia of Blattaria. II. Poeciloderrhis spp. (Blaberidae: Epilamprinae).
Psyche 77:104-119, 1970.

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THE MALE GENITALIA OF BLATTARIA. 11.
POECILODERR3HIS SPP.
(BLABERIDAE : EPILAMPRINAE) .*
BY Louis AI. ROTH
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01760
Although the males of most species currently assigned to Epilainpra lack tergal glands, there are several with dorsal abdominal modifica- tions (Roth. 19693.) which are presumably involved in sexual be- havior. The presence or absence of tergal glands and the differences in the genitalia of these two groups of Epilampra warrant their being placed in different genera.
MATERIALS AND METHODS
The material used in this study were all museum specimens which were softened and had their genitalia removed after slitting the lateral margins of the terminal abdominal segments. The genitalia were treated with 10% KOH, cleared and mounted in Permount. In the illustrations, the hooked right phallomeres are mounted ventral side up, and phallomeres LI and L2d are mounted dorsal side upper- most.
The source of each of the specimens illustrated is given, using the following abbreviations: (AMNH) == American Museum of Natu- ral History, New York; (ANSP) = Academy of Natural Sciences, Philadelphia; (BMNH) = British Museum (Natural History), London; (L) = Zoological Institute, Lund, Sweden; (MCZ) = Museum of Comparative Zoology, Harvard University, Cambridge, Mass.; (USNM) = United States National Museum, Washington, D.C.
"Manuscript received by the editor January 15, 1970. Figs. 1-9. 1-3. Epi/ampra maya Rehn. (158, USNM). Boston Quaran- tine (det, Roth). 4-9. Poeciloderrhis proximo. 4-6. (22, MCZ). Brazil (det. Roth). 7-9. (20, BMNH). Brazil, Rio de Janeiro, Itatiaya (det. Roth). AL =-: accessory lobe; H == hook; LI = first sclerite of left phal- lomere; Lib ÌÔ setal brush of Ll; L2vm = median sclerite (L2 ventro- medial) ; L2d == dorsal sclerite of L2; P :ÌÔ prepuce; R2 = hooked sclerite of right phallomere;
C = cleft of Ll; FC = fused cleft of Ll; SI == subapical incision of R2; SL = subapical lobe. (scale = 0.2 mm). Pn&f 77:104-119 11970). http:ffpsyche.nilclub.ora/77/77-lM html



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19701 Roth - Male Genitalia of Blattaria 105



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i 06 Psyche [March
Figs, 10-15. Poeriloderrhis spp. 10-12. (1501, L). P. paxima. Ornt- kftldsvik, Sweden; tergal glands in Pigs. 48, 49. (Adventive) (de~ Princis). 13-15. (58, USNM). P. vwtica!u. Serta Caraca, Mam Groaso, Brazil; adult and tergal gland4 in Figs, 50-52. (det Gurney). (The tip of L2d waa inadvertantly cut off.) (scale =: 0.2 mm).



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19701 Roth - Mate Genitalia of Blattaria 107 Figs. 16-24. Foeciladerrflis spp, 16-22. P. fmyffwea. 16-18. (110, ANSP. 550 Paulo, Brazil {det Roth). 19-21. (31, AMNH). Corupq
Hansa Humbolt. St Catharina, Brazil. (dct. Roth). 22. (167, USNM). St. Catharinn, Nova Teutonia, Brad (det. Roth). 23-34. (45, ANSP). F. a~riventris. {det. Hebard). (scale = 0.2 nun).



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108 Psyche [March
Pip. 25-33. PaecUsdtt-rhis spp. 25-27. (12. BMNH). P. ferruginea. Rio de Janeiro, Organ Mts. near Tijiica, Brazil. (det. Primis). 28-33, Foedloderrhh ap. A. 28-30. (72, MCZ). Annapolis, Goias, Brazil; tergal gland in. Fig. 40, (det. Roth). 31-33. (109, ANSP). "Vlanoopolis", Goias, Brazil. (det. Roth). (scale = 0.2 m).




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Figs. 34-39. Poedoderrhis spp. 34-36.
(9, BMNH). P. atriwntrisi
ter~al gland3 in Figs. 43-44.
(det. Hebard). 37-39. (183, WSNM). P.
cntharinu.
(Type of dudrek catharim Shelford. St. Catharina, Brad; adult in Figs. 41-42.)
(wale = 0.2 mm).




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110 Psyche [March
Geographical collection data, and the specialists who identified the specimens, 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 museums indicated.
RESULTS AND DISCUSSION
In a previous paper on Efiilauztra, Roth and Gurney (1969) fol- lowed 3IcKittricPs ( 1964) terminology. In her figure 128 (p. 180) of the male genitalia of E9iZampra azteca Saussure there are 2 struc- tures below L2vn1 (median sclerite of left phallomere) one of which she labeled L2d (dorsal sclerite of left phallomere). In most Efii- lanzpra there is a sclerotized plate which lies at the base of L2vm (but separated from it by a thin membrane) (Fig. I ) and Roth and Gurney (1969) named this structure L2bp (basal plate of L2). However, I have studied the male genitalia of a large number of Blaberidae and believe that L2bp is actually L2d md what we and McIGttrick ( 1964) called L2d in Efiilmzpra is in reality a modified prepuce. In BZabems (Princis, 1946; Roth, 1969b) and 0th~ Blaberin~e (Roth, unpublished observations) the prepuce has char- acteristic spines associated with it and partly surrounds the virga (L2d) (See Fig. 121 in AlcKittrick 1964). ~<al (1874) without mentioning any species proposed the name Poeciloderrhis for American EfiiZamfira and separated them from the genus Efii/ampra (which he considered Asiatic-Australian) on the differences of front fen2ai-a1 spines. Kirby ( I 903) split the old genus Ejdan~p-a, proposing EIeteroZamfira for an Australian species, retained Burn~eister's Ejdampra for American species with E. brasiZiensis (Fab-)
as type species, and selected E. wrticalis Burm. (a species of Brazil and Argentina) as the type of Poeciloderrhis SGl. In his
1904 catalogue Kirby listed Poecilodcrrhis as valid, with verticalis as type; he also included under this genus, jv-oxinza Brunner, bivit- tafa Saussure, and maculifrons S&l. In the Genera Insectorum f asck1e an Epilampi-inae Shelford ( I 9 10) listed Poeciloderrhis as a syEonym of Epilampra, as does Pi-incis (1967a) in his catdogue. The species of Ejdmnpra in which males have tei-gal glands have markedly different genitalia than the species which lack these mod- ifications. EfiiZm~jv-a verticalis Burmeister has tergal glands on seg- ments I and 2 and since this species is the type of Poeciloderrhis I use this generic name to represent tergal modified species of Epi- lam pa.




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19701 Roth - itfaze Genitalia of BZattaria III The male genitaliz of species of Epilanzpra (lacking tei-gal glands) consist of 3 principal phallomeres (Roth and Gurney, 1969) which may vary sufficiently in shape to distinguish species and species groups (Roth, 1970). All 3 phallomeres of Efiilamfira differ basically from Poeci10derrhi.r (cf. Figs. 1-3 and 4-6). The two genera may be sep- arated in the following key:
I.
h1ales without tergal modifications on segments I and 2. L2d and L2vm separated from each other) and the prepuce ~~sually distinctly outlined and setose (Fig. I). Hooked portion of R2 usual~y relatively slender and tapering (mmetin~es broad- ened in the middle by a flange), with a subapical incision, and subapical lobe (Fig. 2). LI with 01- without a setal brush ............ (Fig. 3), and cleft not fused together. EpiZampra spp.
2. 3Iales xvith tel-gal glands on abd01nina1 segments I and 2 0111~7. L2d pointed and fused solidly to L2vm; prepuce membranous, not clearly defined by dense setae (Figs. 4, 7) 10, 13, 16, 19, 22, 23, 2.5) 28) 3 I ) 34) 37). R2 short a,nd stout, without apical incision (Figs. 5: 8, 11) 14, 17, 20, 24) 26, 29) 32, 35) 38). LI without a setal brush and the sclerotized cleft fused to- gether (F'igs.6,gj 12) 15, 18,21,27, 30) 33) 36,39). ............ ..................................................................... Poeciloderrhis spp. P 7
1 he tesgal glands on segments one and two of Poeciloderrhis show greater differences between species than do their genitalia and the following key chasacters may be used to distinguish the forms dis- c~ssed in this paper:
I. Glands v~ithout sclerotized horns or deep depressions on eithei- segments one and two. Segment I with a low median ridge flanked on each side by raised mounds bearing setae. Segment 2 with a transverse elevation bearing setae; mediallj7, a small wrtical ridge anterior to the elevation (Fig. 40). ............ sp. A. Tv!lodifications include medial knobs on segments I a,nd 2) or a sclerotized horn on segment 2 only. .................................... 2. 2.
Elroad, anteriorly rounded) relatively flat medial lcnobs on seg- ............
ments I and 2, catharina (Shelford) (Figs. 41, 42). atriwntris ( Saussure) (Figs. 43-44)
Segn~ent 2 with a black sclerotized horn arising from a deep de- ................................................................................... pression, 3-
3. Deep sclerotized pit behind the black horn on segment 2; horn relatively short and stout. Segment I with a well defined ............ medial mound! bearing fine short setae (Figs. 45-46). ferruginea (Brunner) ,
................................................................



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112 Psyche {March
Figs. 40-42. Poecilodcrrhis spp. 40.
A. Tergal modifications on sq&ent~ 3 and 2 (circled numbers). ~nna~olih, Goias, Brazil (det. Roth). (genitalia in Figs. 28-30), 4142. (183, CSNM). P. culbu~ina-
The ter~d gland! are indicated by arrows {Fig. 41 j and circled ni~merals (Fig. 42). (Type of Audrtio cathurin~ Shclford, St, Catharina, Brazil; genitalia in Figs. 37-39).



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Figs, 43-44, Poecdod~rrhis atriwnfri~, Tergal gland3 on segments i and 2 (circled numbers). (9, BMNH). Dorsal (Fig. 43) and laterai (Fig. 44)
views; genitalia in Fi@, 34-36). {det, Hebard).



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114 Psyche [March
Figs. 45-46. Poerdodmrh a ffrrugtnra. Tergal glands on segments I and 2 (circled numbers). (166, USNM). St. Catharim, Nova Teutonia, Brazil. Dorsal (Fig. 45) and lateral (Fig. 46) views. (det. Roth). Mi = pyramidal shaped, setose mound on segment 1; H =: sclerotized horn on segment 2; P = pit.




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19703 Roth -Male Genitalia of BlattarIa 115 Fig*. 47-49, Pocril~dfrrhJi froxima. 47. (22, MCZ). Bras'il. (det. Roth). 48-49. (1501, L). Tergal glands on segments 1 and 2 (circled numbers), OrnskiSldsvik, Sweden (adventive). Dorsal (Pig, 48) and lateral (Fig. 49) views; genitalia in Figs. 10-12. (det. Printis). Ml = pyramidal shaped mound on segment 1; H = sclerotized horn on seg- ment 2 ; M2 ==: pyramidal shaped mound on segment 2.



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116 Psyche [March




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19701 Roth - Male Genitalia of Blattarza 117 Pyramidal shaped elevation on segment 2, behind a slender horn, which is elongate, hook shaped, directed anteriorly, and reaches ........................................ the posterior margin of segment I.
4.
4. Anterior margin of segment 2 roundly indented, semicircular in outline. Medial moundlike elevation on segment I similar in shape though smaller than the one found behind the horn on . ......................... segment 2 ( Figs. 48-49) proximo. ( Brunner )
Anterior margin of segment 2 not as deeply excavated nor as clearly semicircular in outline. Raised medial ridge on segment I not pyramidal in shape but more or less uniform in height for its entire length (Figs. 50-52). .... verticalis (Burmeister) . The subgenital plates of Poeciloderrhis males are highly asym- metrical and more or less similar in shape (see Fig. 8, in Princis, 1967b). Epilampra cribrosa Burm. has tergites I and 2 modified ( Princis, I 967b) and therefore is a. Poeciloderrhis. The status of the genus Audreia Shelford has been questionable and was based principally on a reduction in wings and tegmina in both sexes.
It will be reported on elsewhere. The genitalia of Audreia catharina Shelford are obviously typical of species of "Epilampra" which have tergal glands and I therefore place it in the genus Poecilodcrrhix. The tergal glands and genitalia of catharina are closer to airiventris than they are to the other species of the genus. There may be some question as to the identity of verticalis and proximo. These are species which have lateral pale markings of the pronotum (Figs. 47, 50). The genitalia of proximo shown in Figs. 4-6, came from a specimen (Fig. 47) determined by Rehn as ver1- ticalls; however, its tergal gland was similar to the specimen deter- mined by Princis as proximo.
(Figs. 48, 49). I have seen a third
species belonging to this con~plex (pale, lateral pronotal coloration) whose tergal glands differed from the species here considered to be proxima and verticalis. In the final analysis, only type males can solve questions of identity in these similar appearing species. Epilampra is in the tribe Epilamprini ( McKittrick, I 964). Be- cause of the morphologic (tergal glands, subgenital s late) and geni- talic differences between Epilanzpra and Poeciloderrhis I suggest that the latter genus be assigned to the Poeciloderrhini. Figs. 50-52. Poeciloderrhis verticalis. (58, USNM). Serra Caraqa, Matto Grosso, Brazil; genitalia in Figs. 13-15. The tergal glands are indicated by arrows (Fig. SO), and circled numbers (Fig. 51, dorsal). Fig. 52 is a lateral view. (det. Gurney). Rl = medial ridge on segment 1 ; H = sclerotized horn on segment 2; M2 ==: pyramidal sh,aped horn nn segment 2.




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Psyche
[March
SUMMARY
Based on differences in male genitalia, and the presence or absence of tergal glands, species of Epihpra are divided into 2 genera. Species in which the males have tergal glands on segments I and 2 are placed in the genus Poeciloderrhis Sfal. Species with males lacking glandular modifications on the tergites are restricted to the genus Epilampra Burmeister. Poeciloderrhis is placed in the tribe Poecilo- derrhini.
ACKNOWLEDGEMENTS
I thank the following for the loan of Museum material: Dr. M. G. Emslsey, Academy of Natural Sciences, Philadelphia, Dr. Ashley B. Gurney, U. S. National Museum, Dr. Karl Princis, Zoological Institute, Lund University, Dr. D. R. Ragge, British Museum (Nat- ural History), and Dr. Jerome G. Rozen, Jr., American Museum of Natural History. The type of Audreia catharina Shelford was made available from the Hope Department of Zoology (Entomology), University of Oxford, through Dr. Gurney. I am especially grateful to Dr. Gurney for his helpful suggestions, and Mr. Samuel Cohen for taking the photographs.
REFERENCES
KIRBY, W. F.
1903. Notes on Blattidae, etc., with descriptions of new genera and species in the collection of the British Museum, South Kensington. No. 11. Ann. Mag. Nat. Hist. ser. 7, 12: 273-280. 1904. A synonymic catalogue of Orthoptera. 11. London. 501 pp. MCKITTRICK, F. A.
1964. Evolutionary studies of cockroaches. Cornell Univ. Agric. Expt. Stat. Mem. 389: 197 pp.
PRINCIS, K.
1946. Zur Kenntnis der Gattung Blaberus Serv. (Blatt.) . Opusc. Entomol. 11 : 139-146.
1967a. Orthopterorum Catalogus. Pars 11. Epilamproidea, pp. 616-710. s-Gravenhage.
1967b.Kleine Beitrage zur Kenntnis der Blattarien und ihrer Verbrei- tung. X. Opus. Entomol. 32 : 141-151.
ROTH, L. M.
1969a.The evolution of male tergal glands in the Blattaria. Ann. Entomol. SOC. Amer. 62: 176-208.
1969b. The male genitalia of Blattaria. I. Blaberus spp. Psyche 76: 217- 250.
1970. The male genitalia of Blattaria. 111. Epilampra spp. (Blaberi- dae: Epilamprinae). Psyche (in press)




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19701 Roth - Male Genitalia of Blattaria 119 ROTH, L. M. AND A. B. GURNEY.
1969. Neotropical cockroaches of the Epilampra abdomennigmm com- plex:
a clarification of their systematics. (Di~tyoptera~ Blattaria). Ann. Entomol. SOC. Amer. 62: 617-627.
SHELFORD? R.
1910. Orthoptera? Fam. Blattidae) Subfam. Epilamprinae. Genera In- sectorum, lo 1 : 1-21) Bruxelles.
ST~L? C.
1874. Recherches sur le syst6me des Blattaires. Bihang Sv. Vet. Ak. Hand]. 2. No. 13: 1-18.




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