Louis M. Roth.
The Male Genitalia of Blattaria. VIII. Panchlora, Anthoblatta, Biolleya, Pelloblatta, and Archroblatta. (Blaberidae: Panchlorinae).
Psyche 78(4):296-305, 1971.

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THE MALE GENITALIA OF BLATTARIA. VIII.
PANCHLORA, ANCHOBLATTA, BIOLLEYA,
PELLOBLATTA, AND AClHROBLATTA.
(BLABERIDAE: PANCHLORINAE).
BY Louis M. ROTH
Pioneering Research Laboratory
U. S. Army Natick Laboratories
Natick, Massachusetts 01 760
McKittrick (1964) placed three blaberid genera, Panchlora Bur- meister, Achroblatta Saussure, and Capucina Saussure, in the Panch- lorinae. Princis ( 1960) listed Panchlora, Phortioecoides Rehn, and Proscratea Burmeister under the Panchloridae. My studies of male genitalia of Blaberidae have shown that Capucina and Phortioecoides are members of the Zetoborinae (Roth 19703). The genitalia of Proscratea differ markedly from those of genera I believe should be placed in the Panchlorinae and I do not consider it in this paper. The male genitalia of species of Panchloru, Achroblatta, Ancho- blatta Shelford, Biolleya Saussure, and Pelloblatta Rehn are basically similar and I place these five genera in the Panchlorinae. Unfor- tunately the male genitalia of an undescribed species of Pelloblatta were lost in preparation, due principally to their marked reduction and light sclerotization. Princis (1965) placed Pelloblatta in the Oxyhaloidae but the male genitalia of members belonging to the Oxyhaloinae (Roth, 1971 ) differ markedly from those of the Panch- lorinae. Princis (1960) placed Achroblatta in the Laxtinae, a sub- family which he considered provisional (Roth, 1970a), and Biolleya in the Latindiidae. McKittrick (1964) placed Laxta in the Epilam- prinae, tribe Laxtini. The genus Latindia Stil is an oviparous poly- phagid genus whereas Biolleya is an ovoviviparous member of the Blaberidae. Princis (1960) placed Anchoblatta in the Brachycolinae with a ( ?) but included it under this subfamily in his 1963 Catalogus. I consider Princis' Brachycolinae to be a tribe in the Blaberinae (Roth, igyob) . Anchoblatta signifera ( Scudder) was originally placed in Panchlora, but Kirby assigned it to Achroblatta. Princis (1963) listed Anchoblatta signifera with a query under Achroblatta, but Gurney (personal communication) regards it to be an Ancho- blatta.
MATERIALS AND METHODS
The genitalia were treated with 10% KOH and mounted in Per- mount. Considerable care must be taken with these specimens be- cause they are so lightly sclerotized and small that they can be readily



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Fig. 1. (978 L). Mate genitalia (dorsal view) of Panthtora vosiel~ Shelford. Usambara-Berg, Tanganyika (det. Princis). (LI = first sclerite of left phallnmere; L2d = dorsal sclerite of L2; L2vm = ventromedial aclerite; R2 =: hooked aclerite of right phallomere. To conserve space the photographs of the phallomeres have been placed closer together than they were arranged on the slide.
lost in preparation. Because of light pigmentation, Polaroid Type 51, high-contrast film usually was used to illustrate them (cf. Figs. 2 1-22).
The source of each of the specimens illustrated is given using the following abbreviations ; ( BMNH ) = British Museum (Natural History), London; (CUZM) = Copenhagen University, Zoological Museum, Denmark ; (L) Ì Zoological Institute, Lund, Sweden ; (N)
= U. S. Army Natick Labs.; (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




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298 Psyche [December
refers to the number assigned the specimen and its corresponding genitalia (on a slide) which are deposited in their respective museums. The most characteristic feature of the male genitalia of most Panchlorinae is the absence of the genital hook and light pigmenta- tion and marked reduction of the remaining 2 phallomeres when they are present. McKittrick (1964, p. 72) noted the marked reduction in the genitalia of Panchlora nivea and pointed out that the only sclei-otized structures are the cleft region of LI and the lightly pig- mented median sclerite (L2vm). However, a study of several species of Panchlora shows that some forms have all three phallomeres (Figs. 1-4) ; males that lack certain phallomeres probably evolved from species that had them. Based upon the presence or absence of genital phallomeres, the Panchlora studied here can be divided into the following Species Groups:
Group I (Panchlora vosseleri) . Though lightly sclerotized, all three basic phallomeres are present, including an L2d (Figs. 1-4). Group 2 (Panchlora stanle~ana). L2d is absent; LI, Lavm, and R2 are present (Figs. 5-7).
Group 3 (Panchlora nivea, P. thalassina). L2d and R2 are absent; LI, and Lmm, are present (Figs. 8-15). Group 4 (Panchlora bidentula, P. minor, P. sagax, P. dumicola, P. peruana). Lad, R2, and apparently L2vm are absent; only LI is present ( Figs. I 6-20).
Group 5 (Panchlora exoleta Burrneister). Apparently no sclero- tized structures are present.
Groups I and 2 are African and 3 to 5 are found in Central and/ or South America.
In Groups 4 and 5 it is quite possible that L2vm was so lightly sclerotized and LI sol small, and poorly defined, that they were lost in preparation of the genitalia.
The male phallomeres L2vm of Achro blatta (Figs. 24-25 ) , Biolleya (Figs. 21-23), and Anchoblattn (Figs. 26-29) are very lightly pig- mented. LI is also reduced as in most Panchlora, and Lad and R2 are absent.
The marked reduction in the male genital phallomei-es of this sub- family may be related, in some way, to the mating behavior of these genera. In most species of cockroaches the receptive females respond to male courtship by palpating his doi-sum, which in many genera possesses specialized glands which persumably produce a secretion at-



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Figs. 2-7, Male genitalia of Pauchlora spp. 24. (978 L). f. vasteleri (same specimen as shown in Fig. 1).
5-7. P. sfun~eyuaa Rehn.
5-6. (977 L)
Cameroon Republic (det. Primis). 7. (N). Ivory Coast (det. Ruth). (scale: 0.1 mm).




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Psyche [December
Figs. 8-16. Male genitalia of Pancklora spp. 8-9. (N). P, nivea (det. Gurney). 10-11. (56 BMNH). P. thalassina Saussure and Zehntner. Villa Ana, F.C.S.F. Argentina fdet. Hebard). 12-13, (173 USNM). P. sp. Palenque, Chiapaa, Mexico, 14-15. (172 USNM). P. sp. Paleaque, Chiapas, Mexico. (IS is a ventral view). 16. (57 BMNH). P. bideviula Hebard. Mosqueiro, Rio de Para (det. Hebard). (scale = 0.1 mm).



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h'i~s. 11-25, lviaie gennaiia ui a r n - i ~ ~ ~ ~ ~ ~ . .,. ,,,- -,. - - -- prruona Saussure. Bolivar, Bataral, Colombia (det. Princis). 18. (57
CUZM ) . Paachiara minor Saussure and Zehntner. Brazil (det. Printis) . 19. (973 L). Punchlora sagax Rehn and Hehard. St. Thomas, Lesser An- tilles (det. Princis). 20. (975 L). Punrl~lora dumicola Albuquerque and Gurney, Brazil (de!. Princis). 21-23. (174 USNM). Biolitya ahis Saus- sure.
Paratype. La Palma, Costa Rica.
(Figs. 21 and 22 are the same
L2vm taken with Polaroid Type 51 and 52 film respectively 1, 24-25 Arhro- blafta iuleola (Blanchard). Darien Province, Sante Fe, Panama (det. Fisk), (scale = 0.1 rnm).




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Psyche [December
Figs. 26-29. Male genitalia of ^nchoblalta. 26-27. (170 USNM). &mi- /era (Scudder). San Martin, Peru (det. Gurney). 28-29. (53 USNM). Rio Ucuyali, Pucallpa, Peru (det. Gurney). (L2vm was accidentally broken [arrows indicate the two parts of the ~clerite]). (scale =^ 0.1 mmi.



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19711 Roth - Blattaria 303
tractive to the female (Roth, 1969). When the female is in the proper position above the male he extends the genital hook (R2) and seems to use this structure to pull down the female's subgenital plate ,so that he can insert his genitalia and grasp her genitalia. Ac- cording to Khalifa ( 1958) in Blattella germanica (L.) , when the female is palpating the male's tergal glands, the male fully extends the hooked phallomere, which is on the left side in this genus (not the right as in all Blaberidae) directs it upwards, and inserts it into the female's genital chamber; there it clasps a sclerite situated in front of the ovipositor. Once a secure hold is obtained, the pair assumes an end-to-end position, with their heads facing in opposite directions, and the hook ". . . acquires a hold on the ovipositor." In hiplaneta
americana (L.) the male ". . . prior to getting an actual hold on the female pulls down the gynovalvular portion of the seventh ster- nite of the female by the tip of its protruded titillator." and then inserts its genitalia into the female's genital pouch. (Gupta 1947). Zabinski (1933) showed that the titillator, in Blatta orientalis, is used to seize the female in the initial phase of copulation. The male apparently cannot mate if the titillator (genital hook or R2 in Blaberidae) is surgically removed (Zabinski, I 933, Roth and Willis, 1952).
The mating behavior of PanchZora differs markedly from the above species. In Panchlora nivea (Roth and Willis, 1958) and P. irro- rata Hebard (Willis, 1966) the female does not assume a position above the male ~rior to mating, but the males simply back into the female. Possibly stridulation plays a role in mating behavior of these species (Roth and Hartman, 1967). The difference in precopulatory positions may have had some role in the marked reduction of male genitalic structures, especially the loss of the genital hook. Nothing is known of the precopulatoi-y positions of the genera of Achroblatta, Anchoblatta, and Biolleya, and it would be of interest to see if the mating behavior of these genera is similar to that of Panchlora. It
should be pointed out that in Gromphadorhina portentosa (Schaum) (Oxyhaloinae) the male also backs into the female to assume the copulatory position (Barth, 1968) but the males of this genus have a well-developed, but relatively short, genital hook (Roth, I 97 I ) .
SUMMARY
Five genera, Panchlora, Anchoblatta, BioZZeya, Pelloblatta, and AchrobZatta are included in the blaberid subfamily Panchlorinae. Cer- tain male genitalic phallomeres are usually reduced or absent and structures that are present are very lightly sclerotized. One African




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304 Psyche [December
species of Panchlora has all the phallomeres characteristic of most male blaberid genitalia. Five groups are erected for species of Panchlora, based on the number of phallomeres which are missing. The genital ho>oks (R2) are absent in Panchlora (from Central and/ or South America), A chroblatta, Biolleya, and Anchoblatta. ,- -.
1 he mating behavior of Panchlora differs from most other Blat- taria, and it is discussed in relation to the loss of the genital hook. I thank the following for the loan of museum specimens: Dr. David Ragge (British Museum), Dr. Ashley Gurney (U. S. Na- tional Museum), Dr. Karl Princis (Lund), Dr. S. L. Tuxen (Copenhagen), and Dr. Frank Fisk. I am grateful to Mr. Samuel Cohen for taking the photographs.
REFERENCES
BARTH, R. JR.
1968. The mating behavior of GromphadorJima portentosa (Schaum) (Blattaria, Blaberoidea, Blaberidae, Oxyhaloinae) : an anomalous pattern for a cockroach. Psyche 75: 124-131. GUPTA, P. D.
1947. On copulation and
insemination in the cockroach Periplaneta americana (Linn.).
Proc. Nat. Inst. Sci. India 13: 65-71.
KHALIFA, A.
1950. Spermatophore production in Blattella germmica L. Orthoptera: Blattidae). Proc. Roy. Entomol. Soc. London 25 (A) : 53-61. MCKITTRICK, F. A.
1964. Evolutionary studies of cockroaches. Cornell Univ. Agr. Exp. Sta. Mem. No. 389, 197 pp.
PRINCIS, K.
1960. Zur systematik der Blattarien. Eos 36: 427-449. 1963. Orthopterorum Catalogus. Subordo Polyphagoidea : Fam. Homoe- ogamiidae, Euthyrrhaphidae, Latindiidae, Anacompsidae, Atti- colidae, Attaphilidae. Subordo Blaberoidea : Fam. Blaberidae. s'-Gravenhage, pp. 76-172.
ROTH, L. M.
1969. The evolution of male tergal glands in the Blattaria. Ann. En-
tomol. SOC. Amer. 62: 176-208.
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. VI. Blaberidae: Oxyhaloinae. Psyche 78: 84-106.
ROTH, L. M. AND H. B. HARTMAN.
1967. Sound production and its evolutionary significance in the Blat- taria. Ann. Entomol. Soc. Amer. 60: 740-752.



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19711 Roth - Blattaria 305
ROTH, L. M. AND E. R. WILLIS.
1952.
A study of cockroach behavior. Amer. Midland Nat. 47: 66-129. 1958. The biology of Panchlora nbea with observations on the eggs of other Blattaria. Trans. Amer. Entomol. Soc. 83. 195-207. WILLIS, E. R.
1966. Biology and behavior of Panchlora irrorata, a cockroach ad- ventive on bananas. Ann. Entomol. Soc. Amer. 59: 514-516. ZABINSKI, J.
1933. Fonctionnement des diffkrentes parties des appareils copulateurs chitinks males et femelles de la Blatte (Periplaneta orientalis L.). SOC. de Biol. de Varsorvie 112: 598-602.



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