L. M. Roth.
The Male Genitalia of Blattaria. XI. Perisphaeriinae.
Psyche 80(4):305-348, 1973.

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THE MALE GENITALIA OF BLATTARIA.
XI. PERISPHAERIINAE.*
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 I I genera (indicated with an ** in Table I ) of cockroaches which belong to the Perisphaeriinae. Three recent classifications which include these genera are shown in the table. I follow McKittrick ( 1964) in placing all of those discussed here under the Blaberidae: Perisphaeriinae, and Beier (1961) in using tribes to separate them. Rehn (1922) discussed the characteristics of Pilema, Cyrtotria, Dero- calymma, Zuluia, and Hostilia, and Princis (1963) devised keys to some South African species of the above genera, as well as those of Bantua, Perisphaeria, and Blephamdera.
All genitalia were treated with 10% KOH, dehydrated, cleared, and mounted in Permount. The source of each of the specimens examined is given using the following abbreviations: (ANSP) = Academy of Natural Sciences, Philadelphia; (BMNH) = British Museum (Natural History), London; (CUZM) = Zoological Museum, Copenhagen; (L) = Zoological Institute, Lund, Sweden; (HO) = Hope Department of Entomology, Oxford, England; (MCZ) = Museum of Comparative Zoology, Harvard University, Cambridge, Mass. Locality data and names of specialists who identi- fied the specimens, if known, follow these abbreviations; most of the species were determined by Princis or Rehn. The number preceding the abbreviations refers to the number assigned the specimen and its corrsponding genitalia (on a slide) which are deposited in their re- spective museums.
The normal arrangement of the phallomeres of male genitalia of the Perisphaeriinae, as seen in dorsal view, is illustrated in figure 1. Based on male genitalia, I suggest the erection of 4 tribes for the *Manuscript received by the editor October 10, 1973 Pu&e W:W-M t 1971). http:llpsyche cnlclub ore/SWSOMS html



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306 Psyche [December
Table I.
Three recent classifications of some of the species of Blattaria discussed in this paper (nm-ked with an *) -
Princis ( I 960)
Bciei- ( I 96 I )
McKittrick ( 1964)
Derocalymmidae Blaberidae Blaberidae
Apterinae
Aptera Sauss.
E /lip t ica
Sauss. & Zehn.
Elliptobtatta
Sauss. & Zehn.
Gymnonyx
Sauss. & Zehn.
Thoracopygia
Sauss. & Zehn.
Derocalymminae
l*Bantua Shelf.
Co ?npsagis
Chop.
*Cyrtotria
Stil
*Derocalynz7na
Burm.
Glom erihiatta
Bey-Bienko
*Host ilia
Stil
* Perisp haerus
Serv.
Vilema Sauss.
Platysilp ha
Shelf.
Pronaonota
Sauss. & Zehn.
*Pseudoglomeris
Brunner
l*Zuluia Rehn
Perisphaeriinae
Apterini
A piera
Elliptica
Ellipto blatta
Gymnonyx
Thoracop ygia
Derocalymmini
Perisp haerus
Glomeriblatta
Pseudoglomeris
Fronaonota
Cyrtotria
Hostilia
Co m psagis
Derocalymma
Platysilpha
and others
Perisphaeriini
Perisphaeria
Blepharodera
Poeciloblatta
Parasphaeria
Gy na
Paranaup hoeta
Paraplecta
and others
Bsachycolinic
Brachycola Sei-v.
å´Hormetic Burm.
Parahormetica
Brunner
and others
Perisphaeriinae
Perisp haeria
( = Hyposphaeria)
Derocaly mina
Eustegasta Gerst.61
Gym
Pilema
Perisp haerus
Isoniscus BorgWa




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Princis ( I 960) Bei,er ( I 96 I )
McKittrick ( I 964)
-
Perisphaeriidae
Gyninae
Evea Shelf.
Gyna Brunnerb
Paranuup hoeta
Brunner
Paraplecta Shelf.
Progonogamia
Rmehn
ThZiptoblatta
Sauss. & Zehn.
Perisphaeriinae
*Blepharodera
Burm.
Parasp haeria
Brunnerd
*Peris$ haeria
Burm.
Poeciloblatta
Sauss. & Zehn.
Vrichoblatta
Sauss. & Zehn.
Lanxoblatta Hebard
Phortioeca Sauss.
Schistopeltis Rehn
Schizopilia Burm.
Tribonoidea Shelf.
Tribgonium Sauss.
Zetobora Burm.
Zetoborella Hebard
aPrincis (1960) placed these 2 genera in Blaberidae: Laxtinae. Beier (1961) placed Eustegasta in the Panchlorinae: Laxtini. The $ genitalia of Panchlorinae are characterized by a marked reduction or absence of one or more of the phallomeres
(Roth, 1971; Gurney and Roth, 1972). The genitalia of Eustegasta are quite different from members of the Panchlorinae (unpublished~ observations).
^The genitalia of Gyna spp. have been described (Roth, 1972). c1 (1970a) followed McKittrick (1964) and placed this tribe in Blaberi- nae.
^The genitalia of Parasphueria ovata (Blanchard) (South America) is more typical of members of the Epilamprinae and I placed it in this sub- family (Roth, 1973).
e1 (1970) followed McKittrick (1964) in placing this subfamily closer to the Blaberinae (Blaberoid Complex) than to the Perisphaeriinae (Epilam- proid Complex).




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Psyche [December
Figure 1. (62 BMNH). Male genitalia (dorsal) (rf CyrtotIta lalipennu (Kirby), British East Africa (from & shown in Fig. 37). Left (L) and . right (R) phallomeres numbered according to McKittrick (1964) ; C = cleft of LI; UL and LL-upper and lower lobes of Ll, respectively. genera studied here; additional tribes will undoubtedly be required for other genera of this subfamily (unpublished observations) . Perisphaeriini. - The upper lobe of Li is usually about the same height or slightly larger than the lower lobe and it has a snoutlike extension (accidentally lost in some of the illustrated preparations) ; setae are absent from the lower lobe. The Lad is variable in shape, its surface distinctlv mieular fat least in ~artl, or reticulate (at e
1e
into groups.
Tribal Group I. The entire or at least part of the surface of Lad is spicular (not reticulate) : Perhphaeria, mharodera, Pllema, and



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9731 Roth - Blattaria 309
Figures 2-9. Adult males of Perisphaeria spp. 2. (1424 L). P. virescens (Erunner). (det. Princis). 3. (40 CUZM). P. J ~ ? U (Thunb,). Cap bono Spei, Dec. 1816. 4. (1420 L). P. scahra. Hotub, South Africa (det Princis), 5. (1419 L), P. saxicola (Esch.) Capetown, Table Mountain, 2400' (deb Princis). 6.
(1422 L). P. ruftcotnis (Sauss. and Zebu.), Swart- bergpas, Platberg, Cape Province, South Africa, ca. 5000' (det. Princis). 7. (14-21 L). P. ityltfera. Burm. Manhica fdet. Frincia). 8. (161 ANSI'). P. styliferu. Durban, (det. by Rehn as Hyfosphaeria giebdiana (Sauss, and Zehn.). 9. (1423 I.). P. aspera (Walker). Baberton, Levetzow (det.



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ens (from $ shown
shown in Fig. 3). 16-




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Figures 19-27. Male genital phallomeres of Per'tifhaeria and Cyrtotria spp, 19-21. (1419 L). A saxicola (from $ shown in Fig. 5). 22-24. (1422 L). P. rufirornis (from 8 shown in Fig, 6). 25-27. (IS HO). Gyrtotria sp. (from 8 shown in Fig. 10s). (scale = 0.2 mm).



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312 Psyche [December
^"it^<å´^:
i
Figures 23-36. Ma!e genital phallomeres of Perts$hae~iu spp. 28-30. (1421 L). P. stylifcra (from 8 shown in Fig. 7). 31-33. (161 W). P. styl'ifera (from 8 shown in Fig. 8). 34-36. (1423 L). P. alpera (from 8 shown in Fig. 9). (scales 0.2 mm).




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19731 Roth - Blattaria
Figures 37-42.
Adult males of Cyriotria, Pilema, and Hostilia spp. 37. (62 BMNH). C. LaHfennii (holotype of Thysunoblatta ldifennis Kirby) British East Africa. 38. (53 BMNH). P. dentata Sauss. and Zehn. Pre- toria, South Africa (det. Princi's). 39. (989 L). P. cribrosa (Sauss.). Pretoria, South Africa (det. Princis).
40. (990 L). P. thorarica (Walker).
Mamathes, Basutotand (del:. Princis). 41. (991 L). P. refexa (Walker). Zulutand, Mtubatuba Dist., Dukuduku, South Africa (det. Princis). 42. (1406 L) . H, c!avigera (Kirby). Blouberg, Transvaal (det. Frincis). (scale = Pigs. 37, 41, 42 = 5 mrn ; Figs. 38-40 = 10 mm) .



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314 Psyche [December




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Roth - Blattaria
Figures 43-50.
Adult males of Hostilia, Blepharodera, and Bantua spp. 43. (170 ANSP). H. proterva (st&). Pinetown, Natal, South Africa (det. Rehn). 44. (1405 L). H. proterva. Stanford Hill, Durban, Natal, South Africa (det. Princis). 45. (1426 L) . Blepharodera discoida.5~ (Brunner). Cap. 1872 (det. Princis).
46. (1425 L). B. ciliata Burm. Hopefield, C.P., X. 1883 (det. Princis). 47. (34 CUZM). B. ciliata. Cap. bono Spei, June 1917 (det. Princis). 48. (1404 L). Bantua scabra (Walker). Okahandja, S.W. Africa (det. Princis). 49. (999 L). B. repentina Rehn. Kruger National Park, South African Transvaal (det. Princis).
50. (1403 L). B. robusta. (Shelf.)
Sandspruit, 15 miles northeast of Pietersburg, South African Transvaal (det. Princis). (scale = 5 mm) .




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[December
Figures 51-58. Male genital phallomeres of Hostilia spp. 51-53. (1405 L). H. proterwa (from 3 shown in Fig. 4.4). 54-56. (170 ANSP). H. frotmff (from $ shown in Fig. 43).
57-58. (1406 L). H. fluwigera (from
8 shown in Fig. 43). (scale = 0.2 mm).




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Figures 59-70. Male genital phalhmeres of Pilema spp. 59-61. (989 L). '. cribroia (from 3 shown in Fig. 39). 62-64. (53 BMNH). F. devfata from $ shown in Fig. 38). 65-67. (SO0 L). P. fhoracica (from 8 shown scale == 0.2 mm).




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Figures 71-79. Male genital phallomeres of Blefhurodera spp. 71-73. 1426 L). I?. discotduL)~ (from $ shown in Fig, 45). 74-76, (34 CUZM). I. discoidah (from 3 shown in Fig. 47). 77-79, (1425 u. B. cilkta from 8 shown in Fig. 46). (scale = 0.2 mm).



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19731 Roth - Bhttaria
319
Figures go 83.
Male genital phallomeres of Bftnfua spp, 80-82. (1403 L).
B. robwta (from 8 shown in Fig. SO). 83-85. (148 ANSP). B. robusta (Sheiford), paratype of Banlua armigera Kcho, 86-88. (54 BMNH). 5, war. Distant Coll. (labeled Dewalywma palUpes Kirby). (scale= 0.2 mm),




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Psyche [December
Figures 89-97. Male genital phailomeres of Bastua and Cyrloiria spp. 89-91. (999 L). B. repentinu (from 8 shown in Fig. 49). 92-94. (1404 L). 3. scah (from 8 shown in Fig. 4s). 95-97. (993 L). C. grander (from 8 shown in Fig, 99). (scale = 0.2 tnm).




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In Perisphaeria 2 species groups can be distinguished based on the R2:
Species Group I. R2 with a distinct, deep, subapical incision (Fig. 23). P. stylifera (Figs. 7-8, 28-33), P. aspera (Figs. 9, 34- 36) ; these 2 species also have very similar shaped L2d's (Figs. 28, 34) ; P. ruficornis (Figs. 6, 22-24).
Species Group 2. R2 without a subapical incision, or, if present, weakly represented. P. virescens (Figs. 2, 10-12) has a slight in- cision in the middle of the hook (Fig. 11) and its L2d lacks any extension (Fig. 10). P. scabra (Figs. 3, 13-18) and P. saxicola (Figs. 9, 19-21 ) lack incisions in their R2's (Figs. 14, I 7, 20) ; both species have smooth outgrowths arising from the right side of the spicular L2d (Figs. 13, 16, 19).
In Hostilia (Figs. 42-44, 51-58) the lower margin and left side of L2d is spicular and a smooth hook-like extension arises from the upper right corner (Figs. 51, 54, 57). R2 has a shallow subapical incision (Figs. 52, 55).
In Pilema (Figs. 39-41, 59-70), the oxter margin of R2 is rela- tively straight. Three species, cribrosa, dentata, and thoracica have deep subapical incisions (Figs. 60, 63, 66), whereas reflexa (Fig. 69) has a relatively shallow one. Thoracica and reflexa have smooth outgrowths from the right side of L2d (Figs. 65, 68), which appear to be lacking or weakly indicated in cribrosa and dentata (Figs. 59, 62).
In Blepharodera (Figs. 45-47, 71-79) the spicules on L2d are very minute; the L2d outgrowth is broad in P. discoidalis (Fig. 71) and much narrower in B. ciliata (Figs. 74, 77). R2 of discoidalis has a minute lobe at about the middle of the hook (Fig. 72), whereas in ciliata there is no subapical incision (Figs. 75, 78). Tribal Group II. Part of the Lad surface on the left is reticulate (not spicular) (e.g. Figs. 80, 121), but otherwise very similar to Group I having a lateral extension on the right: Bantua (Figs. 48- 50, 80-94) and Cyrtotria (Figs. 19-21, 94-144). All of the 5 species of Bantua examined have R2's with deep sub- apical i'ncisions that extend almost to the middle of the curved part of the hook (Figs. 81, 84, 87, go, 93). Their L2d's are also basi- cally similar in having a smooth extension on the right side arising from the reticulate portion of the sclerite (Figs. 80, 83, 86, 89, 92). Cyrtotria is very close to Bantua as evidenced by the similarity of the Lad's of some of the species. There are 2 groups of Cyrtotria based on the presence or absence of a subapical incision: Species Group I. Subapical incision on R2 absent. C. pallicomis (Figs. 108, 109-III), C. marshalli (Figs. 106, 107, 121-126). C. latipennis (Figs. I, 3, 7, 101, 136-144).



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Psyche
[December




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Roth - Blattaria
Figures 98-108. Adult males of Cyrtotria spp. 98. (63 BMNH). C. gibbicollis (st;]). Natal. (labeled Perisphaeria linearis Walker, selected as a lectotype by K. Princis). 99. (993 L). C. graniger (Sauss. and Zehn.). Mudugh Prov., Somalia (det. Princis). 100. (997 L). C. tuberculata Shel- ford. Lukuledi, German East Africa (det. Princis). 101. (151 ANSP). C. latipennis (Kirby). N.E. Africa (det. Princis; Rehn det. the specimen as C. capucina, but the genitalia are similar to the type of latipennis (cf. Figs. 142-144 and 136-138) ; the L2d of capucina (Fig. 130) differs from that of latipennis (Fig. 136). 102. (998 L). C. orientalis (Chopard). Kasenyi, East Congo ((det. Princis).
103. (56 CUZM). C. orientalis Kasenyi, East Congo. 104. (996 L). C. capucina (Gerst.). Kakamego (det. Princis). 105. (15 HO). C. sp. Damaraland, South Africa. Shelford selected this specimen as the $ holotype of C. poduriformis (Walker) but Princis (personal communi- cation) claimed that he had no authority to select an additional type to the $ holotype in the British Museum; the L2d surface (Fig. 25) and out- growth differ somewhat from the L2d of the specimen which Princis believes to be poduriformis (Fig. 133). 106. (995 L). C. marshalli Shelford. ES- pungabera (det. Princis). 107. (14 HO) . C. marshalli, holotype, Salisbury, Mashonaland, 5000'. 108. (992 L). C. pallicornis (Kirby). Pretoria, South African Transvaal (det. Princis) . (scale = 5 mm).



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Figures 118-126. Male genital phallomerea of Cjwtotria ~pp. 118-120.
63 BMNH). C. gttbwtlii (from 8 shown in Fig. 98). 121-123. (14 HO). warfhallt (from 8 shown in Pig. 107). 124-126. (995 L). C, marshain A 1'" nirr ini\ /---i- - n - -- --I




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.-
Figures 127-135. Male genital phalkmeres of Cyrtotria app. 127-129. (99g L).
C. orientah (from 5 shown in Fig. 102). 130-132. (996 L). C. tafu&a (from 8 shown in Fig, I#). 133-13s. (55 BMNH). C. $da- riforMis. Natal, Sooth, Africa (k PriacM), (scale = 0.l m}.



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Figurea 136-144. Male genital phallonieres of Cyrtotria latipen& 136- 138. (62 BMNH). (from 3 shown in Fig. 37). 139-141. (995 L). Kitui (defc Primis). 142-I++. (151 ANSF). (from 8 &own in Fig. 101),- (scale ~ 0 . 1 mrn).




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Psyche [December
Figures U5-14s. Adult males of Zulufa spp. 145, (1408 L), 2. lithv- strata Rehn, Letsletle Valley, Gravelotte Djst., South African Transvaal {det. Princis).
146, (154 ANSF). Z. fubeicens (Sausa.) . Durban, paratype of 2. mttella Rehn. 147. (145 ANSP). 2. Abscissa {Walker), Durban (det. Rehn; Princia [1960, p. 1411 states that Rehn's identification of ah- swia was an error and that the specimen was Z. litftastrota Rehn. The L2d of this specimen (Fig. 158) is more like that of abscissa (Fig. 155) than lithostrofa (Fig. 149)). 148. (1407 L). 7,. abscissa (det Princis). . . < ,




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330 Psyche
[December
Species Group 2. Subapical incision on R2 present. C. graniger (Figs. 95-97, 99)) C. orientalis (Figs. 102, 103, 127-1291, C. capu- cina (Figs. 104, 130-1321, C. podurijornzis (Figs. 133-1351, C. tubercuZata (Figs. 100, I 12-1 14), C. gibbicollis (Figs. 98, I 18-120) ) C, sp. (Figs. 25-27, 105).
Zuluiini. - L2d is an unevenly outlined plate, the left side of which is scale-like or smooth; R2 with a subapical incision. LI sim- ilar to Bantua. I place Zuiuia (Figs. 145-160)) an African genus, in this tribe. 2. abscixsa (Figs. 155, 158) has a lobe-like extemion on the left side of L2d which is absent in 2, lithostrota (Fig. 149) and 2. pubescens (Fig. I 52).
Rehn (1932) stated that ZuZuia is a relative of DerocaZymma and Hostilia, but in general, the selationship appeared closer to Hostilia. The d genital phallomeres do not suppo~t this suggestion. Derocalyn~mini. - LI, which is very similar in all the species examined, has a deep cleft which may curve upwards slightly; norn~ally th,e margins of the 2 10,bes are close together (e.g., Figs. 187, ~gg), but in some preparations the phallomere was distorted by pressure of the coverslip so that the sclerotized margins were widely separated (e.g., Figs. 178, 196). The upper lobe is usually somewhat mr~-ower and more elongate than the lower lobe, and the latter lacks setae. L2d is a broad flat sclerotized plate of variable shape, slightly szparated from Lmm, ,and whose surface appears weakly wrinkled or scale-like. The R2, generally similar in all species examined, is mnmdified, lacking an apical notch or subapical incision. I place
Deroculynznza (Figs. I 61-227)) an African genus, in this tribe. Based on the shape of L2d, I separate 2 species groups: Species Group I. The L2d variable in shape, but lacking a finger-like projection on the right side: D. Zampyrina (Figs. 161, I 73-1 78)) D. kalahari (Figs. 163, 182-1 84), D. granuZata (Figs. 164, 185- go), D. porcellio (Figs. 165, 191-1961, D. silphoid~s (Figs. 166, 197-199)~ D. rajiformis (Figs. 167, 200-205)) D. cru- raZis (Figs. 168, 206-208)~ D. versicolor (Figs. 169, 209-2141, D. sp. (Figs. 162, 179-181).
Species Group 2. Distinct finger-like projection extending from the right side of the L2d: D. coftata (Figs. 170, 215-217)~ D. pluteus (Figs. I 7 I, 2 I 8-223), D. scr,uposa scmposa (Figs I 72, 224-226).
Rehn ( 1932) in a, study of Derocalymma, using female characters, arranged the species in what seemed to him to be a natural phylo- genetic ordei-. He considered D. plut,eus as the least specialized of the genus (the shows no traces of caudo-lateral diverging "gut-



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Figures 161-1615,. Adult male~ of Derocu~ymmu spp. 161. (37 CUZM). D. !ampyrinu Gerstaecker, (det. Princi~}. 162. (141 ANSP). R. ap. Ger- man East Africa (det. by Rehn as lamfiyritaa and by Princia as wersico~of; the Ud (Fig. 141) differs from bokh these species (see Fiss. 173, 176 and 209, 212). 163. (1+14 L). D. kdahari Rehn, Khomashochland, S.W. Africa (det. Pxincis).
I&, (1413 L).
R. granulata S~uss. and Zeh. S.W. Africa (det. Princi*). 165. (1416 L), D, porcdio Gerstaecker. Morogoro, German East Africa (det. Primis). 166. (1411 I.). D. d$hoidt$ Bolhar. Abachau3, Darmarfand (det. Primis).
(scale = 5 mm).




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332 Psyche [December
Figures 167-172. Adult males of Demrdymm~ spp. 167. (38 CUZM). D. rujiformh Rehn. Kenya (det. Rrincis). 168. (1410 L). R. mwah (Stal). Bipindi, South Cameroon (det. Prin&), 169. (1412 L). D, versi- rdor Burmeister. Southern Rhodesia (de~ Princh). 170. (1417 L). D. cosiata Rehn, Cameroon (det. Princis), 171- (1409 L). D, phiem Rehn. Haut-Luapula, Kansenia (det, Princis). 172. (1418 L). B. ~crupo~a JCW- posu Ftehn. Tcigo, Misahohi-Mittc (det. Princk). (scale 5 rnm).



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19731 Roth - Biattarb 333
Figures 173-181. Male genital pha~lomeres of Deroculymma spp. 173- 178. (1415 L). D. iampyri~u. Okawngo, Runto, S.W. Africa (det. Printis}. 176-17k (37 CUZM). (from 8 shown in Fig. 161). 179-181. (I41 ANSP). D, sp. German East Africa (from 8 shown in Fig. 162). (scde = 0.1 mm).



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Figum lK!-190. Male genital phallomm of D~~oculymma spp. 182-184, (1414 L). D. kdahari. (from 8 shown in Fig. 163). 185-190. D. randata. 185-187. (1413 L). (from 8 ahown in Fig. 164). 18s-190. (143 AMP). Damara (det. Rehn). (scale = 0.1 nun).




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Figures 191-199. Male genital phallomeres of Dersculymma spp. 191- 196. D. ~o~cdlio, 191-193. (1416 L). (from 8 shown in Pig, 165). 194- 196. (167 ANSP). Tendaguru, Lindi, German East Africa (dct Hebard). 197-199. (141 t L). D. dfhoides (from 8 shown in Fig. 166). (scale= 0.1 rnm).




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[December
Figures 200-208. Male genital phaliomerea of Deracalymma spp. 206- 205. D. miifor& 200-202. (38 CUZM). (from 8 shown in Fig. 167). 203-205. (169 ANSP). Bukama (Muaee du Congo) (det Rehn). 206-308.
(14-10 L). D. cwah (from 4 ahown ID fig, 168). (scale= 0.1 ram),



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Figures 209-217. Male genital phallomeres of Derocalymma spp. 209- 214. D. wer~icolor. 209-211. (142 AMP). Gabeionea, Bechuanaland (del Rehn). 212-214. (14t2 L). (from 8 shown in Fig. 169). 215-217. (1417 L). D. cottda (from 8 shown in Fig. 170). (̤;ale=0. nun).



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Figures 218-226. Male genital phailomerea of Derocalymma 3pp. 218- 23, D. pluteus. 218-220. (157 ANSP). Paratype, Mayurnbe, Congo, 221- Ìö (1409 L). (from $ shown in Fig. 171). 224-326. (1418 L). D. stru- *** *---.A*-," tx--- A -L--- ?- m-- a"'., , * .. - .



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19731 Roth - Blattaria 339
ters", the mediocephalic portion of the disk virtually without a me- dian carinula) and the ventro-lateral faces of the pronotun1 are not inflated or bullate, their surface deplanate). The L2d of pluteus with its finger-like projection falls into group 2 along with costata and scruposa, two species which are almost at the opposite extremes of Rehn's listing. The finger-like extension could represent a "primi- tive" character which was lost in the species placed in group I.
However, it is also possible that the L2d of pluteus represents a more highly evolved form with the extension developing from species which lacked it.
Peri~~haerini. - Unfortunately, the spelling of this tribe differs only slightly from the Perisphaeriini, but Rehn ( 195 I) used Peri- sphaerus to erect the tribe based on wing venation. LI of this tribe is distinctive in that the upper lobe is considerably larger than the lower lobe, with a snoutlike projection over the cleft opening and a patch of numerous, moderately spaced setae on the lower lobe (e.g., Fig. 44). The preputial membrane is almost entirely modified to form the Lad which is weakly separated from the sclerotized L2vm; a small part of the preputial membrane on the right side may not be sclerotized, and minute microtrichia can be seen in this region. The hook (R2) with (Figs. 240, 281, arrows) or without an apical notch; no other genera, here examined, have an apical notch: Perisphaerus (Figs. 227-259)) P~eudo~lomeris (Figs. 260-262)) and Trichoblatta (Figs. 263-289). Three of 16 known species of Trich~oblatta are African, the others, as well as all the species of Perisphaerus and Pseudoglomeris are Asian and adjoining islands.
The genera and species can be grouped according to the degree of development of the apical notch on R2 :
a. Apical notch absent : Pseudoglomeris glomeris (Figs. 260- 262)) Trichoblatta sericea (Figs. 264, 288-289). b. Apical notch poorly defined : Perisphaerus ficwipes (Figs. 257-259) (the shape of Lad of this species (Fig. 257) differs from all the others of the tribe), P. armadillo (Figs. 233-235, 248-25 I )) and P. glomerifornzis (Figs. 231-232, 254-256). c. Apical notch well developed: Perisphaerus contiguus (Figs. 227, 236-238)) P. flavicornis (Figs. 228, 239-241), P. semilunatus (Figs. 230, 242-244), P. aterrimus (Figs. 229, 245-247)) Tricho- blatta magnified (Figs. 268, 271-273)) T. oniscina (Figs. 267, 274- 276)) T. tarsalis (Figs. 264, 277-279)) T. nigra (Figs. 269, 280- 282)) T. guerini (Figs. 266, 283-285).




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Psy the [December
P. contiguus {Sauss.). Mt. Lamington, N.E. Papua (det Hebard). 228. (3F
CUZM). P. flavicornii Burm. Java. 229, (983 L). P. atfrrhti (Herbst). Kangean Island, Oengkokop (det. Princis). 230. P. semilunatus (Hanitsch). Sakaerat, Khorat Province, Thailand (dot. Princis). 231-232. (104 MCZ) . P. ghcriformis Lucas. Samar, Catarman, Philippine Islands (de~ Princis). 232. Tooth (arrow) (in underside of prorioturn. 233-235. P. armadillo Ser- ville. 233. (989 L). Prai jawang, East Sumba (det. Princis). 234-235. (109 MCZ). Zamboanga, Philippine Islands (det, Princia) . 335. Tooth (arrow) on underside of pronoturn. (scale: Figs. 232, 235 = 1 mm; all others== 5 mm).




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Figures 236-244. Male genital phalbmeres of Perisphatrus spp. 236- 238, (176 ANSP). P. co?ztiguus (from 5 shown in Fig. 227). 239-244. (39
CUZM). P. ftawinrnis (from 8 shown in Fig. 228). 242-244. P. semi- Iwiatus (from 8 shown in Fig. 230). (acale = 0,1 mm).



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342 Psyche [December
Figures 24.5-253. Male genital phallomeres of Pfrisfiiuerus spp. 245- 2+7. (983 L). R. derrimus (from 8 shown in Fig. 229). 248-251. P. arma- ditto, 248-249. (109 MCZ). (from 3 shown in Fig. 234). 250-251. (9B2 I,). (from 8 shown in Fig. 233). 252-253, (180 ANSP). P, aeneus (Brunner). Sibuyan Island (det. Hebard). (scale = 0.1 mm),



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344 Psyche [December
Figures 263-270. Adult malea of Trirhblafta spp. 263, (987 L). T. pyfmaea (Karny). Ryukyu (der. Princisj. 264. (985 L}. 7'. tarsalis (Walker). Kambaiti, S.E. Burma, 7000' (det. Princis). 265. (9% L). T. serlcea (Sauss.). Madras, India (det. Princia). 266. (8 BMNH). T. guerini Sauss, and Zehn. "Givalior" (det. Frincis), 267. (9S8 L). T. ontscina (Gerstaecker) . Urwald, Ukaika [det. Princis) . 268. ( 986 L) , T. magnifica (SheHord) . Tonkin, Chapa, Indo-China (det. Princis). 269-270. (103 MCZ). 269. T. nigm (Stnraki). Formosa (det. Princis). 270. Tooth (arrow) oa underside of pronoturn. (scale:
Pigs. 363-269 = 5 mm ; Fig. 270 = 1 mrn).



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Figuw 271-279. Male genital phalbmeres of Trichobtatta spp, 271-271. (986 L). T. magffiftca (from 8 shown in Fig. 268). 274-276. (988 L). T. k i n a (from 8 shown in Fig. 267). 377-279. (985 L), T. turfah (from 8 shown in fig. 264). (scale ==: 0.1 rnm).



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Figures 2B9-289. Male genital phailomerea of Trichblatta spp. 280-282. (103 MCZ), T. niyra (from 8 shown in Fig. 269). 285-285. (8 BMNH)& T. guerki (from 5 shown in Fig. 266), 286-2S7, (987 L). T. pygmum (from $ shown in Fig. 263).
288-289. (9S4 L). T. serka (from 5 shown in Fig. 265). (scale = 0.1 min). .




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19731 Roth - Blattaria 3 47
Some genera of Perisphaeriinae have a characteristic tooth (Figs. 232, 235, 270) on each side underneath the pronotum into which the apex of the tegmina fit. Princis ( 1960) and Beier ( 1961 ) used this structure as a characteristic of Derocalymminae and Derocalym- mini, respectively. The shape of the tooth may have value as a species character (cf. Figs. 232 and 235). Shelford ( I 908) summarized Saussure and Zehntner's ( I 895 ) analysis of the evolution of the form of the pronotum in the Peri- sphaeriinae. The complex type of the pronotum of genera such as Pilema and Cyrtotria appear to have evolved from a more simple form. Pilema thoracica has been found in family groups of a female and nymphs in round holes in the ground about 6 inches in depth. Shelford suggests that the shovel-like pronotum of Pilema is prob- ably used for digging and evolved as a result of a change to this habitat. Derocalymma spp. live under heavy stones and their flat- tened form would enable them to crawl through narrow crevices for security from predators. Some species of C~rtotria also have the pronotum adapted for digging, but others haven't; nothing is known of the habits of this genus. The Bantua pronotum is intermediate in structure between C~rtotria and Derocalymma, but nothing is known of the genus other than it has been collected under a log. Shelford suggests that the Derocalynzma form originated from a form like P i h evolving through a Bantua-like stage. Based on male genitalia, I I genera of Perisphaeriinae are placed in 4 tribes as follows :
I. Perisphaeriini : Perisphaeria, Blepharodera, Pilema, Hostilia, Bantua, and Cyrtotria.
2. Zuluiini : Zuluia.
3. Derocalymmini : Derocalymma.
4. Perisphaei-ini : Perisphaerus, P~eudo~lonzeris, and Trichoblatta. I
thank the following for the loan of museum specimens: Dr. David Rentz, Academy of Natural Sciences of Philadelphia; Dr. David Ragge, British Museum (Natural History) ; Dr. Karl Prin- cis, Zoological Institute, Lund, Sweden; Dr. M. W. R. de V. Graham, Hope Department Entomology, Oxford, England; Dr. Howard Evans, formerly at the Museum of Comparative Zoology, Harvard University; Dr. S. L. Tuxen, Zoological Museum, Copen-



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348 Psyche [December
hagen, Denmark.
I am grateful to Mr. Samuel H. Cohen for taking the photographs.
BEIER, M.
1961. H. G. Bronns Klassen und Ordnungen des Tierreichs. Arthro- poda, Insecta, Blattopteroidea, Blattodea. 5, Div. 111, Book 6. Part 4, pp. 587-848. Leipzig.
GURNEY, A. B. AND L. M. ROTH
1972. A generic review of the cockroaches of the subfamily Panchlori- nae (Dictyoptera, Blattaria, Blaberidae) . Ann. Entomol. Soc. Amer. 65 : 521-532.
MCKITTRICK, F. A.
1964. Evolutionary studies of cockroaches. Cornell Univ. Agric. Exp. Sta. Memoir 389, 197 pp.
PRINCIS, K.
1960. Zur systematik der Blattarien. Eos 36: 427-449. 1963. Blattariae. In South African Animal Life. Stockholm, pp. 1-318. REHN, J. A. G.
1922. Contributions to our knowledge of the Dermaptera and Ortho- ptera of the Transvaal and Natal. Part 1. Dermaptera and Blattidae. Ann. Transvaal Mus. 9: 1-99.
1932. African and Malagasy Blattidae (Orthoptera). Part 11. Proc. Acad. Nat. Sci. Philadelphia 84: 405-511. 1933. Dermaptera and Orthoptera of the De Schauensee South African Expedition, - Part 1. Proc. Acad. Nat. Sci. of Philadelphia 85: 39-66.
REHN, J. W. H.
1951. Classification of the Blattaria as indicated by their wings (Orthop- tera). Mem. Amer. Entomol. Soc. No. 14: 1-134. ROTH, L. M.
1970. The male genitalia of Blattaria. 111. Blaberidae: Zetoborinae. Psyche 77: 217-236.
1970a. The male genitalia of Blattaria. IV. Blaberidae : Blaber,inae. Psyche 77: 308-342.
1971. The male genitalia of Blattaria. VIII. Panchlora, Anchoblatta, Biolleya, Pelloblatta, and Achroblatta (Blaberidae : Panchlorinae). Psyche 78: 296-305.
1972. The male genitalia of Blattaria. IX. Blaberidae. Gyna spp. (Peri~~haeriinae), P/ioraspis, Thorax, and Phlebonotus (Epilam- prinae). Psyche 98 : 185-217.
1973. Paramuzoa (Nyctiborinae), a new cockroach genus previously confused with Parasphaeria (Epilamprinae) . Psyche 80 : 249-264. SAUSSURE, H. DE AND L. ZEHNTNER.
1895. Revision de la Tribu des Peris~haeriens. Insectes Orthoptkres de la famille d'es Blattides. Rev. suisse 2001. 3: 1-59. SHELFORD, R.
1908. Some new genera and species of Blattidae, with notes on the form of the pronotum in the subfamily Peris~haeriinae. Ann. and Mag. Hist. Ser. 8, 1 : 157-177.




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