THE MALE GENITALIA OF BLATTARIA. III. BLABERIDAE: ZETOBORINAE.

In this paper I shall illustra.te and characterize the male genitalia of 9 genera of blaberids which belong in the subfamily Zetoborinae. These genera are Capucina Saussure, Lanxoblatta Hebard, Phortioeca Saussure, Phortioecoides Rehn, Schistopeltis Rehn, Schizopilia Burmeister, Tribonium Saussure, Zetobora Burmeister, and Zetoborella Hebard. Princis (I96O) includes Tribonoidea Shelford in this sub]amily but I have been unable to obtain a male of the one species in this genus. The male genitalia of most species of Blaberidae are characterized by the presence of three phallomeres: a median sclerite, a sclerite on the left, and a retractable hook on the right side. This led McKittrick (I964, p. 72) to conclude that "The male genitalia are extraordinarily uniform throughout the Blaberidae." She also stated (I964, p. 35) that "The male genitalia differ slightly in hook shape and the configuration of the posterior tip of the median sclerite within the Blaberidae, but otherwise show little variation within that group," and on p. IO "The great uniformity of blaberid male genitalia precludes drawing many conclusions regarding subfamily relationships, except in the case of the Blaberinae and Panesthiinae." "The great uniformity" of the male genitalia of Blaberidae is generally true only for the usual presence of three distinct phallomeres; in certain genera one or more of the phallomeres may be greatly reduced and even absent (e.g. R2 in Panchlora). However, the shapes of the phallomeres, especially the retractable hook (R2), prepuce, and L2d are often so distinctive that they may show subfamily, tribal, generic, species Group characteristics, and in some genera may be used for specific determination (Roth and Gurney, I969 Rcth, I969, I97O).

The male genitalia of most species of Blaberidae are characterized by the presence of three phallomeres: a median sclerite, a sclerite on the left, and a retractable hook on the right side. This led McKittrick (I964, p. 72) to conclude that "The male genitalia are extraordinarily uniform throughout the Blaberidae." She also stated (I964, p. 35) that "The male genitalia differ slightly in hook shape and the configuration of the posterior tip of the median sclerite within the Blaberidae, but otherwise show little variation within that group," and on p. IO "The great uniformity of blaberid male genitalia precludes drawing many conclusions regarding subfamily relationships, except in the case of the Blaberinae and Panesthiinae." "The great uniformity" of the male genitalia of Blaberidae is generally true only for the usual presence of three distinct phallomeres; in certain genera one or more of the phallomeres may be greatly reduced and even absent (e.g. R2 in Panchlora). However, the shapes of the phallomeres, especially the retractable hook (R2), prepuce, and L2d are often so distinctive that they may show subfamily, tribal, generic, species Group characteristics, and in some genera may be used for specific determination (Roth and Gurney, I969 Rcth, I969, I97O). /]'ATERIALS AND. METHODS The technique for preparing the genitalia has been described in previous papers (Roth,I969,I97O). The source of each of the specimens illustrated is given, using the ollowing abbreviations: (ANSP) Academy of Natural Sciences, Philadelphia; (BMNH) 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 the museums indicated.

RESULTS AND DISCUSSION
The basic features of the 3 phallomeres o the male genitalia o Zetoborinae are as follows: R2 (hooked right phallomere) is well developed and usually has a subapical incision (Fig. 32). The. preputial membrane has a heavily sclerotized area (L2d) which extends dorsally or dorsolaterally to the right of L2vm (Fig. 28); L2d is flattened in the preparations and it should be remembered that it usually extends dorsally. L is very well developed and various areas are darkly sclerotized (Fig. 3o).
Based on the degree of development of L2d and shape of L2vm, the Zetoborinae may be divided into 3 Tribes as follows: I. Zetoborini (Zetobora and Phortioecoides, Figs. 1-5, 22-39).
L2d is least developed in this tribe and is a thin sclerotization of the preputial membrane which extends dorsally (Figs. 22,25,27,28,31,34,37). The subapical incision of R2 is clearly defined (Figs. 23,29',32,35,38). LI is very similar in all the species in this tribe (Figs. 24,26,30,33,36,39). Hebard (I92I,p. 245) stated that "The differences shown by the forms ecognized by us as Tribonidium [--Zetobora] monasticum, transversum, and amplum, may indicate full specific distinction, geographic racial differentiation or, indeed, mere individual variation in a single exceptionally plastic species." The male genitalia of the above 3 species ( Capucina patula (Fig. IO) was placed in the Panchlorinae by McKittrick (I964) and in the Laxtinae by Princis (I96O). The Laxtinae of Princis is not recognized by McKittrick and she has assigned several of the genera in this grouping to other subfamilies. Princis (personal communcation) regards his subfamily Laxtinae "... as provisional. Further study will probably show that it is not at all a homogeneous group and must be sooner or later split up." McKittrick used proventricular and female genitalic characters in placing Capucina in the Panchlorinae. However many of the character differences she used in comparing ovipositors and proventriculi seem to be very subtle, and I believe the male genitalia offer characters which show more clear cut relationships in the Blaberidae than do the proventriculus or ovipositor. Miller (I969) found that the morphological diversity of the proventriculus in 9 subfamilies of Blaberidae "... often defied the most dedicated efforts to group them into subfamilies."' The male genitalia of Capucina are so similar basically to those of other members of the Phortioecini that I do not hesitate to assign this genus to the Zetoborinae. In Phortioeca peruana (Fig. 87) and P. phorasIoides (Fig. 98), the sclerotization of the preputial membrane, which forms the base of L:d, may be incomplete leaving a lacuna or window. However, it is clear that this character is variable (Figs. 9o,93,(96)(97)(98) and cannot be used for specific determination. In P. phoraspoides the rounded margin of the preputial membrane which surrounds the sclerotized base of L2d has more or less uniformly spaced indentations ( Fig. 93) which are absent in other .species of Phortioeca examined.
Based on the shape of R2, Phortioeca nimbata, maximiliani, al)olinari, and verrucosa can be gro.uped together. In these species the curved hook portion of R: is relatively narrow and its subapical incision is poorly developed or absent (Figs. 73,76,79,8:,85 (1932) stated that the genera Zetobora, Lanxoblatta, Zetobotella, and Schizopilia "... are clearly derivatives o.f a single phylum," whereas Schistopeltis and Tribonium "... typifies a distinct and clearly marked phylogenetic series." In general the male genitalia tend to support Rehn's hypothesis. However, I have separated Zetobora from the other 3 genera of his grouping because of the relatively poorly developed Lad.