The Male Genitalia of Blattaria. V. Epilampra spp. (Blaberidae: Epilamprinae)

"The genus Epilampra is one of those assemblages which have developed wit:hin the tropics of both hemispheres a vast number of species, often quite distinct, again closely related and difficult to distinguish. With a general type o,f coloration the fluctuations of which make definite and exact characterization difficult, if not at times virtually impossible, it combines, a uniformity o.f development in numerous o.ther features, that in general in the amily are sufficiently 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 o.f the most difficult, obscure and generally unsatisfactory to. study in the entire Blattidae." (Rehn and Hebard, I927, p. 2o9). Princis (1967) lists 6o species of Epilampra all of which are found only in the New World. At least five of these species [atriventris (Saussure), cribrosa (Burmeister), ferruginea (Brunner) proxima (Brunner), and verticalis (Burmeister)] have: males with tergal glands, and their genitalia are so distinctly different from the males which lack tergal glands that I (I97O) have placed them in the genus Poeciloderrhis Still. This study of about 3o of the remaining 55 species of Epilampra listed by Princis (I967) shows that the. male genitalia are useful not only for .specific determinations of many species, but they may also indicate species relationships.

data, 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 were depo.sited 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 Blatta.ria disagreed in their determinations, emphasizing the. difficulty in identifying species of E1)ilamlra from literature descriptio.ns. Unfortunately male type mate.rial 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.

RESULTS AND DISCUSSION
The phallomeres characteristic of Elilamlbra male genitalia are shown in Figure . Prepuce Usually distinctively shaped with a definite marginal outline and often covered by microtrichia (Fig. , P). Mexicana Group ); E. ; E. con ]. This group includes species in which most of L.2d 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 let:t side. is separated from and lies above the prepuce, and on the right side the L2d tapers and extends upward towa.rd the L2vm (Figs. 2,5,8,I.O,I,I2). In [allax the lateral extension of the right side of L2d is quite long (Figs. 14,17,2,o,21 ). In some con-[erta the L2d extends well beyond the left side o.f the prepuce. (Figs. 24,27,(29)(30)(31)(32)34) and the extension on the right may vary considerably in length. The prepuce of mexicana is deeply notched and is readily distinguished Cro.m the other members of the group. Psyche . Cockroach male genitalia. EpilamlSra fallax. 14-16. (51 ANSP). Sapucay, Paraguay (det. Roth). [17][18]. Santa Catarina, Brazil. (det. Albuquerque). (SI Subapical Incision). [19][20]. Santa Catarina, Brazil. (det. Albuquerque'). 21-23. (103 USNM). Rio Lujer, Buenos Aires, Argentina (det. Albuquerque). (in Fig. 21 Group,R2 (Figs. 3,6,I5,I8 22,25,28,33,37,40,42, 45) has a. subapical incision, and a setal brush (Figs. 4,7,9,I3,I6,19,23 26,35,38,43,46) occurs on LI. Princis 1958, p. 63) synonymized Elilamlra stigmosa Giglio-Tos with Elilamlra con[erta Walker. The specimens determined by , 3I) as con[erta are similar to a specimen, in the Philadelphia Academy collection, which was labeled (in pencil) E. stigmosa (Fig. 3o). However, the E. con[erta identified byHebard, Albuquerque, and  have entirely different genitalia from Princis' con[erta 3I,(36)(37)(38) It is apparent that (]urney is not convinced that stigmosa and con-[erta are the same species. However, or the present, I am tollowing Princis' conclusion.s.. It is highly probable that more than one species 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 con[erta', as indicated above, is. a female. T'he prepuce and L:d of con[erta specimens from Ecuador (Figs. 3:, 34) differ somewhat tro.m these structure.s in specimens rom Panama. and Costa Rica (Figs. :4,:7,:9, 3o, 3I) and from specimens rom Peru (Figs. 36,39 (Roth and Gurney, 1969). The Puerto Rican record reported here (Figs. 5o-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 locality but it is possible that the specimen bo.arded ship in a Central American port. Rehn (I9O2) 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 Burmeisteri Group. Heba.rd (1929, p.   but is separated from it by the absence of a. setal brush on L I. 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 following 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, wheeler'i, gundlachi, haitensis, hamiltoni, bromeliadarum, gatunae, [ugax) In this subgroup the area of L2d is relatively small in relation to, and covers o.nly a small anterior part of the prepuce. In burmeisteri ( Without the sexes of each species, we do not feel in a positio.n to characterize A udreia more definitely than has been done in the meager description given by Shelford." A few years later Rehn and Hebard (927,p. 204) commented urther on the genus d udreia stating that it "... is composed of a. small number of species described from the tropics and subtropics of both hemispheres, the majority, however, tropical American. The species much resemble, certain forms referred to the genus Epilampra, but all possess reduced or subquadrate tegmina in both sexes, these subquadrate or distally emarginate in most of the forms Until the. genus Epilampra as a whole is critically studied, and our knowledge of the extent to which brachypterism occurs in that assemblage is more complete, it is unwise to do, other than follow Shelford's use of the generic name d udreia. We can say, ho.wever, that the genus Calolampra, to which a number of the species now placed in the more recently described d udreia were originally referred, is well distinct from d udreia of Shelford. The West Indies possess two species which can logically be assigned to d udreia, one from higher mountains of eastern Cuba., the, other rom Blue Mountains of Jamaica. The possibility that these may be. members of an ancient relic fauna forces itself upon one, although the converse argument that tegminal reduction 'has been brought about by adjustment to. a peculiar and restricted montane environment cannot be igno.red." The male genitalia of Calolampra carinulata Saussure (Saussure) of Central America, the geno.type, as selected by Hebard, although it has a number of features of difference." However, the L2d and prepuce o1: hamiltoni (Fig. 66), bromeliadarum (Fig.   I69), gatunae (Fig. 72), fugax (Fig. I75), and exl)loratrix (Fig.  359) are so typical of E1)ilampra that I assign them to this genus. The R2 of fugax (Fig. I76) lacks a subapical incision; this incision is present in the other 4 species of "'dudreid" (Figs. I67, 7o, I73, 36o) but is much reduced in bromeliadarum (Fig. I7O), gatunae (Fig. I73), and exloloratrix (Fig. 36o) 3o2-3o7) are basically those of EI)ilampra and I have. placed it in a separate group (see below). The male of d udreia catharina Shel1:ord has tergal glands on tergites and 2. Its genitalia are basically similar to species of "'Eloilamra'" that possess tergal glands and I placed it in the. genus Poeciloderrhis (Roth I97.o).
For the present I suggest that the genus d udreia, as represented by carinulata, be retained until a more detailed study is. made o1: other morphological characters o1: this species.
According to Princis (I958, p. 62) Walker's E. olaca is a synonym o1: his E. substrigata. Princis (personal communication) examined the types of opaca and substrigata "... and could not find any noteworthy differences. Hebard had never seen the. types and he thought them to be two different species. I supposed that Hebard's records 1:tom French Guiana [ol)aca] could be correct, whereas his record of substrigata 1:rom Colombia evidently relates to another species. This was, however, a pure speculation of mine." The genitalia, o1: Hebard's substrigata from Colombia (Figs. 99-2oo) clearly differ from those of specimens he considered to. be ol)aca from French Guiana (Figs. I78-I8O). Princis' substrigata (Fig. 2o5) is similar to Hebard's substrigata specimens (Fig. I99) For the present I am considering ot)aca and substrig'ata to be distinct. In addition to the marked differences in L:d and prepuce these 2 species, the. hook (R2) of substrigata (Figs.. 197,200,2Ol, 2o4, 2o7) is usually distinctly wider than that of oloaca (Figs. 179,182,I85,188). Hebard (1926, p. 2OI) stated that E. opaca may prove to be a synonym of the Brazilian E. maculicollis (Serville), and the Ecuadorean E. stigmosa Giglio-Tos may fall in the same synonymy. Numerous distinct, though easily confused, species are known to. belong to this group and, until a better concept of the distribution and individual variation within these is formed, we believe it best to use the name opaca." According to Princis stigmosa is a synonym of E. conferta (see discussion under Mexicana Croup). Hebard (I9:I, p. I36) stated that substrigata is closely related to grisea "... though separable by numerous features." The. genitalia of substrigata relate it more. closely to columbiana (Figs.. :o8-:I9) and oaca , than to. grisea (Figs. 68-76).
The difficulty in identifying some of these species is shown by the fact that two, specimens (Figs. I84-I80, I87-I89) determined as closely related to "'berlandi'" (cf. Figs. IIS-II7) by Albuquerque and Gurney (I96:, p. :43) are similar to Hebard's ol)aca. Princis determined o.ne of these specimens (Figs. 84-I86.) as well as one from Surinam (Fig. I9 O) aS E. sagitta. (]urney examined these specimens and in the absence of a careful study of types and genitalia felt that external features suggested the occurrence of more than one species. There can be little doubt that the genitalia of the Type of E. sagitta  and what is here considered to, be. opaca are distinctly different. In sagitta L has a setal brush (Figs. 61, 64, 67) (dbdomennigrum (]ro.up) and the tip of the prepuce is directed more posteriorly (Figs. 59,'6:,65). In oloaca there is no setal brush on LI (Figs. 8o, I83, I86, I89) (Burmeisteri Group) and the tip of the prepuce, is directed laterally (Figs. 178 181 (Fig. 299) are very similar to. those of abdomennigrum (Figs. 5o,(53)(54)(55); the L of latifrons lacks a setal brush (Fig. 3Ol ), whereas these setae are present in abdomennigrum (Fig. 52).
The unique tarsal-clawlike shape of the L2d of basistriga (Figs. 22o,223,(226)(227)(228) distinguishes this species from any other E1)ilampra. Hebard I929, p. 369) believed that E. delicata (Fig. 227) (-basistriga) seemed to be near E. berlandi, and E. jorgenseni and apparently even more closely related to ztudr'eia catharina; this conclusion is not supported by genitalia. E. berlandi (Figs. II4-II7) and jorgenseni    (Princis,I967). This specimen is from the Fry Collection in BMNH and Hanitsch in reporting it commented that "No particulars are available concerning the specimens from the late Mr. Alexander Fry's collection. Some of the material seems Malayan, but the rest is almost certainly Neotropical." Fry lived at one time in Rio, de Janeiro and this specimen most probably came from there.
Subgroup C (azteca, crossea): The prepuce is usually poorly developed or indefinitely outlined; if the prepuce is clearly outlined, its surface, sculpturing is scalelike.
Psyche [D ecembe r The genitalia of E. azteca from different localities suggest that two. species and possibly three may be included in this taxon. In specimens from Mexico (Fig. 247), Costa Rica (Fig. 25o), and Panama , L2d is irregular in outline and lies above the prepuce; the prepuce is irregular in outline and some portions tend to blend into the. surrounding membrane. The L2d in specimens from Trinidad (Fig. 269), Surinam (Figs. 26o,263,265,267), Venezuela (Fig. 272),and Ecuador (Figs. 275,278), is knoblike in shape and appears to be an outgrowth o.f the prepuce.  Princis), and o.ne from Panama (Fig. 287) differs distinctly from the azteca just discussed; the prepuce is more clearly defined and its scalelike surface sculpturing also differs from the other azteca. This is probably an undescribed species and I am tentatively calling it El)ilampra sp. D. Gurney (personal communication) has commented on E. sp. D. and states "... they (Nos. 113, I5, 139) are very much like. azte'ca from Central America (Nos. 14o, 141, I43, I44) but differ in face markings. However, No. 131 from Ecuador is darker in general, has darker and larger face markings, and the. ventral surface o.f the abdomen is mostly blackish, unlike 15 et al. The. type of azteca is a male from Mexico. because., though Cuba and Mexico were both mentioned originally, Cuba was eliminated as type consideration by comments in the Biologia. We have a male from "Mexico" which in face markings is more like the Princis det. specimen than like I4o, et. al., so perhaps Princis is right. Consulting the type' o.f azteca should solve the. problem." According to, Albuquerque and (Burney I96:, p. 244), E. colorata is related to the "maculifrons" Sfftl group. The genitalia (Figs. 290-292) of the. Type of colorata are very similar to those of E. azteca fro.m Central America (e.g. Figs. 25o-252). Gurney (personal communication) re-examined the Type and stated that colorata is quite likely a synonym o.f azteca. The Type is smaller than many azteca but probably within the normal .size range.. With Gurney's concurrence I consider colorataa synonym of azteca.  (Figs. 3o4, 3o7). The battleaxe-shaped L2d is continuous with a sclerotized portion of the prepuce and is not separated from the prepuce by a thin clear membrane (as is usual in the ztbdom'ennigrum and Burmeisteri Groups). In the Mexicana Group, L2d is not a distinct sclerite lying above the prepuce but is a flat sclero.tization lying on the same plane as the prepuce.
There has been some question about the placement of Et)ilaml)ra heusseriana Saussure. According to. Hebard (I92I) this species "... has been assigned to the genera Calolaml)ra and ztudreia, due mainly to the fact that the type female had short truncate, tegmina. Though this is true for the female sex, the male. before us is seen to have fully developed organs of flight, and shows no. characters which would warrant its being placed other than in the genus Eloilaml)ra." Princis (1967) lists heusseriana under d udreia apparently following Albuquerque (I964). The male genitalia of heusseriana are typical of many other Et)ilaml)ra and tend to support Hebard's placement of the species.
8oddis  In this group which includes E. sodalis and po.ssibly several undetermined species, there is no. distinctive prepuce and L2d is a single more. or less clawlike sclerotization. In a recent paper I indicated that the male, of E. sodalis had tergal modifications, on segments 3 and 4 (Roth,I969a,p. 2o, (Princis,958,p. 6), and found that the. males lack tergal glands. I also examined Walker's Type 9 of sodalis and concur with Princis tha.t cinnamomea i.s a synonym of this species. The male genitalia of sodalis (Figs. 3o'8-328) are distinctive; Lad resembles a single, tarsal claw, and R2 is rather stout with a subapical "incision" represented by a fine. line which is best .seen in a lightly sclerotized specimen (Fig. 327', arrow). The genitalia. (Figs. 329-33) of the specimen previously reported by me as sodalis (Roth, ,I969a) (here designated as Epilampra sp. A) are only subtely different from those of sodalis; L2d (Fig. 329) and R2 (Fig. 33o) are slightly more slender in sp. A. However, there is a striking difference in the pronotal markings of these two species. In sodalis the microspots are, all small and more. or less the same size whereas in sp. A there are distinctly large, spots, interspersed among small ones. These two forms are probably sibling species. Species A is the only Epilaml)ra I knov in which male tergal syche    In E. yersiniana the prepuce, when flattened, extends obliquely to the right of L2d (Fig. 353). The hook (R2) lacks a subapical in-Genitalic differences indicate that E. oaca is a distinct species and not a synonym of E. substrigata.
The male genitalia of 5 species of d udreia (hamiltonl, bromeliadarum, exloratrix, gatunae, and ]ugax) are so characteristic of E15i-larnI)ra that I have transferred them to this genus. Epilampra colorata is synonymized with E. azteca.