Part 1: the Male Genitalia and Indo-Malayan Podoscirtini

Part 1: the Male Genitalia and Indo-Malayan Podoscirtini

© Zoological Institute, St.Petersburg, 2002 Taxonomy of Podoscirtinae (Orthoptera: Gryllidae ). Part 1: the male genitalia and Indo-Malayan Podosci1iini A.V. Gorochov Gorochov, A.V. 2002. Taxonomy ofPodoscirtinae (Orthoptera: Gryllidae). Part I: the male genitalia and Inda-Malayan Podoscirtini. ZoosystematicaRossica, 10(2), 2001: 303-350. The morphology of the male genitalia ofPodoscirtinae and the nomenclature of their struc­ tures are discussed. Six Inda-Malayan genera of Podoscirtini are characterized; 4 new genera, 2 new subgenera, 20 new species, and 5 new subspecies are described; several other species are revised (including the descriptions of some unknown females and new data on distribution). One generic name (Calyptotrypus Sauss.) and one specific name (Platydactylus helvolus Serv.) are considered nomina dubia. New synonymy (Zvenella nigrotibialisLiu, Yin & Wang, syn. n. = Z. geniculata; Madasumma obscuripennis Chop., syn. n. = Z. parcevenosa; Calyptotrypusjlavomarginatus Hsia & Liu, syn. n. = Valia pulclzra) is established. The lectotype for Truljalia lzofmanniis designated. A. V. Gorochov, Zoological Institute, Russian Academy ofSciences, Universitetskayanab. 1, St.Petersburg 199034, Russia. At present, thestudy of taxonomy and biology of Sonotrella collected by me in Sumatra (appar­ Podoscirtinae is in an initial stage. Only the first ently, hermaphroditism is caused by the parasi­ steps towards thetribal and generic classifications tosis, as this specimen was parasitized, possibly of this group are made. The key to Indo-Malayan by Strepsiptera). This specimen has almost nor­ genera of Podoscirtinae recently published by mal male genitalia (evidently, it is a genetic ma­ Ingrisch ( 1997) includes 19 genera, but their num­ le), small rudiments of ovipositor, and female ber is much greater in reality. In this very good tegminal venation. In connection with the above­ paper, Ingrischcorrectly returned the generaNocti­ mentioned reasonings, the Ingrisch's interpreta­ trellaGor., PhyllotrellaGor., SonotrellaGor., Zve­ tion of the tribeAphonoidini as the Indo-Malayan nellaGor., and TrelleoraGor. to the tribe Podoscir­ group without tegminal stridulatory apparatus in tini. He considered that this tribe is characterized male is too simplified (numerous convergences by the developed tegrninalstridulatory apparatu s in are possible). male, but the male of the monotypic type genus I described this tribe and some other tribes of PodoscirtusServ. fromMadagascar is lacking this Podoscirtinaeon the basis of the most important apparatus! changes in the modes offunction of male genita­ The disappearance of this apparatus in male is lia (Gorochov, 1986). I consider that it is verydif­ a very usual phenomenon in different branches ficult to understand the taxonomic significance of Podoscirtinae, and this process is probably cor­ of morphological structures without any func­ related with the feminizationof males (genetic tional interpretations. Therefore, ii is reasonable change in the hormonal balance or similar phe­ to dwell upon the questions of functional mor­ nomena). The reduction of stridulatory appara­ phology of the male genitalia in Podoscirtinae. tus in male as a result of feminization may pass very quickly and without important genetic Nomenclature of the male genital structures changes. For example, the male tegmina of dif­ ferent species of the Australian genus Riatina In the study of taxonomy of Grylloidea, I use Otte & Alex. may have a normal stridulatory the modifiednomenclature by English language apparatus, strongly reduced one (almost female­ authors (Randell, 1964; Alexander & Otte, 1967). like tegmina), and numerous intermediate vari­ The Randell's idea of functionally-basedtermi­ ants (e.g., Otte & Alexander, 1983: Fig. 258). The nology is more suitable in comparison with all other indirect evidence of this hypothesis is an others, as it allows one to use a few terms for anomalous hermaphrodite specimen of the genus numerous convergent structures of more or less 304 A. V. Gorochov: Taxonomy of Podoscirtinae. Part I • ZOOSYST. ROSSICA Vol. 10 similar origin. Presently, this nomenclature is 6) Ectoparameres: the paired sclerotized and most elaborated in Russian (Gorochov, 1995). distinctly articulated processes oflateral parts of Therefore,it is reasonable to give here the mod­ dorsal fold, epiphallus, or guiding rod (Figs I: 7- em variant of this nomenclature in English. 9) (these structures may be absent or more numer­ 1) Dorsaland ventralfolds ( or lobes): two main ous). The ectoparameres together with epiphallus membranous folds around the gonopore ( dorsal usually participate in grasping of the sclerotized foldabove gonopore and ventral one under it) in copulatory papilla of female (Figs I: 15, 16), or the most primitive (forrecent Ensifera) Hagloid in a special kind of anchor-like fixation in the type of genitalia (all recent Hagloidea, most of genital chamber of female (Figs I: 11, 13). Stenopelmatoidea, numerous Tettigonioidea, and 7) Endoparameres:the sclerotized paired partsof only Mogoplistini among Grylloidea). These lower membrane of the dorsal fold baseprovided folds are divided into differentlobes; all struc­ with special apodemes (Figs I: 7-9). Probably, these tures of spe1matophore are formedin the special apodemes are always homologous; usually, they are cavity between these folds (Figs I: 1, 2). very long, but sometimes slightly or strongly re­ 2) Epiphallus: the sclerotized part of dorsal duced. The endoparameres may be short or long, foldconsisting of one sclerite or several slightly connected with each other by a sclerotized ribbon movable sclerites (not separated fromeach other, (Figs I: 8-11) or isolated fromeach other (Figs I: 14, i.e. not articulated or almost not articulated). In 15). They ensure movable connection betweenthe the Grylloidtype of genitalia, the epiphallus oc­ above-mentioned apodemes ( and their muscles)and cupies the most part of dorsal fold; in the the guiding rod, ectoparameres,or some otherstru c­ Tettigonioid typeof genitalia, the epiphallus is tures. represented by small or rather large, paired or 8) Mold ofspermatoplwre attachment plate: unpaired sclerites on the upper surfaceof dorsal the special partly sclerotized structure (or com­ fold(hooks, plate with denticles, etc.); sometimes plex of sclerites) on lower membrane of the dor­ these sclerites are named titillators, but the geni­ sal foldbase. This structure is situated proximally tal terminology of Tettigonioidea is not elabo­ to the base of guiding rod; this mold participates rated up to now (Figs I: 3-9). in formationof the attachment plate ofspermato­ 3) Ampulla, neck, attachment plate, and tube: phore (Figs I: 8, 10, 14). the parts of spermatophore of Grylloidea. The 9) Rami: the paired elongated sclerites form­ large ampulla includes a capsule with sperm. All ing an interrupted or solid sclerotized ribbon other structures of spermatophore are situated around the base of ventral fold(rami may be ar­ around the long and narrow spermatophore ca­ ticulated or fusedwith epiphallus, but sometimes nal (this canal leads from capsule to apex of they are strongly reduced) (Figs I: 7-9). tube): the narrow neck, which may be reduced, 10) Spermatophore sac: the rounded or later­ the widened attachment plate with lobes for the ally compressed invagination of the basal part of adherence to female, and the thin tube usually guiding rod ( or an invagination of lower surface inserted into the femalespermathecal canal (Figs of the dorsal fold base between the guiding rod I: 10, 14). The ampulla may be covered with and the mold ofspermatophore attachment plate). spermatophylax (special solidified secret), pro­ The spermatophore sac participates in formation tecting the spermatophore from the premature of the long spermatophore tube and appears for destruction by female. lengthening of spermatophore tube without 4) Valvae: the ventral foldof the Grylloid type lengthening of guiding rod (Fig. I: 14). of genitalia (Figs I: 5, 6). This foldusually con­ sists of a pair of membranous lobes (Figs I: 8, 9); Characteristic featuresof the male genitalia in these lobes participate in formation of a sper­ Podoscirtinae matophore ampulla in the cavity between them only (Figs I: 14, 15), or between them and the Initially, the male genitalia of Podoscirtinae special membranous mold on upper surfaceof the were possibly adapted for the special kind of genital plate (Figs I: 10-13). anchor-like fixation in the genital chamber of 5) Guiding rod: the membranous or partly female (Figs I: 11-13), as such structure of the sclerotized process of lower membrane of the genitalia is presumably, inherited from the pos­ dorsal foldbase. This process participates in for­ sible ancestors belonging to the subfamily Penta­ mation of the spermatophore tube and helps to centrinae. Types of this fixation in recent forms insert it into the female spermathecal canal (if are rather diverse. guiding rod is changed into a thin and long (1) The guiding rod together with endo­ sclerite, it is named virga) (Figs I: 8-16). parameres may strongly deviate from the rest ZOOSYST. ROSSI CA Vol. 10 • A. V. Gorochov: Taxonomy ofPodoscirtina e. Part l 305 -- (�\-:.:\\Jll{.t>:::,,,:'ii"iC/.,'.';,,•.· , , �3, e'p / �a '"i{i{i?·· \ 5 d 1 d ,v , ep s vI I• \ , I s � I s � . 4 \�6, . � � 12 Figs I (1-16). Scheme of male genitalia. 1, 2, Hagloid type; 3, 4, Tettigonioid type; 5, 6, primitive Grylloid type; 7-16, derivative Grylloid type (Podoscirtinae) (7-13, Podoscirtini:

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