(Heteroptera, Pentatomidae): III. Subfamily Asopinae D
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ISSN 0013–8738, Entomological Review, 2006, Vol. 86, No. 7, pp. 806–819. © Pleiades Publishing, Inc., 2006. Original Russian Text © D.A. Gapon, F.V. Konstantinov, 2006, published in Entomologicheskoe Obozrenie, 2006, Vol. 85, No. 3, pp. 491–507. On the Structure of the Aedeagus of Shield Bugs (Heteroptera, Pentatomidae): III. Subfamily Asopinae D. A. Gapon and F. V. Konstantinov St. Petersburg State University, St. Petersburg, 199034 Russia Received December 24, 2003 Abstract—This communication is the third one in a series of papers treated the morphological review of the aedeagi of Pentatomidae and its possible taxonomic significance. Fully inflated internal structures of the aedeagus of 18 species of Asopinae were studied. The examination of the Asopinae confirmed the monophyly of this taxon and revealed three distinct types of the aedeagal structure within the subfamily. These types differ in the structure of the theca, median penal plates, and ventral, ventrolateral, and apical conjuctival lobes. These characters can be used in phylogenetic analysis as well as in a tribal classification of asopines. DOI: 10.1134/S0013873806070050 The present communication is the third one in posterior ends, the male genitalia combining a well- a series of papers treated the morphological review of developed apical part of the theca with genital plates the aedeagi of shield bugs (Pentatomidae) and reveling (parandria) of the pygophore, and the predatory way of the characters significant for the supergeneric classifi- life. Many authors [Leston, 1953, 1954 (cited from cation of the group. The structure of the aedeagus of Gapud, 1981); McDonald, 1966; Gross, 1975] point to the family Pentatomidae and subfamilies Discocepha- the families Podopinae and Pentatominae as closely linae and Phyllocephalinae was described in the first related to the subfamily Asopinae. Gapud (1991), de- communication (Gapon and Konstantinov, 2005). The nying relationship between Asopinae and Podopinae, present communication deals with the subfamily As- treats the Eurydema-Murgantia group within Penta- dopinae. All the aedeagi are described in their natural tominae as a sister group of Asopinae. At present, the (stretched) state, obtained by the method of hydraulic supergeneric classification of the subfamily Asopinae is absent. Amyot and Serville (1843) and Schouteden stretching with subsequent drying under air pressure (1907) subdivided Asopinae into two tribes: Asopini (Gapon, 2001). Amyot and Serville, 1843 and Discocerini Schouteden, The following abbreviations are used for designat- 1907. The majority of species was included into the ing morphological structures in the figures: a. c, apical nominotypical tribe, and the genera Discocera and part of the conjunctiva; a. th, apical part of the theca; Stiretrus, into the tribe Discocerini. Later, however, b. th., basal part of the theca; d. s, seminal duct; the type species of the genus Asopus appeared to be- g. s, secondary gonopore; l. a, apical lobe of the con- long to the genus Discocera. Representatives of the junctive; l. v, ventral lobe of the conjunctive; l. vl, subfamily are distributed nearly in all the zoogeo- ventrolateral lobe of the conjunctive; pr. a, apical out- graphical regions. Recently, Thomas (1992, 1994) growths of median plates of the penis; p. tr, pons revised species of the subfamily Asopinae of the Old transversus; ram. d, distal branches of ventrolateral and New World, mentioning 187 species of 43 genera lobes; ram. l, lateral branches of ventrolateral lobes; and 110 species of 26 genera, respectively. The tr. pr, longitudinal filaments of median plates of the aedeagi of 18 species of this group are described be- penis; tub. v, ventral tubercles of theca; ves, vesica. In low. all the figures, the scale corresponds to 0.2 mm. Macrorrhaphis acuta Dallas, 1851 (Figs. 1, 2) The subfamily Asopinae Amyot et Serville, 1843 is Material. Madagascar, Nanghoa, Itasy, III.1930 a holophyletic group, characterized by the presence of (Olsufjev), 1 specimen; Abessinia, Sekuala—lac. Suai, a thickened labium, the base of which is closely situ- H. Dzhalla maki, Hanash, 3–11.V.1903 (Sedov), ated to the base of the labrum, the parallel anterior 1 specimen; Daressal. D. O. Afr. (collection of ends of the proboscal (bukkal) plates with adjoining I. Gudim), 1 specimen. 806 ON THE STRUCTURE OF THE AEDEAGUS ... III 807 Description. Phallobase about as long as wide. rotized filaments and oval apical outgrowths. Longitu- Dorsal processes absent. Ventral processes in form of dinal fibers running in parallel along entire length, wide and short rectangular plates with rounded angles, narrowed proximally and somewhat widened medially; strongly reflexed proximally, with outer margins pro- running along ventral surface of lobe, only slightly jecting slightly beyond margins of basal. Ventral con- curved at its sides. Distally, they turning into apical nectives ill-defined. Dorsal connectives about as long outgrowths at apex of ventral lobe. Apical outgrowths as basal plates, with small elongate conical sclerotized running nearly in parallel, with apices strongly curved mushroom-like bodies at ends. Suspensory apodemae toward base of conjunctive and with longitudinal axis about as thick and long as dorsal connectives. running nearly in parallel to those of ventral lobe; their outer walls membranous, continuing membranous Theca uniformly heavily sclerotized, subdivided lateral walls of ventral lobe. Sclerotized inner walls of into two unequal parts. Both parts divided by distinct apical outgrowths connected by concave sclerotized constriction; their longitudinal axes nearly perpendicu- bridge at base. Membranous part of ventral wall of lar to each other. Basal part of theca small, tubular, ventral lobe convex, not separated from wall of con- narrowed at base and slightly widened at apex. Its junctive, narrowing toward apex, turning into rather ventral wall forming pair of small membranous ventral high sclerotized carina fused at apex with pons trans- tubercles at base. Ejaculatory reservoir occupying versus. Sperm duct fine and weakly sclerotized, wid- nearly entire cavity of basal part of theca. Apical part ened and C-curved at apex. Vesica running from of theca large, widened, strongly convex, enveloping membranous wall near pons transversus between api- base of conjunctive laterally and dorsally. Dorsoapical cal lobes, appearing as short curved sclerotized tube margin of apical part of theca forming rather deep with secondary gonopore at apex. V-shaped emargination. Lateral margins of apical part of theca projecting to form widely rounded angles. Dorycoris pavoninus (Westwood, 1837) Wall of conjunctive running from wall of theca at level of base of dorsal emargination, making lateral Material. Ankober, Shoa, Abessinia, VIII.1899 apical angles of apical part of theca free. Dorsal wall (Luk’yanov), 1 specimen; Harrar, Abessinia or., of apical part of theca nonsclerotized along median 8.VI.1913 (Svertshkov, expedition of Gumilev), line, appearing as narrow membranous stripe. 1 specimen; Madagascar, Analamazotra, XI.1930 (Ol- sufjev), 1 specimen. Conjunctive large, about twice as long as theca, Description. Aedeagus similar to that of preceding with paired membranous ventrolateral lobes, unpaired species. Phallobase slightly widened dorsally. Dorsal basal lobe bearing sclerotized median plates of penis, connectives slightly shorter than basal plates, with and unpaired apical lobe. Ventrolateral lobes of con- small, fine, conical mushroom-like bodies at ends. junctive divided into two branches: proximal and dis- Ventral processes in form of small rectangular plates, tal. Proximal branch fine, running nearly perpendicu- strongly reflexed proximally, projecting slightly be- lar to axis of common base and slightly curved toward yond lateral margins of phallobase. apex of conjunctive at apex. Distal branch thicker, appearing to be continuation of common base of lobes, Dorsal emargination of apical margin of apical part directed toward, and nearly reaching base of theca. of theca rounded and moderately deep. Distal branches diverging beginning from base of Proximal branches of ventrolateral lobes of con- lobes, forming apical hooks, with fine sclerotized tips junctive straight, directed ventrolaterally and slightly directed toward each other. Apical lobe long, with toward apex of conjunctive. Distal branches slightly shallow median constriction and with apex divided by longer and finer, directed ventrolaterally and toward shallow emargination into two short lateral branches base of aedeagus. Fine sclerotized hook directed to- strongly widened laterally. Fine long digitiform mem- ward base of branch situated at apex of each distal branous outgrowth running downwards from ventral branch. Apical lobe with constriction at base, deeply surface of each branch. divided into two long and thick arcuate branches with Ventral lobe directed downwards and obliquely to- apices directed toward each other. Pair of membranous ward base of aedeagus; ventral lobe situated between digitiform outgrowths situated on ventral surface of ventrolateral lobes. Median plates of penis situated on apical lobe at base of its branches, similarly to that of sides of ventral lobe, consisting of longitudinal scle- Macrorrhaphis acuta. ENTOMOLOGICAL REVIEW Vol. 86 No. 7 2006 808 GAPON, KONSTANTINOV Figs. 1–8. (1, 2) Macrorhaphis acuta, aedeagus [(1) ventral, (2) lateral view]; (3–5) Friarius alluaudi (Schouteden), aedeagus [(3) ven- tral lobe, lateral view; (4)