On the Structure of the Aedeagus in Shield Bugs...

ISSN 0013–8738, Entomological Review, 2006, Vol. 86, No. 2, pp. 149–167. © Pleiades Publishing, Inc., 2006. Original Russian Text © D. A. Gapon, F. V. Konstantinov, 2006, published in Entomologicheskoe Obozrenie, 2006, Vol. 85, No. 1, pp. 13–34.

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On the Structure of the Aedeagus in Shield Bugs (Heteroptera, Pentatomidae): II. Subfamily Podopinae

D. A. Gapon and F. V. Konstantinov

St. Petersburg State University, St. Petersburg, 199034 Russia

Received April 15, 2004

Abstract—This paper is the second communication in a series of papers that treats the structure of the aedeagus in the family Pentatomidae and its taxonomic applications. Inner structures of the aedeagus were examined in the completely inflated state. The structure of the aedeagus was studied on 25 species. It is impossible to offer a diag-nosis of the subfamily Podopinae on the basis of the structure of the aedeagus only, because this structure is very diverse. Nevertheless, some distinguishing characters were found in each of the three tribes, currently recognized within the subfamily. On the basis of the structure of the aedeagus, the tribe Graphosomatini can be subdivided into four groups, partly corresponding to the previously recognized tribes Graphosomatini (sensu Puchkov, 1965), Ancyrosomatini, and Trigonosomatini. Three different types of the structure of the aedeagus were revealed in the tribe Podopini. Special types of the structure of the aedeagus were found in the genera Bolbocoris and Testrica with unclear taxonomic position.

DOI: 10.1134/S0013873806020047

The present communication is the second one in a series of papers treated the morphological review of the aedeagi of shield bugs (Pentatomidae) and reveling characters significant for the supergeneric classifica-tion of the group. The structure of the aedeagus in the family Pentatomidae and in the subfamilies Disco-cephalinae and Phyllocephalinae was described in the first communication. This paper deals with the struc-ture of the aedeagus in shield bugs of the subfamily Podopinae. All the aedeagi are described in their natu-ral (stretched) state, obtained by the method of hydrau-lic stretching with subsequent drying under air pres-sure (Gapon, 2001).

The following abbreviations are used for designat-ing morphological structures in the figures: a. c, apical part of the conjunctiva; a. s, suspensory apodema; a. th, apical part of theca; b. th., basal part of theca; c. f., mushroom-like bodies; con. d, dorsal connectives; con. v, ventral connectives; d. end, endophallic duct; d. s, seminal duct; g. s, secondary gonopore; l. b, basal lobe of conjunctive; l. d, dorsal lobe of conjunctive; l. v, ventral lobe of conjunctive; l. vl, ventro-lateral lobe of conjunctive; lam. m, median parts of penis; lam. b, basal plates of phallobase; pr. a, apical out-growths of median plates of penis; pr. d, dorsal proc-esses of conjunctive; pr. v, ventral processes of phallo-base; r. ej, ejaculatory sack; th, theca; tr. pr, longitudi-nal filaments of median plates of penis; tub. v, ventral

tubercles of theca; val. b, basal carina of conjunctive; ves, vesica. In all the figures, the scale corresponds to 0.2 mm. The material from the basic collection of the Zoological Institute RAS ZIN, and also from the Mu-seum of the Entomology Department Rostov State University, marked by an asterisk in the text, were used in the present work.

Subfamily PODOPINAE Amyot et Serville, 1843

The subfamily Podopinae is a widespread taxon comprising 255 species of 64 genera (Schuh and Sla-ter, 1995). Representatives of this group are character-ized by the following morphological characters: presence of visible antennal tubercles; 4- or 5-segmen-ted antennae; lateral angles of pronotum looking like a tooth or a tubercle; 3-segmented tarsi; long scutellum reaching membrane or even apex of abdomen and concealing most part of elytra; genital segment bearing processes at posterior-lateral margin. Different authors treat Podopinae as a monophyletic taxon (McDonald, 1966; Davidova-Vilimova and McPherson, 1995) or as a polyphyletic one (Leston, 1953; Gross, 1975; Gapud, 1991). At the same time, the majority of authors point to the fact that Podopinae are closely related to Penta-tominae and Asopinae. Different authors distinguish from two to five groups within this group. According to the modern classification proposed by Davidova-Vilimova and Štys (1994), the family Podopinae in-

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cludes the tribes Podopini Stål, 1872, Graphosomatini Mulsant et Rey, 1865, Tarisini Stål, 1872, and two more yet undescribed groups. Some genera and the tribe Procleticini Pennington, 1920 [taxonomic posi-tion of this tribe in the family Pentatomidae and its relation to Podopinae are discussed by Gapon (2005)] were excluded by these authors from the subfamily Podopinae. Aedeagi of representatives of 25 species of this family are given below.

Tribe GRAPHOSOMATINI Mulsant et Rey, 1865

Derula longipennis Oshanin, 1871 (Figs. 5, 6)

Material. Ak-Terek area, 5 km N Gava, Ferghana Mt. Range, 28.VII.1937 (Kiritshenko), 1 specimen; Yavan-Su River, nearly Porchi Sai, Tajikistan, 15.V. 1943 (Kiritshenko), 1 specimen; Kondara, Kvak, Var-zob River valley, Tajikistan, 5.VI.1943 (Kiritshenko), 1 specimen.

The phallobase is transverse from the ventral side, slightly dilating dorsally. Dorsal connectives are very short; they terminate as short and wide sclerotized mushroom-shaped conical processes. Ventral proc-esses look like short triangular diverged plates directed somewhat proximally. Ventral connectives are weakly developed. Suspensory apodemes are thick and slightly shorter than the basal plates.

The theca is subdivided into two parts. The basal part of the theca is large, more convex dorsally, bear-

ing developed membranous ventral tubercles at the base. Ejaculatory reservoir is large, occupying nearly the entire cavity of the theca. The apical part of the theca is significantly shorter and narrower than its basal part and weakly separated from the latter. Lateral apical angles of the apical part of the theca are nar-rowly rounded; dorsal emargination of its apical mar-gin is V-shaped, narrow, and deep. The endophallic duct is wide, elastic, weakly sclerotized, and Z-shaped. Departing from the semen reservoir, this duct runs to the apex of the theca and then strongly bends back-wards and again turns to the apex of the theca in its distal part. The endophallic duct gradually narrows toward the place of its fusing with the wall of the con-junctive.

The conjunctive bears a membranous paired ventro-lateral and unpaired ventral lobes. Ventro-lateral lobes are swollen, with two low rounded apices. One of these apices is directed toward the apex of the con-junctive, nearly contacting a similar apex of another lobe; the other apex is directed toward the base of the aedeagus. The apical part of the conjunctive looks like a low swollen and flattened cupola.

The ventral lobe is narrow at the base; it is situated between ventro-lateral lobes and directed ventrally. Sclerotized median plates of the penis, situated on the ventral lobe, consist of very short, narrow, and strongly converging longitudinal filaments and apical outgrowths, diverging apically. Apical outgrowths are

Figs. 1–4. Podopinae. Phallobase in distal view: (1) Sternodontus ampliatus Jakovlev; (2) Graphosoma lineatum (Linnaeus); (3) Vento-coris fischeri (Herrich-Schäffer); (4) Scotinophara lurida (Burmeister).

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long and narrow; they are sclerotized at both sides and flattened laterally; their pointed and bent apices are directed toward the base of the aedeagus. From the dorsal side, median plates of the penis are connected by a membranous wall approximately as far as their

middle. The vesica, looking like a very fine and long (approximately 0.4 times as long as theca) sclerotized tube, directed ventrally-apically, departs from a mem-branous wall at the apex of the ventral lobe. The se-condary gonopore is situated at the apex of the vesica.

Figs. 5–13. Podopinae. Aedeagus in (5, 7, 9, 12) ventral, (6, 8, 10, 13) lateral, and (11) dorsal view. (5, 6) Derula longipennis Oshanin; (7, 8) D. flavoguttata Mulsant et Rey; (9) Ancyrosoma leucogrammes (Gmelin); (10, 11) Sternodontus ampliatus Jakovlev; (12, 13) Oplistochilus pallidus Jakovlev.

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In resting state, the vesica and the conjunctive fill the cavity of the theca; bending of the endophallic duct facilitates this filling.

Derula flavoguttata Mulsant et Rey, 1856 (Figs. 7, 8)

Material. Kerch, Crimea, 20.VII.1918 (Kiri-tshenko), 2 specimens; Tarsi Vill. near Makhachkala, Daghestan, 1.VI.1941 (Ryabov), 1 specimen.

The phallobase and theca are structurally similar to those of Derula longipennis. The phallobase is slightly wider than long; basal plates run in parallel. Dorsal connectives are slightly longer and mushroom-like bodies are smaller than in the preceding species. Di-verging ventral processes are triangular and short.

The apical part of the theca is indistinctly separated from its basal part. It is short, with smoothed lateral apical angles and a narrow and shallow dorsal emargi-nation.

Ventro-lateral lobes of the conjunctive are single-pointed and rather narrow; they are flattened laterally and directed ventrally. Their outer walls are weakly sclerotized; the inner walls are membranous with api-ces directed toward the base of the aedeagus. When the conjunctive is filled with a liquid causing erection, ventro-lateral lobes weakly bloat. The apical part of the conjunctive looks like a low cupola with a convex dorsal wall at the base.

The ventral lobe is long (only slightly shorter than ventro-lateral lobes) and directed ventrally. It bloats during impregnation by the erecting fluid. Median plates of the penis are rudimentary, they are not subdi-vided into longitudinal fibers and apical outgrowths. They look like weakly sclerotized narrow strips run-ning along lateral margins of the ventral lobe, being somewhat raised ventrally and forming a grove in the distal half. Apically, the ventral lobe gradually nar-rows, being divided into two short tongue-shaped out-growths (apparently, homologous to the apical proc-esses of median plates of penis). In the proximal half of the dorsal part, lateral parts of the ventral lobe fuse with the inner walls of ventro-lateral lobes. The vesica similar to that in D. longipennis departs from the ven-tral wall of the ventral lobe in its basal part.

Ancyrosoma leucogrammes (Gmelin, 1790) (Fig. 9)

Material. Shurabdara near Chubek, E Bukhara, 5.VII.1910 (Zarudnij), 3 specimens; Yangi-Bazar at Kafirnigan, Bukhara, 12.VII.1931 (Gussakovski), 1 specimen.

The phallobase is slightly wider than long and rounded ventrally. Free ends of the basal plates (i.e., their parts situated dorsally to place of fusing of the basal plates and connected with a bridge proximally) are rather short, somewhat diverging dorsally. Dorsal connectives are very short, terminating as small coni-cal mushroom-like bodies. Ventral processes look like diverging narrow triangular plates. The suspensory apodema is slightly shorter than basal plates.

The theca is subdivided into two disproportional parts. The basal part of the theca is large and wide, bearing a pair of oblong ventral tubercles at the base. The apical part of the theca is indistinct and weakly separated from its basal part, looking like small rounded lateral plates widely separated by a dorsal emargination of the apical margin of the theca. The ejaculatory reservoir is large, occupying nearly the entire cavity of the theca excluding its apical part.

Ventro-lateral lobes of the conjunctive are single-pointed, wide basally and narrowing apically. They run ventro-laterally and their apices are slightly bent toward the base of the aedeagus. The apical part of the conjunctive looks like a low and widely rounded cu-pola.

The ventral lobe is well-developed and directed ventrally. Diverging (especially basally) longitudinal fibers of the median plates of the penis are short and narrow. Rather long apical outgrowths of the median plates are strongly dilated in the median plane. Apical outgrowths are diverged ventrally at the base; more distally, they are pointed toward each other, adjoining by margins along a small distance. They diverge api-cally, and their margins are again strongly converging at the dorsal side, without touching. The vesica goes in ventro-apical direction from the membranous wall of the conjunctive between apical outgrowths. It looks like a short and slightly bent sclerotized tube lying ventrally to the place of adjoining of the apical out-growths.

Sternodontus ampliatus Jakovlev, 1887 (Figs. 1, 10, 13)

Material. Right bank of Iskander Darya River near source, 5.VIII.1947, 9.VIII.1947 (Kiritshenko), 2 spe-cimens; Khozor-Mech River near Lake Iskander Kul, 15.VIII.1947 (Kiritshenko), 1 specimen.

The aedeagus is structurally similar to that of Ancy-rosoma leucogrammes. The phallobase (Fig. 1) is slightly wider than long, rounded ventrally. Free ends



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of basal plates are short. Dorsal conjunctives are short, mushroom-like bodies are conical, oblong, and nar-row. In comparison with the preceding species, ventral processes are situated somewhat closer to each other.

The basal part of the theca is large, bearing a pair of developed ventral tubercles. The apical part of the theca is indistinct and weakly separated from its basal part, looking like narrow lateral plates, divided by a wide dorsal emargination of the apical margin of the theca.

Ventro-lateral lobes of the conjunctive are directed ventrally, adjoining median plates of the penis. Ven-tro-lateral lobes and penis are of the same length. Bases and outer walls of ventro-lateral plates are swol-len; the inner walls are flattened. The apical part of the conjunctive is rounded.

The ventral lobe is longer than that of the preceding species. Rather long longitudinal fibers of the median plates of the penis run in parallel. Bases of the longi-tudinal fibers laterally continue as narrow sclerotized fibers passing between bases of the apical part of the conjunctive and ventro-lateral lobes and nearly reach-ing apical margin of the theca. Apical outgrowths are pointed toward each other apically, adjoining by mar-gins along a short distance. Further on, their margins diverge, being again strongly converged dorsally.

Opistochilus pallidus Jakovlev, 1887 (Figs. 12, 13)

Material. Stalinabad, Tajikistan, 9.VI.1943 (Nikol-skaya), 1 specimen; Kondara, Kvak, Varzob River valley, Tajikistan, 5.VI.1943 (Kiritshenko), 1 speci-men; lower Yavan Su River, Kuibyshevsk, Tajikistan, 28.VII.1943 (Kiritshenko), 1 specimen.

The aedeagus is structurally similar to that of the two preceding species. The phallobase is slightly wider than long, rounded ventrally. Dorsal conjunc-tives are short, mushroom-like bodies are small. Ven-tral processes are triangular and narrow.

The basal part of the theca bears a pair of developed ventral tubercles at the base. The apical part of the theca is slightly wider than in preceding species, but is also weakly separated from its basal part. It looks like a ring that is wider laterally. The ring possesses nar-row and shallow dorsal emargination of the apical margin.

Ventro-lateral lobes of the conjunctive are dilated at the base and narrowed apically; they are directed ven-trally, running nearly in parallel. The apex of the con-

junctive is slightly higher than that of Ancyrosoma leucogrammes; its dorsal wall is less convex at the base.

Rather long and wide longitudinal fibers of the me-dian plates of the penis are weakly sclerotized, running in parallel. Apical outgrowths are stronger sclerotized along margins and apically. Apically, their margins are turned toward each other and adjoined; then they di-verge, being again strongly converged dorsally. The vesica is several times as long as that of the two pre-ceding species, it is slightly S-shaped and directed ventro-apically. Margins of the vesica are diverging apically; in this area, the vesica looks like a groove.

Graphosoma lineatum (Linnaeus, 1758) (Figs. 2, 14, 15)

The phallobase (Fig. 2) is slightly wider than long and rounded ventrally. Free ends of the basal plates are rather short, running in parallel. Dorsal connec-tives are slightly shorter than free ends of the basal plates, terminating as short and wide mushroom-like bodies. Ventral processes look like narrow short trian-gular plates. The suspensory apodema is slightly shorter than basal plates.

The basal part of the theca is oblong, possessing thick walls; apically, it is very weakly dilated and slightly bent ventrally. Ventral tubercles at the base are developed. The ejaculatory reservoir is large, oc-cupying nearly the entire cavity of the theca excluding its apical part. The apical part of the theca is very weakly separated as a narrow ring without distinct dorsal emargination of its apical margin.

The conjunctive is small and rounded. Ventro-lateral and ventral lobes of the conjunctive are absent. Lateral walls of the conjunctive are slightly swollen; they are bordered by the median plates of the penis apically. In non-swollen conjunctive, taken from the cavity of the theca, these areas look like small folds; they can be treated as rudiments of the ventro-lateral lobes. The apical part of the conjunctive looks like a low cone, shallowly bifurcating at the apex.

Median plates of the penis are large, arcuate, and narrow at the base; they dilate distally and their apices are directed toward the base of the conjunctive. Their bases are situated on the lateral walls of the conjunc-tive; they can be treated as the homologues of the lon-gitudinal fibers of some species described above. Free ventral margins of the median plates are roundly bent toward each other, but they do not adjoin. They can be

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homologized to the apical outgrowths of some species described above. Margins of the median plates, di-rected to the base of the conjunctive, are connected with its projecting membranous wall. The vesica goes

in ventral direction from the membranous wall of the conjunctive between the median plates. It looks like a very short and weakly sclerotized tube, bearing the secondary gonopore at the end.

Figs. 14–23. Podopinae. Aedeagus in (14, 16, 20, 22) ventral, (15, 17, 19, 23) lateral, and (18, 21) dorsal view. (14, 15) Graphosomalineatum (Linnaeus); (16, 17) Tholagmus flavolineatus (Fabricius); (18, 19) Vilpianus galii (Wolff); (20, 21) Ventocoris fischeri (Her-rich-Schäffer); (22, 23) Crypsinus angustatus (Bärensprung).



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Tholagmus flavolineatus (Fabricius, 1798) (Figs. 16, 17)

Material. Upper Dneprovka River, left bank of Ural River upstream of Orenburg, 23.VI.1932 (L. Zimin), 1 specimen; Ershov, Saratov Prov., 15–16.VI.1939 (Lukyanovich), 1 specimen; Rostov Province, environs of Tsimlyansk, Kuchugury, 10.VIII.1999 (Yu.G. Ar-zanov), 1 specimen*.

The aedeagus is structurally similar to that of Gra-phosoma lineatum. The phallobase is slightly wider than long and rounded ventrally. Free ends of the basal plates are short. Dorsal conjunctives are shorter than free ends of the basal plates. Mushroom-like bodies are short and wide. Ventral processes are short and triangular. The suspensory apodema is approximately as long as basal plates.

The basal part of the theca is oblong; it narrows apically and is slightly convex dorsally, bearing rather large ventral tubercles at the base. The apical part of the theca is indistinctly separated from its basal part, looking like a ring, which is narrow ventrally and somewhat wider dorsally. The dorsal emargination of the apical margin of the theca is shallow and narrow.

The conjunctive is small, swollen, spherically rounded, and slightly elongate dorso-ventrally.

Median plates of the penis are rather long and wide. Their narrowed bases are situated on the lateral walls of the conjunctive. Free apices of the median plates are strongly dilated, possessing margins widely bent toward each other; these margins adjoin apically. Dor-sally, their margins are strongly converged but do not adjoin. Abovementioned proposals on the homologies between parts of the median plates in G. lineatum can also be applied in this case. The vesica looks like a short but rather wide tube.

Vilpianus galii (Wolff, 1802) (Figs. 18, 19)

Material. Veliko-Anadolskaya dacha, 7.V.1905, 1 specimen*; Verkhnyaya Dneprovka, left bank of Ural River upstream of Orenburg, 28.VIII.1934 (L. Zimin), 1 specimen; Yanvartsevo, right bank of Ural River, Kazakhstan, 4.IX.1949 (Kiritshenko), 1 specimen; Rostov Prov., Sholokhov Distr., Elan-skaya stanitsa, 25.VI.1999 (E.A. Khachikoiv), 1 speci-men*; Rostov Prov., Oktyabrskii Distr., Persianovskii Vill., 8.VI.2000 (D.A. Gapon), 1 specimen*.

The phallobase is approximately square and rounded ventrally. Free ends of the basal plates are

rather wide. Dorsal connectives are very short; they terminate as short discoid sclerotized mushroom-like bodies. Ventral processes look like rather long triangu-lar plates. Suspensory apodemes are slightly longer than the basal plates.

The theca is rather large, not subdivided into parts; it is narrowed at the base and dilated apically, bearing small rounded ventral tubercles. The ejaculatory reser-voir occupies nearly the entire cavity of the theca ex-cept for its apical part.

The conjunctive is small. Its base is distinctly sepa-rated as a basal carina, especially wide and convex from the dorsal side and laterally. Ventro-lateral lobes are rather short, widely widened basally and narrowed apically; they run in parallel. They are directed ven-trally and their apices are slightly bent toward the base of the aedeagus. Laterally, the base of each lobe forms a small and weakly convex tubercle. The apical part of the conjunctive is developed very weakly, looking like a flat area at the level of the bases of the median plates of the penis, smoothly passing into the basal carina dorsally. Bases of the sclerotized dorsal processes are situated on the wall of the conjunctive laterally to its apex. These processes run in parallel; their free fine ends are directed dorsally and strongly bent toward the base of the aedeagus, reaching (but not touching) the walls of the theca.

Median plates of the penis mainly look like widely diverging longitudinal fibers, situated laterally to the apex of the conjunctive and strongly diverging in ven-tral direction. Distally, they look like small plates situ-ated on the inner walls of the ventro-lateral lobes in the median plane. Distal lamellar parts of the median plates can be homologized to apical outgrowths of the median plates of some other species; outer walls of these plates have become a part of the ventro-lateral lobes. Apical margins of the distal lamellar parts of the median plates are turned up toward each other. More distally, median plates look like fine arcuate fibers directed toward the base of the conjunctive. The wall of the conjunctive between the median plates is very weakly sclerotized ventrally. The vesica, looking like a small sclerotized cone with apical gonopore, departs from the wall of the conjunctive.

Ventocoris (Selenodera) fischeri (Herrich-Schäffer, 1851) (Figs. 3, 20, 21)

Material. Changyr near Khatyrchi, NW Bukhara, 13.VI.1930 (L. Zimin), 1 specimen; Ankara, Asia Mi-nor, 16.XII.1933 (Zhenzhurist), 1 specimen.

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The aedeagus is structurally similar to that of Vilpi-anus galii. The phallobase (Fig. 3) is strong, wider than long, and rounded ventrally. Free ends of the basal plates are rather long. Dorsal conjunctives are slightly shorter than the basal plates, terminating as wide discoid mushroom-like bodies. Ventral processes look like triangular plates, approximately equal in length to free ends of the basal plates; they are slightly bent proximally. Ventral conjunctives are rather strong. The suspensory apodema is approximately as long as the basal plates.

The theca is not subdivided into parts; it is massive, long, narrowed at the base and slightly dilated api-cally. It is somewhat C-shaped bent ventrally, possess-ing a narrow mouth. Ventral tubercles are well-developed. The ejaculatory reservoir is large, occupy-ing nearly the entire cavity of the theca.

The conjunctive is large. Its main part is represented by strongly swollen and rounded ventro-lateral lobes, adjoining apically and strongly projecting above the wall of the conjunctive. Each lobe forms small median and lateral tubercles, directed toward the base of the aedeagus, on the ventral side closer to the base of the conjunctive. The apical part of the conjunctive is weakly convex, bearing a small unpaired tubercle situ-ated dorsally to the base of the conjunctive. Dorsal processes are somewhat stronger than in Vilpianus galii.

Longitudinal fibers of the median plates of the penis are rather long and widely diverged. They are com-pletely concealed by overlapping ventro-lateral lobes. Distally, apical margins of the median plates, situated on the walls of the ventro-lateral lobes, are very widely bent toward each other, nearly adjoining. Api-cal ends of the median plates look like fine fibers bent toward the base of the conjunctive. The vesica looks like a rather long tube directed ventro-apically. It be-comes finer immediately after the base and is slightly dilated at the apex.

Ventocoris (Selenodera) oschanini (Horvath, 1889)

Material. Persia sept.-or., Shachrud, 14.V.1914, 31.V.1914 (Kiritshenko), 2 specimens; Ak Tau Mts. near Tamda, Kyzyl Kum Desert, 29.VI.1966, 1 speci-men (I. Kerzhner).

The aedeagus is structurally similar to that of Ven-tocoris fischeri. Free ends of the basal plates of the phallobase are slightly shorter than those of the pre-ceding species. Ventral processes look like small tri-

angular projections. Ventral conjunctives are less dis-tinct. The theca is strongly sclerotized. Ventro-lateral lobes of the conjunctive are less swollen, not adjoining apically. The apical part of the conjunctive is less con-vex, without tubercle at the base of the conjunctive. Median plates of the penis are less bent toward each other.

Ventocoris (Ventocoris) trigonus (Krynicki, 1871)

Material. Rostov Prov., Nedvigovka, 10.VII.1975 (Dotdtseva, Kurtser), 1 specimen*; Rostov Prov., W Nedvigovka, 26.VI.2003 (Gavritskov, Mirzoyan), 1 specimen*.

The aedeagus is structurally similar to that of the three abovementioned species. Free ends of the basal plates of the phallobase are rather long (as in V. fischeri). Ventral processes look like long and nar-row triangular plates. Ventral conjunctives are dis-tinct. The theca is strongly sclerotized. Ventro-lateral lobes of the conjunctive do not adjoin apically, their lateral tubercles are oblong and directed laterally; ven-tral tubercles are small and directed ventrally. The apical part of the conjunctive is slightly convex, with a small unpaired tubercle, situated ventrally at the base of the conjunctive. Dorsal processes are absent. Me-dian plates of the penis are nearly parallel at the base and are widely bent toward each other in the distal half.

Crypsinus angustatus (Bärensprung, 1859) (Figs. 22–24)

Material. Odessa, Khadzhibeiskii firth, salt marsh, 6.VII.1922 (Aleksei Kiritshenko), 1 specimen; Odessa, Khadzhibeiskii-Kuyalnitskii firths, salt marsh, 17.VI. 1923 (Aleksei Kiritshenko), 1 specimen; Elanets, NE Voznesenks, 10.VII.1923 (Aleksei Kiritshenko), 1 specimen; Privolnoe, upper Ingul River, 12.VII.1923 (Aleksei Kiritshenko), 1 specimen; Rostov Prov., Donskoi Chulek dean, Armenian pond, 8.V.2000 (D.A. Gapon), 1 specimen*.

The aedeagus is structurally similar to that of Vilpi-anus and Ventocoris. The phallobase is approximately square and rounded ventrally. Free ends of the basal plates are long. Dorsal conjunctives are very short, terminating as small discoid mushroom-like bodies. Ventral processes are very short and wide, with rounded angles; they are bent proximally. The suspensory apodema is slightly shorter than the phallo-base.



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Figs. 24–31. Podopinae. Aedeagus. (24) Crypsinus angustatus (Bärensprung), phallobase and theca in ventral view; (25) Scotinopharalurida (Burmeister), lateral view; (26, 27) Gambiana aspera (Walker): (26) lateral view, (27) parasagittal section through the conjunc-tive); (28, 29) Amaurochorus cinctipes (Say) in (28) ventral and (29) lateral view; (30, 31) Podops incertus Horvath in (30) ventral and (31) lateral view.

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The theca is cylindrical, not subdivided into parts. It is very slightly narrowed apically, possessing devel-oped ventral tubercles at the base.

The conjunctive is rather large. Its main part is rep-resented by strongly swollen paired ventro-lateral lobes, adjoining along the median line. Each lobe forms a pair of small median tubercles directed toward each other and towards the base of the conjunctive and a pair of lateral tubercles. The basal carina of the con-junctive is large and convex, overlaps the apex of the theca. Dorsal processes are long and strong, with rounded apices directed toward the base of the aedeagus. Their bases are situated at a long distance from each other, resting on the bases of the ventro-lateral lobes.

Median plates of the penis are strongly reduced, looking like very small triangular plates situated on the ventral side of the walls of the ventro-lateral lobes. They partly diverge apically and are connected by a weakly sclerotized surface at the base. The vesica looks like a short, fine, and bent tube, situated some-what more apically of the bases of the median plates.

Tribe PODOPINI Amyot et Serville, 1843

Scotinophara lurida (Burmeister, 1834) (Figs. 4, 25)

Material. Nagasaki, Kiu-Shiu, 1889 (Krupenin), 1 specimen; Sichuan, Yachzhou-Yunchzhinsyan, 8.IV. 1983 (Potanin), 1 specimen; Vietnam, 50 km W Khan Hoa, 10.I.1989 (Korotyaev), 1 specimen; China, Tshou-San, 1 specimen.

The phallobase (Fig. 4) is large, strongly scle-rotized, narrowed ventrally and strongly dilated dor-sally. Free ends of the basal plates are long. Dorsal connectives are very short; they terminate as large mushroom-like bodies. Their base looks like rather long sclerotized stem; the distal part is wide, weakly sclerotized triangular plate. Laterally, this plate is strengthened by strongly sclerotized ridges converging to the stem and forming a Y-shaped structure. Ventral processes are rather long, looking like narrow triangu-lar plates. Their external angles are less sclerotized than the body of the phallobase; these angles strongly project above margins of the basal plates. Suspensory apodemes are very short and thick.

The theca is subdivided into well-developed basal and apical parts. The basal part of the theca is small and strongly sclerotized, slightly narrowed apically and convex dorsally. Ventral tubercles are undevel-

oped. The apical part of the theca is approximately of the same length as its basal part, but is sclerotized to a significantly lesser extent; it is weakly convex and slightly dilates toward the apex. The dorsal emargina-tion of its apical margin is rectangular and rather wide; its base smoothly passes into the membranous wall of the conjunctive. The endophallic duct is rather thick, weakly sclerotized, and elastic.

The conjunctive is large, flattened laterally, and, in the resting state, is situated totally outside the theca. The apical part of the conjunctive is very low from the dorsal side; it is situated virtually at the same level with the base of the dorsal emargination of the apical margin of the theca. Ventrally, it gradually and weakly elevates. The largest part of the conjunctive is repre-sented by a convex ventral wall and long sclerotized median plates of the penis.

Median plates are situated nearly entirely on the lat-eral walls of the conjunctive and smoothly elevate from its apex dorsally. They are narrowed basally and dilated apically; their apices adjoin dorsally and are set apart ventrally, where their margins are connected by the ventral wall of the conjunctive. Over the rest of the length, they are connected at the dorsal side by a membranous convex wall passing into the wall of the apex of the conjunctive. Similarly to those in Grapho-soma lineatum, narrowed basal parts of the median plates can be homologized to the longitudinal fibers of some other species mentioned here; the dilated apical areas can be homologized to the apical outgrowths. The ventral aperture, formed by bent apical margins of the median plates of the penis, passes into a very deep cylindrical depression of the membranous wall of the conjunctive between median plates; the vesica departs from the bottom of this depression. The latter looks like a very long and fine tube; in resting state, the vesica is folded spirally and entirely lies in the mem-branous depression in the cavity of the conjunctive. The secondary gonopore looks like a small aperture on the apex of the vesica. When the aedeagus is erected, owing to the pressure of pumped on fluid affecting the membranous depression, the vesica emerges out of this aperture between the apices of the median plates of the penis. At the same time, it makes a wide circle; as a result, its apex is situated above the dorsal margin of the apices of the median plates.

Scotinophara fibulata (Germar, 1839)

Material. Poundou, Haute Volta, Afrique Occiden-tale Francaise, III.1928 (Olsufjev), 2 specimens.



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In this case, the aedeagus is described in the resting state. It is structurally similar to that of the preceding species.

Dorsal ends of the basal plates of the phallobase very widely diverge apically. Dorsal connectives are slightly longer than those of the preceding species; mushroom-shaped bodies are also structurally similar to those of S. lurida, but with less developed scle-rotized ridges laterally to the lamellar part. Ventral processes look like narrow and wide rectangular plates.

The basal part of the theca is small. The apical part of the theca consists of 2 long separate plates (more than twice as long as basal part of theca), wide at the base and narrow, slightly bent ventrally in the apical half.

The conjunctive is very large, slightly flattened lat-erally and strongly dilated and swollen dorsally. Short median plates of the penis are situated on its ventral wall closer to the apex. They are weakly sclerotized at the base, gradually passing into the wall of the con-junctive. Apically and from the ventral side, the me-dian plates are widely bent toward each other and form emarginations along the line of adjoining; these emar-ginations form a pair of apertures; the basal aperture is larger. A fine sclerotized fiber departs from each me-dian plate along its ventro-lateral wall to its base. The depression of the wall of the conjunctive between median plates is wide and very long, forming 11 coils in the cavity of the conjunctive; these coils are situated on the right side. Walls of five basal coils are scle-rotized; sclerotization is especially strong on the dor-sal side. Other coils are membranous, gradually nar-rowing in diameter. The vesica, looking like a fine sclerotized tube approximately 3 times as long as the body lies in the cavity of the depression along its en-tire length. It terminates as a long and slightly dilated groove exposed dorsally. The endophallic duct turns right in the cavity of the conjunctive and goes through the space surrounded by coils of the depression of the wall of the conjunctive, reaching the base of the vesica. Probably, only a part of the vesica can project from the conjunctive during copulation; this part cor-responds to no more than six coils of the depression of the wall of the conjunctive in resting state.

Gambiana aspera (Walker, 1867) (Figs. 26, 27)

Material. Ouagadougou, Afrique Occidentale Fran-caise, X.1926 (Olsufjev), 2 specimens; Poundou, Hau-

te Volta, Afrique Occidentale Francaise, 16.X.1927, 18.X.1927 (Olsufjev), 2 specimens.

The phallobase is large, narrowed ventrally and di-lated dorsally. Free ends of the basal plates are rather long. Dorsal connectives are very short. Mushroom-like bodies are structurally similar to those of S. lurida, but are smaller. Ventral processes look like small and narrow rectangular plates, not projecting beyond the margin of the phallobase. Suspensory apo-demes are slightly longer than in S. lurida.

The theca and the conjunctive are similar to those of S. lurida. The basal part of the theca is long and wide, narrowing apically. Ventral tubercles are undeveloped. The apical part of the theca is small, separated from the basal part by an indistinct band. The dorsal emar-gination of its apical margin is narrow and rather deep. Lateral apical angles are acute. The elastic endophallic duct is S-shaped.

The conjunctive is small. The apical part of the con-junctive is low on the dorsal side; ventrally, it forms a small cone bent dorsally. The ventral wall of the conjunctive is significantly swollen. Median plates of the penis are wide; their triangular bases (probably, homologous to longitudinal fibers of other species) are situated on the sides of the conjunctive. Apices (also, probably, corresponding to the apical outgrowth of other species) of the median plates are widened and cupulate, adjoining each other dorsally. Apices of each plate possessing an emargination on the ventral mar-gin. Emarginations of both plates form an oval aper-ture. A shallow but rather capacious depression of the median wall of the conjunctive is situated between median plates (Fig. 27); the vesica departs from the bottom of this depression. The latter looks like a not very long sclerotized tube, slightly dilated basally and weakly arcuate. The distal knee of the S-shaped endo-phallic duct adjoins the described depression of the wall of the conjunctive. During erection of the aedeagus, pumped erective fluid presses on the wall of the membranous depression; as a result, the vesica projects between apices of the median plates of the penis and slightly bends its apex towards the base of the aedeagus.

Amaurochorus cinctipes (Say, 1828) (Figs. 28, 29)

Material. Roselle Park, New Jersey, 16.IV.1911 (collected by sifting), 1 specimen; Brigantine, New Jersey, 5.IX.1930 (J.C. Lutz), 2 specimen.

The phallobase is distinctly transverse, rounded ventrally. Free ends of the basal plates are rather long.

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Dorsal connectives are very short. Mushroom-like bodies are rather large, with very short proximal stem. Distally, they are divided into two flattened branches, situated at the right angle toward each other. Ventral processes look like small and widely diverged promi-nences. Suspensory apodemes are shorter than the phallobase.

The theca is distinctly divided into two parts. The basal part of the theca is short and wide, with a large basal orifice, strongly convex dorsally. Ventral tuber-cles are undeveloped. The apical part of the theca is slightly shorter than the basal part and is divided into two widely diverged narrow plates with narrowly rounded apices. The wall between them is entirely membranous, without visible border passing into the wall of the conjunctive.

The conjunctive is rather large, with developed ven-tro-lateral lobes. The latter are weakly sclerotized at the dorsal side, with wide base and two short apices each. Apices of one pair are very widely rounded and directed laterally; apices of another pair are finer and directed toward the base of the conjunctive. The apical part of the conjunctive looks like a low and wide dome projecting beyond the apical angles of the plates of the apical part of the theca.

Median plates of the penis are wide, situated from the ventral side of the conjunctive in the frontal plane of the aedeagus (in non-erected state, they are situated nearly perpendicular to this plane. Along the line of adjoining, the inner margins of the median plates form emarginations. These emarginations together form a narrow oval aperture. The membranous wall of the conjunctive between the median plates forms a shal-low depression. The vesica, looking like a short and weakly sclerotized tube with oblong secondary gonopore at the end, departs from the wall of this de-pression. Probably, it extends from the aperture be-tween the median plates during copulation.

Podops (Opocrates) rectidens Horvath, 1883 (Figs. 30, 31)

Material. Tavricheskaya Prov., Kinburn, salted lake, 19.V.1938 (S.I. Medvedev), 1 specimen; Poti, W Georgia, 10.VIII.1938 (D. Kobakhidze), 1 specimen; Othkara, Gudauta Distr., Abkhazia, marsh, 1.V.1958 (Kurnakov), 1 specimen.

The phallobase is strongly transverse, rounded ven-trally. Free ends of the basal plates are short. Dorsal connectives are very short. Mushroom-like bodies are

rather large, with a long proximal stem. Distally, they are divided into two flattened sclerotized branches, T-shaped. Ventral processes look like small, weakly sclerotized triangular plates, not projecting beyond the margin of the phallobase. Suspensory apodemes are as long as basal plates.

The theca is distinctly divided into two parts by a narrow band. The basal part of the theca is rather large, strongly convex dorsally and flat ventrally. Ven-tral tubercles are developed; they are partly scle-rotized. The apical part of the theca is slightly shorter than basal part and is less convex; however, it is wider than the basal part at the apex. The dorsal emargina-tion of the apical margin of the apical part of the theca is indistinct; the wall of the theca near this emargina-tion is nearly membranous. Latero-apical margins of the apical part of the theca are widely rounded.

The conjunctive is rather large and complexly fis-sured. Ventro-lateral lobes of the conjunctive are rather long and wide; they are directed laterally. The apex of each lobe looks like a dome, elongated dorso-ventrally and narrowed apically. A pair of small di-verged spherical swellings is situated at the apex of the conjunctive on its dorsal side. A pair of basally con-verged and proximally fine long ventral branches, dilating apically, directed apically, and diverging lyre-like, are situated at the apex of the conjunctive from its ventral side. Small oblong tubercle directed api-cally is situated between them. Rather large sections of the lateral walls of the conjunctive are strongly scle-rotized, looking like triangular concave plates con-nected with the apical angles of the apical part of the theca.

The ventral lobe is developed. Median plates of the penis are subdivided into longitudinal fibers and apical outgrowths. Longitudinal fibers are long and short; they are diverged rather widely. Apical outgrowths are long and wide, most sclerotized at the apex; in this region, they are slightly bent toward each other. Dor-sally, apical outgrowths are connected by a weakly sclerotized and flattened wall approximately in the middle of their length. The sclerotized vesica, looking like a bent tube slightly projecting beyond apical out-growths, is situated between these outgrowths closer to the ventral wall of the ventral lobe.

Podops (Podops) inunctus (Fabricius, 1775)

Material. Magenta, Italy, 17.V.1934 (Passaura), 2 specimens; Gallia, 1 specimen.



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The aedeagus is structurally similar to that of P. rectidens, differing in the following characters. Ventro-lateral lobes of the conjunctive possess narrow bases directed ventrally and also widened carina-like distal parts directed laterally. Apices of the ventro-lateral lobes look like strongly smoothed tubercles directed dorsally. Ventral branches of the apex of the conjunctive are significantly longer than of Podops incertus; they are fine basally and widened and slightly bent apically in the distal part; they are di-rected ventrally. Apical plates of the median lobes of the penis are situated closer to each other.

Kayesia parva Schouiteden, 1903

Material. Ouagadougou, Afrique Occidentale Fran-çoise, X–XI.1926 (Olsufjev), 2 specimens; Sudan, Upper Nile, nr. Malakal, 5–20.I.1963 (Linnavuori), 2 specimens.

The phallobase is square. Free ends of the basal plates are rather long. Dorsal connectives are very short; they terminate as mushroom-like bodies, look-ing like large triangular and weakly sclerotized plates situated on a very short proximal stem. Ventral proc-esses look like short and rather wide rectangular plates. Suspensory apodemes are as long as the phallo-base.

The theca is distinctly divided into two parts. The basal part of the theca is oblong, with thick walls and dilated prominence at the base of the ventral wall. It is narrow at the base, dilating apically. Ventral tubercles are indistinct. The apical part of the theca is short; at the base, it is wider than the apex of the basal part. The dorsal emargination of the apical margin of the apical part of the theca is rather deep. Latero-apical angles are acute.

The conjunctive is small. Ventro-lateral lobes are rather long, smoothly narrowing towards the apex. Their proximal parts are directed ventro-laterally and apices are directed ventrally and slightly bent toward the base of the aedeagus at the end. A small lateral swelling is formed by the conjunctive at the base of each ventro-lateral lobe. The wall between the ventro-lateral lobes is flat. The dorsal wall of the conjunctive smoothly rises and forms a small tubercle at the apex; the median plates of the penis are situated ventrally to this tubercle. The plates look like small parallel plates with free margin gradually rising from the base on the dorsal side and, forming an arch, gradually lowering to the base of the ventro-lateral plates. The vesica, look-

ing like a very short sclerotized tube, is situated be-tween the median plates of the penis.

Tribe TARISINI Stål, 1872

Tarisa subspinosa (Germar, 1839) (Figs. 32–35)

Material. Khanskaya Stavka, Astrakhan Prov., 26.V–16.VI.1887 (Plyushchevskii), 4 specimens; Bu-chara mer., Termez, 27.VI.1912 (Kiritshenko), 2 spe-cimens.

The phallobase (Fig. 32) is wide and oval. It is weakly sclerotized ventrally and stronger at sides. Dorsal connectives are very short; they are attached to the margin of large lamellate and oval mushroom-shaped bodies. Ventral processes look like very small and virtually indistinct prominences. Ventral conjunc-tives are weakly developed. The pons is rather wide and weakly sclerotized. The basal aperture of the phallobase is wide. Suspensory apodemes are slightly shorter than the phallobase.

The basal part of the theca (Fig. 33) is represented by a wide and convex membrane, making the theca very mobile in relation to the phallobase. Ventral tu-bercles (in sense as they treated in some abovemen-tioned species). The theca is not subdivided into parts. It is cylindrical and narrowed only at the base on the dorsal side. The theca is weakly sclerotized. A single longitudinal and strongly sclerotized ridge is situated on each of the dorso-lateral surfaces of the theca.

The conjunctive (Figs. 33, 34) is small and nar-rowed laterally. Ventro-lateral lobes look like flat scle-rotized trapezoid plates with longer margin directed toward the base of the conjunctive and with acute an-gles. A small basal lobe (probably, a derivative of ventral lobe), looking like a weakly sclerotized lingui-form process, directed obtusely toward the base of the aedeagus, is situated between the ventro-lateral lobes closer to the base of the conjunctive. Median plates of the penis are reduced. The apical part looks like a rather long cone directed virtually strictly apically. The dorsal wall of the conjunctive is bent and convex. The vesica is situated at the base of ventro-lateral lobes on the wall of the conjunctive closer to its apex. It looks like a sclerotized tube, bent at the base and directed toward the base of the aedeagus. The secon-dary gonopore opens at its apex. The ejaculatory res-ervoir (Fig. 35) is small, flattened laterally; it is situ-ated in the conjunctive. The semen duct is long and elastic; it adjoins the ventral wall of the theca. The endophallic duct is very short.

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Tarisa pallescens Jakovlev, 1871

Material. Sarepta, 1865 (Bekker), 2 specimens; Kerch, Crimea, 9.VII.1918 (Kiritshenko), 1 specimen; Kumak, Kattakurgan near Samarkand, 10–11.VI.1929 (L. Zimin), 2 specimens; Vakhsh River valley, 6 km W Kuibyshev, 19.VII.1943, 20.VII.1943, 22.VII.1943 (Kiritshenko), 3 specimens; Frunze, Kirghizia, 18.VII. 1945 (Ljubishchev), 1 specimen.

The aedeagus is structurally similar to that of T. subspinosa. The pons of the phallobase is scle-rotized stronger than of the preceding species. Ventro-lateral lobes of the conjunctive are lanceolate, possess-

ing pointed apices, slightly bent toward the apices of the conjunctive narrow bases directed ventrally and also widened carina-like distal parts directed laterally. The basal lobe is sclerotized weaker than the ventro-lateral lobes; it is rectangular and slightly bifurcated apically. The base of the vesica is strengthened by narrow short fibers, running laterally along the wall of the conjunctive. The apical part of the conjunctive is conical and directed ventro-apically.

Tarisa virescens Herrich-Schäffer, 1847

Material. Nork near Yerevan, Armenia, 26.IV.1948 (Rikhter, Vartikyan), 2 specimens.

Figs. 32–39. Podopinae. Aedeagus. (32, 35) Tarisa fraudatrix Horvath: (32) phallobase and theca in ventro-lateral, (33) lateral, and (34) ventral view; (35) ejaculatory reservoir in lateral view; (36, 37) Bolbocoris variolosus (Germar) (according to Gapon, 2005): (36) in ventral and (37) lateral view; (38, 39) Testrica antica Walker: (38) ventral and (39) lateral view.



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The aedeagus is structurally similar to that of the two preceding species. Ventro-lateral lobes of the con-junctive are lanceolate, narrow at the base, slightly bent toward each other directly before the apex and bent toward the apex of the conjunctive. These lobes are diverged to a greater extent than in the preceding species. The basal lobe slightly dilated distally, rounded, and slightly incised at the end. The apical part of the conjunctive is conical, slightly bent dor-sally at the apex.

Genera with Undetermined Taxonomic Position

Bolbocoris variolosus (Germar, 1839) (Figs. 36, 37)

Material. Ouagadougou, Afrique Occidentale Fran-çoise, 1926 (Olsufjev), 2 specimens.

The phallobase is slightly narrowed and rounded ventrally; dorsally, it is wider. Free ends of the basal plates are rather long and wide. Dorsal connectives are very short; they are attached near the margin of wide and weakly sclerotized oval mushroom-like bodies. Ventral processes look like widely set apart and slightly visible triangular prominences. Ventral con-nectives are indistinct. Suspensory apodemes are ap-proximately as long as basal plates.

The theca is not subdivided into parts; it is elon-gated, approximately twice as long as wide, and strongly sclerotized. Its ventral wall forms dilated prominence at the base. Ventral tubercles are undevel-oped; only a narrow weakly convex membranous strip is found basally to the prominence of the ventral wall of the theca. The ejaculatory reservoir is large and oblong; it occupies nearly the entire cavity of the theca except for its apical part.

The conjunctive is small. Its base is distinctly sepa-rated as a narrow and weakly convex carina. Paired ventro-lateral lobes and an unpaired dorsal lobe form the largest part of the conjunctive. Ventro-lateral lobes are short, dilated at the base; they are directed api-cally, bent arcuate, and adjoin each other above the apical part of the conjunctive. The dorsal lobe is wide at the base; this base adjoins bases of the ventro-lateral lobes. It is directed dorsally and its pointed apex is bent toward the base of the conjunctive. Its apical wall forms a small membranous carina. The apex of the conjunctive looks like a small flattened plate at the level of the bases of the longitudinal fibers of the me-dian plates of the penis between bases of the ventro-lateral and dorsal lobes.

Median plates of the penis are rather long. Their proximal parts look like narrow and somewhat set apart converging longitudinal fibers situated between ventro-lateral lobes. Very fine sclerotized fibers run from bases of longitudinal fibers along ventro-lateral surfaces of the conjunctive in the basal direction. Free apical outgrowths of the median plates are widened and less sclerotized at ends; they diverge, projecting beyond the ventro-lateral lobes ventrally. Their mar-gins, facing the base of the aedeagus, are connected by a fine weakly sclerotized wall. The vesica, looking like a short and rather thick bent tube, pointed ventro-apically, is situated between the median plates.

Bolbocoris inaequalis (Germar, 1838)

Material. Zanzibar, Africa or., 4.1812, nos. 143–192, 1 specimen; RSA, Skukuza, Kruger’s Nat. Park, 14–16.III.1994 (V. Belozerov), 1 specimen.

The aedeagus is similar to that of B. variolosus, but the phallobase is more narrowly rounded ventrally; free ends of the basal plates are finer; mushroom-like bodies are significantly narrower; the apex of the dor-sal lobe of the conjunctive is widely rounded at the end; longitudinal membranous carina on its apical wall is undeveloped.

Tetrica antica Walker, 1867 (Figs. 38, 39)

Material. Australia, Ayers Rock, 5.V.1978 (To-bias), 1 specimen.

The phallobase is narrowed and rounded ventrally. Free ends of the basal plates are rather long and di-verging. Dorsal connectives are very short; they termi-nate as short and wide conical mushroom-like bodies. Ventral processes look like rather long (slightly shorter than free ends of basal plates) and narrow tri-angular plates, slightly projecting beyond the sides of the phallobase by apical angles. Ventral connectives are indistinct. Suspensory apodemes are approximately as long as basal plates.

The theca is not subdivided into parts; it is rather short and wide; it is strongly convex dorsally. Ventral tubercles at its base are undeveloped. The basal aper-ture of the theca is small and somewhat shifted ven-trally.

The conjunctive is approximately as long as the theca; it is narrowed laterally. Its ventro-lateral lobes are rudimentary, looking like small convex membra-nous tubercles, situated baso-laterally. The apical part

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of the conjunctive is low; it is divided into two long and narrow branches, pointed ventrally and running in parallel. The ventral lobe is rather short, wide, and very convex, especially at the dorsal side.

The median plates of the penis are subdivided into two parts. Their proximal parts are very strongly scle-rotized, wide and convex. They are rather long, stretching along the ventro-lateral walls of the con-junctive near to the bases of branches of its apex. Their distal parts are situated at the level of the bases of the ventral lobe; they form free outgrowths, slightly bent toward each other. Distal parts of the median plates run along the perimeter of the ventral lobe ven-trally. They are immovably connected with proximal parts and look like rather narrow flat fibers, narrowing toward the apical lobe and passing into each other in that region. They are less sclerotized than proximal parts of the median plates. The vesica is situated on the ventral wall of the ventral lobe at its base. It looks like a very long and pointed apically S-shaped tube, strongly sclerotized, widened basally, and narrowing apically. Its distal part is elongated into a rather long and fine, weakly sclerotized elastic thread.

DISCUSSION

In the examined species of the subfamily Podopi-nae, the aedeagi are so structurally diverse that it is impossible to compose the basic structural scheme, differing from that in the family Pentatomidae on the whole. The absence of a common scheme, typical of all the aedeagi, is one more argument for the poly-phyletic origin of Podopinae. At the same time, com-mon structural characters typical of the aedeagi of representatives of separate tribes of the family Po-dopinae, could be distinguished. These common char-acters concern the phallobase and theca, whereas the structure of the endosoma allows distinguishing super-generic groups within tribes.

For example, the ventrally rounded phallobase with small triangular ventral processes and conical or flat-tened conical, i.e., discoid mushroom-like bodies (Figs. 1–3) are characteristic of examined species of the tribe Graphosomatini. In representatives of this tribe, the theca possesses well-developed ventral tubercles. The apical part of the theca is weakly devel-oped; it is short and indistinctly separated from the basal part. Occasionally, the apical part is absent (Figs. 5–23).

The examined species of the tribe Podopini possess the phallobase with rectangular or triangular ventral

processes (Fig. 4). In these species, mushroom-like bodies possess a stem at the base and a weakly scle-rotized lamellar distal part with lateral margins strengthened by strongly sclerotized ridges converging toward the stem. Different modifications of this struc-ture are found in different species. In the all examined species of this tribe, the apical part of the theca is well-developed and distinctly separated from its basal part (Figs. 25–31). Ventral tubercles are developed or absent.

Species of the tribe Tarisini differ from all the other Podopinae most strongly, which is evident from the structure of all the parts of the aedeagus (see below).

In species of the subfamily Podopinae, the de-scribed aedeagi form several stable structural types within tribes. These types are characterized below.

Tribe GRAPHOSOMATINI Mulsant et Rey, 1865

(A) The phallobase is slightly transverse. Free ends of the basal plates are rather long. Mushroom-like bodies are conical. The theca is subdivided into two parts (Figs. 5–8). The basal part of the theca possesses developed ventral tubercles. The apical part of the theca is weakly separated from its basal part. The con-junctive possesses well-developed ventro-lateral and ventral lobes. The apical part of the conjunctive looks like a low dome. The apical outgrowths of the median plates of the penis are not turned toward each other by their margins. The vesica is fine and long; it is directed ventro-apically. This basic scheme is characteristic of species of the genus Derula. In some characters, these species rather significantly differ from each other. For example, D. longipennis (Figs. 5, 6) is characterized by a long apical part of the theca with the double apex and developed median plates of the penis, subdivided into longitudinal fibers and apical outgrowths. In D. flavoguttata (Figs. 7, 8), the aedeagus possesses the lower and less convex apical part of the theca, directed ventrally (as in Sternodontus ampliatus) ventro-lateral lobes with single apices, weakly sclerotized outer walls, and fused with the ventral lobe basally. The median plates of the penis are rudimentary; they are not subdivided into longitudinal fibers and apical out-growths.

(B) The phallobase is slightly wider than long, with short free ends of the basal plates (Fig. 1). Mushroom-like bodies are conical. The theca is subdivided into two parts. The basal part of the theca possesses devel-oped ventral tubercles (Figs. 9–13). The apical part of



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the theca is weakly developed, short, and indistinctly separated from its basal part. The conjunctive pos-sesses ventro-lateral lobes with single apices; they are rather long and narrowing apically. The apical part of the conjunctive looks like a low dome. The ventral lobe is situated between the ventro-lateral lobes di-rected ventrally. The median plates of the penis are rather large, with longitudinal fibers and wide apical outgrowths. The apical outgrowths are set apart at the base ventrally; apically, they are widely turned over each other, adjoining at a short distance; then they diverge and again converge on the dorsal side. This basic scheme includes the aedeagi of Ancyrosoma leucogrammes, Sternodontus ampliatus, and Oplisto-chilus pallidus.

(C) The phallobase is slightly transverse, with short free ends of the basal plates (Fig. 2). Mushroom-like bodies are conical. The theca is subdivided into two parts. The basal part of the theca possesses developed ventral tubercles. The apical part of the theca is very weakly developed. The conjunctive is small, with re-duced ventro-lateral lobes (rudiments are found in Graphosoma lineatum); its apex looks like a low dome or cone. The median plates of the penis are rather large, with bases (which can be homologized to longi-tudinal fibers of other species) situated on the sides of the conjunctive. Their distal parts (also, probably, homologous to apical outgrowth of other species) are directed ventrally; they are free and their margins are bent toward each other (in Tholagmus flavolineatus they adjoin). The vesica is very short. This basic scheme includes the aedeagi of Graphosoma lineatum and Tholagmus flavolineatus.

(D) Free ends of the basal plates of the phallobase are rather long (Figs. 3, 24). Mushroom-like bodies are discoid. The theca is not subdivided into parts, with well-developed ventral tubercles (Figs. 19, 20, 24). Dorsally and laterally, the base of the conjunctive is separated as a wide and convex, especially dorsally, basal carina (Figs. 18, 19, 21–23). Bases of long and fine sclerotized processes, pointed toward the aedeagus, are situated on its dorsal wall (these proc-esses are absent in Ventocoris trigonus). Ventro-lateral lobes are large (Figs. 18–23), bearing a pair of tuber-cles (in Vilpianus galii, narrowing apices of the lobes correspond to median tubercles and small tubercles at the base of lobes, to lateral tubercles). The lateral walls of the ventral lobe are evident parts of the ven-tro-lateral lobes. The apex of the conjunctive is strongly smoothed, dorsally passing into the basal

carina. Longitudinal fibers of the median plates of the penis are narrow; they are situated in the same plane with the apex of the conjunctive; the distal parts of the median plates look like plates lying on the inner walls of the ventro-lateral lobes (Figs. 18, 20, 21). The api-cal margins of these plates are bent toward each other. More distally, median plates pass into fine fibers pointed to the base of the conjunctive (in Crypsinus, the median plates are reduced to very small triangular plates) (Fig. 22). The vesica looks like a small scle-rotized tube pointed ventrally. This basic scheme in-cludes aedeagi of Vilpianus galii, Ventocoris fischeri, V. oschanini, V. trigonus, and Crypsinus angustatus.

As seen from the descriptions, the structural scheme (A) is closer to scheme (B) [mushroom-like bodies, apical part of theca, ventro-lateral lobes of conjunc-tive, and vesica]; scheme (B) is, in its turn, rather simi-lar to scheme C [free ends of basal plates, mushroom-like bodies, and apical outgrowths of median plates of penis]. Scheme (D) is the most remote from the other schemes of the structure of the aedeagus in this group.

Tribe PODOPINI Stål, 1872

(A) The phallobase is narrowed ventrally, with long free ends of the basal plates and rectangular ventral processes (Fig. 4). Mushroom-like bodies possess a stem at the base and lamellar distal part; laterally, this plate is strengthened by strongly sclerotized ridges converging to the stem and forming a Y-shaped struc-ture. Ventral tubercles on the basal part of the theca are undeveloped (Fig. 25). The apical part of the theca is well-developed. Ventro-lateral and ventral lobes of the conjunctive are absent. The median plates of the penis are rather large, they are situated on the lateral walls of the conjunctive; their apices are pointed ven-trally, forming emarginations along the line of adjoin-ing; these emarginations form apertures. The membra-nous wall between the median plates forms a depres-sion, where a long or not very long vesica is situated; this vesica shifts outwards during erection of the aedeagus. This basic scheme includes the aedeagi of Scotinophara lurida, S. fibulata, and Gambiana as-pera.

The aedeagus of Amaurochorus cinctipes (Figs. 28, 29) is very similar to this structural scheme, differing only in the presence of double-pointed ventro-lateral lobes; flat median plates of the penis are situated on the ventral walls of these plates. If we accept the re-duction of these lobes, then median plates will occupy the position characteristic of this structural scheme. In

GAPON, KONSTANTINOV


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A. cinctipes, mushroom-like bodies possess the re-duced distal lamellar part; only flattened lateral ridges, forming the right angle, are left.

(B) The phallobase is strongly transverse, rounded ventrally. Free ends of the basal plates are short, ven-tral prominences are triangular. Mushroom-like bodies possess a long stem at the base, forming T-shaped structure with its two branches. The basal part of the theca possesses well-developed ventral tubercles (Figs. 30, 31). The apical part of the theca is large. The conjunctive possesses a complicated structure. Ventro-lateral lobes are large; they are directed later-ally and their apices are slightly bent dorsally. The apex of the conjunctive bears paired membranous ven-tral branches and small dorsal tubercles. The ventral lobe is developed. Median plates of the penis are sub-divided into longitudinal fibers and apical outgrowths. Apical outgrowths are connected by a weakly scle-rotized wall; they are pointed toward the base of the aedeagus. The vesica is rather long. This basic scheme includes aedeagi of Podops incertus and P. inunctus.

Apparently, the aedeagus of Kayesia parva demon-strates a separate structural scheme.

Tribe TARISINI Stål, 1872

All the species examined in this tribe possess the aedeagus of the same structural type, characterized by the following features. The phallobase is wide and oval, stronger sclerotized laterally (Fig. 32). Mush-room-shaped bodies are large, oval, lamellar; dorsal conjunctives are attached to them laterally. Ventral processes look like very small prominences. The theca is weakly sclerotized, its base looks like a wide convex membrane. Ventral tubercles are undeveloped. Dorso-lateral walls of the theca are strengthened by a pair of strongly sclerotized longitudinal fibers (Fig. 33). The ejaculatory reservoir is situated in the cavity of the conjunctive, adjoining the ventral wall of the theca. The endophallic duct is very short. Ventro-lateral lobes look like weakly sclerotized flattened plates, directed ventrally (Figs. 33, 34). The ventral lobe (in the common sense) is absent. The weakly sclerotized basal lobe, directed toward the aedeagus, is situated at the base of the conjunctive on its ventral side. This lobe appears to be a derivative of the ventral lobe. The vesica is situated on the wall of the conjunctive slightly apically to the bases of ventro-lateral lobes. It is rather long, bent basally, and pointed to the base of the aedeagus. The apex of the conjunctive is conical. This structural scheme is typical of all the species of Tarisa examined.

In species with an unclear taxonomic position, the aedeagi are also characterized by clearly distinguished structural schemes, testifying to the natural character of these groups.

(A) The phallobase is rounded ventrally. Mush-room-like bodies look like oval plates attached by their margin to dorsal connectives. Ventral processes are very small and triangular. The theca is not subdivided into parts; ventral tubercles at its base are undeveloped (Figs. 36, 37). The base of the conjunctive forms a small carina. Ventro-lateral lobes are single-pointed; they are directed apically and their apices are bent toward each other. The dorsal plate, directed dorsally, is well-developed. Median plates of the penis are sub-divided into longitudinal fibers and apical outgrowths. Fine sclerotized fibers run from the bases of the longi-tudinal fibers along the dorso-lateral walls of the con-junctive. Apices of the apical outgrowths of the me-dian plates are directed ventrally; these outgrowths are connected by a weakly sclerotized wall. The vesica looks like a short sclerotized tube, directed ventro-apically. This structural scheme is typical of Bolboco-ris variolosus and B. inaequalis.

(B) The structural scheme corresponds to that typi-cal of the aedeagus of Testrica antica (Figs. 38, 39).

It should be noted that groups of genera, mentioned within the tribes concerned, correspond to some taxa isolated previously. However, this correspondence is incomplete in relation to the volume of groups. For example, the group of species of the tribe Graphoso-matini, with the aedeagi corresponding to the type B of the present publication, corresponds to the tribe Ancy-rosomatini unaccepted by modern authors. At the same time, the genus Tholagmus included into this tribe by Puchkov (1965) possesses the aedeagus of the struc-tural type C. A similar structural scheme is character-istic of the group of genera corresponding to the tribe Graphosomatini sensu Puchkov (1965). At the same time, however, the genus Derula is characterized by the aedeagi belonging to a separate structural scheme A. The group of genera with the aedeagi be-longing to type D correspond to the tribe Trigonoso-matini unaccepted by modern authors. The taxonomic position of other genera included into the tribe Trigonosomatini needs clarification.

Probably, the structural schemes of the aedeagi of the tribe Podopini also characterized natural groups within the tribe.



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The above-described groups of genera possess dis-tinct sinapomorphies in the structure of the aedeagus, making it necessary to treat some previously described taxa, unaccepted by modern authors, as natural groups, needing some changes in their volume. The data ob-tained testify to the prospective character of the use of the structure of completely erect aedeagi in the taxon-omy of the subfamily Podopinae and to the necessity of the continuing of investigations in this field.

ACKNOWLEDGMENTS

The authors are sincerely grateful to Dr. I.M. Ker-zhner (ZIN RAS) for constant help and to Dr. David A. Raider (North Dakota State University) for consul-tations. The work was financially supported by the Russian Foundation for Basic Research, project no. 02–04–49138 and federal programs “Integration of Science and Higher Education in Russia in 2002–2006”; “Russian Universities,” project no. 07.01.056; and “Leading Scientific Schools” (2232.2003.4).

REFERENCES

1. Davidová-Vilimová, J. and Štys, P., “Diversity and Variation of Trichobothrial Patterns in Adult Podopinae (Heteroptera, Pentatomidae),” Acta Univ. Carolinae 33, 33–72 [1993 (1994)].

2. Davidova-Vilimova, J. and McPherson, J.E., “History of the Higher Classification of the Subfamily Podopinae

(Heteroptera, Pentatomidae), a Historical Review,” Acta Univ. Carolinae 38, 99–144 [1994 (1995)].

3. Gapon, D., “Inflation of Heteropteran Aedeagi Using Microcapillaries (Heteroptera, Pentatomidae),” Zoosyst. Ross. 9 (1), 157–160 [2000 (2001)].

4. Gapon, D.A., “On the Taxonomic Position of the Tribe Procleticini Pennington (Heteroptera: Pentatomidae),” Kavkaz. Entomol. Bull. 1 (1), 4–18 (2005).

5. Gapud, V.P., “A Generic Revision of the Subfamily Asopinae, with Consideration of its Phylogenetic Posi-tion in the Family Pentatomidae and Superfamily Penta-tomoidea (Hemiptera-Heteroptera): Parts 1, 2,” Philip-pine Entomologist 8 (1), 865–961 (1991).

6. Gross, G.F., Handbook of the Flora and Fauna of South Australia. Plant-Feeding and Other Bugs (Hemiptera) of South Australia. Heteroptera: Part 1 (Adelaide, Handbooks Committee, South Australian Government, 1975).

7. Leston, D., “The Wing Venation, Male Genitalia and Spermatheca of Podops inuncta (F.) with a Note on the Diagnosis of the Subfamily Podopinae Dallas (Hem. Pentatomidae),” J. Soc. British Entomol. 4, 129–135 (1953).

8. McDonald, F.J., “The Genitalia of North American Pentatomoidea (Hemiptera: Heteroptera),” Quaest. En-tomol. 2, 7–150 (1966).

9. Puchkov, G.V. Shield Bugs of Middle Asia (Ilim, Frunze, 1966) [in Russian].

10. Schuh, R.T. and Slater, J.A. True Bugs of the World (Hemiptera: Heteroptera): Classification and Natural History (Cornell University Press, Ithaca, 1995).

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