Russian Entomol. J. 13(3): 127149 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2004 Revision of the taxonomic structure of the tribe Dorcadionini (Coleoptera: Cerambycidae) on the base of endophallic morphology Ðåâèçèÿ òàêñîíîìè÷åñêîé ñòðóêòóðû òðèáû Dorcadionini (Coleoptera: Cerambycidae) íà îñíîâå àíàëèçà ñòðîåíèÿ ýíäîôàëëþñà M.L. Danilevsky1, D.G. Kasatkin2, A.A. Rubenyan3 Ì.Ë. Äàíèëåâñêèé1, Ä.Ã. Êàñàòêèí2, À.À. Ðóáåíÿí3 1 A.N. Severtzov Institute of Ecology and Evolution, Russian Academy of Science, Leninsky prospect 33, Moscow 119071 Russia; e-mail: [email protected] 1 Èíñòèòóò ïðîáëåì ýêîëîãèè è ýâîëþöèè ÐÀÍ, Ëåíèíñêèé ïðîñïåêò 33, Ìîñêâà 117071 Ðîññèÿ. 2 Rostov branch of FGU Rosgoskarantin, 20th line str. 43/16, Rostov-on-Don, Russia; e-mail: [email protected] 2 Ðîñòîâñêîå îòäåëåíèå "Ðîñãîñêàðàíòèí", óë. 20-ÿ ëèíèÿ, 43/16, Ðîñòîâ-íà-Äîíó, Ðîññèÿ 3 Moscow State Pedagogical University, Kibalchich str. 6, building 5, 129278 Moscow, Russia; e-mail: [email protected] 3 Ìîñêîâcêèé ïåäàãîãè÷åñêèé ãîñóäàðñòâåííûé óíèâåðñèòåò, óë. Êèáàëü÷è÷à, ä.6, êîðï. 5, 129278 Ìîñêâà, Ðîññèÿ KEY WORDS: Cerambycidae, Dorcadionini, Dorcadion, Iberodorcadion, Neodorcadion, Eodorcadion, endophallus, morphology, taxonomy, phylogeny, new subgenera. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Cerambycidae, Dorcadionini, Dorcadion, Iberodorcadion, Neodorcadion, Eodorca- dion, ýíäîôàëëþñ, ìîðôîëîãèÿ, òàêñîíîìèÿ, ôèëîãåíèÿ, íîâûå ïîäðîäû. ABSTRACT: The morphology of everted and in- proposed on the base of endophallic characters. D. flated endophallus is described and figured on the base klavdiae is transferred from D. (Carinatodorcadion) to of dry constant samples of 127 species and subspecies D. (Cribridorcadion). D. turkestanicum is placed in D. of four genera: Neodorcadion, Eodorcadion, Iberodor- (Cribridorcadion). The endophallus morphology of D. cadion and Dorcadion of all subgenera. The homology tschitscherini, D. mystacinum rufogenum and D. optatum of different endophallus parts is established. The origi- matthieseni (all three taxa were sometimes regarded as nal terminology is proposed. All genera and subgenera Pedestredorcadion) is typical for D. (Acutodorcadion, of Dorcadionini are clearly delimited on the base of subgen.n.). D. danczenkoi, stat.n. is raised to the species endophallic structures. New compositions of Dorcadi- rank. The phylogenetic relations inside the tribe are on (s.str.) and Eodorcadion (s.str.) are proposed. Two discussed. Several taxons are proposed to be accepted new subgenera are described on the base of endophallic as subspecies: Eodorcadion carinatum blessigi (Gangl- structures: Eodorcadion (Humerodorcadion, subgen.n.) bauer, 1884), E. c. bramsoni Pic, 1901, E. c. altaicum type species: Dorcadion humerale Gebler, 1823; (Suvorov, 1909), stat.n., Dorcadion cinerarium cauca- Dorcadion (Acutodorcadion, subgen.n.) type spe- sicum Küster, 1847, stat.n., D. sareptanum euxinum cies: D. acutispinum Motschulsky, 1860. The unique Suvorov, 1915, stat.n., D. sulcipenne goektschanum taxonomical position of D. (Politodorcadion) is dem- Suvorov, 1915, stat.n. onstrated; possible generic level of the taxon is sup- posed. Five synonyms are proposed: Dorcadion (s.str.) ÐÅÇÞÌÅ: Îïèñàíî è ïðîèëëþñòðèðîâàíî ñòðî- = D. (Compsodorcadion); D. (Cribridorcadion) = D. åíèå âûâåðíóòîãî è íàäóòîãî ýíäîôàëëþñà ïî ïî- (Pedestredorcadion), syn.n. = D. (Dzhungarodorcadi- ñòîÿííûì ñóõèì ïðåïàðàòàì 127 âèäîâ è ïîäâèäîâ on), syn.n. Dorcadion (s.str.) consists of 8 species: D. 4 ðîäîâ Neodorcadion, Eodorcadion, Iberodorcadion glicyrrhizae, D. crassipes, D. cephalotes, D. gebleri, D. è Dorcadion ñî âñåìè ïîäðîäàìè. Óñòàíîâëåíà ãî- ganglbaueri, D. alakoliense, D. abakumovi, D. tenue- ìîëîãèÿ ðàçëè÷íûõ ÷àñòåé ýíäîôàëëþñà, äëÿ îáî- lineatum; other species, which were traditionally in- çíà÷åíèÿ êîòîðûõ ïðèìåíåíà îðèãèíàëüíàÿ òåðìè- cluded in Dorcadion (s.str.), are placed in D. (Acutodor- íîëîãèÿ. Âñå ðîäû è ïîäðîäû òðèáû Dorcadionini cadion subgen.n.). Eodorcadion (Humerodorcadion èìåþò î÷åíü ÷¸òêèå ãðàíèöû ïî ñòðîåíèþ ýíäîôàë- subgen.n.) consists of two species: E. humerale and E. ëþñà. Ïðåäëîæåí íîâûé ñîñòàâ ïîäðîäîâ Dorcadion lutshniki. E. quinquevittatum, E. leucogrammum, E. (s.str.) è Eodorcadion (s.str.). Íà îñíîâå ñòðóêòóð tuvense, E. ptyalopleurum and E. maurum, as well as E. ýíäîôàëëþñà îïèñàíû äâà íîâûõ ïîäðîäà: Eodorca- sifanicum and E. glaucopterum are placed in Eodorca- dion (Humerodorcadion, subgen. n.) ñ òèïîâûì âè- dion (s.str.). A key for 4 genera and all subgenera is äîì: Dorcadion humerale Gebler, 1823 è Dorcadion 128 M.L. Danilevsky, D.G. Kasatkin, A.A. Rubenyan (Acutodorcadion, subgen.n.) ñ òèïîâûì âèäîì: D. We have studied 127 Dorcadionini species and sub- acutispinum Motschulsky, 1860. Ïîêàçàíà îáîñîá- species of four genera (Eodorcadion, Iberodorcadion, ëåííàÿ òàêñîíîìè÷åñêàÿ ïîçèöèÿ D. (Politodor- Neodorcadion, Dorcadion), choosing the representa- cadion), çàñëóæèâàþùåãî, âåðîÿòíî ðîäîâîãî ðàí- tives of different subgenera and group of species. En- ãà. Ïðåäëîæåíî ïÿòü ñèíîíèìîâ: Dorcadion (s.str.) = dophallic structures in Dorcadionini clearly show the D. (Compsodorcadion); D. (Cribridorcadion) = D. limits of genera, subgenera and species groups and (Pedestredorcadion), syn.n. = D. (Dzhungarodor- allow to clarify their natural connections. Usually sever- cadion), syn.n. Dorcadion (s.str.) ðàññìàòðèâàåòñÿ â al specimens of each species were used for preparation ñîñòàâå 8 âèäîâ: D. glicyrrhizae, D. crassipes, D. to realize the rate of individual variability, which was cephalotes, D. gebleri, D. ganglbaueri, D. alakoliense, not very high. D. abakumovi è D. tenuelineatum; äðóãèå âèäû, òðà- Usually we studied more than one specimen of each äèöèîííî îòíîñèâøèåñÿ ê Dorcadion (s.str.), ïîìå- taxon (up to 20), so our morphology conclusions are ùåíû â D. (Acutodorcadion subgen.n.). Eodorcadion well representative. But some species were poorly rep- (Humerodorcadion subgen.n.) âêëþ÷àåò äâà âèäà: E. resented in our collections, and we were not able to humerale è E. lutshniki. E. quinquevittatum, E. leuco- study more than one specimen (Dorcadion glabrofas- grammum, E. tuvense, E. ptyalopleurum è E. maurum, ciatum, Eodorcadion sifanicum and some others). Indi- êàê è E. sifanicum è E. glaucopterum ïîìåùåíû â vidual variability of endophallus morphology is not Eodorcadion (s. str.). Ñîñòàâëåíà îïðåäåëèòåëüíàÿ significant, so it was not always necessary to use many òàáëèöà äëÿ âñåõ 4 ðîäîâ ñî âñåìè ïîäðîäàìè íà specimens of a taxon for study. The general form of îñíîâå ñòðóêòóð ýíäîôàëëþñà. D. klavdiae ïåðåíå- everted endophallus more or less depends on the air- ñ¸í èç D. (Carinatodorcadion) â D. (Cribridorcadion). pressure during preparation. D. turkestanicum ïîìåù¸í â D. (Cribridorcadion). Ñòðîåíèå ýíäîôàëëþñà D. tschitscherini, D. mysta- Morphology of endophallus cinum rufogenum è D. optatum matthieseni (âñå òðè òàêñîíà èíîãäà ðàññìàòðèâàëèñü â Pedestredorca- dion) âïîëíå òèïè÷íî äëÿ D. (Acutodorcadion, Dorcadionini endophallus is represented by thin, subgen.n.). D. danczenkoi, stat.n. ïðèçíàí ñàìîñòîÿ- transparent membrane partly covered with different òåëüíûì âèäîì. Îáñóæäàþòñÿ ðîäñòâåííûå îòíî- cuticula elements: very small and short spicules (Fig. øåíèÿ òàêñîíîâ âíóòðè òðèáû. Íåñêîëüêî òàêñîíîâ 87), strongly elongated microtrichiae (Fig. 100), rela- ïðèíÿòû â êà÷åñòâå ïîäâèäîâ: Eodorcadion carinatum tively wide spines (Figs. 55, 112) and with more or less blessigi (Ganglbauer, 1884), E. c. bramsoni Pic, 1901, distinct sclerits. Endophallus generally is relatively long, E. c. altaicum (Suvorov, 1909), stat.n., Dorcadion cine- usually much longer than abdomen, sometimes longer rarium caucasicum Küster, 1847, stat.n., D. sarep- than body, specially, in Dorcadion (Carinatodorcadi- tanum euxinum Suvorov, 1915, stat.n., D. sulcipenne on) (Figs. 43, 47, 49) and D. (Maculatodorcadion) (Fig. goektschanum Suvorov, 1915, stat.n. 110), usually if long then more or less S-shaped; or relatively short, shorter than abdomen, specially in Eodorcadion (Humerodorcadion, subgen.n.) (Figs. 25 Introduction 28). In the most complicated cases: Dorcadion (Cari- natodorcadion) (Figs. 4349), D. (Maculatodorcadi- The utilization of endophallic structures in Ceram- on) (Fig. 110), D. (Megalodorcadion) (Fig. 109), D. bycidae taxonomy is traditionally limited because of (Cribridorcadion, sensu nov.) (Figs. 50108) it con- exceptionally big problems with preparation of con- sists of several distinctly limited parts of different length stant, dry patterns of everted and inflated very thin and shape. In others groups: Neodorcadion (Figs. 13), membranous tube in the position close to the natural Eodorcadion (Figs. 429), Iberodorcadion (Figs. 30 erection of endophallus. Several works used Ceramby- 38), Dorcadion (Politodorcadion) (Figs. 3942) differ- cidae males, which were collected during copulation ent elements of endophallus can be partly or totally with naturally everted endophallus [Danilevsky, 1987, fused. We distinguish two main endophallus divisions, 1988a, b; Danilevsky, Dzhavelidze, 1990], were pub- which are very distinct in all groups of Dorcadionini. lished. M. Kuboki [1980, 1981] described everted en- Basal phallomer bp (consists of basal tube bt, dophallus of some Lepturinae as well as S.W. Lin- medial tube mt, central trunk ct and preapical gafelter and R.E. Hoebeke [2002] studied everted Ano- bulb pb) and apical phallomer ap (consists of plophora Hope, 1839 endophallus. Both investigations apical bulb ab and apical bubble bb) with did not use the inflation of endophallic tube, so its