Revision of the Poecilimon Ornatus Group (Orthoptera: Phaneropteridae) with Particular Reference to the Taxa in Bulgaria and Macedonia*
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Eur. J. Entomol. 107: 647–672, 2010 http://www.eje.cz/scripts/viewabstract.php?abstract=1575 ISSN 1210-5759 (print), 1802-8829 (online) Revision of the Poecilimon ornatus group (Orthoptera: Phaneropteridae) with particular reference to the taxa in Bulgaria and Macedonia* DRAGAN P. CHOBANOV 1 and KLAUS-GERHARD HELLER2 1 Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 1 Tsar Osvoboditel blvd., 1000 Sofia, Bulgaria; e-mail: [email protected] 2Grillenstieg 18, 39120 Magdeburg, Germany; e-mail: [email protected] Key words. Orthoptera, Phaneropteridae, Poecilimon, P. jablanicensis sp. n., systematics, bioacoustics, morphology, communication system, phylogeny, evolutionary trends, Balkan Peninsula Abstract. The Poecilimon ornatus group has an exclusively European distribution and includes the largest species in the genus. A revision of the taxa belonging to this group in Bulgaria and Macedonia (Central and Eastern Balkan Peninsula) is presented. Nine taxa described from Bulgaria are synonymised with 3 previously known species, as follows: Poecilimon ornatus (= P. mistshenkoi marzani, syn. n., P. mistshenkoi tinkae, syn. n., P. mistshenkoi vlachinensis, syn. n.), P. affinis s. str. (= P. mistshenkoi mistshenkoi, syn. n., P. affinis ruenensis, syn. n., P. affinis rilensis, syn. n., P. affinis medimontanus, syn. n., P. harzi, syn. n.) and P. hoelzeli (= P. kisi, syn. n.). The synonymy of P. poecilus with P. affinis and the subspecific status of P. affinis komareki are confirmed. One species, Poecilimon jablanicensis, sp. n., is described as new to science. A tabulated key, lists and maps of all known localities and oscillograms of the songs of all the species in this group are presented. The phylogenetic relationships and evolutionary trends in the Poecilimon ornatus group are discussed. INTRODUCTION by large or moderate body size, posteriorly widened and more or less upcurved pronotum, well visible tegmina and Poecilimon Fischer, 1853 is a group of short-winged moderately incurved cerci with a single tooth. The Poe- herbivorous bushcrickets, distributed from the Apennines cilimon ornatus-group after Heller (1984), consists of 8 to Eastern Siberia and Central Tien-Schan (Bey-Bienko, taxa, namely P. nobilis Brunner von Wattenwyl, 1878, P. 1954), and the largest genus of the suborder Ensifera in obesus obesus Brunner von Wattenwyl, 1878, P. obesus the Palaearctic. Its centre of speciation and diversity, artedentatus Heller, 1984, P. affinis affinis (Frivaldszky, however, is restricted to the Pontic region with only a few 1867), P. affinis komareki ýejchan, 1957, P. affinis hoel- taxa occuring outside the Balkans, Asia Minor and the zeli Harz, 1966, P. ornatus (Schmidt, 1850) and P. pan- Caucasus region. They have poor dispersal ability due to cici Karaman, 1958, distributed mainly in the Balkans. their microptery and this along with the diverse geomor- Later on, P. artedentatus and P. hoelzeli was given spe- phological and climatic aspects and history of the area cific status (Willemse, 1985; Willemse & Heller, 1992), where they occur has resulted in rapid morphological and P. pancici synonymised (Willemse, 1985) and a few new behavioural evolution. Within the genus there are many taxa, somewhat resembling the latter species, described groups of apparently related taxa. Despite the few full or (P. pindos F. Willemse, 1982, P. soulion L. Willemse, partial revisions (e.g. Ramme, 1933; Bey-Bienko, 1954; 1987 and P. gracilioides F. Willemse et Heller, 1992). Willemse, 1982; Heller, 1984; Heller & Lehmann, 2004; The last species connect the group morphologically with Heller & Sevgili, 2005; Heller et al., 2006; Heller et al., P. gracilis (Fieber, 1853). P. schmidtii (Fieber, 1853), 2008) the phylogeny and systematics of Poecilimon is forming together with P. gracilis the second section of only partly resolved. Even a recent partial review of the Ramme’s (1933) group I, is not closely related to the genus based on molecular data (Ullrich et al., 2010) has others (e.g. Ullrich et al., 2010) and is not included in this not clarified all questions. study. In this group, however, the following 9 taxa, One group for which the phylogeny and systematics is described by Peshev (1980) from Bulgaria should be poorly known is the Poecilimon ornatus group. The spe- included: P. mistshenkoi mistshenkoi, P. mistshenkoi cies belonging here were listed in group I (“Gruppe I”; marani, P. mistshenkoi tinkae, P. mistshenkoi vlachinen- containing two sections) by Ramme (1933) in the first sis, P. affinis ruenensis, P. affinis rilensis, P. affinis medi- revision of the genus. The group’s taxa are characterized * While this paper was in press, a study by Ingrisch and Pavicevic [2010; Seven new Tettigoniidae (Orthoptera) and a new Blattel- lidae (Blattodea) from the Durmitor area of Montenegro with notes on previously known taxa. Zootaxa 2565: 1–41] was published, also dealing with taxa of the P. ornatus group. The authors describe some new forms and change the rank of others. In contrast to the ideas presented in that paper we still consider the form P. rumijae as an infrasubspecific variation of P. affinis komareki. Judging from the extent of the differences we do not exclude the possibility that some of the newly described forms are local variants of P. affinis (compare with the morphological and bioacoustic variation described here). 647 montanus, P. kisi and P. harzi. These forms, though and 4135 microphones (frequency response flat up to 40, described in detail, lack a well defined differential diag- respectively 70 kHz), were used. nosis and their taxonomic status and systematic relation- Bioacoustic terminology: calling song – the song produced by ships are still doubtful. After studying the type material an isolated male; syllable – the sound produced by one opening- and-closing movement of the tegmina; syllable duration – the and investigating the morphology, behaviour and bioa- time measured from the beginning of the first to the end of last coustics of topotypic populations, some changes in the impulse in a syllable; syllable period – the span including a syl- systematics and taxonomy of the group from Bulgaria lable and the following interval; impulse – a undivided transient became necessary. These taxa are the subject of the pre- train of sound waves produced by the stridulatory tooth striking sent work. The rest of the species in this group are the plectrum (the anal edge of the opposite tegmen). restricted to Greece. Their distribution and systematic Appendix with detailed distribution of individual species is position have been studied by Willemse and co-authors available online at: (Willemse, 1982, 1985; Willemse & Heller, 1992) and <http://www.eje.cz/supplfiles/eje1575suppl.pdf> only the most important facts are repeated here for com- RESULTS parison. Due to the lack of song recordings for the two subspecies of P. affinis described by Karaman (1974), P. A. Species of the Poecilimon ornatus group occurring affinis serbicus Karaman, 1974 and P. affinis hajlensis in Bulgaria and Macedonia Karaman, 1974, these taxa are not discussed in detail. In All species in this group on the Central and Eastern addition, during field studies, a new taxon from the Poe- Balkan Peninsula and adjacent territories in the North fall cilimon ornatus group was discovered. While all other into two groups: either large and bulky animals or small species of the group have a bidirectional sound- and slender ones (see Table 2). The first group can mor- communication system (ancestral condition in Poeci- phologically (and by song) be separated into three sub- limon), this species possesses a unidirectional acoustic groups, which are considered here to be three separate communication system (loss of female tegmina together species (although they rarely occur sympatrically or syn- with stridulatory organs). This new species, Poecilimon topically), P. ornatus, P. affinis and P. hoelzeli. The jablanicensis, sp. n., is described and some conclusions second is made up of two distinct species, one widespread about the evolutionary trends in the group are presented (P. gracilis) and one very local, which is a new species. in this paper. Poecilimon ornatus (Schmidt, 1850) MATERIAL AND METHODS Ephippigera ornata Schmidt, 1850 (184), type locality Slovenia [northern part = S Kärnten, Kamnik (see Willemse, 1985)]. During the present study material from several collections Barbitistes fieberi Fieber, 1853 (175), type locality Italy, were surveyed, including the National Museum of Natural His- Trieste; synonymized with P. ornatus by Krauss (1878); tory (at that time containing also the former collection of Brunner von Wattenwyl (1878, 1882) used the younger name Orthoptera of the Institute of Zoology), Bulgarian Academy of fieberi including P. ornatus. Sciences, Sofia; collection of the Natural History Section of the Odontura ornata; Schmidt, 1866 (81). Regional History Museum in Blagoevgrad; collection of the Poecilimon ornatus; Krauss, 1878 (497); Ramme, 1933 (511, Pl. Macedonian Museum of Natural History, Skopje; collection of 6, Fig. 2., Pl. 7, Fig. 1, Pl. 8, Fig. 1, Pl. 11, Fig. 1, Pl. 12, Fig. Dragan Chobanov; collection of Klaus-Gerhard Heller, abbrevi- 1); Bey-Bienko, 1954 (270–271); Harz, 1969 (119–120, Figs ated as NMNH, HMB, MMNH, CC and CH, respectively. In 32, 268, 270, 271, 319, 322); Willemse, 1982 (partim; addition, material from some private collections was examined. 156–158, Figs 1, 2, 5, 9); Heller, 1984 (83, 85, Figs 5–D, Concerning the systematics and distribution of species only 7–E–G, 9–B); Willemse, 1985 (11–12, Figs 1–5, 34); Heller, references with original or supplementary information are pre- 1988 (50–51, Pls 34–G, 88, 120–D). sented. For each taxon a distributional map is prepared Poecilimon pancici Karaman, 1958 (36, Figs 1–7), type locality including both literature (localities revised when material avail- Macedonia, Gradishtanska Planina Mt., Gabresh vill.; Harz, able) and unpublished data. Data on altitude were added, if 1969 (122); Willemse, 1982 (158, Fig. 10). Synonym of P. missing, using the co-ordinates and the GPSVisualizer online ornatus after Heller (1988).