Abstract
The Chorthippus biguttulus group distributed in the west Palaearctic, while intensively examined in Europe, is poorly known in the glacial refugia such as Anatolia. This produces constraints in making accurate statements about evolution and the biogeography of the group. The C. brunneus subgroup of this lineage is examined using large amounts of morphological and song data from Anatolia (Asian Turkey) and representatives from Europe. Song and morphology in combination suggested three species to be found in Anatolia. The first is C. bornhalmi Harz which is also known from south-east Europe. The other two are new species: Chorthippus antecessor sp. n. and Chorthippus relicticus sp. n.. Morphologically, C. antecessor sp. n. is the most aberrant species of the C. brunneus subgroup, but is similar to C. bornhalmi in song. The specific song and morphology (the aberrant number of stridulatory pegs) define C. relicticus as a new species and both also indicate that it is closely related to C. brunneus and C. jacobsi. A song and morphology based phyloylogenetic assumption for C. brunneus subgroup suggests C. antecessor, C. bornhalmi and C. miramae to constitute one clade and C. brunneus, C. jacobsi and C. relicticus another. The scenario suggested for their evolution assume the following steps: (i) divergence of C. bornhalmi from a C. antecessor like ancestor, (ii) derivation of an ancestral population (which later give rise to C. brunneus + C. jacobsi + C. relicticus) from a C. bornhalmi like ancestor, and (iii) later fragmentation of this ancestral population to result in the present three species (C. brunneus + C. jacobsi + C. relicticus). All of these events seem to be correlated with the climatic cycles during Pleistocene. The conclusion is that the two new species are range-restricted, vulnerable species as is the case for many other taxa present in the Mediterranean Taurus biodiversity hotspot.References
Bailey, R.I. Thomas, C.D. & Butlin, R.K. (2004). Premating barriers to gene Exchange and their implications for the structure of a mosaic hybrid zone between Chorthippus brunneus and C. jacobsi (Orthoptera: Acrididae). Journal of Evolutionary Biology 17, 108–119.
Bei-Bienko, G.J. & Mistshenko, L.L. (1951). The grasshopper of the fauna of the USSR and adjacent countries. Academy Nauk., Moskova-Leningrad. Volume II. 667 pp.
Beharav, A. & Nevo, E. (2003). Predictive validity of discriminant analysis for genetic data. Genetica 119, 259–267.
Bridle, J.R. Saldamando, C.I. Koning, W. & Butlin, R.K. (2006). Assortative preferences and discrimination by females against hybrid male song in the grasshoppers Chorthippus brunneus and Chorthippus jacobsi (Orthoptera: Acrididae). Journal of Evolutionary Biology, 19, 1248–1256.
Bukhvalova, M.A. (1995). Acoustic signals and morphological features of some grasshoppers of the Chorthippus biguttulus group (Orthoptera, Acrididae) of Russia and adjacent territories. Entomological Review, 74, 56–67.
Challis, R.J. Mutun, S. Nieves-Aldrey, J.L. Preuss, S. Rokas, A. Aebi, A. Sadeghi, E. Tavakoli, M. & Stone, G.N. (2007). Longitudinal range expansion and cryptic eastern species in the western Palaearctic oak gallwasp, Andricus coriarius. Molecular Ecology, 16(10), 2103–2114.
Ciplak, B. (2003). Distribution of Tettigoniinae (Orthoptera, Tettigoniidae) bush-crickets in Turkey: the importance of the Anatolian Taurus Mountains in biodiversity and implications for conservation. Biodiversity and Conservation, 12, 47–64.
Ciplak, B. (2004a). Biogeography of Anatolia: the marker group Orthoptera. Memorie della Societa Entomologica Italian, 82, 357–372.
Ciplak, B. (2004b). Systematics, phylogeny and biogeography of Anterastes (Orthoptera, Tettigoniidae, Tettigoniinae): evolution within a refugium. Zoologica Scripta, 33, 19–44.
Ciplak, B. (2008). The analogy between glacial cycles and global warming for the glacial relicts in a refugium: a biogeographic perspective for conservation of Anatolian Orthoptera. In: Fattorini, S. (Ed), Insect Ecology and Conservation (Chapter 6). Research Signpost. Fort P.O., Trivandrum, Kerala, India, pp. 135–163.
Ciplak, B. & Demirsoy, A. (1996). Caelifera (Orthoptera, Insecta) alttakımının Türkiye'de endemizm durumu. Turkish Journal of Zoology, 20, 241–246.
Ciplak, B. Yalım, B. & Demirsoy, A. (1999). Türkiye Orthoptera (=Düzkanatlılar=Çekirge) Faunası. In: Demirsoy, A. (Ed): Genel Zoocoğrafya ve Türkiye Zoocoğrafyası, 2nd ed. Meteksan Yayınları, Ankara. pp 748–772.
Ciplak, B. Heller, K.G. & Willemse, F. (2009). Review of the genus Eupholidoptera (Orthoptera, Tettigoniidae): different genitalia, uniform song. Zootaxa (in pres).
Cooper, S.J.B. Ibrahim, K.M. & Hewitt, G.M. (1995). Postglacial expansion and genome subdivision in the European grasshopper Chorthippus parallelus. Molecular Ecology, 4, 49–60.
Demirsoy, A. (1977). Türkiye Caelifera (Insecta, Orthoptera) faunasının tespiti ve taksonomik olarak incelenmesi. Atatürk Üniversitesi Basımevi, Erzurum, Türkiye, 252 pp.
Elsner, N. (1974). Neuroethology of sound production in gomphocerine grasshoppers (Orthoptera: Acrididae) I. Song patterns and stridulatory movements. Journal of Comparative Physiology, 88,67–102.
Gottsberger, B. & Mayer, F. (2007). Behavioral sterility of hybrid males in acoustically communicating grasshoppers (Acrididae, Gomphocerinae). Journal of Comparative Physiology A, 193, 703–714.
Gunduz, I. Jaarola, M. Tez, C. Yeniyurt, C. Polly, P.D. & Searle, J.B. (2007). Multigenic and morphometric differentiation of grounds quirrels (Spermophilus, Scuiridae, Rodentia) in Turkey. Molecular Genetics and Evolution, 43, 916–935.
Hanzelova, V. Kuchta, R. Scholz, T. & Shinn A.P. (2005). Morphometric analysis of four species of Eubothrium (Cestoda: Pseudophyllidae) parasites of salmonid fish: An interspecific and intrasepecific comparision. Parasitology International, 54, 207–214.
Harz. (1971). Orthopterologische Beitrage. Atalanta 3, pp. 331–338.
Harz, K. (1975). The Orthoptera of Europe. Göttingen, The Hague. Volume II. 749 pp.
Heller, K.G. (2006) Song evolution and speciation in bushcrickets. In: Drosopoulos, S. & Claridge, M.F. (Eds): Insect sound and communication. Taylor & Francis, Boca Raton, US.. pp. 137–153.
Helversen, D. von & Helversen, O. von. (1975a). Verhaltensgenetische Untersuchungen am akustischen Kommunikationssystem der Feldheuschrecken (Orthoptera, Acrididae) I. Der Gesang von Artbastarden zwischen Chorthippus biguttulus und Ch. mollis. Journal of Comparative Physiology, 104, 273–299.
Helversen, D. von & Helversen, O. von. (1975b). Verhaltensgenetische Untersuchungen am akustischen Kommunikationssystem der Feldheuschrecken (Orthoptera, Acrididae) II. Das Lautschema von Artbastarden zwischen Chorthippus biguttulus und Ch. mollis. Journal of Comparative Physiology, 104, 301–323
Helversen, O.von & Elsner, N. (1977). The stridulatory movements of acridid grasshoppers recorded with an opto-electronic device. Journal of Comparative Physiology, 122, 53–64.
Helversen O. v. and Helversen D. v. (1994). Forces driving coevolution of song and song recognition in grasshoppers. Fortschritte der Zoologie, 39, 253–284.
Hewitt, G.M. (1996). Some genetic consequence of ice ages, and their role in diverging and speciation. Biological Journal of Linnaean Society, 58, 247–276.
Hewitt, G. (1999). Post-galacial re-colonisation of European biota. Biological Journal of Linnean Society, 68, 87–112.
Hewitt, G.M. (2000). The genetic legacy of the Quarternar ice ages. Nature, 405, 907–913.
Hill, T. & Lewicki, P. (2005). Statistics: Methods and Aplications. StatSoft, Inc, 800 pp.
Ingrisch, S. (1995). Evolution of the Chorthippus biguttulus group (Orthoptera, Acrididae) in the Alps, based on morphology and stridulation. Revue Suisse de Zoologie, 102, 475–535.
Izenman (2008). Modern multivariate statistical techniques. Springer Science and Business Media, New York, 731 pp.
Kalayci, S. (2005). Faktör Analizi ve Ayırma (Discriminant) Analizi, In: S. Kalayci (Ed.), SPSS Uygulamalı Çok Değişkenli İstatistik Teknikleri, Asil Yayın Dağıtım Limited Sirketi Ankara, pp. 321–344,.
Karabağ, T. (1958). A synonymic and distributional catalogue of Turkish Orthoptera (Türkiye’nin Orthoptera faunası). Ankara Üniversitesi Fen Fakültesi Yayınları Umumi: 81, Zooloji 4, Ankara. (in Turkish and English). 198 pp.
Klappert, K. & Reinhold, K. (2007). Indirect benefits for choosy female grasshoppers (Chorthippus biguttulus)? Zoology, 110(5), 354–359.
Kleukers, R. Odé, B. & Willemse, F. (2004). Hybridization of Glyptobothrus brunneus and Glyptobothrus bornhalmi in northeastern Italy. Memorie della Societa Entomologica Italian, 82, 547–556.
Mason, D.J. Butlin, R.K. & Gacesa, P. (1995). An unusual mitochondrial DNA polymorphism in the Chorthippus biguttulus species group (Orthoptera: Acrididae). Molecular Ecology, 4, 121–126.
Mayr, E. (1982). Systematics and the origin of species. Columbia University Press. New York. 334 pp.
Mclachlan, G.J. (2004). Discriminant analysis and statistical pattern recognition. John Wiley & Sons, Inc. New Jersey, 526 pp.
Médail, F. & Quézel, P. (1997). Hot-spots analysis for conservation of plant biodiversity in the Mediterranean basin. Annals of Missouri Botanical Garden, 84, 112–127.
Mol, A. Ciplak, B. & Sirin, D. (2003). Song and morphology of the three related Anatolian endemic species of genus Chorthippus (Orthoptera, Acrididae, Gomphocerinae): C. ilkazi, C. bozdaghi and C. helverseni sp. n.. Annales de la société entomologique de France 39, 121–128.
Perdeck, A.C. (1957). The isolating value of specific song patterns in two sibling species of grasshoppers (Chorthippus brunneus Thunb. and C. biguttulus L.). E.J. Brill, Leiden. 75 pp.
Radloff, S.E., Hepburn, R. & Fuchs, S. (2005). The Morphometric affinities of Apis cerana of the Hindu Kush and Himalayan regions of western Asia. Apidologie, 36, 25–30.
Ragge, D.R. (1976). A putative hybrid in nature between Chorthippus brunneus and C. biguttulus (Orthoptera: Acrididae). Systematic Entomology, 1, 71–74.
Ragge, D.R. & Reynolds, W.J. (1988). The songs and taxonomy of the grasshoppers of the Chorthippus biguttulus group in the Iberian Peninsula (Orthoptera: Acrididae). Journal of Natural History, 22, 897–929
Ragge, D. R. & Reynolds, W.J. (1998). The songs of the grasshoppers and crickets of western Europe. Harley Books, London. 221 pp.
Ragge, D.R. Reynolds, W.J. & Willemse, F. (1990). The songs of the European grasshoppers of the Chorthippus biguttulus group in relation to their taxonomy, speciation and biogeography. Bolletin de Sanidad Vegetal Plagas, 20, 239–245.
Reinhold, K. Reinhold K. & Jacoby K.J. (2002). Dissecting the repeatability of female choice in the grasshopper Chorthippus biguttulus. Animal Behaviour, 64, 245–250.
Saldamando, C.I. Miyaguchi, S. Tatsuta, H. Kishino, H. Bridle, J.R. & Butlin, R.K. (2005a). Inheritance of song and stridulatory peg number divergence between Chorthippus brunneus and C. jacobsi, two naturally hybridizing grasshopper species (Orthoptera: Acrididae). Journal of Evolutionary Biology, 18, 703–712.
Saldamando, C.I. Tatsuta, H. & Butlin, R.K. (2005b). Hybrids between Chorthippus brunneus and C. jacobsi (Orthoptera: Acrididae) do not show endogenous postzygotic isolation. Biological Journal of Linnaean Society, 84, 195–203.
Salman, S. (1978). Ağrı, Kars ve Artvin illerinin Orthoptera (Insecta) faunası üzerine taksonomik araştırmalar. Atatürk Üniversitesi Fen Fakültesi Yayınları No: 82, Erzurum, Türkiye. 184 pp.
Schmidt, G.H. (1990). Notes on the Chorthippus (Glyptobothrus) species (Orthoptera: Acrididae) in Greece and the calling songs of their males. Bolletin de Sanidad Vegetal Plagas, 20, 247–253.
Sevgili, H. & Ciplak, B. (2000). The Orthoptera of Şanlıurfa province from the Mesopotomian part of the Turkey. Italian Journal of Zoology, 67, 229–240.
Sychyov, M.M. (1996). Taxanomic structure of the biguttulus-group of the genus Chorthippus (Orthoptera Acrididae Gomphocerinae) and phylogenetic interrelations within the group. Russian Entomological Journal, 5, 3–11.
Taberlet, P. Fumagalli, L. Wust-Saucy, A. G. & Cosson, J. F. (1998). Comparative phylogeography and postglacial colonization routes in Europe. Molecular Ecology, 7,453–464.
Tarknisvili, D.N. Torphe, R.S. & Arntzen, J.W. (2000). Pre-Pleistocene refugia and differantiation between populations of the Caucasian salamander (Mertensiella caucasica). Molecular Phylogenetics and Evolution, 14, 414–422.
Ustinova, J. & Mayer, F. (2006). Alternative starts of transcription, several paralogues, and almost-fixed interspecific differences of the gene fruitless in a hemimetabolous insect. Journal of Molecular Evolution, 63(6), 788–800.
Veith, M. Schmidtler, J.F. Kosuch, J. Baran, I. & Seitz, A. (2003) Palaeoclimatic changes explain Anatolian mountain frog evolution: a test for alternating vicariance and dispersal events. Molecular Ecology, 12(1), 185–199.
Yalim, B. & Ciplak, B. (2002). Termessos Milli Parki (Antalya) Orthoptera (Insecta) faunasi: Fauna elemenlarinin zoocografyalari ve vejetasyona gore dagilislari. Türkiye Entomoloji Dergisi, 26, 267–276.
Weidner, H. (1969). Beiträge zur Kenntnis der Feldheuschrecken (Caelifera) Anatoliens. Mitteilungen Hamburger Zoologisches Museum und Institut, 66, 145–226.
Willemse, F. (1984). Catalogue of the Orthoptera of Greece: Fauna Graeciae I. Helenic Zoological Society, Athens, Greece. 275 pp.
Willemse, F. (1985). A key to the Orthoptera species of Greece: Fauna Graeciae II. Helenic Zoological Society, Athens, Greece. 288 pp.
Willemse, F. & Willemse L. (2008). An annotated checklist of the Orthoptera-Saltatoria from Greece including an updated bibliography. Articulata Beiheft, 13, 1–90.
Willemse, F.M.H., von Helversen, O. & Ode, B. (2009). A review of Chorthippus species with angled pronotal lateral carinae from Greece with special reference to transitional populations between some Peloponnesean taxa (Orthoptera, Acrididae). Zoologische Mededelingen, 83(2), 319–507.