Battal Çiplak

Systematics, phylogeny and biogeography of Anterastes (Orthoptera, Tettigoniidae, Tettigoniinae): evolution within a refugium

Çıplak, B. (2004). Systematics, phylogeny and biogeography of Anterastes (Orthoptera, Tettigoniidae, Tettigoniinae): evolution within a refugium. —Zoologica Scripta, 33, 19–44.

FRUIT MORPHOLOGY OF SECTIONS OF THE GENUS ONOBRYCHIS MILLER (FABACEAE) AND ITS PHYLOGENETIC IMPLICATIONS

ABSTRACT In this study, fruit morphology of sections of the genus Onobrychis were examined. Fruit morphology origin, taxonomic position, and phylogeny of the genus are discussed. The hypothesis obtained from a phylogenetic analysis based on fruit morphology suggests the polyphyly of the genus excluding section Dendrobrychis. However, the result supports monophyly of four generic sections, Hymenobrychis + (Heliobrychis + (Lophobrychis + Onobrychis)).

Altitudinal size clines, species richness and population density: case studies in Orthoptera

Abstract We examined size clines for various external body structures in two grasshopper species (Chorthippus vagans and Oedipoda miniata) and one katydid species (Poecilimon birandi), along an altitudinal gradient ranging from sea level to 1,980 m in southwest Anatolia (Turkey). In O. miniata and P. birandi all measured structures (body, tegmina, pronotum, and hind femur) were smallest at the highest altitude for both sexes. Hence, in these species, different structures covaried in the same direction, with increasing altitude. In contrast, there was no clear relationship between altitude and size in C. vagans, and different structures covaried in different directions in males and females (i.e., some structures became larger at higher elevations, whereas others became smaller, and this differed with sex). For some other C. vagans traits, there was no significant intraspecies variation. Hence, O. miniata and P. birandi followed the converse Bergmanns Rule, whereas there was no consistent pattern for C. vagans. For C. vagans, small body size was not associated with either local population density (measured as relative abundance) or local species richness, suggesting that neither intra- nor interspecific competition determined body size for this species. For O. miniata, the smallest individuals were found at the site with the highest grasshopper diversity, suggesting that interspecific competition could have influenced O. miniata body sizes at this altitude. However, the largest O. miniata individuals generally occurred at the sites with the highest relative abundance of O miniata. Hence, in this species, body sizes were generally larger at sites with high population densities — consistent with sites that were most favorable and so able to support high densities of O miniata also producing the largest individuals of this species. Observational data on P. birandi similarly suggested that, for this katydid, there is little relationship between local diversity/abundance and altitudinal size-clines.

Systematics and phylogeny of Parapholidoptera (Orthoptera: Tettigoniidae: Tettigoniinae)

Genus Parapholidoptera is revised and three species, P. yoruka sp.n., P. bolkarensis sp.n. and P. salmani sp.n., are described. Parapholidoptera delineata Stolyarov is placed in synonymy with P. ziganensis Karabağ. Parapholidoptera bodenheimeri Karabağ and P. distincta bodenheimeri Karabağ are placed in synonymy with P. distincta (Uvarov). Parapholidoptera flexuosa Karabağ, previously a subspecies of P. castaneoviridis (Brunner von Wattenwyl), is recognized as a separate species. A key to world species is provided. Cladistic analysis revealed the monophyly of the genus with two major clades. A short account of distribution is presented.

Phylogeny and biogeography Eupholidoptera Mařan (Orthoptera, Tettigoniidae): morphological speciation in correlation with the geographical evolution of the eastern Mediterranean

Recently, the systematics and biogeography of the Mediterranean biota have received much attention. This paper deals with Eupholidoptera Mařan, a Mediterranean lineage of Tettigoniidae. The genus is restricted to the northern and eastern basin of the Mediterranean, with a significant number of species found on the Aegean islands. To produce a phylogeny and use it to make assumptions about the historical biogeography of Eupholidoptera, material of 46 species from several collections was studied. A phylogenetic analysis based mainly on morphological characters suggested two lineages in the genus: the E. chabrieri and the E. prasina groups. Based on the consistency between historical geographical events and branching events on the phylogenetic tree, Eupholidoptera is assumed to have evolved from an ancestor present in the Aegeid plate in the Mid‐Miocene. The division of the Aegeid plate into Anatolia and Greece in the Tortonian, the reoccurrence of terrestrial corridors between these mainlands in the Messinian, the regression of the Aegean area in the Pliocene and sea level changes in the Pleistocene are assumed to have been the main palaeogeographical events directing speciation in Eupholidoptera. As most of the species are allopatric, vicariance is suggested to be the main pattern. By combining the nature of the characters used in the phylogenetic analysis, the phylogenetic tree produced and the biogeographical assumptions, four tentative conclusions can be made: (i) radiation in the genus is a result of divergence in morphology; (ii) because the main character source is male genitalia, there has possibly been intensive sexual selection, which leads to morphological speciation; (iii) as the difference in temporal parameters of the song is prominent in sympatric/parapatric species pairs only, co‐occurrence is suggested to be the main reason driving divergence in the song; (iv) there seems to be a negative correlation between the size of the distribution range and the evolutionary rate in speciation; this may be the reason why the E. prasina group (restricted to a small part of the range of the genus) is more diverse than the E. chabrieri group, which is distributed over the entire range.

Phylogeography of Anterastes serbicus Species Group (Orthoptera, Tettigoniidae): Phylogroups Correlate with Mountain Belts, but not with the Morphospecies

Abstract Ten species of the genus Anterastes Brunner von Wattenwyl (Orthoptera, Tettigoniidae) show insular distribution in mountain meadows of Anatolia and the Balkans. Current understanding of the taxonomy and species relationships within the genus is based on morphological characters. However, the extent to which morphological characters are phylogenetically informative, when used to define taxonomic groups or to elucidate detailed evolutionary relationships within Anterastes, is in need of further examination. Moreover, because little is known about the historical biogeography and diversification factors in members of this genus, additional datasets are necessary to test the robustness of species, relationship hypotheses and associated biogeographic patterns, Here we specifically examined, using 16S rDNA sequences, the evolutionary relationships and species boundaries of three closely related species of Anterastes (i.e., the A. serbicus group, comprising A. serbicus, A. burri and A. antitauricus). Additionally A. tolunayi, a species not in the A. serbicus group, but morphologically very similar to members of the group, was included in the molecular analysis to locate the species of the A. serbicus complex within a phylogenetic frame. Hence, the phylogenetic relationships and taxonomic interpretation of the species complex appear more intricate than previously hypothesized, The current molecular data do not allow us to identify A. serbicus, A. burri and A. antitauricus as distinct phylogenetic species, but rather suggest that these morphospecies are themselves a complex of cryptic taxa. Despite the incongruencies among the phylogenetic trees and nonmonophyly of each the three morphospecies, the median joining network resulted in haplotype grouping consisting of four clusters that are definable by geography. Thus, based on the congruency between geography and gene clusters, and the molecular clock estimate, it can be interpreted that 1) a strong correlation between the radiation of the group and the topography of their ranges may exist, 2) the radiation of the group dates back to Late Pliocene or Early Pleistocene and 3) there is a break between the Anatolian and the European lineages, in respect to range change of cold-preferring forms, dating back prior to the last four glacial periods.

The Orthoptera of Şanliurfa province from the Mesopotamian part of Turkey

Abstract Previous records and 1920 newly collected specimens formed the basis of this survey of the Orthoptera of Şanliurfa (Urfa) province in southeastern Turkey. Eighty‐six species and subspecies were recorded, both as the result of the field study and use of published data; they belong to seven families and fifty genera. Eleven genera and thirty‐two species were recorded for the first time from Şanliurfa. Seven species were recorded as new for the southeastern Anatolia. Eleven species with subspecies are endemic to Anatolia. Eremial faunal elements are dominant in the area.

Review and key to species of Platycleis from Turkey (Orthoptera: Tettigoniidae) with descriptions of Yalvaciana subgen. n. and two new species

Species of Platycleis from Turkey are reviewed. Twenty-three species are recorded: Platycleis affinis (Fieber, 1853), P. albopunctata (Goeze, 1778), P. escalerai (Bolívar, 1899), P. intermedia (Serville, 1838), P. incerta Brunner von Wattenwyl, 1882, P. persica Uvarov, 1917, P. ankarensis (Karabaǥ, 1950), P. armeniaca Ramme, 1951, P. elegans (Uvarov, 1934), P. schereri Werner, 1901, P. taurica Bolívar, 1899, P. uvarovi Karabağ, 1950, P. sepium (Yersin, 1854), P. sporadarum (Werner, 1933), P. kurmana (Ramme, 1951), P. melendisensis sp. n., P. sinuata Ramme, 1951, P. salmani sp. n., P. weidneri (Demirsoy, 1974), P. nigrosignata (Costa, 1863), P. tessellata holoptera Ramme, 1951, P. veyseli Koçak, 1984, P. yalvaci (Demirsoy, 1974). Keys to subgenera and species, necessary figures and distribution maps are provided. A new subgenus Yalvaciana subgen. n. is established to include Decorana yalvaci Demirsoy. Two new species, P. salmani sp. n. and P. melendisensis sp. n. are described. Montana bifoveolata Karabağ syn. n. is placed in synonymy with Platycleis schereri Werner. Metrioptera brevipes Uvarov is placed in synonymy with P. sporadarum Werner in subgenus Sporadiana Zeuner. Platycleis (Squamiana) sinuata Ramme, 1951 is transferred from subgenus Platycleis. Song oscillograms from P. armeniaca and P. taurica are presented. Platycleis (Montana) armeniaca Ramme is a new record for Turkey.

Mountainous genus Anterastes (Orthoptera, Tettigoniidae): autochthonous survival across several glacial ages via vertical range shifts

Although the high‐latitude range margins in Europe and North America are intensively studied, attention is gradually turned towards the taxa/populations inhabiting glacial refugia. Here, we evaluate the genealogical history of the cold‐adapted Anatolio‐Balkan genus Anterastes especially to test the possible effects of intrarefugial vertical range shifts during climatic oscillations of the Quaternary. Using concatenated data from sequences of COI+16S and ITS1–5.8S–ITS2, intrageneric relationships and the time of speciation events were estimated. Thirteen different demographic analyses were performed using a data set produced from sequences of 16S. Different phylogenetic analyses recovered similar lineages with high resolution. The molecular chronogram estimated speciation events in a period ranging from 5.60 to 1.22 Myr. Demographic analyses applied to 13 populations and five lineages suggested constant population size. Genetic diversity is significantly reduced in a few populations, while not in others. Fixation indices suggested extremely diverged populations. In the light of these data, the following main conclusions were raised: (i) although glacial refugia are the biodiversity hotspots, species level radiation of the cold‐adapted lineages is mainly prior to the Mid‐Pleistocene transition; (ii) heterogeneous topography provides refugial habitats and allows populations to survive through vertical range shifts during climatic fluctuations; (iii) prolonged isolation of refugial populations do not always result in reduced intrapopulation diversity, but in high level of genetic differentiation; (iv) the cold‐adapted lineages with low dispersal ability might have not colonised the area out of Anatolian refugium during interglacial periods; and (v) populations of invertebrates may have restricted ranges, but this does not mean that they have small effective population size.

Estimating effects of global warming from past range changes for cold demanding refugial taxa: A case study on South-west Anatolian species Poecilimon birandi

Although changes in biodiversity and in ecosystems are surely caused by a range of interacting drivers, such as natural or human-induced factors, one of the important drivers having major impacts on climate and biodiversity and leading to range changes and fragmentation is global warming. Defining past range changes/fragmentations during interglacial periods may provide tools to understand possible impacts of global warming on present biodiversity. To test this assumption we studied a marker gene in the bush-cricket Poecilimon birandi, a species confined to South-west Anatolia that demands a cold climate. Haplotypes of P. birandi constituted three main phylogroups,West, East and Demre. All haplotypes are unique to the respective phylogroup. An AMOVA suggested considerable divergence at all hierarchical levels. Though there is a strong isolation between phylogroups, the East and West groups harbour considerable haplotype diversity. Most of the demographic analyses suggest stable historical populations for the West and East phylogroups, but a coalescent-based demographic analysis indicates a bottleneck for the West phylogroup. The main conclusions are; (i) P. birandi contains considerable phylogenetic signal in 16S rDNA, (ii) there were at least three contemporaneous radiations, which might have originated from isolated refugial populations during Pleistocene, (iii) within a refugium, range changes induced by climatic shifts may be only vertical through an altitudinal gradient, (iv) significant genetic structure can arise in a small heterogeneous area, if the species requires particular habitats and has weak dispersal ability, (v) climatic shifts may cause fragmentation or extinction of populations, but can also lead to divergence of populations suffering from fragmentation, and (vi) altitudinal heterogeneity plays a buffering role, allowing for survival of the refugial biodiversity.