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The Phlesirtes Complex (Orthoptera, Tettigoniidae, Conocephalinae, Conocephalini) Reviewed: Integrating Morphological, Molecular, Chromosomal and Bioacoustic Data

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The Phlesirtes Complex (Orthoptera, Tettigoniidae, Conocephalinae, Conocephalini) Reviewed: Integrating Morphological, Molecular, Chromosomal and Bioacoustic Data Systematic Entomology (2010), DOI: 10.1111/j.1365-3113.2009.00512.x The Phlesirtes complex (Orthoptera, Tettigoniidae, Conocephalinae, Conocephalini) reviewed: integrating morphological, molecular, chromosomal and bioacoustic data CLAUDIA HEMP1, KLAUS-GERHARD HELLER2,SIEGFRIED KEHL1,ELZBIETA˙ WARCHAŁOWSKA-SLIWA´ 3, JOHANN WOLFGANG WAGELE¨ 4 and ANDREAS HEMP5 1Department of Animal Ecology II, University of Bayreuth, Bayreuth, Germany, 2Institute of Biology, University of Erlangen-Nurnberg,¨ Germany, 3Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow, Poland, 4Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany and 5Institute Biochemistry and Biology, University of Potsdam, Potsdam, Germany Abstract. The tettigoniid genus Phlesirtes Bolivar and its allies are reviewed. Morphological, ecological and molecular data prompt the erection of the new genus Chortoscirtes gen.n. with type species Xiphidion meruense Sjostedt.¨ The genera Phlesirtes, Chortoscirtes, Karniella and Naskreckiella are characterized by morphological characters supported by molecular, acoustic, ecological and chromosomal data. Four species, Chortoscirtes pseudomeruensis sp.n., C. masaicus sp.n., C. puguensis sp.n. and C. serengeti sp.n., are described from localities in northern and coastal Tanzania and one Karniella, K. crassicerca sp.n., is described from Uganda. The following comb n. are proposed: Phlesirtes kibonotensis (Sjostedt)¨ and Phlesirtes kilimandjaricus (Sjostedt).¨ Subtribal status is proposed for the four investigated African genera. A key to the Chortoscirtes species is provided. Introduction young volcanoes such as Mts Kilimanjaro and Meru or Mt Hanang (estimated ages of 1–2 Ma, Marek, 2001) also reveals Montane East Africa has high biotic diversity and endemism high diversity and endemism (Lambrechts et al., 2002; Hemp, (Lovett, 1988; Behangana et al., 2009). The Eastern Arc moun- 2006). The geological age of these higher volcanoes may allow tains, geologically ancient formations with an estimated age of calibration for speciation processes. With typical wide savanna 30 Ma (Burgess et al., 1998) and extending throughout Tan- plains and the climatically isolated mountains looming to con- zania and southern Kenya, are well investigated for some siderable heights, the area can serve as a model system to taxa (Howell, 1993; Emberton et al., 1997; Newmark, 2002; investigate mechanisms of evolution of tropical species diver- Robertson, 2002; Burgess et al., 2007; Poynton et al., 2007) sity. The flightless Saltatoria are ideal for investigating mech- and are regarded as hotspots of diversity and endemism. How- anisms to explain the biogeographical distribution patterns. Many genera show arrays of closely related species isolated ever, more intensive recent research on the high mountains of on the high mountains. The study of distribution patterns and the East African rift valley system, especially the geologically of the ecology, habitat demands, acoustics and molecular rela- Correspondence: Claudia Hemp, Department of Animal Ecology tionships of species groups contributes to an understanding of II, University of Bayreuth, D-95440 Bayreuth, Germany. E-mail: the climatic and vegetational history of East Africa. Such stud- [email protected] ies require basic taxonomic work because the fauna remains poorly known. Unpublished for the purposes of zoological nomenclature (Art. 8.2, That allopatric speciation generated the biogeographical pat- ICZN) terns found amongst some East African lentulid genera has © 2010 The Authors Journal compilation © 2010 The Royal Entomological Society 1 2 C. Hemp et al. been shown for example for Rhainopomma species (Hemp Table 1. Members of the Phlesirtes genus complex. et al., 2007; Schultz et al., 2007), a genus presently contain- ing seven species. All are restricted to the montane zones P. bilineatus Chopard, 1954 P. brachiatus Uvarov, 1923 of mountains along the Eastern Arc chain of Tanzania and P. kevani Chopard, 1954 southern Kenya, and all occupy similar ecological niches (for- P. kibonotensisa (Sjostedt,¨ 1909) est edge and clearings) at corresponding altitudes of differ- P. kilimandjaricus (Sjostedt,¨ 1909) ent mountains. The morphological similarity suggests a recent P. latifrons Chopard, 1954 radiation, probably as a result of past climatic fluctuations. The P. meruensisa (Sjostedt,¨ 1909) molecular phylogenetic analysis confirms that related species P. merumontanus (Sjostedt,¨ 1909) of Rhainopomma occur on neighbouring mountains (e.g. East K. bullata Rehn, 1914 N. kalamboi Unal,¨ 2005 and West Usambara, South Pare, North Pare) and that species are genetically and geographically separated. Another flight- aAccording to Bolivar’s nomenclature (Bolivar, 1922) the genus name less genus with an array of morphologically closely related Phlesirtes demands a masculine ending of the species name. Therefore species is the pyrgomorphid genus Parasphena Bolivar, with the species names ending with -se have to have the ending–is. 17 so-far known species found on almost every high moun- tain from southern Tanzania to northern Kenya and eastern Here we compile knowledge of the hitherto described Uganda. Species of Parasphena are restricted to grasslands of species of Phlesirtes, study their relationships, including the the montane and afroalpine zone. Morphology, ecology and description of a new genus, characterize the genera Karniella molecular data suggest allopatric speciation as the most plau- and Naskreckiella, and clarify the taxonomic position of these sible explanation for the biogeographical history (Rehn, 1942; genera within Conocephalinae. Kevan, 1948, 1956; Hemp et al., 2009). The Phlesirtes genus complex is a poorly investigated group Methods and materials of Conocephalinae bush crickets. Because of their small size and nymphal appearance they have been overlooked and poorly Depositories sampled since the erection of Phlesirtes by Bolivar (1922), for Xiphidion merumontanum Sjostedt¨ from northern Tanzania. MNB: Museum fur¨ Naturkunde, Zentralinstitut der New sampling in the past 15 years has revealed over 30 species Humboldt-Universitat¨ zu Berlin. NHML: Natural History in the highlands of East Africa. Museum London, U.K. EDNMK: Entomological Depart- Sjostedt¨ (1909) described four Xiphidion (Tettigonioidea: ment National Museums of Kenya, Nairobi. NHRS Stock- Conocephalinae) species (merumontanum, meruense, kilimand- holm: Naturhistoriska Riksmuseet, Stockholm, Sweden. All jaricum, kibonotense) from the Kilimanjaro-Meru area of Tan- other material remains in the collections of C. Hemp and zania. Although Bolivar (1922) founded the genus Phlesirtes K.-G. Heller. on Xiphidion merumontanum Sjostedt,¨ he did not transfer the other three species described by Sjostedt¨ to his newly Molecular analysis erected genus. Karny (1921) stated that Xiphidion kibonotense should be placed into the genus Conocephalus because of its DNA was extracted from the muscles of one hind leg using unarmed prosternum, and noted that the other three species the QIAamp® DNA mini kit (Qiagen, Germany, following the from the Kilimanjaro area should be placed into Conocephalus standard protocol for blood and tissue) and the NucleoSpin® if the prosternum was unarmed. Obviously unaware of Karny’s tissue kit (Machery & Nagel, Germany, following the standard (1921) study, Uvarov (1924) described Phlesirtes brachiatus. protocol for human and animal tissue). He transferred Phlesirtes, also mentioning the three species For the analysis of Orthoptera species groups we selected Xiphidion meruense, kilimandjaricum and kibonotense as prob- genes for which data already exist and that show sufficient ably closely related, into Decticinae. Chopard & Kevan (1954) variation between closely related species (mitochondrial genes added another three species from northern Kenya (Table 1). and especially the barcoding gene cytochrome oxidase subunit Ragge (1977) and Rentz (1988) doubted that Phlesirtes should I (COI)) and a more conserved gene for deeper nodes in be placed under Decticinae, transferring the genus back to the phylogenetic tree (histone gene H3). Preliminary work the subfamily Conocephalinae. Rehn (1914) erected the genus confirmed the suitability of the selected genes. Karniella with the sole species K. bullata, quite similar in gen- The mitochondrial 16S rRNA gene, the mitochondrial COI eral morphology to Sjostedt’s¨ three Phlesirtes species, but dif- gene and nuclear histone H3 gene (H3) were amplified by fering strongly in the shape of the pronotum and tegmina. Unal¨ polymerase chain reaction (PCR) using the primers 16a: 5- (2005) described the new Conocephalinae genus Naskreck- CGC CTG TTT ATC AAA AAC AT-3 and 16b: 5-CCG GTC iella, also listing Phlesirtes as a genus under Conocephalini. TGAACTCAGATCACGT-3 for the 16S rDNA (Kocher A list of Conocephalini genera is given in Unal¨ (2005). In the et al., 1989); H3fwd: 5-ATG GCT CGT ACC AAG CAG Orthoptera Species File online (Eades & Otte, 2009) the three ACG GC-3 and H3rev: 5-ATA TCC TTG GGC ATG ATG Phlesirtes species kilimandjaricus, kibonotensis and meruensis GTG AC-3 for the histone H3 gene (Colgan et al., 1998); and are erroneously listed under Conocephalus (Anisoptera). LCO1490: 5-GGT CAA CAA ATC ATA AAG ATA TTG G-3 © 2010 The Authors Journal compilation © 2010 The Royal Entomological Society, Systematic Entomology, doi: 10.1111/j.1365-3113.2009.00512.x The Phlesirtes complex reviewed 3 and HCO2198: 5-TAA ACT TCA GGG TGA CCA AAA AAT Cytotaxonomic
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