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Orthoptera, Tettigoniidae, Hexacentrinae): Chromosomes, Morphological Relations, Phylogeographical Patterns and Description of a New Species

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Orthoptera, Tettigoniidae, Hexacentrinae): Chromosomes, Morphological Relations, Phylogeographical Patterns and Description of a New Species Org Divers Evol (2013) 13:521–530 DOI 10.1007/s13127-013-0133-7 ORIGINAL ARTICLE The genus Aerotegmina (Orthoptera, Tettigoniidae, Hexacentrinae): chromosomes, morphological relations, phylogeographical patterns and description of a new species Claudia Hemp & Klaus-Gerhard Heller & Elzbieta Warchalowska-Sliwa & Andreas Hemp Received: 20 December 2012 /Accepted: 4 March 2013 /Published online: 28 March 2013 # Gesellschaft für Biologische Systematik 2013 Abstract The genus Aerotegmina Hemp is common on East Keywords Orthoptera . Hexacentrinae . Aerotegmina . African mountains. Two species are known and a third, A. Euryastes .TaitaHills .EasternArcMountains .EastAfrica . taitensis n. sp., is described in this paper. A. kilimandjarica New species . Bioacoustics . Chromosomes . Morphology . Hemp is widespread while A. shengenae Hemp is endemic to Phylogeography the South Pare Mountains and A. taitensis n. sp. is known only from the Taita Hills. Morphologically, and from their song, A. shengenae and A. taitensis n. sp. are closely related. In Introduction chromosome number A. kilimandjarica (2n=33) differs clear- ly from A. shengenae (2n=27). Data presented on other flight- The genus Aerotegmina was erected on A. kilimandjarica less Orthoptera suggest that the South Pare Mountains and the Hemp, 2001, from Mt. Kilimanjaro, Tanzania (Hemp 2001). Taita Hills, both belonging to the geologically old mountain Another Aerotegmina species was described by Hemp chain of the Eastern Arc, show a faunistic similarity not (2006) restricted to montane forests of the South Pare shared by any other mountain range in the area. The mech- Mountains of northern Tanzania. Gorochov (2007)re- anisms that led to this phylogeographic pattern in flightless established the subfamily Hexacentrinae Karny transferring Orthoptera in the Eastern Arc Mountains of northern Aerotegmina from Listrocelidinae to this subfamily. Tanzania and southern Kenya and the inland volcanoes The genus Aerotegmina is characterized by highly mod- are discussed. A key to the three Aerotegmina species is ified wings and the shape of the pronotum. The tegmina are presented, as well as bioacoustical data of all species com- inflated strongly to form an acoustic chamber, closed by the pared to the phaneropterine species Euryastes jagoi. flap-like alae completing the chamber ventrally. The pronotum is saddle-shaped. Inflation of the tegmina results in an unusually loud song C. Hemp (*) at a low frequency (Heller et al. 2010). Low-frequency Department of Animal Ecology and Tropical Biology songs have important advantages in acoustic communica- (Zoology III), University of Würzburg, Am Hubland, tion, increasing the range between sender and receiver. 97974 Würzburg, Germany Aerotegmina are flightless insects with low mobility. Thus, e-mail: [email protected] the need to be heard by females could have been the evolu- K.-G. Heller tionary pressure leading to development in the male sex of Institute of Biology, University of Erlangen-Nürnberg, this effective communication to attract females. Nürnberg, Germany Hemp (2010) suggested that the origin of the genus E. Warchalowska-Sliwa Aerotegmina lies in the geologically ancient chain of the Institute of Systematics and Evolution of Animals, Polish Eastern Arc Mountains of Tanzania und southern Kenya. Academy of Sciences, Kraków, Poland Areas obviously generating a high biodiversity in this region were found to be the South Pares and West Usambaras due A. Hemp Department of Plant Systematics, University of Bayreuth, to the special topography and geographical location of these Bayreuth, Germany massive and high mountain chains. The record of another 522 C. Hemp et al. Aerotegmina species in the Taita Hills of Kenya sheds more were obtained from adult male gonads. Testes were ex- light on the evolution of this taxon and the faunistic rela- cised, incubated in a hypotonic solution (0.9 % sodium tionship of the mountain ranges of the northern branch of citrate), and then fixed in ethanol:acetic acid (3:1). The the Eastern Arc Mountains. fixed material was squashed in 45 % acetic acid. Cover In this paper we describe a new species of Aerotegmina slipswereremovedbythedryiceprocedureandthe restricted to the Taita Hills of Kenya. We compare song and preparations were air-dried. C-banding was carried out the acoustic apparatus of the three species, present the chromo- using a slightly modified version of Sumner’s(1972) somal set of two species and discuss their phylogeographical technique. The silver staining method (Ag-NO3) for relationships. localisation of the nucleolus organiser regions (NORs) was performed as previously reported (Warchałowska- Śliwa & Maryańska-Nadachowska 1992). Chromosomes Materials and methods were classified on the basis of the criteria proposed by Levan et al. (1964). Collection localities Depositories A. taitensis n. sp. Fig. 1 was collected at night in the two largest remaining pieces of montane forest of the Taita Hills, MNB: Museum für Naturkunde, Zentralinstitut der on Mt. Vuria and the Ngangao forest reserve. Humboldt-Universität zu Berlin. Measurements Results Total body length refers to the body length of the insect, disregarding tegmina but not genitalia. Tegminal length is Key to Aerotegmina species (males) determined as viewed from above. 1 Alae surpassing abdominal apex by about one-third of its Song length; alae elongate with pointed tips (Figs. 2, 3a and 4a). A. kilimandjarica Hemp Recordings of song in the field and in captivity (20 °C) were 1′ Alae shortened, surpassing abdomen by only a few mm. made with a Pettersson X1000 bat detector using a sampling 2 rate of 100 kHz. The sounds were analysed using the program 2 Alae broadly rounded (Figs. 3c and 4b). A. taitensis n. sp. Amadeus II (Martin Hairer; http://www.hairersoft.com). 2′ Alae spatulate (Fig. 3b). A. shengenae Hemp Frequency data were taken from one syllable per specimen Aerotegmina taitensis n.sp.HempC.(Figs.1, 2, 3c, 4b (mean in sliding window of 512 points; hamming window). and 5c) For A. shengenae a Sony ECM-121 microphone was also used. Holotype Male: Kenya, Taita Hills, Mt. Vuria, 1950 m, Chromosomes UTM zone 37 M, 421900 E, 9623400S, canopy of under- growth tree, November 2010, C. Hemp coll.; depository, Seven males of Aerotegmina kilimandjarica (CH6825, CH6827, MNB. CH6850-54) and one male of A. shengenae (CH7268) were used for cytotaxonomic analyses. Chromosome preparations Fig. 2 Aerotegmina taitensis n. sp.: ventral view of male abdominal Fig. 1 Habitus of male Aerotegmina taitensis n. sp. apex with subgenital plate and styli The genus Aerotegmina (Orthoptera, Tettigoniidae, Hexacentrinae) 523 Fig. 3 a–c Aerotegmina species, ventral view on tegmina and alae. a A. kilimandjarica. b A. shengenae. c A. taitensis n. sp. Bars 3mm Paratype 1 male, Ngangao forest reserve, 1900 m, canopy Genitalia. Subgenital plate symmetrical (Fig. 2), of same of undergrowth tree, February 2011, collection Hemp. milky white colour as venter of abdomen, with smooth surface, flattened ventrally, lobes with scattered hairs. Description Male. Color. Light green. Head and antennae. Cerci robust with rugose and hairy surface; in the living Antenna more than twice as long as length of body, whitish specimen stretched out from the body. with irregular annulate dark markings (Fig. 1). Cuticle of head smooth. Conical and laterally compressed fastigium of Measurements Males (N=2); Total length of body 13.5 mm; vertex situated before antennal sockets in dorsal aspect. Length of pronotum 3.5–3.6 mm; Length of hind femur 10.8– Space between eyes about 2.5 times as wide as diameter 11 mm; Length of tegmina (as seen from above) 11–14.5 mm. of eye. Eyes almost circular, prominent, whitish in living Female. Not known. insect, brown in preserved insect. Thorax. Green with rugu- lose callosities. Anterior and posterior margins of pronotum Distribution A. taitensis n. sp. was collected in montane yellow. Abdomen. Abdomen milky white (Fig. 2). Tegmina forests of Mt. Vuria ranging between 1,950 and 2,200 m and alae. Acoustical chamber imperfectly closed by alae; and in montane forest of the Ngangao forest reserve at about alae with rounded apices, leaving a gap posteriorly (Figs. 3c 1,800 m. and 4b). Tympanum (mirror of right tegmen) oval, about 4. 2 mm long and 3.9 mm wide. Legs. Fore and mid femora Habitat Closed montane forest with a upper tree canopy of with four stout yellowish spines on the outer sides, and four 12–15 m height and 80–90 % coverage on Mt Vuria. The green smaller stout spines on the inner sides of the fore, and forest is heavily disturbed and probably logged for certain three on the mid femora. Apically with a pair of short spurs. timber species. The Taita forests in which A. taitensis n. sp. Hind femur with six stout yellow outer and four small inner was recorded belongs to the same forest type as on the South green spines, distally one spur at each side. Fore and mid Pare Mountains where A. shengenae occurs (Hemp 2006). tibiae with five pairs of light green predatory spines at each The montane forest of the Ngangao forest reserve is less side, apically with a pair of spurs. Hind tibiae with four rows heavily impacted in its centre thus harbouring still intact of minute spines. With well developed tarsal arolium. afromontane forest communities. A. taitensis males were located by their calling songs starting at nightfall. Individuals were collected perched under broad leaves. Most individuals were heard from considerable heights up in the canopy. Only two specimens were encoun- tered at lower heights and were shaken from branches. Song The calling song of the species was heard in the evening and night hours only. It consisted of long sequences of single syllables produced at a rate of 2–5 Hz (Fig. 6, Table 1). Differences between the recordings are very likely mainly an effect of different temperature. However, the ambient temperature in the field could not be measured close to the animals. Fig. 4 a,b Dorsal aspect of male Aerotegmina alae.
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