The Phlesirtes complex (Orthoptera,Tettigoniidae, Conocephalinae,Conocephalini) reviewed: integrating morphological,molecular, chromosomal and bioacoustic data

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 Sjöstedt. 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 (Sjöstedt) and Phlesirtes kilimandjaricus (Sjöstedt). Subtribal status is proposed for the four investigated African genera. A key to the Chortoscirtes species is provided.     

Ecological adaptations of grassland‐inhabiting flightless Orthoptera: Fulvoscirtes and Acanthoscirtes,two new genera of African Karniellina (Orthoptera,Tettigoniidae, Conocephalinae,Conocephalini)

Two new genera, Fulvoscirtes n.gen. and Acanthoscirtes n.gen. , are established within the subtribe Karniellina of Conocephalini. Fulvoscirtes is based on Xiphidion kilimandjaricum Sjöstedt, 1909 and Acanthoscirtes on Phlesirtes kevani Chopard from northern Kenya. The majority of Fulvoscirtes spp. are confined to open grasslands in the submontane zone of mountains. Fulvoscirtes contains eight species, seven of which are newly described in this paper. Three species and one subspecies occur on Mt Kilimanjaro. These are F. kilimandjaricum (Sjöstedt) constricted to the southern slopes, F. legumishera n.sp. confined to the northern side and F. sylvaticus n.sp. occurring on the western side of Kilimanjaro and on the eastern slopes of Mt Meru. Fulvoscirtes fulvus n.sp. is divided into two subspecies, F. fulvus fulvus n.ssp. found in the submontane zone of east Kilimanjaro and F. fulvus parensis n.ssp. in submontane to montane localities of the North and South Pare mountains. Fulvoscirtes fulvotaitensis n.sp. occurs in the Taita Hills of southern Kenya. Fulvoscirtes viridis n.sp. is described from savannah habitats between Mts Longido and Meru. Fulvoscirtes laticercus n.sp. is found in the Kenyan highlands, while the most southerly occurring species, Fulvoscirtes manyara n.sp. , is found on Mt Hanang and the Mbulu highlands of northwestern Tanzania. Acanthoscirtes contains three species, of which A. albostriatus n.sp. is described newly from savannah habitas of eastern Kilimanjaro. Information is given on the ecology and the acoustic behaviour of some of the species together with keys to the genera of the Karniellina and the species of Fulvoscirtes and Acanthoscirtes. The genera of Karniellina probably evolved at a time when grasslands spread in East Africa due to an increasing aridification of the climate. The earliest lineage, the genus Karniella, is adapted to more forested habitats while the majority of the genera of Karniellina prefer open grasslands. Major splits within Karniellina probably occurred with the emergence of savannah grasslands due to the ongoing fragmentation of forest habitats several millions years ago, but most species within the genera are geologically young, their radiation being boosted by climatic fluctuations of the past 1–2 Ma.     

Dispersal and speciation of East African mountain bushcrickets of the genus Phlesirtes (Orthoptera: Tettigoniidae: Conocephalinae,Conocephalini) related to climate fluctuations

A phylogeny of the genus Phlesirtes Bolivar is presented, based on new sequence data of three genes (16S rDNA, COI, H3). Species of the genus Phlesirtes (subtribe Karniellina of the Tribe Conocephalini) occupy habitats of montane to afroalpine grasslands in East Africa. Phlesirtes is the most species‐rich genus of the subtribe Karniellina, a group of small flightless Ensifera restricted to eastern Africa. Taken together, the biogeographical patterns seen in Phlesirtes and its molecular phylogeny suggest a migration scenario: the mountain ranges acting as stepping stones, enabling a spread of Phlesirtes ancestors during periods of favourable climatic conditions in the past. The Pleistocene inland volcanoes, such as Mt Kenya or Mt Kilimanjaro, allow us to date speciation processes within the genus Phlesirtes. It is suggested that cooler humid periods of the past 3 Ma boosted speciation of Phlesirtes in East Africa.     

A new genus and species of African Phaneropterinae (Orthoptera: Tettigoniidae), with data on its ecology,bioacoustics and chromosomes

A new genus is proposed for a new East African Phaneropterinae species, Lunidia viridis, occurring on Mt. Kilimanjaro, Tanzania. Based on 33 records, notes on distribution and habitat are given, as well as acoustical data provided. Climate and vegetation parameters obtained along several transects on Mt. Kilimanjaro were evaluated describing the ecological niche of the new species. This interdisciplinary approach allows not only a profound characterisation of the ecological demands of the new genus but also predictions of the potential distribution area, which is tested for the first time for an African bush cricket species. Lunidia viridis n. gen. n. sp. occurs within humid and perhumid forests and Chagga home gardens, avoiding subhumid conditions on the mountain. It is found from 1,330 m upwards on the southern slopes, whereas the same ecological conditions are expressed from 1,930 m upwards on the drier northern slopes. Lunidia viridis has an unusually complex and variable song, which is described from field and laboratory recordings. The FISH technique for characterizing chromosomes is applied for the first time for an African species; L. viridis exhibits a karyotype typical for most Tettigoniidae.     

A molecular phylogeny of East African Amytta (Orthoptera: Tettigoniidae,Meconematinae) with data on their cytogenetics

This is the first study to address the molecular phylogeny of East African Meconematinae. We used DNA sequence data from the three genes cytochrome oxidase subunit I (COI ), 16S rRNA and histone H3 to reconstruct the relationships within the genus Amytta using Bayesian Inference and maximum likelihood. We included 45 individuals representing 19 Meconematinae taxa from Africa, Europe and Australia, and three outgroup species to explore taxonomic limits at different levels. Members of the genus Amytta belong to three related lineages within Meconematinae. Species of Amytta were found to be monophyletic with Meconema and Phlugidia as sister groups. The molecular results are in accordance with previous morphological and ecological studies. Chromosome numbers and the type of sex determination systems found in Amytta agrees with the three lineages also found in the phylogenetic tree. We conclude that the radiation of Amytta species is geological young. Biogeographical patterns caused by climatic fluctuations of the past seen also in other nonrelated Orthoptera taxa such as the coptacrine genus Parepistaurus are the drivers of biodiversity in the area.     

A new genus of Decticinae (Orthoptera: Tettigoniidae) from the Middle East

A new genus, Raggeana, is erected for Metrioptera bodenheimeri Uvarov, and the hitherto unknown female of this species is described.     

Taxonomic Study of Tribe Copiphorini in Korea (Orthoptera: Tettigoniidae: Conocephalinae)

ABSTRACT Five species of the tribe Copiphorini (Orthoptera, Tettigoniidae, Conocephalinae) including a newly recorded species Xestaphrys javanicus Redtenbacher are recognized in Korean fauna. The key to species, characteristic figures, and diagnosis for the newly recorded species are presented.     

Taxonomic Study of the Genus Conocephalus Thunberg in Korea (Orthoptera: Tettigoniidae: Conocephalinae)

ABSTRACT The genus Conocephalus Thunberg in Korea was taxonomically studied. As the result, five species were confirmed to Korean fauna including a newly recorded species, Conocephalus bambusanus Ingrisch. The key to species, distributional data and comparative figures for identification are given.     

The genus Aerotegmina (Orthoptera, Tettigoniidae, Hexacentrinae): chromosomes, morphological relations, phylogeographical patterns and description of a new species

The genus Aerotegmina Hemp is common on East African mountains. Two species are known and a third, A. taitensis n. sp., is described in this paper. A. kilimandjarica Hemp is widespread while A. shengenae Hemp is endemic to 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 chromosome number A. kilimandjarica (2n?=?33) differs clearly from A. shengenae (2n?=?27). Data presented on other flightless Orthoptera suggest that the South Pare Mountains and the Taita Hills, both belonging to the geologically old mountain chain of the Eastern Arc, show a faunistic similarity not shared by any other mountain range in the area. The mechanisms that led to this phylogeographic pattern in flightless Orthoptera in the Eastern Arc Mountains of northern Tanzania and southern Kenya and the inland volcanoes are discussed. A key to the three Aerotegmina species is presented, as well as bioacoustical data of all species compared to the phaneropterine species Euryastes jagoi.     

Cytological and morphological differences between two species of the genus Tettigonia (Orthoptera,Tettigoniidae) from Korea

Tettigonia ussuriana and T. dolichopoda maritima differ in the length of tegmina, details in venation, and in females in details of the subgenital plate. The two species of the genus Tettigonia have the same number and morphology of autosomes but a different morphology of the X chromosome: in T. ussuriana it is metacentric, whereas in T. dolichopoda maritima acrocentric. In both species, euchromatic zones and breaks of one or to chromatids during meiosis and mitosis in the X chromosome were observed. Additionally, B chromosomes were noted in most individuals of both species.     

A new genus of African Acrometopini (Tettigoniidae: Phaneropterinae) based on morphology,chromosomes, acoustics,distribution, and molecular data,and the description of a new species

A new genus, Altihoratosphaga, is erected for species formerly assigned to Horatosphaga Schaum, 1853, and a new species is described. Four species are included in Altihoratosphaga: Altihoratosphaga nomima (Karsch, 1896), Altihoratosphaga montivaga ( Sjöstedt, 1909 ), Altihoratosphaga nou (Hemp, 2007) and Altihoratosphaga hanangensis sp. nov. All four species are restricted to Tanzanian localities, and, except for A. nomima, for which no ecological data are available, are confined to montane forest habitats. Data on ecology, acoustics, chromosomes, and molecular relationships are provided, as well as a key to Altihoratosphaga species. The present‐day distribution of Altihoratosphaga species suggests former migration events at times when wetter and colder climatic fluctuations favoured connections between montane forest communities, which today are isolated, enabling flightless taxa such as Altihoratosphaga and Monticolaria to spread. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 66–82.     

Chromosomal diversification in the flightless Western Mediterranean bushcricket genus Odontura (Orthoptera: Tettigoniidae: Phaneropterinae) inferred from molecular data

We used molecular characters to infer the phylogenetic position of the Western Mediterranean bushcricket genus Odontura and to trace its high karyotype diversity. Analysis of 1391 base pairs of two mitochondrial genes (COI and ND1) and one nuclear sequence (ITS2) was conducted. Phylogenetic topologies were estimated using maximum parsimony, maximum likelihood and likelihood‐based Bayesian inference. The genus Odontura is a phylogenetic outlier in respect of all other European Phaneropterinae genera and has been proposed to have originated from a hitherto unknown ancestor. Our results support the monophyly of the genus Odontura and the recognition of two subgenera: Odontura and Odonturella. We found that both Sicilian taxa of the subgenus Odontura have a completely identical morphology and song patterns. Combining these results, we proposed that both should be treated as subspecies: O. (Odontura) stenoxypha stenoxypha and O. (O.) st. arcuata. Bioacoustic data also proved to support independent markers, with song characteristics reflecting the molecular topology. Mapping the karyotypic characters onto the phylogenetic tree allows a reconstruction of the directions and transitional stages of chromosome differentiation. The number of autosomes within the genus Odontura ranges from 26 to 30. In addition to the ancestral X0 sex determination mechanism, neo‐XY and neo‐X₁X₂Y sex chromosomes have evolved independently.     

A new genus and species of larval mite (Acari: Prostigmata: Microtrombidiidae) parasitising Orthoptera (Tettigoniidae) from the Sierra Nevada, Spain

Nevada capileirarum n. g., n. sp. (Acari: Microtrombidiidae: Microtrombidiinae) is described from ectoparasitic larvae parasitising two endemic species of Orthoptera (Tettigoniidae), Baetica ustulata (Rambur) and Pycnogaster inermis (Rambur) from the Sierra Nevada mountain range, Granada, Spain. A key to the larvae of microtrombidiine genera with three dorsal scuta and a coxal setal formula of 2-1-1 is presented.     

The biology of the West African bush-cricket,Euthypoda acutipennis Karsch (Orthoptera: Tettigoniidae)

This paper presents the results of the first comprehensive study of the biology of a tropical bush-cricket. The eggs were laid without any external protective structures and lost water readily in unsaturated air; losses of more than 16% of the original fresh weight were usually fatal. Development and hatching took place only if the eggs were in contact with water. The water content increased by about 500% during development. The mean incubation period was 171 and 116 days at constant temperatures of 20^oC and 28^oC, respectively, but at temperatures fluctuating by±3°C about a mean of 28°C this period was reduced to a mean of 88 days. The eggs failed to develop if exposed to freezing temperatures for more than five days or kept at a constant temperature of 307deg;C or above. The duration of the nymphal period varied from 83 to 131 days at a temperature range of 22–30°C (mean 26° C). There were four or five nymphal instars in the male and five or six in the female. The linear growth of the hind femur conformed to Dyar's law. There was a conspicuous colour change during development, all the first-instar nymphs being leaf green and the adults predominantly brown. In nature the species lives exclusively on the forest floor and is markedly nocturnal in habits, hiding underneath litter during the day. The eggs are dormant during the dry season, hatching at the onset of the rains (March/April at Ibadan). The nymphs reach maturity from July onwards and the adults have mostly died off by the end of the rainy season (October). In the laboratory the species was found to have a temperature preference of 26–32°C, immobilization setting in at 18°C and 42°C, and a humidity preference of 60–80% R.H.; this corresponds with the prevailing conditions in its forest habitat. The nocturnal pattern of activity persisted for several days in continuous darkness, with no marked acceleration. Movement was inhibited for several days by artificial illumination. Reversed illumination reversed the rhythm of activity.     

Quality calls: phylogeny and biogeography of a new genus of neotropical katydid (Orthoptera: Tettigoniidae) with ultra pure-tone ultrasonics

This paper describes Artiotonus, a new genus of tropical katydid from Colombia and Ecuador. These acoustic ensiferans are represented by three species with a geographic distribution generally restricted to the rainforest of the Bolivar geosyncline of northwestern South America (Pacific). A phylogenetic analysis based on 28 morphological and six behavioural characters produced a tree (A. artius, (A. tinae, A. captivus)) with a consistency index of 0.9. All species are defined by a few autapomorphic changes. The most reliable character for identification is the temporal pattern of the calling song, suggesting a recent genetic divergence. Biogeographic analysis indicates that such genetic divergence began with geographic isolation produced before the Holocene transgression. Species of this genus are remarkable in that the calling song of males exhibits a narrow spectrum with a high quality factor (37–42), unusual values for such ultrasonic signals. A broad discussion on the evolution of tonal calls and pulse trains is offered.     

Testing concordance in species boundaries using acoustic,morphological, and molecular data in the field cricket genus Itaropsis (Orthoptera: Grylloidea,Gryllidae: Gryllinae)

In most taxa, species boundaries are inferred based on differences in morphology or DNA sequences revealed by taxonomic or phylogenetic analyses. In crickets, acoustic mating signals or calling songs have species‐specific structures and provide a third data set to infer species boundaries. We examined the concordance in species boundaries obtained using acoustic, morphological, and molecular data sets in the field cricket genus Itaropsis. This genus is currently described by only one valid species, Itaropsis tenella, with a broad distribution in western peninsular India and Sri Lanka. Calling songs of males sampled from four sites in peninsular India exhibited significant differences in a number of call features, suggesting the existence of multiple species. Cluster analysis of the acoustic data, molecular phylogenetic analyses, and phylogenetic analyses combining all data sets suggested the existence of three clades. Whatever the differences in calling signals, no full congruence was obtained between all the data sets, even though the resultant lineages were largely concordant with the acoustic clusters. The genus Itaropsis could thus be represented by three morphologically cryptic incipient species in peninsular India; their distributions are congruent with usual patterns of endemism in the Western Ghats, India. Song evolution is analysed through the divergence in syllable period, syllable and call duration, and dominant frequency. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 285–303.     

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.     

Colour polymorphism in Ephippiger ephippiger (Orthoptera, Tettigoniidae)

Colour in the Ephippiger ephippiger complex varies with the geographical distribution and with rearing density. In the W European subspecies E. e. vitium , wild insects from northern and eastern parts of the range tend to have green tergites with relatively inconspicuous banding, whereas those from S France have conspicuously pale banded dark-green or brown tergites. These forms represent the ends of a continuum and there is no zone of distinction. However, the E Pyrenean subspecies, E. e. cunii , lacks the broad banding of its neighbouring E. e. vitium. The tergites of laboratory cultured insects of these stocks when reared in isolation are distinctly green but when reared together become progressively brown. The darkness of the brown is density dependant, so that at high densities the tergites may become almost black. These colour changes are irreversible. Adult insects also become darker with sexual maturity and after contact on mating. Wild-caught E. e. ephippiger from Italy and Yugoslavia have green tergites without obvious banding. Colour changes have not been found or induced in this subspecies. The significance of the colour change is considered and although firm data are wanting, circumstantial evidence suggests a deliberate shift to an aposematic or pseudoaposematic colour pattern.