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.     

A new genus of African Karniellina (Orthoptera,Tettigoniidae, Conocephalinae,Conocephalini): integrating morphological,molecular and bioacoustical data

Melanoscirtes gen.n. is established within Karniellina. The members of this subtribe are small conocephaline bush crickets, confined to Africa. Melanoscirtes is erected on Phlesirtes kibonotensis, a species restricted to forest clearings and forest edge in the submontane and montane zones of Mt. Kilimanjaro. A subspecies, M. kibonotensis uguenoensis, is described from the North Pare mountains, a mountain range of the Eastern Arc adjacent to Mt. Kilimanjaro. Further species of Melanoscirtes occur on other mountain ranges of the northern branch of the Eastern Arc mountains of northern Tanzania and southern Kenya. The South Pare mountains harbour M. shengenae; the West Usambaras, M. usambarensis, and the Taita Hills, M. taitensis. All species and subspecies of Melanoscirtes exhibit a similar morphology and occupy analogous habitats on the respective mountains. The song patterns for all species found within this genus are very similar, and this, together with evidence from molecular data, suggests that allopatric speciation is the reason for the biogeographic pattern found in this genus. A key for the subspecies and species of Melanoscirtes is provided.     

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.     

Broken bridges: The isolation of Kilimanjaro's ecosystem

Biodiversity studies of global change mainly focus on direct impacts such as losses in species numbers or ecosystem functions. In this study, we focus on the long‐term effects of recent land‐cover conversion and subsequent ecological isolation of Kilimanjaro on biodiversity in a paleobiogeographical context, linking our findings with the long‐standing question whether colonization of African mountains mainly depended on long‐distance dispersal, or whether gradual migration has been possible through habitat bridges under colder climates. For this, we used Orthoptera as bioindicators, whose patterns of endemism and habitat demands we studied on about 500 vegetation plots on Kilimanjaro and Mt. Meru (Tanzania) since 1996. Land‐cover changes in the same area were revealed using a supervised classification of Landsat images from 1976 to 2000. In 1976, there was a corridor of submontane forest vegetation linking Kilimanjaro with Mt. Meru, replaced by human settlements and agriculture after 2000. Until recently, this submontane forest bridge facilitated the dispersal of forest animals, illustrated by the large number of endemic submontane forest Orthoptera shared by both mountains. Furthermore, the occurrence of common montane endemics suggests the existence of a former forest corridor with montane vegetation during much earlier times under climatic conditions 2–7°C cooler and 400–1,700 mm wetter than today. Based on the endemicity patterns of forest Orthoptera, negative consequences are predicted due to the effects of isolation, in particular for larger forest animals. Kilimanjaro is becoming an increasingly isolated ecosystem with far reaching consequences for diversity and endemism. Forest bridges between East African mountains acted as important migratory corridors and are not only a prehistoric phenomenon during periods with other climatic conditions but also disappeared in some places recently due to increasing and direct anthropogenic impact.     

Ecology of the pteridophytes on the southern slopes of Mt. Kilimanjaro – I. Altitudinal distribution

140 taxa of 61 genera in 24 families of pteridophytes were recorded on the southern slopes of Mt. Kilimanjaro. These represent about one third of the entire pteridophyte flora of Tanzania. The families richest in species are the Aspleniaceae, the Adiantaceae, the Dryopteridaceae, the Thelypteridaceae and the Hymenophyllaceae. Due to its luxuriant montane rain forest, which receives a precipitation of up to over 3000 mm, Mt. Kilimanjaro is distinctly richer in pteridophyte species than other volcanoes in East Africa. However, compared with the mountains of the Eastern Arc, the number of pteridophytes on Mt. Kilimanjaro is smaller. This can be explained by the widely destroyed submontane (intermediate) forest rather than by the higher age of the Eastern Arc Mts.The altitudinal distribution of all ferns was investigated in 24 transects. On the southern slopes of Mt. Kilimanjaro they were found in an altitudinal range of 3640 m. Cyclosorus quadrangularis, Azolla nilotica, Azolla africana andMarsilea minuta are restricted to the foothills, while Polystichum wilsonii, Cystopteris nivalis and Asplenium adiantum-nigrum are species found in the highest altitudes.Based on unidimensionally constrained clustering and on the analysis of the lowermost and uppermost occurrence of species, floristic discontinuities within the transects were determined. From these data and from an evaluation of the distribution of ecological groups and life forms, 11 altitudinal zones could be distinguished: a colline zone (–900 m asl), a submontane zone (900–1600 m asl) with lower and upper subzones, a montane zone (1600-2800 m asl) divided into 4 subzones, a subalpine zone (2800–3900 m asl) with lower, middle and upper subzones, and finally a (lower) alpine zone above 3900 m. The highest species numbers were observed in the lower montane forest belt between 1600 and 2000 m altitude. The zonation of ferns found at Mt. Kilimanjaro corresponds well with the vegetational zonation described by other authors using bryophytes as indicators in different parts of the humid tropics.     

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.     

Phylogeny and revision of the ‘Cubitermes complex’ termites (Termitidae: Cubitermitinae)

The phylogeny of the genus Cubitermes Wasmann was reconstructed using two mitochondrial genes (COI and COII) and a fragment of the nuclear 28S rDNA, including samples of Apilitermes Holmgren, Crenetermes Silvestri, Megagnathotermes Silvestri and Thoracotermes Wasmann. Based on our analyses, we recovered these genera within a paraphyletic Cubitermes clade. Cubitermes species are distributed between five main clades, highly reflective of their enteric valve armatures (EVAs). Consequently, Cubitermes is here divided into five monophyletic genera based on phylogenetic analyses and EVAs: Cubitermes (sensu stricto), Isognathotermes Sjöstedt, Nitiditermes Emerson, Polyspathotermes Josens & Deligne, gen.n. , and Ternicubitermes Josens & Deligne, gen.n. Moreover, the two species of Megagnathotermes included in this study, exhibiting different EVAs, are phylogenetically distant: M. katangensis Sjöstedt belongs in the genus Polyspathotermes, while M. notandus Silvestri remains as a monotypic genus. During the evolution of the Cubitermes complex, sclerotized EVA spatulae appeared twice independently: three or six spatulae in Polyspathotermes (sulcifrons and oblectatus patterns) and two jaw‐like spatulae in Nitiditermes (sankurensis pattern), which is absent in some species (oculatus pattern). Zoobank registration: http://zoobank.org/urn:lsid:zoobank.org:pub:576AAFF8‐63C6‐4962‐A8F5‐75C9317AEA7B .     

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.     

Three new genera of Braconini from Australasia and Malaysia (Insecta, Hymenoptera, Braconidae)

Ancilibracon gen.n. (type species: Ancilinracon townesi sp.n.). Ancilibracon bakeri sp.n., Gelasinibracon gen.n. (type species: Gelasinibracon sedlaceki sp.n.), Gelasinibracon simplicicaudatus sp.n., Esengoides gen.n. (type species: Esengoides crentdatus sp.n.) and Esengoides fulvus sp.n. are each described and illustrated. Ancilibracon towensi is from Malaysia. A. bakeri is from Borneo, both Gelasinibracon species are from Guinéa, E. crenulatus is from the solomon Islands and E. fulvus is from Australia. These three new genera belong to the Plesiobrscon Cameron group of the Braconina. Their relationship are discussed.     

New and little known crickets of the subfamily Phalangopsinae (Orthoptera,Gryllidae): 6. Neotropical taxa of the tribes Phalangopsini and Paragryllini

Two new genera, ten new species, and two new subspecies: Uvaroviella izerskyi sp. n., U. morona sp. n., U. ucayali sp. n., U. pastaza sp. n., U. affinis sp. n., U. bolivia sp. n., U. bora atalaya subsp. n., Kevanacla orientalis contraria subsp. n., Peruacla solitaria gen. et sp. n., Ecuadoracla propria gen. et sp. n., Adelosgryllus spurius sp. n., and A. phaeocephalus sp. n. are described. The tribe Paragryllini is divided into three subtribes: Paragryllina Desutter, 1988 stat. n., Neoaclina Desutter, 1988 stat. n., and Strogulomorphina Desutter 1988, stat. n. The composition of the genera Uvaroviella Chop. and Neoacla Desutter is discussed. The species from Costa Rica and the Antilles are supposed to belong to these genera and subgenera. New synonymy [U. trinidadi Gorochov, 2007 = U. enodos Otte et Perez-Gelabert, 2009, syn. n. and N. multivenosa (Chopard, 1937) = Selvacla choreutes Otte, 2006, syn. n.] is proposed.     

Diversity of geometrid moths (Lepidoptera: Geometridae) along an Afrotropical elevational rainforest transect

Geometrid moths were investigated at 26 sites on 9 elevational levels along an elevational transect at Mt. Kilimanjaro (Tanzania), stretching from the fine‐grained mosaic of small agroforest plots with combined cultivation of trees, shrubs and crops at 1650 m through mountain rainforest to heathland at 3300 m. We sampled moths manually at light between 19 : 00 and 22 : 00 in the rainy seasons of March to May and October to January in the years 2000, 2001 and 2002. Along the transect, the composition of moth communities changed from a domination by Sterrhinae and Ennominae to a dominance of Larentiinae with increasing elevation. Overall, alpha diversity was very low compared to other tropical mountain regions. Fisher's alpha showed a maximum of 30 in the agroforest mosaic at 1650 m and decreased to values around 12 in the mountain rainforest. Communities of geometrid moths within the forest belt were significantly dissimilar from communities outside the forest. The diversity patterns on Mt. Kilimanjaro can be related to the young age, island‐like position and history of the mountain. These factors have led to the formation of a homogeneous upper mountain rainforest habitat which in turn houses homogeneous moth communities with a low diversity compared to habitats at lower elevations. Here, a heterogeneous habitat mosaic allowing the intrusion of savannah species into this former forest habitat may account for an increased diversity. In the heath zone above the forest, climatic conditions are very harsh, permitting only few specialists to thrive in this ericaceous woodland. Edge effects were discernible at the forest–heathland boundary where some moth species from heathland invaded the closed forest. At the boundary between agroforest and a forest mosaic of exotic Acacia and Eucalyptus forest plantations and natural mountain forest, diversity values remained low as the dominant species Chiasmia fuscataria accounted for far higher proportions than other dominant species in any of the other habitats.     

New and little known phalangopsinae (Orthoptera,Gryllidae): 7. Neotropical taxa of the tribes Paragryllini and Luzarini

Three new genera, one new subgenus, and 13 new species (Strogulomorpha separata sp. n., S. davidi sp. n., S. proxima sp. n., Luzarida lata sp. n., Luzara venado sp. n., L. sapani sp. n., Ucayacla pulchella gen. et sp. n., Peruzara atalaya gen. et sp. n., P. loreto sp. n., Amazonacla imitata gen. et sp. n., A. primitiva sp. n., Leptopsis (Leptopsis) ucayali sp. n., and L. (Aberracla subgen. n.) morona sp. n.) are described. The composition of the tribes Paragryllini and Luzarini and of the genera Luzarida Heb. and Luzara Walk is discussed. The new synonymies Luzarida Hebard, 1928 = Ecuazarida Nischk et Otte, 2000, syn. n.; Lerneca Walker, 1869 = Doposia Otte et Perez-Gelabert, 2009, syn. n.; Nemoricantor Desutter-Grandcolas et Hubbell, 1993 = Kumalorina Otte et Perez-Gelabert, 2009, syn. n. are given. The systematic position of some previously described species is corrected.     

Biogeography, ecology, acoustics and chromosomes of East African Eurycorypha Stål species (Orthoptera, Phaneropterinae) with the description of new species

Although Eurycorypha is the most species-rich Phaneropterinae genus in Africa, little is known about the distribution and the ecology of the species. In this study data on distribution, ecology, song and on chromosomes of some East African species are provided. The nymphs of some species are shown, famous for their ant-like appearance and behaviour. The male of E. punctipennis Chopard and three species of Eurycorypha are newly described. These are E. resonans n. sp. and E. combretoides n. sp. occurring in different habitats on Mount Kilimanjaro, and E. conclusa n. sp from forest habitats along the Tanzanian coast. Bioacoustically, the four recorded species are unusually divers, presenting non-resonant and resonant songs. As four Eurycorypha species occur syntopically on geological young Mount Kilimanjaro, the biogeographical pattern and possible speciation mechanisms in the genus Eurycorypha are discussed in context of the climatical history of eastern Africa.     

Remarkable Rhinophoridae in a growing generic genealogy (Diptera: Calyptratae,Oestroidea)

Four new genera (Apomorphyto gen.n. from Costa Rica, Bixinia gen.n. from Australia, Rhinodonia gen.n. from New Caledonia, Rhinopeza gen.n. from Papua New Guinea) and nine new species (Apomorphyto inbio sp.n. , Bixinia collessi sp.n. , B. solitaria sp.n. , B. spei sp.n. , B. variabilis sp.n. , B. winkleri sp.n. , Rhinodonia antiqua sp.n. , R. flavicera sp.n. , Rhinopeza gracilis sp.n.) of Rhinophoridae (Diptera: Calyptratae, Oestroidea) are described. All new species were included in a morphology‐based phylogenetic analysis to provide arguments for the justification and monophyly (when nonmonotypic) of the new genera and for including these in the Rhinophoridae. The New Caledonian Rhinodonia is a candidate sister taxon to all other rhinophorids, and the Australasian ‘axiniine’ species emerge inside a clade of all Neotropical taxa thus suggesting migration from South America across Antarctica into Australia. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:51C1F448‐DDD0‐4F14‐8173‐B8C687F7E841 .     

Diverging diversity patterns of vascular plants and geometrid moths during forest regeneration on Mt Kilimanjaro, Tanzania

Aim This study investigates diversity patterns of vascular plants and plant‐feeding geometrid moths during montane rain forest regeneration in relation to the biogeographical and historical conditions of Mt Kilimanjaro. Location Investigations were undertaken on the south‐western slopes of Mt Kilimanjaro at altitudes between 2075 and 2265 m. Methods Thirteen plots were selected for this study. Four of these were situated in the middle of large clearings (> 1000 m²), three in secondary forest, two in mature forest remnants surrounded by secondary forest and four plots within continuous closed mature forest. Vascular plant species were recorded in an area of 20 × 20 m². Geometrid moths were attracted using lamps placed inside reflective gauze cylinders. Results Ninety‐three species of vascular plants were recorded on the plots. Plant diversity increased in the course of forest regeneration from clearings and secondary forest to mature forest remnants and mature forest. This increase was visible in all vegetation strata as well as in the species number of Dicotyledoneae. The diversity of geometrid moths conversely decreased from early to late successional stages. A total of 2276 Geometridae representing 114 morphospecies were included in the study. Local values of Fisher's α varied from 10.3 to 18.3 on clearings and in secondary forest, whereas they remained below 8.0 in mature forest and mature forest remnants. There was a significant negative correlation between the diversity of Geometridae and the number of dicots, and of plant species in the shrub layer. Main conclusions Contrary to an expected positive correlation between the diversity of vascular plants and herbivorous geometrid moths, diversity patterns of these two groups are strongly diverging due to biogeographical and ecological factors differently affecting the two groups. The increase in plant diversity can chiefly be explained with an increase in epiphyte diversity which is related to the occurrence of suitable habitats in extensive moss layers on huge Ocotea usambarensis (Engl.) trees in the mature forest. The low diversity of geometrid moths in these forests may be connected to the isolation and relatively young age of the montane rain forests on Mt Kilimanjaro. Hence only a small number of moth species adapted to the cool and perhumid conditions within moist mature forest have so far immigrated into these habitats, and time was insufficient for the evolution of many new species.     

Small but not ephemeral: newly discovered species of Aphelinidae and Trichogrammatidae (Insecta: Hymenoptera: Chalcidoidea) from Eocene amber

New species of fossil Aphelinidae and Trichogrammatidae are described from middle Eocene (Lutetian) Baltic amber (41.3–47.8 Ma). A new subfamily, two new genera and three new species of Aphelinidae are described, with comments on their placement: Phtuaria fimbriae gen.n. , sp.n. in Phtuariinae subf.n. , Glaesaphytis interregni gen.n. , sp.n. and Centrodora brevispinae sp.n. These represent the first described true fossil Aphelinidae. Four new species of Trichogrammatidae are described: Mirufens illusionis sp.n. , Palaeogramma eos gen.n. , sp.n. , Pterandrophysalis plasmans sp.n. and Szelenyia terebrae sp.n. , thus expanding our knowledge of fossil Trichogrammatidae beyond the single previously described species. The presence of recognizable extant genera of Aphelinidae and Trichogrammatidae in the Eocene suggests that the morphology of these genera has been relatively invariant despite highly variable conditions during and since the Eocene. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:E9AD60B6‐3D56‐4E74‐AA54‐F7B91F4FDC79 .     

Discheramocephalini,a new pantropical tribe of featherwing beetles (Coleoptera: Ptiliidae): description of new taxa and phylogenetic analysis

Abstract Two new genera and eight new species of featherwing beetles (Coleoptera: Ptiliidae) possessing a remarkable horizontal perforation of the mesoventral keel are described: Skidmorella vietnamensis sp.n. (Vietnam), S. memorabilis sp.n. (Indonesia), S. serrata sp.n. (Vietnam), Fenestellidium capensis gen. et sp.n. (South Africa, type species), F. kakamegaensis sp.n. (Kenya), Cissidium okuensis sp.n. (Cameroon), Dacrysoma usambarensis gen. et sp.n. (Tanzania, type species) and D. felis sp.n. (Madagascar). A phylogenetic analysis of 24 taxa and 37 parsimony‐informative characters supports the hypothesis of a single origin of the mesoventral perforation, thus uniting Discheramocephalus, Skidmorella, Africoptilium, Fenestellidium, Cissidium and Dacrysoma into a pantropically distributed clade, for which a new tribe Discheramocephalini (type genus Discheramocephalus) is proposed. Identification keys to Discheramocephalini genera and, in some cases, to species are provided. Each new species is illustrated with scanning electron microscopy images.     

Chromosome variability in the Lemuridae

Chromosome studies have been conducted in Lemur catta, L. macaco, L. mongoz, L. fulvus fulvus, and a hybrid L. fulvus fulvus x L. fulvus albifrons. Comparative analysis shows that inter- and intraspecific chromosome variability is a common finding in Lemurs. The Lemuridae are divided into three groups according to the characteristics of their chromosome complement. A reclassification of L. catta with the Hapalemurs is suggested, based on chromosomal and nonchromosomal data.     

The high‐Andean Jivarus Giglio‐Tos (Orthoptera,Acridoidea, Melanoplinae): systematics,phylogenetic and biogeographic considerations

The high‐Andean genus Jivarus Giglio‐Tos from Ecuador, Colombia and Peru is revised. Morphological cladistic analysis indicated that Jivarus montanus and the new species digiticercus sp.n. and rugosus sp.n. must be treated as a separate genus, Maylasacris gen.n. The remaining species included in the analysis are assigned to the genus Jivarus, for which the following six species groups are identified: americanus group, antisanae group, carbonelli group, cohni group, pictifrons group and jagoi group. Twenty‐nine species are recognized for Jivarus, with ten described as new: J. rectus sp.n. , J. megacercus sp.n. , J. spatulus sp . n. , J. auriculus sp.n. , J. riveti sp.n. , J. sphaericus sp.n. , J. discoloris sp . n. , J. profundus sp.n. , J. ronderosi sp.n. and J. guarandaensis sp.n. The following new synonymies are proposed: Jivarus albolineatus Ronderos with J. antisanae (Bolivar) syn.n. , J. cerdai Ronderos and J. osunai Ronderos with J. alienus (Walker) syn.n. , and J. rubriventris Ronderos with J. ecuadorica (Hebard) syn.n. ; the new combinations Jivarus ecuadorica (Ronderos) comb.n. and Maylasacris montanus (Ronderos) comb.n. are proposed. Keys to the species of the genera and a review of the morphological characters defining the taxa are provided. Patterns of distribution of the clades coincide with the geography of the northern Andes of Colombia and Ecuador. Areas of endemism of the Jivarus species groups and Maylasacris are delimited by both the high‐altitude curves, including transverse zones, and the drier climates of the intra‐Andean valleys, clearly indicating recent, post‐glacial palaeogeography, as shown also in vegetation distributions. This paper has been formatted with many embedded links to images of type and paratype specimens, maps based on geo‐referenced specimen data and species keys available on the Orthoptera Species file online ( http://orthoptera.speciesfile.org ).