Beata Grzywacz

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.

Changes in the numbers of chromosomes and sex determination system in bushcrickets of the genus Odontura (Orthoptera: Tettigoniidae: Phaneropterinae)

Chromosomes of the males of five species of Odontura, belonging to the subgenera Odontura and Odonturella, were ana- lyzed. Intensive evolution of the karyotype was recorded, both in terms of changes in the numbers of chromosomes (from 2n = 31 to 27) and the sex chromosome system (from X0 to neo-XY and X0 to neo-X1X2Y). Karyotype evolution was accompanied by tandem autosome fusions and interspecific autosomal and sex chromosome differentiation involving changes in the locations of nucleolar organizer regions, NORs, which were revealed by silver impregnation and confirmed by FISH using an 18S rDNA probe. O. (Odon- turella) aspericauda is a polytypic species with X0 and neo-X1X2Y sex determination. The latter system is not common in tettigo- niids. It possibly originated by a translocation of a distal segment of the original X chromosome onto a medium sized autosome, resulting in a shortened neo-X1 and a metacentric neo-Y. The remaining autosome homologue became the neo-X2 chromosome. This shift from X0 to neo-X1X2Y is supported by the length of the X chromosome and location of the NOR/rDNA.

A comparative study of genome organization and inferences for the systematics of two large bushcricket genera of the tribe Barbitistini (Orthoptera: Tettigoniidae: Phaneropterinae)

BackgroundPoecilimon and Isophya are the largest genera of the tribe Barbitistini and among the most systematically complicated and evolutionarily intriguing groups of Palearctic tettigoniids. We examined the genomic organization of 79 taxa with a stable chromosome number using classical (C–banding, silver and fluorochrome staining) and molecular (fluorescence in situ hybridization with 18S rDNA and (TTAGG)n telomeric probes) cytogenetic techniques. These tools were employed to establish genetic organization and differences or similarities between genera or species within the same genus and determine if cytogenetic markers can be used for identifying some taxonomic groups of species.ResultsDifferences between the karyotypes of the studied genera include some general changes in the morphology of the X chromosome in Isophya (in contrast to Poecilimon). The number of major rDNA clusters per haploid genome divided Poecilimon into two main almost equal groups (with either one or two clusters), while two rDNA clusters predominated in Isophya. In both genera, rDNA loci were preferentially located in the paracentromeric region of the autosomes and rarely in the sex chromosomes. Our results demonstrate a coincidence between the location of rDNA loci and active NORs and GC-rich heterochromatin regions. The C/DAPI/CMA3 bands observed in most Poecilimon chromosomes suggest the presence of more families of repetitive DNA sequences as compared to the heterochromatin patterns in Isophya.ConclusionsThe results show both differences and similarities in genome organization among species of the same genus and between genera. Previous views on the systematics and phylogenetic grouping of certain lineages are discussed in light of the present cytogenetic results. In some cases, variation of chromosome markers was observed to correspond with variation in other evolutionary traits, which is related to the processes of ongoing speciation and hybridization in zones of secondary contact. It was concluded that the physical mapping of rDNA sequences and heterochromatin may be used as an additional marker for understanding interspecific relationships in these groups and their routes of speciation.

Bacterial symbionts of the leafhopper Evacanthus interruptus (Linnaeus, 1758) (Insecta, Hemiptera, Cicadellidae: Evacanthinae).

Plant sap-feeding hemipterans harbor obligate symbiotic microorganisms which are responsible for the synthesis of amino acids missing in their diet. In this study, we characterized the obligate symbionts hosted in the body of the xylem-feeding leafhopper Evacanthus interruptus (Cicadellidae: Evacanthinae: Evacanthini) by means of histological, ultrastructural and molecular methods. We observed that E. interruptus is associated with two types of symbiotic microorganisms: bacterium ‘Candidatus Sulcia muelleri’ (Bacteroidetes) and betaproteobacterium that is closely related to symbionts which reside in two other Cicadellidae representatives: Pagaronia tredecimpunctata (Evacanthinae: Pagaronini) and Hylaius oregonensis (Bathysmatophorinae: Bathysmatophorini). Both symbionts are harbored in their own bacteriocytes which are localized between the body wall and ovaries. In E. interruptus, both Sulcia and betaproteobacterial symbionts are transovarially transmitted from one generation to the next. In the mature female, symbionts leave the bacteriocytes and gather around the posterior pole of the terminal oocytes. Then, they gradually pass through the cytoplasm of follicular cells surrounding the posterior pole of the oocyte and enter the space between them and the oocyte. The bacteria accumulate in the deep depression of the oolemma and form a characteristic ‘symbiont ball’. In the light of the results obtained, the phylogenetic relationships within modern Cicadomorpha and some Cicadellidae subfamilies are discussed.

Review of the Plangia graminea (Serville) complex and the description of new Plangia species from East Africa (Orthoptera: Phaneropteridae, Phaneropterinae) with data on habitat, bioacoustics, and chromosomes

The Plangia graminea complex is partly reviewed and two new species of Plangia are described, Plangia multimaculata n. sp. from savanna habitats and Plangia satiscaerulea n. sp. from the submontane zones in northern Tanzania. Plangia compressa (Walker 1869) is synonymized with P. graminea (Serville, 1838). Data on habitat, bioacoustics, and chromosomes are provided. Both analyzed Plangia species showed the same chromosomal number. Compared to other investigated African Phaneropterinae, Plangia had a reduced chromosome number and thus shows a derived condition in its genome. The sex chromosomes differed morphologically in both investigated species of Plangia suggesting different mechanisms leading to these differences. Niche specialization and the male calling song are discussed as drivers for speciation in fully alate and thus mobile taxa.

The Anatolio‐Balkan phylogeographic fault: a snapshot from the genus Isophya (Orthoptera, Tettigoniidae)

Due to active tectonic evolution of the Aegean Area during Miocene and Pleistocene, the Balkans and Anatolia have repeatedly connected and disconnected causing isolation and secondary contact of populations along the present Dardanelles – Sea of Marmara – Bosphorus waterway. This has led to an outstandingly rich fauna and a reticulate biogeography in the area. A typical example is Orthoptera having here possibly highest diversity within the Western Palaearctic. With the present study, we concentrate on the bush‐cricket genus Isophya, which is characteristic with a large share of endemics in the Balkans and Anatolia. We aim to understand when and how the isolation of marine or other barriers in the region of the Turkish Straits System influenced the evolution of the morpho‐acoustic groups of species found on both sides of the strait. For this purpose, sequences of two mitochondrial (COI and ND2) and two nuclear (ITS1 and ITS2) markers were obtained and used for phylogenetic reconstructions, time estimations for lineage divergence and automatic species delineation (statistical parsimony, GMYC, ABGD) tests. The phylogenetic analyses did not support all the existing morphogroups and was in favour of a geographical subdivision for the young lineages. Automatic species delineation tests confirmed most of the present morpho‐acoustic species and suggested further cryptic species, at the same time unifying some phenetic species. Time estimation analyses suggested time to most recent common ancestor of the genus as 8.28 Ma corresponding to the Late Tortonian. As a result of the study, we reached to the following conclusions: (i) most of the studied phenetic species are monophyletic, but some earlier suggested morpho‐species groups are not, (ii) three main episodes dominate the evolutionary history of Isophya (7.32–5.84 Myr in the Messinian, 2.2–2.5 Myr in the beginning of the Pleistocene and at around 0.75 Ma corresponding to the end of the Mid‐Pleistocene transition), all of which well correlate with episodes of existing terrestrial connections between Anatolia and the Balkans, and (iii) there are several faunal exchanges in both directions between Anatolia and the Balkans.

Review of the East African species of the phaneropterine genus Parapyrrhicia Brunner von Wattenwyl, 1891 (Insecta: Orthoptera): secret communication of a forest-bound taxon

The East African species of Parapyrrhicia Brunner von Wattenwyl, 1891, are reviewed. Beside the description of five new species from the West Usambara, Udzungwa, Uluguru Mountains, and coastal Tanzania and the male of P. zanzibarica Brunner von Wattenwyl, 1891, data on habitat, morphology, acoustics, and the chromosomes are provided. A key to the species of Parapyrrhicia is provided. Eulioptera zanzibarica Brunner von Wattenwyl, 1891, is synonymized with Parapyrrhicia zanzibarica Brunner. All species have stridulatory files with intertooth intervals decreasing from anal to basal and low duty cycle calling songs (four species recorded) consisting of very short, narrow-banded, resonant syllables. Parapyrrhicia species showed a plesiomorphic karyotype with a diploid chromosome number of 2n = 31 and a X0 sex chromosome system in males. All species are restricted to closed wet forest areas and thus ideal organisms to be used for monitoring habitat quality of an area since they act as indicators for indigenous wet forest communities. Morphological sister pairs suggest recent speciation processes in coastal and lowland wet forests of East Africa.

Relationship between Chromosomal Races/Subraces in the Brachypterous Grasshopper Podisma sapporensis (Orthoptera: Acrididae) Inferred from Mitochondrial ND2 and COI Gene Sequences

Abstract Podisma sapporensis Shiraki, 1910 (Orthoptera: Acrididae) is distributed on the islands of northern Japan and Far East Russia (Hokkaido, Sakhalin, and Kunashiri), and it exhibits a unique diversity of chromosomal races (X0 and neo-XY) on the island of Hokkaido. To infer the history of geographical isolation and chromosome rearrangements (main races and subraces), we used mitochondrial DNA (mtDNA) sequence variation in two regions, NADH dehydrogenase subunit 2 (ND2) and cytochrome oxidase subunit II (COII), among 55 individuals representing eight X0/XX and three neo-XY/XX populations. The molecular phylogenetic tree revealed a high level of overall mitochondrial diversity, but the clustering of the examined population is in most cases closely related to their geographic distribution and associated with chromosomal races and subraces. Together with cytogenetic observations, we discuss the origin of the polymorphism within the X0 and XY races as well as differences between both northern and eastern races.

Topography and climatic fluctuations boosting speciation: biogeography and a molecular phylogeny of the East African genera Afroanthracites Hemp & Ingrisch and Afroagraecia Ingrisch & Hemp (Orthoptera, Tettigoniidae, Conocephalinae, Agraeciini)

Based on faunistic lists documenting the distribution of taxa and molecular data, a phylogenetic tree of African Agraeciini is presented and detailed hypotheses of speciation patterns discussed. It is discussed that the observed radiation in the geologically old Eastern Arc chain is young since Afroanthracites montium endemic to Mts. Kilimanjaro and Meru is of the same age as species of the East and West Usambara Mountains. A molecular phylogeny on nine Afroanthracites and four Afroagraecia species is presented in this study prepared on the molecular markers 16S rRNA and histone 3. The molecular results confirmed geographical patterns and morphological relationships in Afroanthracites as well as in Afroagraecia.

New genera and new species of Acrometopini (Orthoptera: Tettigonioidea Phaneropterinae) from East Africa and a review of all known stridulatory organs, songs and karyotypes of the tribe

Two new genera of East African Acrometopini are erected: the monotypic Peronurella n. gen. with P. centralis n. sp. from Miombo woodlands in central Tanzania, and Tenerasphaga n. gen. for a morphological uniform group of species formerly included in Horatosphaga, T. chyuluensis n. sp., T. mbulu n. sp. T. nanyuki, T. nuda n. comb, T. meruensis n. comb, T. tenera n. comb and T. bazeletae n. comb. Additional species are described in the genera Altihoratosphaga, Lamecosoma and Peronura. Keys to African genera of Acrometopini and species of the genus Tenerasphaga are provided. We review the male calling songs and the morphology of the acoustical apparatus and discuss how the morphology of the stridulatory files explains the complicated song patterns and elements. All Acrometopini species showed a plesiomorphic karyotype with a diploid chromosome number of 2n = 31 and a X0 sex chromosome system in males.