Host associations and origin in the formation of the Caucasian fauna of fleas (Siphonaptera)

Results of analysis of the Caucasian fauna of fleas and their association with mammal and avian hosts are reported. The Caucasian fauna of potential flea hosts comprises about 130 species of mammals and about 470 species of birds. Most of the flea species in the Caucasian fauna (88 out of 155) parasitize rodents, 51 species of which are permanent hosts of different flea species; 13 flea species occur on 11 species of insectivores; 13 flea species, on 13 species of chiropterans; 14 flea species, on 20 species of carnivores. Only 2 flea species parasitize artiodactyles. 54 species of birds are permanent hosts of 23 species of fleas from 4 genera in the Caucasus. Ten types of ranges of flea species are distinguished; host associations of the Caucasian flea species from these groups are discussed. The greatest numbers of hosts from the families Cricetidae, Muridae, and Sciuridae are associated with fleas with Euro-Asian (extra-Siberian), European, Turanian, and Iranian ranges. Soricidae are known as hosts of flea species with European and Euro-Turanian ranges. Four major groups of flea taxa are represented in the Caucasian fauna. The distribution of the first group is determined by the influence of the palaeofauna of the ancient European continent in the early Cenozoic; that of the second group, by the influence of the fauna of the ancient Asian continent during the Paleogene and part of the Neogene; the third, by the influence of the fauna of southern Europe starting with the Miocene. The fourth group comprises the species which immigrated from northern Europe and Asia in the Late Neogene (2–3 mln years ago).     

Patterns of formation of the flea (Siphonaptera) fauna in the Caucasus

Fleas of the Caucasus belong to 155 species of 40 genera, constituting 17% and 43% of the species and generic composition of the Palaearctic fauna, respectively. The Caucasian fauna includes 23 endemic species but no endemic genera or subgenera. In the number of species, the Caucasian fauna is similar to that of the Mediterranean Subregion and is significantly poorer than the faunas of the Euro-Siberian (by 2.2 times) and Irano-Turanian (by 1.7 times) Subregions. Based on taxonomic diversity, we can propose a hypothesis on the West and East Palaearctic sources of the Caucasian fauna. The West Palaearctic source has determined the distribution of pulicomorph fleas of the families Pulicidae and Coptopsyllidae from Africa, on the one hand, and of fleas of the genera Ctenopthalmus and Palaeopsylla from Europe, on the other hand. Fleas of the Holarctic genera, such as Ceratophyllus and Megabothris, entered the Caucasus by the north Asian route; fleas of the genera Neopsylla, Rhadinopsylla, and Hystrichopsylla migrated to the Caucasus from east and central Asia by the south Asian route, through Middle and Western Asia.     

Taxonomic composition and zoogeographic characteristics of the flea fauna (Siphonaptera) of Russia

255 species and 59 subspecies of fleas from 55 genera of 7 families are known from Russia, which is 30% of the Palaearctic fauna. Additionally, over 187 species of 47 genera from 7 families are known from the neighboring territories of Central and Southern Europe, Transcaucasia, Kazakhstan, Middle Asia, Mongolia, Northeast China, and Japan. 13 species of 12 genera are known only from Russia. Noteworthy is the low percent of endemic species (not more than 4%) and genera (one genus) in the Russian fauna. The principal centers of taxonomic diversity in the Palaearctic, including many endemic species and genera, lie in the Eastern Asian, Central Asian, and Turano-Iranian Subregions, outside Russia and the Euro-Siberian Subregion. The bulk of the Russian fauna is formed by the species and genera of the three largest flea families: Hystrichopsyllidae, Ceratophyllidae, and Leptopsyllidae. The family Ceratophyllidae has the greatest number of genera in the Russian fauna, and Hystrichopsyllidae, the greatest number of species. Western (Western and Western-Central Palaearctic; 84 species from 41 genera of 7 families) and Eastern (Central-Eastern and Eastern Palaearctic; 78 species from 42 genera of 6 families) species are nearly equally represented in the Russian fauna.     

At least some meiofaunal species are not everywhere. Indication of geographic,ecological and geological barriers affecting the dispersion of species of Ototyphlonemertes (Nemertea,Hoplonemertea)

Most meiofaunal species are known to have a broad distribution with no apparent barriers to their dispersion. However, different morphological and/or molecular methods supported patterns of diversity and distribution that may be different among taxa while also conflicting within the same group. We accurately assessed the patterns of geographic distribution in actual genetic species of a marine meiofaunal animal model: Ototyphlonemertes. Specimens were collected from several sites around Europe, Northern and Central America, Southern America, Pacific Islands and Asia. We sequenced regions of two mitochondrial and two nuclear genes. Using single‐gene, a concatenated data set, multilocus approaches and different DNA taxonomy methods, we disentangled the actual diversity and the spatial structures of haplotypes and tested the possible correlation between genetic diversity and geographic distance. The results show (i) the importance of using several genes to uncover both diversity and highlight phylogeographic relationships among species and that (ii) independent genetic evolutionary entities have a narrower distribution than morphological species. Moreover, (iii) a Mantel test supported a positive correlation between genetic and geographical distance. By sampling from the two sides of Isthmus of Panama, we were additionally able to identify lineage divergence times that are concordant with vicariance mechanisms caused by the geological closure of the seaway across the Isthmus. We therefore propose that in addition to distance, other geological and ecological conditions are also barriers to the dispersion of and gene flow in marine meiofaunal organisms.     

On the Distribution and Origin of Salix in the World

1. The distribution of Salix species among the continents. There are about 526 species of Salix in the world, most of which are distributed in the Northern Hemisphere with only a few species in the Southern Hemisphere. In Asia, there are about 375 species, making up 71.29 percent of the total in the world, including 328 endemics; in Europe, about 114 species, 21.67 percent with 73 endemics; in North America, about 91 species, 17.3 percent with 71 endemics; in Africa, about 8 species, 1.5 percent, with 6 endemics. Only one species occurs in South America. Asia, Europe and North America have 8 species in common (excluding 4 cultivated species). There are 34 common species between Asia and Europe, 14 both between Europe and North America and between Asia and North America, 2 between Asia and Africa. Acording to the Continental Drift Theory, the natural circumstances which promoted speciation and protected newly originated and old species were created by the orogenic movement of the Himalayas in the middle and late Tertiary. Besides, the air temperature was a little higher in Asia than in Europe and North America (except its west part) and the dominant glaciers were mountainous in Asia during the glacial epoch in the Quaternary Period. Then willows of Europe moved southwards to Asia. During the interglacial period they moved in opposite direction. Such a to-and-fro willow migration between Asia and Europe and between and North America occurred so often that it resulted in the diversity of willow species in Asia. Those species of willows common among the continents belong to the Arctic flora. 2. The multistaminal willows are of the primitive group in Salix. Asia has 28 species of multistaminal willows, but Europe has only one which is also found in Asia. These 28 species are divided into two groups, “northern type” and “southern type”, according to morphology of the ovary. The boundary between the two forms in distribution is at 40°N. The multistaminal willows from south Asia, Africa and South America are very similar to each other and may have mutually communicated between these continents in the Middle or Late Cretaceous Period. The southern type willows in south Asia are similar to the North American multistaminal willows but a few species. The Asian southern type willows spreaded all over the continents of Europe, Asia and North America through the communication between them before the Quaternany Period. Nevertheless, it is possible that the willows growing in North America immigranted through the middle America from South America. The Asian northern type multistaminal willows may have originated during the ice period. The multistaminal willows are more closed to populars in features of sexual organs. They are more primitive than the willows with 1-3 stamens and the most primitive ones in the genus. 3. The center of origin and development of willows Based on the above discussion it is reasonable to say that the region between 20°-40°N in East Asia is the center of the origin and differentiation of multistaminal willows. It covers Southern and Southwestern China and northern Indo-China Pennisula.     

The ecology and prevalence of arboviruses on the territory of the Saratov Region

The territorial spread of Tahyna, Batai, Sindbis, West Nile fever and Crimean-Congo hemorrhagic fever viruses throughout the Saratov region in 1998 - 2000 was analyzed. The characteristics of the epizootic activity of the natural foci of these arboviruses in different landscape zones (temperate forest-steppes, steppes and semi-deserts) were calculated. The species composition of small mammals, the natural reservoirs of the causative agents of arbovirus infections, was determined.     

Fossil representatives of the family Greenideidae (Hemiptera, Aphidoidea) from the Miocene of Europe

Four new aphid species (Hemiptera, Aphidoidea, Greenideidae) are described on the basis of imprints in oil-shales from the Miocene of Europe (Rubielos de Mora, Spain; Vishnevaya Balka ?Stavropol?, Russia): Eutrichosiphum europaeum nov. sp., Greenidea hispanica nov. sp., Greenideoida (Pentatrichosiphum) turolensis nov. sp. and Mollitrichosiphum rubusensis nov. sp. All the taxa are placed within recent genera of the subfamily Greenideinae. Miocene representatives of this subfamily have been found in the south of Europe. Later changes of climatic conditions limited its geographic range to south-eastern Asia, where it is now represented by over 130 species of seven genera.     

Evolutionary and biogeographical patterns within Iberian populations of the genus Squalius inferred from molecular data

Origins and biogeography of the chub, genus Squalius (formerly within the genus Leuciscus), in the Iberian Peninsula were inferred from comparison between patterns of geographic distribution and phylogenetic relationships among populations belonging to 14 European Squalius species. The phylogeny recovered was based on the complete sequence of the mitochondrial cytochrome b gene. Squalius species were grouped into three major clades. The basal clade included species distributed across the ancient Paratethys Sea. The second clade included species from Central and East Europe and the northern areas of the Mediterranean basin towards Minor Asia. The third clade included species from the Mediterranean Peninsulas (Iberian, Italy, and Balkans). The Iberian Squalius species do not constitutes a monophyletic group. Our data indicate that the Iberian Peninsula was colonized at least twice by two different monophyletic lineages, a meridional group and a Central Europe group. The amount of species diversity found in the Iberian Peninsula and the phylogenetic relationships among these species, together with their geographic distribution, suggest that the Central Europe lineage colonized the Iberian Peninsula at a latter time. Our data indicate that the northeastern Iberian lineage is phylogenetically close to Greek populations of Squalius cephalus, while the second lineage formed a monophyletic group including Squalius pyrenaicus, Squalius carolitertii, Squalius aradensis, and Squalius torgalensis. The speciation process that generated these species and the geographic structure of their populations, principally in S. pyrenaicus, can be attributed to paleogeographical events like the ancient endorrheism and the development of hydrographic basins.     

The Palaearctic steppe biome: a new synthesis

The Palaearctic steppes range from the Mediterranean basin towards China, forming one of the largest continuous terrestrial biomes. The literature on steppe ecology and conservation is vast but scattered and often not available in English. We provide a review of some key topics based on a new definition of steppes, which includes also Mediterranean steppes and alpine rangelands of the Asian Highlands. Revisiting the terrestrial ecoregions of the world, we estimate that the Palaearctic steppe biome extends over ca. 10.5 million km². Major chorological regions differ in their macroclimatic niche with a clear distinction between Middle Asia with its winter precipitation and the Central Asian summer-rain regions of the Mongolian plateau and of Tibet. Steppe soils store large amounts of carbon, yet the sequestration potential is debated and depends on land use. Major physiognomic-ecological steppe types include forest-, typical-, desert-, and alpine-steppe, which vary in the importance of grasses, mainly C3 species. The steppes host a specialised fauna, and Middle Asia, Tibet, and especially Mongolia, have large herds of migrating ungulates. The share of pristine and protected sites is low in the steppe regions, with conversion into croplands being the most important land use impact in Europe, Middle Asia, and the Mediterranean, while grazing has a severe impact in some parts of Mongolia and Tibet. There are major gaps in our knowledge on: (1) the effects of climate change on the crucial seasonal patterns; (2) the role of steppe soils in the global carbon budget; and (3) the ecology and distribution of most animal groups except vertebrates.     

The geographical parallels of lithorheophil biotic community organisation of small streams of eastern Europe and northern Asia

The influence of geographical position on taxonomical composition and structure of communities inhabited stony riffles is considered on a material from 79 small streams situated in four regions of Eastern Europe (Moscow region, Southern Ural, Northern Caucasus and Mountain Crimea), as well as on publishing data on Asia and Northern Ural regions. Litoreophil communities of mentioned regions include different but closely allied species with similar habits. Eight basic living forms of litoreophils are common practically everywhere irrespective of geographical position and taxonomical composition of the community. The average number of species in a sample is also constant and includes 13-14 species. Local relief and climate influence the abundance of living forms and species diversity.     

Genetics of Caucasian population: distribution of various immunological and biochemical markers in western Georgia

The distribution of genetic markers of blood groups (ABO, Rhesus, MNSs, P, Duffy, Kell-Cellano), plasma proteins (Hp, Gc, Tf, C'3) and red-cell enzymes (AcP, EstD, GLO-1), and also ABH secretion among 10 populations of Western Georgia has been studied. The common characteristic of distribution of gene frequencies for the markers studied was obtained as a whole in Georgia. The Georgians were compared for these markers with some populations of the Caucasus, Europe and West Asia. Among Caucasian populations, Georgians are most similar to Abkhasians. According to some systems, Georgians are close to European groups (ABO, Dubby, GLO-1, EstD), while they are similar to West-Asian groups, as judged by other systems (ABH secretion, AcP). According to Rhesus and MNSs systems, Georgians differ both from populations of Europe and from populations of West Asia.     

Towards resolving the evolutionary history of Caucasian pears (Pyrus,Rosaceae)—Phylogenetic relationships,divergence times and leaf trait evolution

With approximately 25 endemic species, the genus Pyrus (pears) is highly diverse in the Caucasus ecoregion. The majority of Caucasian pears inhabit xerophytic open woodlands or similar habitats, to which they display morphological adaptations, such as narrow leaves. The other species, both Caucasian and non‐Caucasian taxa, mainly inhabit mesophytic forests and display broad leaves. Using a representative taxon sampling of Pyrus from the Caucasus, Europe and Asia, we reconstruct phylogenetic relationships in the genus based on multiple plastid regions. We also estimate the divergence times of major clades in Pyrus, reconstruct the evolution of leaf shapes, and discuss the emergence of xeromorphic leaf traits. Our results confirm the monophyly of Pyrus and the existence of two major clades: (a) an E Asian clade with a crown group age of 15.7 (24.02–8.37 95% HPD) My, and (b) a W Eurasian clade that comprises species from Europe, SW Asia and the Caucasus and that displays a slightly younger crown group of 12.38 (19.02–6.41 95% HPD) My. The existing infrageneric classification of Pyrus was found partially incongruent with the inferred phylogenetic trees. Several currently accepted species were not recovered as monophyletic, indicating that current species limits require re‐evaluation. Ancestral character state reconstructions revealed several independent transitions from broad‐ to narrow‐shaped leaves in Pyrus, probably via intermediate‐shaped leaves.     

The ants of the Chuya depression and the Yustyd River basin in Southeastern Altai

The ant fauna of the Chuya depression numbers 17 species, of which four species inhabit desert steppes, eight species occur in the near-water saline steppes, and eleven, in valley meadows and forests on mountain slopes. Of the 17 registered species, 14 are common in adjacent Mongolia and Tuva, which testifies to a high faunal commonness of these territories. The population pattern of ants is determined by two species with very low abundance (up to 0.01 nest/100 m²), Proformica mongolica Ruzsky and Cataglyphis aenescens Nylander, and in shallow hollows Formica subpilosa Ruzsky is added to them. Two zones with a higher density of population have been revealed in the vertical transzonal direction, in alpine desert steppes on the southern slopes (Yustyd River basin) and in the saline biotopes of the Chuya River floodplain.     

Step(pe) up! Raising the profile of the Palaearctic natural grasslands

Palaearctic steppes are primary grasslands dominating the landscape of the Eurasian Grassland Belt from Central and Eastern Europe to Northern China across the temperate zone of Eurasia. We also include structurally and floristically similar habitats in North Africa, Anatolia, and Iran. The biota of the steppes are diverse, including many endemic species. As a result of the high rate of anthropogenic conversion and widespread degradation, the Palaearctic steppes have become one of the most endangered terrestrial biomes of the world. These facts underline the importance of sustaining landscape-scale biodiversity in steppes and stress the necessity of their conservation and restoration. Literature about the ecology, biodiversity, and conservation of Palaearctic steppes is not easily accessible for an international audience. Therefore, summarising the current state of knowledge as well as knowledge gaps is very timely. This Special Issue on “Palaearctic steppes: ecology, biodiversity and conservation”, comprises 17 research papers from many different regions throughout the biome, as well as a broad review synthesising current knowledge.     

Modern analogues from the Southern Urals provide insights into biodiversity change in the early Holocene forests of Central Europe

Aim The diversity changes that occurred in Central European forests during the early Holocene can be better understood using ecological knowledge of modern analogues of these forests, which occur in far Eastern Europe. Here we compare the diversity of vascular plants, bryophytes and snails among different forest types of the Southern Urals to provide insights into the palaeoecology of the vanished Central European forests. Location Southern Ural Mountains, Bashkortostan, Russia. Methods We recorded all species of vascular plants, ground-dwelling bryophytes and land snails, and measured biotic and abiotic variables, in 100-m² plots located in four forest types dominated by different trees: Pinus sylvestris–Larix sibirica, Betula pendula–Populus tremula, Quercus robur and Acer platanoides–Tilia cordata–Ulmus glabra. These types correspond to the chronosequence of forest communities that occurred in Central Europe in the early Holocene. Results The species richness of herb-layer plants was most affected by the canopy-transmitted light. The former three forest types had open canopy and were rich in species, whereas the Acer–Tilia–Ulmus forests were darker and poor in species. The species richness of ground-dwelling bryophytes decreased from coniferous to broad-leaved deciduous forests. In contrast, the highest species richness of snails was in the Acer–Tilia–Ulmus forests owing to the higher calcium content in the litter of these trees. Main conclusions Based on the modern analogue of the Southern Ural forests, we propose the hypothesis that the late-glacial open-canopy pine and larch forests of Central European lowlands were rich in light-demanding species of vascular plants, many of which were also typical of tall-grass steppes or mesic grasslands. They also contained several species of ground-dwelling bryophytes. The spread of birch, aspen and oak in the early Holocene reduced the local species richness of ground-dwelling bryophytes but not of vascular plants. The subsequent spread of elm, lime, maple and ash caused canopy closure, a retreat of the light-demanding herbs and a decline in the local species richness of vascular plants. Besides the increased shading by these tree species, their litter enriched soils in calcium, and enhanced decomposition and nutrient cycling. This supported an increase in the species richness of land snails.     

Bark beetles and pinhole borers (Curculionidae, Scolytinae, Platypodinae) alien to Europe

Invasive bark beetles are posing a major threat to forest resources around the world. DAISIE's web-based and printed databases of invasive species in Europe provide an incomplete and misleading picture of the alien scolytines and platypodines. We present a review of the alien bark beetle fauna of Europe based on primary literature through 2009. We find that there are 18 Scolytinae and one Platypodinae species apparently established in Europe, from 14 different genera. Seventeen species are naturalized. We argue that Trypodendron laeve, commonly considered alien in Europe, is a native species; conversely, we hypothesize that Xyleborus pfeilii, which has always been treated as indigenous, is an alien species from Asia. We also point out the possibility that the Asian larch bark beetle Ips subelongatus is established in European Russia. We show that there has been a marked acceleration in the rate of new introductions to Europe, as is also happening in North America: seven alien species were first recorded in the last decade.We present information on the biology, origins, and distributions of the alien species. All but four are polyphagous, and 11 are inbreeders: two traits which increase invasiveness. Eleven species are native to Asia, six to the Americas, and one is from the Canary Islands. The Mediterranean is especially favorable for invasives, hosting a large proportion of the aliens (9/19). Italy, France and Spain have the largest numbers of alien species (14, 10 and 7, respectively). We point out that the low numbers for at least some countries is likely due to under-reporting.Finally, we discuss the difficulties associated with identifying newly invasive species. Lack of good illustrations and keys hinder identification, particularly for species coming from Asia and Oceania.     

Allozymic and morphometric analysis of populations in the Zygaena purpuralis complex (Lepidoptera, Zygaenidae)

In Central Europe and Asia Minor Zygaena purpuralis and Z. minos are two sibling species exhibiting a considerable geographic variation in several features. In this study we examine some populations from the Central and Southern Apennines (Italy) by comparing multivariate analyses of individual allozymic multilocus profiles and of individual morphometric profiles of male genitalia in order to assess their genetic and phenetic relationships.
Results indicate that populations of Z. purpuralis in the Apennines behave in rather different ways with respect to the occurrence of gene exchange with other gene pools. No gene flow appears to occur between two geographically close southern populations; however, extensive introgression, revealable at both the allozymic and morphometric level, seems to occur between some purpuralis and minos -like sympatric populations from Abruzzi. Evolutionary relationships among these peninsular Italy populations are discussed with reference to historical factors, that is, the occurrence of multiple Pleistocene refugia, which could have determined different degrees of isolation among gene pools. This situation is rather different from that previously outlined for Central Europe and present results suggest that the Z. purpuralis complex might include more than two biological species.     

Ways of the fleas (Siphonaptera) fauna formation in the Caucasus

Fleas fauna of the Caucasus is considered, possible ways of its formation are discussed. Caucasian fleas belong to 155 species and 40 genera; 23 species are endemics. Hypothesis on Western Palearctic and Eastern Palearctic sources of the Caucasian fleas' fauna formation are proposed.     

Les hommes de Néandertal du Caucase du Nord : entre l’Ouest et l’Est

A geographical position of the Caucasus in the border between Europe and Asia defines a complex character of the Middle Paleolithic in the region as a whole and in the Northern Caucasus in particular. Today, we can recognize three major cultural areas existed during the Middle Paleolithic in the Northern Caucasus: (1) a local North Caucasian variant of Eastern Micoquian, which is closely related to Eastern Micoquian of Central and Eastern Europe, in the Northwestern Caucasus; (2) a specific Caucasian industry type (called Khostinian Mousterian), which penetrated during later MIS 3 to the Northwestern Caucasus from the Northeastern Black sea cost in the Southern Caucasus; and (3) a similar to Zagros Mousterian industry, which is presented in Weasel Cave in the Northeastern Caucasus.