Factors influencing the natural reproduction decline in the beluga (Huso huso,Linnaeus, 1758), Russian sturgeon (Acipenser gueldenstaedtii,Brandt & Ratzeburg, 1833), and stellate sturgeon (A. stellatus,Pallas, 1771) of the Volga–Caspian basin: A review

The review describes the changes in natural reproduction of three important sturgeon species in the Volga–Caspian basin: (a) the beluga (Huso huso Linneaus, 1758), (b) the Russian sturgeon (Acipenser gueldenstaedtii Brandt &Ratzeburg, 1833), and (c) the stellate sturgeon (Acipenser stellatus Pallas, 1771). Since the past 60 years, these species responded to severe influences of natural and anthropogenic factors. On the basis of original and published data, an analysis has been made of (a) the numbers of larvae migrating from spawning sites (according to plankton net survey), (b) fecundity and histological anomalies in gonad development, (c) the numbers of adult sturgeons in the Caspian Sea and of spawners migrating to the Volga River (according to trawl and beach seine survey), and (d) foraging resources for the sturgeons. The results show that their natural reproduction in the Volga–Caspian basin has declined drastically during the past decades under the impact of (a) fluctuations of the Caspian Sea level and flow discharge from the Volga River, (b) blockage of sturgeon migration routes and loss of spawning sites because of dam construction, (c) water pollution in the lower reaches of the Volga River and in the Caspian Sea, and (d) intensive and selective illegal and unreported fishing. The relative significance of these factors has been changing during the study period.     

The return of the steppe large-scale restoration of degraded land in southern Russia during the post-Soviet era

Comparing satellite images from 1989 and 1998, we analysed environmental changes in the North Caspian lowland during the period of post-Soviet transformation. Research was carried out in four representative test areas, situated in the Ilmen Lake Region and the adjacent Kalmykien Steppe west of the Volga delta. In all test areas significant changes have taken place during the last decade, most of them induced by the abandonment and extensification of former agricultural land use. Most obvious is the sharp decline of irrigated arable land, to less than 5% of its former extent. Reduced livestock grazing has led to a recovery of formerly degraded desert steppe vegetation in all test areas.In contrast to recent scientific publications and statements of regional and international environmental authorities (UNEP), which assume that desertification risk has even increased after the break down of the Soviet system, our results clearly show the widespread regeneration of formerly altered and degraded land between 1989 and 1998.The dramatic decay of agriculture observed, was induced by the breakdown of the Soviet state farm system, which heavily depended on huge subsidies from the central government. Our findings prove that environmental impacts caused by agriculture, like destruction of natural habitats, salinisation of soils, pasture degradation and wind erosion, have diminished considerably in the last decade, while ecological rehabilitation and the preservation of bio-diversity values are favoured by this development. This seems to apply to large areas in the southern part of the former USSR that have undergone a comparable decline of state ruled agriculture.     

Historic speciation and recent colonization of Eurasian monkey gobies (Neogobius fluviatilis and N. pallasi) revealed by DNA sequences,microsatellites, and morphology

Aim Hidden diversity within an invasive ‘species’ can mask both invasion pathways and confound management goals. We assessed taxonomic status and population structure of the monkey goby Neogobius fluviatilis across Eurasia, comparing genetic variation across its native and invasive ranges. Location Native populations were analysed within the Black and Caspian Sea basins, including major river drainages (Dnieper, Dniester, Danube, Don and Volga rivers), along with introduced locations within the upper Danube and Vistula river systems. Methods DNA sequences and 10 nuclear microsatellite loci were analysed to test genetic diversity and divergence patterns of native and introduced populations; phylogenetic analysis of mtDNA cytochrome b and nuclear RAG‐1 sequences assessed taxonomic status of Black and Caspian Sea lineages. Multivariate analysis of morphology was used to corroborate phylogenetic patterns. Population genetic structure within each basin was evaluated with mtDNA and microsatellite data using F_ST analogues and Bayesian assignment tests. Results Phylogenetic analysis of mitochondrial and nuclear sequences discerned a pronounced genetic break between monkey gobies in the Black and Caspian Seas, indicating a long‐term species‐level separation dating to c. 3 million years. This pronounced separation further was confirmed from morphological and population genetic divergence. Bayesian inference showed congruent patterns of population structure within the Black Sea basin. Introduced populations in the Danube and Vistula River basins traced to north‐west Black Sea origins, a genetic expansion pattern matching that of other introduced Ponto‐Caspian gobiids. Main conclusions Both genetic and morphological data strongly supported two species of monkey gobies that were formerly identified as subspecies: N. fluviatilis in the Black Sea basin, Don and Volga Rivers, and the Kumo‐Manych Depression, and Neogobius pallasi in the Caspian Sea and Volga River delta. Genetic origins of introduced N. fluviatilis populations indicated a common invasion pathway shared with other introduced Ponto‐Caspian fishes and invertebrates.     

Range expansion of quagga mussels Dreissena rostriformis bugensis in the Volga River and Caspian Sea basin

In 1992, we discovered populations of the nonindigenous quagga mussel Dreissena rostriformis bugensis in the middle reaches of the Volga River. The same species was found in samples collected between 1994 and 1997 in the Volga delta and in shallow areas of the Northern Caspian Sea. D. r. bugensis always co-occurred with its more widespread congener, the zebra mussel D. polymorpha (Pallas 1771). The quagga mussel's contribution to total Dreissena abundance increased over time in the middle Volga reservoirs and Volga River delta. D. r. bugensis was common in the Volga portion of Rybinsk Reservoir during 1997 and, by 2000, it was in Uglich, Rybinsk and Gorky Reservoirs on the Upper Volga River. D. r. bugensis was neither found in Ivankov Reservoir, nor in terminal sections of the Volga-Baltic corridor including the eastern Gulf of Finland. Presently, all but the northern-most regions of the Volga River have been colonized by D. r. bugensis. We hypothesize that its introduction into the Volga River and Caspian basin occurred no later than the late 1980s via commercial shipping that utilized the Volga-Don waterway to navigate between the source Black-Azov Sea region and recipient areas on the Volga River. Larval drift likely contributed to establishment of populations at downstream sites, while human-mediated vectors may be responsible for introductions to upstream locations on the Volga River. We anticipate continued northward dispersal in conjunction with shipping activities.     

High‐throughput SNP‐genotyping analysis of the relationships among Ponto‐Caspian sturgeon species

Legally certified sturgeon fisheries require population protection and conservation methods, including DNA tests to identify the source of valuable sturgeon roe. However, the available genetic data are insufficient to distinguish between different sturgeon populations, and are even unable to distinguish between some species. We performed high‐throughput single‐nucleotide polymorphism (SNP)‐genotyping analysis on different populations of Russian (Acipenser gueldenstaedtii), Persian (A. persicus), and Siberian (A. baerii) sturgeon species from the Caspian Sea region (Volga and Ural Rivers), the Azov Sea, and two Siberian rivers. We found that Russian sturgeons from the Volga and Ural Rivers were essentially indistinguishable, but they differed from Russian sturgeons in the Azov Sea, and from Persian and Siberian sturgeons. We identified eight SNPs that were sufficient to distinguish these sturgeon populations with 80% confidence, and allowed the development of markers to distinguish sturgeon species. Finally, on the basis of our SNP data, we propose that the A. baerii‐like mitochondrial DNA found in some Russian sturgeons from the Caspian Sea arose via an introgression event during the Pleistocene glaciation.     

New data on the historical and expanded range of Proterorhinus marmoratus (Pallas, 1814) (Teleostei: Gobiidae) in eastern Europe

A review of historical and recent area of distribution of tubenose goby Proterorhinus marmoratus was undertaken using critical analysis of published literature, data from two large fish collections (Natural History Museum, Kiev, Ukraine, and Zoological Institute, St Petersburg, Russia), personal collections of the authors sampled during extensive field activities in 1998–2004, and unpublished reports of the Institute of Lake and River Fisheries in Volgograd. Historically, riverine P. marmoratus from the Black Sea–Azov basin were distributed in the upper reaches of the rivers Dnieper and the Don even before the construction of water retention structures; this form at present has demonstrated only limited upstream range expansion. The native ‘Caspian tubenose goby’ is mostly connected with sea and delta habitats. We adduce some arguments for the tubenose goby's recent invasion of the Middle and Upper Volga as a result of introductions of the Don–Azov riverine form rather than upstream dispersal of the Caspian marine form.     

On the increasing threat of extermination of the unique Caspian sturgeon populations and the urgent measures required to save them.

Sturgeon populations in the Caspian Sea are now in serious danger of extinction. This is primarily a result of reduced natural recruitment following a deterioration in the quality of habitats in both the river and the sea, combined with irrational fishing activities. Within this context, the desire of the newly-arisen Republics bordering the Caspian to re-open the sturgeon fishery in the sea has absolutely no scientific foundation and will prove to be an ecological disaster. There is an urgent need to conclude the International Agreement between Russia, Kazakhstan, Azerbaijan, Turkmenistan and Iran on the conservation, reproduction and rational exploitation of the Caspian sturgeon. The bases for this agreement must be (1) recognition of the fact that sturgeon populations are sustained by an ecosystem that comprises the entire Caspian Sea and the rivers that flow into it (primarily the Volga); (2) an absolute ban on uncontrolled fishing for sturgeon in the sea, and (3) that national fishing quotas must reflect the real contribution of a particular state to overall sturgeon stocks. Without adequate measures to save them, it is to be expected that the unique runs of Caspian sturgeons will be exterminated within 5–1 years.     

Invasion genetics of a freshwater mussel (Dreissena rostriformis bugensis) in eastern Europe: high gene flow and multiple introductions

In recent years, the quagga mussel, Dreissena rostriformis bugensis, native to the Dnieper and Bug Limans of the northern Black Sea, has been dispersed by human activities across the basin, throughout much of the Volga River system, and to the Laurentian Great Lakes. We used six published microsatellite markers to survey populations throughout its native and introduced range to identify relationships among potential source populations and introduced ones. Mussels from 12 sites in Eurasia, including the central Caspian Sea and one in North America (Lake Erie), were sampled. Field surveys in the Volga River basin suggested that the species first colonized the middle reach of the river near Kubyshev Reservoir, and thereafter spread both upstream and downstream. Evidence of considerable gene flow among populations was observed and genetic diversity was consistent with a larger, metapopulation that has not experienced bottlenecks or founder effects. We propose that high gene flow, possibly due to multiple invasions, has facilitated establishment of quagga mussel populations in the Volga River system. The Caspian Sea population (D. rostriformis rostriformis (=distincta)) was genetically more distinct than other populations, a finding that may be related to habitat differences. The geographical pattern of genetic divergence is not characteristic of isolation-by-distance but, rather, of long-distance dispersal, most likely mediated by commercial ships' ballast water transfer.     

History of aquatic invertebrate invasions in the Caspian Sea

Incorporation of the fossil record and molecular markers into studies of biological invasions provides new historical perspectives on the incidence of natural and human-mediated invasions of nonindigenous species (NIS). Palaeontological, phylogeographic, and molecular evidence suggests that the natural, multiple colonizations of the Caspian basin via transient connections with the Black Sea and other basins played an important role in shaping the diversity of Caspian fauna. Geographically isolated, conspecific Ponto-Caspian lineages that currently inhabit fragmented habitats in the Ponto-Caspian region show limited genetic divergence, implying geologically recent episodes of gene flow between populations during the Pliocene to Pleistocene. Several molluscan lineages in the Caspian Sea may have descended from Lake Pannon stock before the Late Miocene isolation of the Caspian depression, about 5.8 million years ago. Anthropogenic activities during the 20th century were responsible for a 1800-fold increase in the rate of establishment of new aquatic species in the Caspian Sea compared to the preceding two million years of natural colonization. The observed success of NIS invasions during the 20th century was due primarily to human-mediated transport mechanisms, which were dominated by shipping activities (44%). Human-mediated species transfer has been strongly asymmetrical, toward the Volga Delta and Caspian Sea from or through Black and Azov Seas. Global and regional trade, particularly that mediated by commercial ships, provides dispersal opportunities for nonindigenous invertebrates, indicating that future invasions in the Caspian Sea are anticipated.     

Evidence of mitochondrial DNA clones of Siberian sturgeon, Acipenser baerii, within Russian sturgeon, Acipenser gueldenstaedtii, caught in the River Volga

Eleven of 34 sturgeons caught in the River Volga classified morphologically as Acipenser gueldenstaedtii were identified as Acipenser baerii from sequence analysis of the mitochondrial cytochrome- b gene. The Caspian Sea and its tributaries including the Volga are not native habitats of A. baerii . No A. baerii haplotype was observed in A. gueldenstaedtii from the Sea of Azov or the South Caspian Sea. Genetic contamination of A. gueldenstaedtii with A. baerii or A. baerii hybrids has occurred in the Volga. Crosses and backcrosses of these specimens with native A. gueldenstaedtii resulted in the loss of the morphological diagnostic A. baerii features. These findings are of special concern for conservation and management programmes, as well as for specimen identification for caviar trading control.     

Range expansion of quagga mussels Dreissena rostriformis bugensis in the Volga River and Caspian Sea basin

In 1992, we discovered populations of the nonindigenous quagga mussel Dreissena rostriformis bugensis in the middle reaches of the Volga River. The same species was found in samples collected between 1994 and 1997 in the Volga delta and in shallow areas of the Northern Caspian Sea. D. r. bugensis always co-occurred with its more widespread congener, the zebra mussel D. polymorpha (Pallas 1771). The quagga mussel's contribution to total Dreissena abundance increased over time in the middle Volga reservoirs and Volga River delta. D. r. bugensis was common in the Volga portion of Rybinsk Reservoir during 1997 and, by 2000, it was in Uglich, Rybinsk and Gorky Reservoirs on the Upper Volga River. D. r. bugensis was neither found in Ivankov Reservoir, nor in terminal sections of the Volga-Baltic corridor including the eastern Gulf of Finland. Presently, all but the northern-most regions of the Volga River have been colonized by D. r. bugensis. We hypothesize that its introduction into the Volga River and Caspian basin occurred no later than the late 1980s via commercial shipping that utilized the Volga-Don waterway to navigate between the source Black-Azov Sea region and recipient areas on the Volga River. Larval drift likely contributed to establishment of populations at downstream sites, while human-mediated vectors may be responsible for introductions to upstream locations on the Volga River. We anticipate continued northward dispersal in conjunction with shipping activities.     

Phylogeography,genetic structure,and conservation of the endangered Caspian brown trout, <Emphasis Type="Italic">Salmo trutta caspius</Emphasis> (Kessler, 1877), from Iran

The Caspian Sea, the largest inland closed water body in the world, has numerous endemic species. The Caspian brown trout (Salmo trutta caspius) is considered as endangered according to IUCN criteria. Information on phylogeography and genetic structure is crucial for appropriate management of genetic resources. In spite of the huge number of studies carried out in the Salmo trutta species complex across its distribution range, very few data are available on these issues for S. trutta within the Caspian Sea. Mitochondrial (mtDNA control region) and nuclear (major ribosomal DNA internal transcribed spacer 1, ITS-1, and ten microsatellite loci) molecular markers were used to study the phylogeography, genetic structure, and current captive breeding strategies for reinforcement of Caspian trout in North Iranian rivers. Our results confirmed the presence of Salmo trutta caspius in this region. Phylogenetic analysis demonstrated its membership to the brown trout Danubian (DA) lineage. Genetic diversity of Caspian brown trout in Iranian Rivers is comparable to the levels usually observed in sustainable anadromous European brown trout populations. Microsatellite data suggested two main clusters connected by gene flow among river basins likely by anadromous fish. No genetic differences were detected between the hatchery sample and the remaining wild populations. While the current hatchery program has not produced detectable genetic changes in the wild populations, conservation strategies prioritizing habitat improvement and recovering natural spawning areas for enhancing wild populations are emphasized.     

Phylogeography and origin of freshwater populations of tubenose gobies of genus Proterorhinus (Gobiidae: Pisces) in Ponto-Caspian Basin

Representatives of the genus Proterorhinus (tubenose gobies) from the water bodies of the Ponto-Caspian Basin were examined for sequence polymorphism of the mitochondrial DNA fragment containing the cytochrome b (Cyt-b) gene. A total of ten haplotypes were discovered, which formed two groups. Thus, the data obtained indicated the existence of only two taxonomically valid phylogenetic lineages, represented by (1) marine and brackish-water populations of the Black Sea and (2) freshwater populations of the whole Ponto-Caspian Basin, along with the brackish-water population of the Caspian Sea. Based on an analysis of the tubenose goby haplotype distribution patterns, the colonization patterns of this group of fishes (phylogeography) in the freshwater drainages of Ponto-Caspian Basin are examined. It was established that the basin of the ancient Caspian Sea was the major donor area for the formation of the freshwater and brackish-water populations. In the Ponto-Caspian Basin, two centers of the tubenose gobies speciation and distribution are distinguished. One of these centers is associated with the modern northwestern part of the Black Sea, and another one is confined to the Caspian Sea. It is suggested that the modern colonization of the Volga River by tubenose goby occurred from the Caspian Sea.     

Diversity of life strategies of pikepech <Emphasis Type="Italic">Sander lucioperca</Emphasis> (L.) in the Lower Volga (by the data of microelement composition of otolithes)

Strontium (Sr) and calcium (Ca) contents and Sr/Ca ratios in otoliths of pikeperch Sander lucioperca have been studied by X-ray fluorescent microanalysis. A sample of 95 specimens from the Lower Volga region are studied. The fish were obtained in a section of the Akhtuba River at a distance of 250–279 km upstream from the Caspian Sea. Some specimens in the sample have low strontium content throughout the otolith. The other fish have increased both Sr content and Sr/Ca ratios in core zones of otolith or near its outer edge: the Sr/Ca ratio is over 6.00 × 10^–3, with a maximum at 10.02 × 10^–3. The fish with a low Sr/Ca ratio live in freshwater and demonstrate a resident life history strategy. Pikeperch that have increased ratios of Sr/Ca in outer zones of otolith demonstrate a migratory life history strategy. These fish leave the freshwater of the Volga and feed in brackish water of the Northern Caspian Sea. Some migratory fish migrate to the sea as underyearlings, stay in a brackish water for 1 year or less, and then return back to fresh water for the rest of its life. The other group migrates back and forth from freshwater to the sea several times during their life. Finally, some fish leave freshwater for 1–2 years, spending this time in the Northern Caspian Sea. The conclusion is that the pikeperch stock in the Lower Volga has a complicated composition and consists of freshwater (residents and rheodromic ones) and migratory (anadromous and semianadromous) fish.     

Present status of commercial stocks of sturgeons in the Caspian Sea basin

Catches for the last 25 years are analyzed for beluga Huso huso, stellate sturgeon A. stellatus and Russian sturgeon Acipenser gueldenstaedtii, which are the three commercially important species of sturgeons found in the Caspian Sea Basin. Population sizes for generations born between 1961 and 1970 are estimated, and found to depend on natural reproduction and the number of young fish stocked annually from sturgeon hatcheries located in the Volga River Delta. A ban on sea fishing from 1962 to 1991 positively impacted the number and total biomass of commercial stocks. Sturgeon growth rates depend on water levels in the Caspian Sea. In order to preserve Caspian Sea sturgeon populations, it will be necessary to coordinate efforts of all countries surrounding the Caspian Sea to achieve rational harvests, preserve juveniles, and produce at least 100 million juveniles annually from hatcheries.     

An invasive gull displaces native waterbirds to breeding habitats more exposed to native predators

The effect of invasive opportunistic predators may include population changes in both native prey and native predators as well as alteration of predator–prey interactions. We analyzed the activity of native magpie Pica pica and changes in population, nest sites and nesting success probability of native waterbirds (namely: grebes, ducks, rails and native gulls) in response to the population growth of the invasive Caspian gull Larus cachinnans. The study was carried out at a reservoir in southern Poland and at a similar control reservoir where the Caspian gull was absent. Both the invasive gulls and the native magpie are opportunistic predators of nests of native waterbirds. The population increase of the invasive gull led to a decline in the population of native black-headed gulls Larus ridibundus only. However, the invasive gull displaced all the native species from the breeding islets located in the central part of the reservoir to islets located close to the shoreline. The latter were frequently visited by magpies, which depredated on nests along the shores, leading to an up to threefold decrease in nesting success as compared with nests located in the central area of the invaded reservoir. Predation by Caspian gulls was rarely observed. Thus, the invasion of Caspian gull caused complex direct and indirect effects on the waterbird community that included competition for breeding sites, changes in the spatial distribution of nests and alteration of predation rate by native predators. Moreover, the effects of invasion may not be reflected by changes in population size of native species.     

Chlorophyll content and factors affecting its spatial distribution in the Middle Volga reservoirs

On the basis of the data obtained during field observations in the summer low water period of 2001, the patterns of chlorophyll distribution and its relation to hydrological and hydrochemical factors in two eutrophic reservoirs of the Middle Volga are studied. The hydrological structure of the Gorky Reservoir, where the Volga water mass dominates, is homogeneous, while in the Cheboksary Reservoir along with the eutrophic Volga waters, the mesotrophic Oka water masses can be distinguished keeping their abiotic and biotic features over a long distance. Phytoplankton in the two contiguous reservoirs with different flow regimens and anthropogenic loads responds differently to the external influence. An autotrophic community in the Gorky Reservoir is more stable and depends little on abiotic factors which account for ∼63% of the explained chlorophyll variation. In the Cheboksary Reservoir under maximal for the Volga cascade flow velocity and anthropogenic load, the development and distribution of phytoplankton are almost completely (R ² = 0.93) controlled by these factors. The trophic state of the reservoirs has not changed as compared to the beginning of the 1990s.     

Contemporary status of Caspian sturgeon (Acipenseridae) stock and its conservation

The paper describes the contemporary status of sturgeon stocks in the Caspian Sea. The effect of natural reproduction and hatchery production on sturgeon population restocking is estimated. Variations in the status of spawning populations of sturgeons migrating into the Volga River were analysed. Recommendations on appropriate measures for conservation of the Caspian sturgeon stocks are presented.     

Lack of mtDNA variation among remote middle Volga and upper Ural brown trout suggests recent and rapid recolonization

Brown trout populations from the middle Volga and upper Ural drainages in the Russian Federation were analyzed with respect to their inter‐relationships and phylogenetic status using complete mtDNA control region sequences and 10 microsatellite loci. Among 56 individuals from eight populations (six from the middle Volga and two from the upper Ural drainages) only one haplotype was detected: Iran1 from the Danube phylogenetic lineage. This haplotype is very frequent in the southern Caspian basin in Iran. Given the high diversity of haplotypes in northern Iran, in contrast to the complete domination of the Iran1 haplotype in our study, it seems likely that Iran1 reached the middle Volga and upper Ural drainages as a consequence of colonization from the southern Caspian basin. In contrast to the mtDNA analysis, a more pronounced differentiation was observed among the studied populations on the basis of microsatellite DNA data. Based on this result, it is proposed that each population should be treated as a distinct management unit in the context of future conservation activities.     

Does the introduced polychaete Alitta succinea establish in the Caspian Sea?

Members of the polychaete family Nereididae do not occur naturally in the Caspian Sea but are documented intentional human introductions that commenced in 1939. However, the identity of the species involved has been uncertain; material has been identified as both Alitta succinea and Hediste diversicolor. In this study, we confirm the presence of both species in the Caspian Sea. We discuss the distribution of both species, based on tolerance to physical factors, especially salinity. Although establishment of this partially predatory Nereidid, as a food reservoir could facilitate the recreation of commercially exploited fish stock, the consequences for native benthic communities are unclear and may be subject to unforeseen negative impacts.