The problem of invaders in the Caspian Sea in the context of the findings of new zoo- and phytoplankton species from the Black Sea

In April 2004, an expedition to the Caspian Sea discovered the species in the Middle Caspian not previously described in this sea. These species preliminary identified as Oithona similis Claus 1963, Sagitta setosa Muller 1847, and Calanus euxinus (Hulsemann) are the main representatives of edible plankton of the Black Sea; most likely they were introduced into the Caspian Sea with ballast waters of ships passing through the Volga-Don Canal. At present, it is difficult to say if these species will be established in the Caspian Sea; if so, they will clearly become a valuable food resource for planktophagous fish in the Caspian Sea. In addition, we discovered two diatom species typical for the Black Sea phytoplankton, Pseudo-nitzschia seriata (Cleve) H. Peragallo and M. Peragallo and Cerataulina pelagica (Cleve) Hendey which invaded the Caspian Sea in recent years. Pseudo-nitzschia seriata synthesizes a toxin (domoic acid) and can become harmful for the basin in the case of mass bloom. We discuss the problem of spontaneous invasion of alien species from the Black Sea and their effect on the Caspian ecosystem in the context of these findings.Translated from Izvestiya Akademii Nauk, Seriya Biologicheskaya, No. 1, 2005, pp. 78–87.Original Russian Text Copyright © 2005 by Shiganova, Musaeva, Pautova, Bulgakova.     

Genetic diversity of sharpnose mullet <Emphasis Type="Italic">Liza saliens</Emphasis> introduced in southern Caspian Sea in comparison with one native Aegean Sea population

Sharpnose mullet, Liza saliens (Risso, 1810) is one of the valuable species in Caspian shoreline and it was first introduced to Caspian Sea from Black Sea between 1930 and 1934. In the present study, we used six microsatellite markers to obtain genetic information for L. saliens from four localities of southern Caspian Sea and for one native population from Greece coastline (northern Aegean Sea). Results showed lower number of alleles (Na = 5.7–6) and higher observed heterozygosity (Ho = 0.74–0.83) in Caspian Sea samples, related to the native Aegean population (Na = 8, Ho = 0.68). Significant deviations from Hardy-Weinberg expectations were detected in 19 out of 28 tests. Low genetic differentiation (provided Fst values) among samples was detected, but Behshahr sample was significantly differentiated from all the others. The UPGMA tree revealed three major clusters: the Behshahr sample and Greek population were clustered in two different clades, and the remaining three samples formed another cluster. The low genetic variation and spatial differentiation among Caspian Sea samples may be justified by the recent establishment.     

Bryozoans are returning home: recolonization of freshwater ecosystems inferred from phylogenetic relationships

Bryozoans are aquatic invertebrates that inhabit all types of aquatic ecosystems. They are small animals that form large colonies by asexual budding. Colonies can reach the size of several tens of centimeters, while individual units within a colony are the size of a few millimeters. Each individual within a colony works as a separate zooid and is genetically identical to each other individual within the same colony. Most freshwater species of bryozoans belong to the Phylactolaemata class, while several species that tolerate brackish water belong to the Gymnolaemata class. Tissue samples for this study were collected in the rivers of Adriatic and Danube basin and in the wetland areas in the continental part of Croatia (Europe). Freshwater and brackish taxons of bryozoans were genetically analyzed for the purpose of creating phylogenetic relationships between freshwater and brackish taxons of the Phylactolaemata and Gymnolaemata classes and determining the role of brackish species in colonizing freshwater and marine ecosystems. Phylogenetic relationships inferred on the genes for 18S rRNA, 28S rRNA, COI, and ITS2 region confirmed Phylactolaemata bryozoans as radix bryozoan group. Phylogenetic analysis proved Phylactolaemata bryozoan's close relations with taxons from Phoronida phylum as well as the separation of the Lophopodidae family from other families within the Plumatellida genus. Comparative analysis of existing knowledge about the phylogeny of bryozoans and the expansion of known evolutionary hypotheses is proposed with the model of settlement of marine and freshwater ecosystems by the bryozoans group during their evolutionary past. In this case study, brackish bryozoan taxons represent a link for this ecological phylogenetic hypothesis. Comparison of brackish bryozoan species Lophopus crystallinus and Conopeum seurati confirmed a dual colonization of freshwater ecosystems throughout evolution of this group of animals.     

Biogeographic zonation of the Black Sea and Caspian Sea basin based on Mysid fauna (Crustacea: Mysidacea)

A biogeographic analysis of the Black Sea and Caspian Sea basin and adjacent Aegean Sea and the Sea of Marmara was conducted based on the distribution of 55 mysid species. The Black and Caspian Sea basin proper is inhabited by 35 mysid species belonging to 12 genera, among them 26 species and 3 genera are endemic to the region. Ponto-Caspian and Caspian species are predominant. The explored areas were included into two provinces of the Mediterranean subrealm of the East-Atlantic Subtropical realm and three provinces of the Ponto-Caspian realm.     

Migration and isolation during the turbulent Ponto‐Caspian Pleistocene create high diversity in the crustacean Paramysis lacustris

The Ponto‐Caspian brackish‐water fauna inhabits estuaries and rivers of the Black, Azov and Caspian seas and is fragmented by higher salinity waters and a major interbasin watershed. The fauna is known for the high levels of endemism, complex zoogeographic histories, and as a recent source of successful invasive species. It remains debated whether the Black and Azov Sea brackish‐water populations survived unfavourable Pleistocene conditions in multiple separate refugia or whether the two seas were (repeatedly) recolonized from the Caspian. Using microsatellite and mtDNA markers, we demonstrate deep among‐ and within‐basin subdivisions in a widespread Ponto‐Caspian mysid crustacean Paramysis lacustris. Five genetic clusters were identified, but their relationships did not reflect the geography of the region. The Azov cluster was the most distinct (4–5% COI divergence), despite its geographic position in the corridor between Black and Caspian seas, and may represent a new species. In the northern Black Sea area, the Dnieper cluster was closer to the Caspian cluster than to the neighbouring Danube–Dniester–Bug populations, suggesting separate colonizations of the Black Sea. Overall, the data implied a predominant gene flow from the east to the Black Sea and highlight the importance of Caspian Sea transgressions in facilitating dispersal. Yet, the presence of distinct lineages in the Black Sea points to the persistence of isolated refugial populations that have gained diagnostic differences under presumably high mutation rates and large population sizes. The unfavourable Pleistocene periods in the Black Sea therefore appear to have promoted diversification of the brackish‐water lineages, rather than extirpated them.     

Impact of the Alien Species <Emphasis Type="Italic">Mnemiopsis leidyi</Emphasis> A. Agassiz, 1865 on Fish of the Caspian Sea

The data of long-term ecological studies (1999–2014) are used for assessing the impact of a new Caspian invader Mnemiopsis leidyi A. Agassiz, 1865 on marine and anadromous fish species. The introduction of the ctenophore and its massive development in the Caspian Sea have mainly affected planktophagous fish, especially the most abundant species anchovy sprat (Clupeonella engrauliformis Borodin, 1904) and sturgeons (Family Acipenseridae). Mnemiopsis leidyi is one of the main negative factors preventing the restoration of the populations of Caspian fishes, especially anchovy tyulka. The critical state of the resources of the spawning parts of sturgeon populations (Family Acipenseridae) has been reinforced as a result of the massive development of M. leidyi.     

Phylogeography and taxonomic status of trout and salmon from the Ponto‐Caspian drainages,with inferences on European Brown Trout evolution and taxonomy

Current taxonomy of western Eurasian trout leaves a number of questions open; it is not clear to what extent some species are distinct genetically and morphologically. The purpose of this paper was to explore phylogeography and species boundaries in freshwater and anadromous trout from the drainages of the Black and the Caspian Seas (Ponto‐Caspian). We studied morphology and mitochondrial phylogeny, combining samples from the western Caucasus within the potential range of five nominal species of trout that are thought to inhabit this region, and using the sequences available from GenBank. Our results suggest that the genetic diversity of trout in the Ponto‐Caspian region is best explained with the fragmentation of catchments. (1) All trout species from Ponto‐Caspian belong to the same mitochondrial clade, separated from the other trout since the Pleistocene; (2) the southeastern Black Sea area is the most likely place of diversification of this clade, which is closely related to the clades from Anatolia; (3) The species from the Black Sea and the Caspian Sea drainages are monophyletic; (4) except for the basal lineage of the Ponto‐Caspian clade, Salmo rizeensis, all the lineages produce anadromous forms; (5) genetic diversification within the Ponto‐Caspian clade is related to Pleistocene glacial waves; (6) the described morphological differences between the species are not fully diagnostic, and some earlier described differences depend on body size; the differences between freshwater and marine forms exceed those between the different lineages. We suggest a conservative taxonomic approach, using the names S. rizeensis and Salmo labrax for trout from the Black Sea basin and Salmo caspius and Salmo ciscaucasicus for the fish from the Caspian basin.     

Composition and Distribution of Caspian Fauna in the Light of Modern Data

The composition of the Caspian fauna is considered in the light of modern data. The number of autochthonous species known for the Caspian Sea has now reached 435 and the Caspian species in the Pontoazov basin and Lake Aral number 178 and 41 species, respectively, i. e. 25–30% more than stated in the lists in my previous review (1960, 1964). The distribution of the Caspian fauna outside the Caspian Sea is discussed and it is shown that the ranges of many species have diminised owing to unfavourable conditions. However, in some species, especially mysids, the ranges have become substantially larger on account of their artificial acclimatization.     

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.     

The biology of terns nesting at Aldabra Atoll, Indian Ocean, with particular reference to breeding seasonality

Five species of tern breed on Aldabra Atoll (09° 24' S; 46° 20' E). The Caspian tern Sterna caspia and Crested tern S. bergii feed exclusively in very shallow reef/lagoon water, the Fairy tern Gygis alba and Brown noddy Anous stolidus feed out at sea, and the Black-naped tern Sterna sumatrana is intermediate in its foraging. Both of the shallow-water species lay during the south-east monsoon season, the Caspian tern from April to August and the Crested tern from June to August, but the Crested tern also lays in December and January. The remaining three species have extended laying periods largely circumscribed by the north-west monsoon season from November to March. Breeding population size of the Caspian tern is in the low tens and of the Brown noddy in the low thousands, with the other species each numbering in the hundreds. The distribution and abundance of the nine species of tern breeding within the Seychelles ( sensu lato ) vary on the different island groups in a manner interpretable in terms of depth of the surrounding waters. Systematic differences between the central Seychelles and Aldabra groups in timing of breeding by terns which feed out at sea may be associated with seasonal latitudinal movement of the divergence zone between the South Equatorial Current and the Equatorial Counter-current, acting via correlated latitudinal shifts of prey species and game-fish abundance.     

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.     

Length–weight and length–length relationships for four shad species along the western Black Sea coast of Turkey

This study investigates length–weight and length–length relationships of four shad species (Alosa caspia caspia, Alosa immaculata, Alosa maeotica, Alosa tanaica) captured in the western Black Sea coast of Turkey. A total of 686 specimens, 355 females and 331 males, were caught by gill net, trammel net and longline from 2006 through 2007 along the western Black Sea coast of Turkey. The relationships of total length (TL), fork length (FL) and standard length (SL) are presented for inhabiting in the Black Sea four shad species and the relationships between total length (TL) and body weight (BW) from the two localities, Sile and Karasu in the western Black Sea coast of Turkey. The Length–weight relationships generally indicated positive allometric growth in Alosa species for both males and females and an isometric growth for A. immaculata females. The coefficient b ranged from a minimum 2.97 for females of A. immaculata to a maximum 3.75 for both sexes A. c. caspia. Results indicated that the length–length relationship between the three length measurements were highly correlated (r² > 0.99, P < 0.001).     

Diet composition of the Persian sturgeon <Emphasis Type="Italic">Acipenser persicus</Emphasis> at the coast of the Guilan Province,Caspian Sea

Sturgeons are one of the most important and valuable species in the Caspian Sea. This investigation (2000–2002) was aimed at determining the feeding habits and the main and secondary prey organisms consumed by the Persian sturgeon (Acipenser persicus Borodin, 1897) living at depths less than 10 m in the Caspian Sea (Guilan Province). The food spectrum of immature A. persicus was comprised of benthic invertebrates of seven genera and fishes of three genera. The main prey items of immature A. persicus were polychaete worms and the secondary prey were crustaceans. The study of stomach contents showed that polychaetes were the most important component of the diet of immature A. persicus of the length class up to 25 cm.     

Sturgeon abundance and distribution in the Caspian Sea.

The number of sturgeons and their distribution on feeding grounds in the Caspian Sea were assessed. Data were collected between 1991 and 1996 on environmental factors, species composition, and population sizes in the Northern, Middle and Southern Caspian Sea except for Iranian waters. The results show a drastic decline of the sturgeon populations on the feeding grounds at sea. In order to preserve the Caspian Sea sturgeon populations, efforts of all States surrounding the Caspian Sea must be increased and harmonised to serve a single system of fishery management, stock conservation and enhancement measures based on natural reproduction and artificial propagation.     

Differentiation and allozyme variability of shads genus alosa (Clupeiformes,Alosiinae) from Azov-Black Sea basin

Biochemical genetic marking of Sea of Azov-Black Sea shads (Alosa immaculata, A. caspia, and A. maeotica) of three local stocks—Danube, Dnepr, and Azov—was performed based on an analysis of 19 enzyme loci and a series of loci coding muscle structural proteins. The high degree of monomorphism and the absence of any differences in the allele pools are shown, thus confirming their conspecificity. It seems that Caspian, Brazhnikov, and Pontic shads are only morphologically discrete forms and their species rank is unjustified. In truth, they are local and migratory ecomorphs that are almost always present in anadromous fish species.     

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.     

Intraspecific genetic polymorphism of Russian sturgeon <Emphasis Type="Italic">Acipenser gueldenstaedtii</Emphasis>

Three populations (Azov, Caspian, and Black Sea) of Russian sturgeon Acipenser queldenstaedtii were tested for polymorphism at nuclear (RAPD and microsatellites) and mitochondrial (PCR identification of two mitotypes) markers. In addition, morphometric analysis of he representatives of Azov population was carried out. According to the morphological characters, the Black Sea population occupied an intermediate position between the Caspian and Azov populations, reflecting the phylogeography of this species. In agreement with the morphometric data, genetic distances (the data of STR analysis) also placed the Black Sea population between the Caspian and Azov populations (F _ST = 0.058 and 0.043). The genetic distance between the Azov and Caspian population was somewhat higher (F _ST = 0.070). The highest allelic polymorphism at four microsatellite loci was found observed in Caspian population, while the lowest polymorphism was in the Sea of Azov. RAPD analysis distinguished high polymorphism within the populations, although it was not feasible for interpopulation analysis. Using the method differentiating the “baerii-like” and typical “gueldenstaedtii” mitotypes, the absence of the “baerii-like” marker in the Black Sea population was demonstrated. The frequency of this marker in Caspian and Azov populations constituted 31.1 and 1.8%, respectively. Possible evolutionary reasons for the interpopulation differences observed are discussed.     

Marine species in the algal flora of the Anzali Swamp (Iran)

The composition of marine species in the algal flora of the Anzali Swamp, which is situated parallel to the Caspian Sea coast, is examined. Sampling was carried out at seven stations. A total of 74 species with 89 infraspecific taxa were identified: Cyanophyta, 20 species; Cryptophyta, 2 species; Bacillariophyta, 51 species (66 infraspecific taxa); and Chlorophyta, 1 species.