Ancient Lake Ohrid: biodiversity and evolution

Worldwide ancient lakes have been a major focal point of geological, biological, and ecological research, and key concepts in, for example, evolutionary biology are partly based on ancient lake studies. Ancient lakes can be found on most continents and climate zones with most actual or putative ancient lakes in Europe being restricted to the Balkan Region. The arguably most outstanding of them is the oligotrophic and karstic Lake Ohrid, a steep-sided graben of rift formation origin situated in the central Balkans. Here, an attempt is made to summarize current knowledge of the geological, limnological, and faunal history of Lake Ohrid. Additionally, existing data on biodiversity and endemism in Lake Ohrid are updated and evaluated, and patterns and processes of speciation are reviewed in the context of the Ohrid watershed, including its sister lake, Lake Prespa. Whereas the geological history of the Ohrid Graben is relatively well studied, there is little knowledge about the limnological and biotic history of the actual lake (e.g., the age of the extant lake or from where the lake first received its water, along with its first biota). Most workers agree on a time frame of origin for Lake Ohrid of 2–5 million years ago (Mya). However, until now, the exact limnological origin and the origin of faunal or floral elements of Lake Ohrid remain uncertain. Two largely contrasting opinions either favour the theory of de novo formation of Lake Ohrid in a dry polje with a spring or river hydrography or a palaeogeographical connection of Lake Ohrid to brackish waters on the Balkan Peninsula. Whereas neither theory can be rejected at this point, the data summarized in the current review support the de novo hypothesis. An assessment of the fauna and flora of Lake Ohrid confirms that the lake harbours an incredible endemic biodiversity. Despite the fact that some biotic groups are poorly studied or not studied at all, approximately 1,200 native species are known from the lake, including 586 animals, and at least 212 species are endemic, including 182 animals. The adjusted rate of endemicity is estimated at 36% for all taxa and 34% for Animalia. In terms of endemic biodiversity, Lake Ohrid is with these 212 known endemic species and a surface area of 358 km² probably the most diverse lake in the world, taking surface area into account. Preliminary phylogeographical analyses of endemic Lake Ohrid taxa indicate that the vast majority of respective sister taxa occurs in the Balkans and that therefore the most recent common ancestors of Ohrid- and non-Ohrid species likely resided in the region when Lake Ohrid came into existence. These data also indicate that there is relatively little faunal exchange and overlap between Lake Ohrid and its sister lake, Lake Prespa, despite the fact that the latter lake is a major water supplier for Lake Ohrid. Studies on selected species flocks and scatters, mostly in molluscs, point towards the assumption that only few lineages originally colonized Lake Ohrid from the Balkans and that the majority of endemic species seen today probably started to evolve within the lake during the early Pleistocene. Within the Ohrid watershed, endemism occurs at different spatial and taxonomic scales, ranging from species endemic to certain parts of Lake Ohrid to species endemic to the whole watershed and from subspecies to genus level and possibly beyond. Modes of speciation in the Ohrid watershed are largely affected by its degree of isolation. Observational evidence points towards both allopatric (peripatric) and parapatric speciation. Though sympatric speciation within a habitat is conceivable, so far there are no known examples. Today, the lake suffers from increasing anthropogenic pressure and a “creeping biodiversity crisis”. Some endemic species presumably have already gone extinct, and there are also indications of invasive species penetrating Lake Ohrid. The comparatively small size of Lake Ohrid and the extremely small range of many endemic species, together with increasing human pressure make its fauna particularly vulnerable. It is thus hoped that this review will encourage future research on the ecology and evolutionary biology of the lake’s taxa, the knowledge of which would ultimately help protecting this unique European biodiversity hot spot. Guest editors: T. Wilke, R. V?in?l? & F. Riedel Patterns and Processes of Speciation in Ancient Lakes: Proceedings of the Fourth Symposium on Speciation in Ancient Lakes, Berlin, Germany, September 4–8, 2006     

Genotypic and phenotypic evaluation of Rutilus spp. from Skadar, Ohrid and Prespa Lakes supports revision of endemic as well as taxonomic status of several taxa

One mtDNA gene (cytochrome b), one nuclear DNA fragment, five microsatellites and a suite of morphological characters were evaluated in samples of Rutilus spp. from Skadar, Ohrid and Prespa Lakes. Both genetic and phenotypic data supported two sympatric taxa in Lake Skadar, whereby Prespa and Ohrid Lakes revealed only a single taxon each. One of the taxa from Lake Skadar was similar to samples from Lake Prespa, whereas the second taxon was the most divergent in the data set. The estimated time to the most recent common ancestor of these two sympatric taxa in Lake Skadar was between 125 000 and 500 000 years. The data did not support existing taxonomic schemes for Rutilus in these lakes. This study poses the following working hypothesis: (1) Rutilus prespensis lives both in Lake Prespa and Lake Skadar and therefore is not endemic to Lake Prespa, (2) Rutilus ohridanus lives in Lake Ohrid only and therefore could be considered an endemic if its species status is retained and (3) a third recently described taxon (Rutilus albus) sympatric to R. prespensis lives in Lake Skadar.     

Anatomical evidence of close affinity between endemic species of Pisidium (Bivalvia, Sphaeriidae) from some ancient lakes, and the widely distributed taxa

Anatomical characters of three endemic taxa from the Balkan lakes Ohrid and Prespa and one species from Lake Baikal were studied and compared with those of the widely distributed species of Pisidium . A close affinity of Pisidium edlaueri Kuiper 1960 from Ohrid and Pisidium maasseni Kuiper 1987 from Prespa to the Holarctic Pisidium nitidum Jenyns 1832 was confirmed; Pisidium raddei Dybowski 1902 from Baikal is anatomically identical to the Boreo-alpine Pisidium conventus Clessin 1877. Similarity in anatomical characters between the Ohridan subspecies Pisidium subtruncatum recalvum Kuiper 1960 and the nominal subspecies was also shown. These findings are compared with the situation in other ancient lakes. The similarity in some shell characters between the nonrelated taxa inhabiting these lakes is interpreted as being the result of convergent evolution.     

Rapid evolution of an ancient lake species flock: Freshwater limpets (Gastropoda: Ancylidae) in the Balkan Lake Ohrid

Ancient lakes have long been recognized as evolutionary theatres and hot spots of endemism; the evolution of their morphologically often highly diverse species flocks has received much attention. However, as each ancient lake has its own geological and evolutionary history, modes of speciation may differ from system to system. Ancient lakes can act as evolutionary reservoirs that assure the survival of relict species, but at the same time extant species may evolve through intralacustrine speciation. Other aspects of interest are the actual rates of immigration, diversification or extinction as well as the temporal framework of morphological change. Many of these questions have been addressed in the African (e.g. Lake Tanganyika) and Asian (e.g. Lake Baikal) ancient lakes. For an European ancient lakes (e.g. Lakes Ohrid and Prespa), such studies are largely missing. In the present paper, extraordinarily shaped endemic freshwater limpets of the genus Ancylus from the Balkan Lake Ohrid are used in a phylogeographic and phylogenetic context to test whether they represent an ancient lake species flock, to study the mode of speciation, and to assess the timing of morphological change. Based on DNA data from two mitochondrial genes (COI, LSU rDNA), it has been found that the Lake Ohrid Ancylus species form an endemic monophyletic group. In addition, the lake's feeder springs are inhabited by another, undescribed Ancylus species. All other studied waterbodies within the watershed do not support their own Ancylus lineages but are inhabited by a widespread Mediterranean taxon. The split between the species endemic to the lake and its sister taxon is dated to 1.4±0.6 million years ago. The study presents the first genetic confirmation for the existence of a species flock in a European ancient lake. Contrary to the prevailing opinion it shows that, concerning Ancylus, Lake Ohrid represents a site of intralacustrine speciation rather than an evolutionary reservoir. Moreover, it provides the first evidence for rapid morphological change in an European ancient lake species flock. See also Electronic Supplement at: http://www.senckenberg.de/odes/06-12.htm.     

Patterns of genetic differentiation and population history of endemic isopods (Asellidae) from ancient Lake Ohrid: combining allozyme and mtDNA data

Ancient lakes as places of extensive speciation processes have been characterized by a high degree of endemicity and biodiversity. The most outstanding European ancient lake is the oligotrophic and karstic Balkan Lake Ohrid. The lake is inhabited by a number of endemic species, but their evolutionary history is largely unresolved. in the present study, the genetic structure, gene genealogy and demographic history of the representatives of the Ohridian endemic Proasellus species were studied using both biparentally (allozyme loci) and maternally (partial mitochondrial cytochrome oxidase subunit I gene) inherited markers. Both data sets gave similar results and supported discrepancies among genetic differentiation, the current morphology-based taxonomy and bathymetric segregation. Horizontal distribution of endemic Proasellus species (Lake Ohrid vs adjacent feeder springs) within the lake presumably promote parapatric speciation whereas the main role of vertical barriers into diversification processes was not fully supported. The analyses of demographic history suggested the decline of endemic isopod populations. The radiation of endemic Proasellus populations within the lake could have started from the sublittoral/profundal zone towards the littoral or in the opposite direction — from the littoral to the profundal. Our analyses did not exclude both possibilities.     

Concurrent evolution of ancient sister lakes and sister species: the freshwater gastropod genus <Emphasis Type="Italic">Radix</Emphasis> in lakes Ohrid and Prespa

Ancient sister lakes are considered to be ancient lakes lying in close geographic proximity, sharing a related origin and significant time of co-existence, usually having hydrological connection as well as a balanced degree of faunal overlap and distinctness. A paradigm for studying sister lake relationships are the ancient lakes Ohrid and Prespa in the Balkans, which are characterized by high degrees of endemicity. Three general patterns of endemic species can be distinguished for these lakes: (1) taxa that are endemic to either lake, with no close relatives in the respective sister lake, (2) closely related but distinct endemic taxa in both lakes (sister species) and (3) shared endemic taxa occurring in both lakes. In the present paper, two endemic freshwater pulmonate gastropod species, Radix relicta (Lake Ohrid) and R. pinteri (Lake Prespa), are used to study the evolution of presumed sister species based on biogeographical and comparative DNA data from world-wide Radix taxa. Phylogenetic, phylogeographical and parametric bootstrap analyses all suggest a sister group relationship of R. relicta and R. pinteri (pattern 2 of endemic diversity). Sister to these two taxa is the widespread R. ampla, which does not occur in the vicinity of lakes Ohrid and Prespa. The southern feeder spring complexes of Lake Ohrid are inhabited by another lineage (Radix sp. 1), which resembles Radix relicta in morphology/anatomy. For Lake Prespa, the widespread R. auricularia was reported in addition to the endemic R. pinteri. Comparative phylogenetic data favour a western Adriatic zoogeographical affinity of lakes Ohrid and Prespa over an Aegean-Anatolian faunal connection. The status of lakes Ohrid and Prespa as sister lakes is evaluated in the light of current knowledge on gastropod speciation and endemism in these hotspots of biodiversity.     

Is Lake Prespa Jeopardizing the Ecosystem of Ancient Lake Ohrid?

Lake Prespa and Lake Ohrid, located in south-eastern Europe, are two lakes of extraordinary ecological value. Although the upstream Lake Prespa has no surface outflow, its waters reach the 160 m lower Lake Ohrid through underground hydraulic connections. Substantial conservation efforts concentrate on oligotrophic downstream Lake Ohrid, which is famous for its large number of endemic and relict species. In this paper, we present a system analytical approach to assess the role of the mesotrophic upstream Lake Prespa in the ongoing eutrophication of Lake Ohrid. Almost the entire outflow from Lake Prespa is found to flow into Lake Ohrid through karst channels. However, 65% of the transported phosphorus is retained within the aquifer. Thanks to this natural filter, Lake Prespa does not pose an immediate threat to Lake Ohrid. However, a potential future four-fold increase of the current phosphorus load from Lake Prespa would lead to a 20% increase (+0.9 mg P m⁻³) in the current phosphorus content of Lake Ohrid, which could jeopardize its fragile ecosystem. While being a potential future danger to Lake Ohrid, Lake Prespa itself is substantially endangered by water losses to irrigation, which have been shown to amplify its eutrophication.     

The neglected side of speciation in ancient lakes: phylogeography of an inconspicuous mollusc taxon in lakes Ohrid and Prespa

The morphologically remarkable endemic fauna within ancient lakes has received much attention in the literature. More inconspicuous taxa, however, often lack detailed molecular and morphometrical examination, although their proportion of the endemic fauna of an ancient lake must not be underestimated. Consequently, a better understanding of evolutionary patterns and processes within these lakes requires more knowledge about the often-neglected inconspicuous taxa. In the present study, we focus on the notoriously cryptic pea clam genus Pisidium (Bivalvia: Sphaeriidae). Though the genus is widely distributed, most endemic species are reported only from ancient lakes, including the European ancient sister lake system of Ohrid and Prespa on the Balkan Peninsula. Here we test for the first time hypotheses on the evolution of the endemic pea clams in this European biodiversity hotspot by molecular means. Combining a broad 16S phylogeny (comprising most European pea clam species), network analyses and morphometrical analyses, we found interesting biogeographical patterns and provide evidence for cryptic species in both lakes. Furthermore, we confirmed the proposed sister-species relationship of the endemics P. edlaueri in Lake Ohrid and P. maasseni in Lake Prespa, and we suggest scenarios of the endemic pea clam evolution within both lakes. The patterns of speciation found in the genus Pisidium are compared to patterns in morphologically distinct molluscan groups in lakes Ohrid und Prespa.     

Testing two contrasting evolutionary patterns in ancient lakes: species flock versus species scatter in valvatid gastropods of Lake Ohrid

Ancient lakes have long been recognized as “hot spots of evolution” and “evolutionary theatres” and they have significantly contributed to a better understanding of speciation and radiation processes in space and time. Yet, phylogenetic relationships of many ancient lake taxa, particularly invertebrate groups, are still unresolved. Also, the lack of robust morphological, anatomical, and phylogeographical data has largely prevented a rigorous testing of evolutionary hypotheses. For the freshwater gastropod genus Valvata—a group with a high degree of endemism in several ancient lakes—different evolutionary scenarios are suggested for different ancient lakes. Lake Baikal, for example, is inhabited by several endemic Valvata taxa that presumably do not form a monophyletic group. For such an evolutionary pattern, the term ancient lake species scatter is introduced here. In contrast, for the Balkan Lake Ohrid, workers previously suggested the presence of a monophyletic group of endemic Valvata species, that is, an ancient lake species flock. Sequence data of the mitochondrial cytochrome oxidase c subunit I gene (COI) from worldwide taxa, with a strong emphasis on Balkan species, are here used to test whether the putative Ohrid Valvata endemics represent an ancient lake species flock and to study patterns of speciation both on the Ohrid and the Balkan scale. The study reveals three distinct clades of endemic Valvata in Lake Ohrid. Monophyly of these taxa, however, is rejected, and they therefore do not represent an ancient lake species flock, but rather an ancient lake species scatter. Also, in contrast to many other gastropod groups in Lake Ohrid, the valvatids apparently did not radiate. Many Valvata taxa in ancient lakes are characterized by enhanced levels of shell complexity. However, it remains unclear whether these patterns are associated with ancient lake environments per se. It is here suggested that similarities in shell structure between North American and Balkan taxa might simply be due to convergent evolution.     

Invaders versus endemics: alien gastropod species in ancient Lake Ohrid

Species invasions into ancient lakes are an important but little understood phenomenon. At ancient Lake Ohrid, a systematic assessment of invasive mollusc species using morphological and genetic data was conducted from 2003 to 2012. Two globally invasive gastropod species, Physa acuta and Ferrissia fragilis, have recently been discovered at 4 out of 386 sites. These sites are anthropogenically impacted. The invasive species co-occur with endemics. Phylogenetic analyses of populations from native and invaded ranges of both species confirmed their identities and provided insights into their invasion histories. Accordingly, P. acuta is genetically more diverse than F. fragilis. Both species are currently present in a considerable number of lakes on the Balkan Peninsula. Possible future trends in Lake Ohrid and the Balkans are discussed and further spread of both species is likely. Given the ongoing environmental change in Lake Ohrid, the number of observations of non-indigenous or other widespread species will probably rise in the coming years and such species and their impact on native species should be carefully monitored. Moreover, ancient lakes with recurrent invasions of alien species might serve as interesting model systems for the study of important topics of invasion biology.     

A tale of time and depth: intralacustrine radiation in endemic Gammarus species flock from the ancient Lake Ohrid

Tentatively dated, the Plio‐/Pleistocene origin of the ancient Lake Ohrid on the Balkan Peninsula makes it the oldest ancient lake in Europe. Given the surface area of the lake and the adjusted endemicity rate, it may be also defined as the most diverse of all the ancient lakes in the world. From all the animal groups endemic to this lake, gammarids are amongst the most scarcely known in terms of their diversity and phylogenetic relationships. Partial DNA sequences of two mitochondrial genes, cytochrome oxidase subunit I (cox1) and 16S ribosomal RNA (16S rRNA) of eight known endemic Gammarus species from the Lake Ohrid valley were analysed. Phylogenetic analyses showed that endemic Gammarus species comprise an ancient species flock, with Gammarus sketi from the feeder springs being their sister taxon outside the lake. Amongst the species inhabiting the lake, Gammarus solidus and Gammarus salemaai are morphologically and molecularly well defined. By contrast, Gammarus ochridensis, Gammarus parechiniformis, Gammarus lychnidensis, and Gammarus stankokaramani revealed high discrepancy between morphological and genetic data. None of these morphospecies form a monophyletic clade and a significant degree of apparent gene flow occurs between them. This could be caused by incomplete lineage sorting and/or hybridization events. Two novel mtDNA lineages were found within the lake, possibly constituting two new species (Gammarus sp. 1 and Gammarus sp. 2). Molecular clock analysis showed that the split between G. sketi and the Gammarus species flock from the lake occurred approximately 5–7 Mya, whereas within the flock there were at least two intralacustrine radiations: one estimated at 2–3 Mya and the second at less than 1 Mya. The first one could be associated with the origin of the lake and the second with the lake water‐level fluctuations during Pleistocene. © 2013 The Linnean Society of London     

Comparative biocenological analysis of Gastropoda on the Macedonian part of Lake Ohrid and its watershed

Lake Ohrid represents a refugial ecosystem which harbors a great number of endemic and relics living forms. Though the whole Lake’s fauna characterizes high biodiversity and endemism, this is most obvious in the class of Gastropoda. Unlike the Lake, the fauna of the adjacent waters fairly differs from the Lake’s fauna, i.e., it is poorer both in term of diversity and endemism. The main goal of our study was to perform comparative biocenological researches on the gastropod fauna from Lake Ohrid and the adjacent waters in the watershed of the Lake. Based on the results we have obtained, it could be clearly noted that different habitats are characterized with different qualitative composition of the gastropod fauna not only when compared the gastropod settlement between the Lake and its surrounding waters, but also in the Lake for itself. Total of 50 species of gastropods have been recorded in the littoral regions of the lake and its coastal waters during 2009/2010. They belong to subclass Orthogastropoda (50 taxa). 21 species out of 50 are recorded in the adjacent waters: 13 of them settle both the adjacent waters and the Lake, while 8 strictly inhabit the adjacent waters. In terms of endemism, 17 are endemic and 4 cosmopolitan. The remainig 29 (out of 50 recorded) settle up only the littoral zone of the Lake: 25 are endemic and 4 are cosmopolitan. The percentage of endemism based on the recorded species for the class Gastropoda is 84%.     

Historical demography of brown trout (Salmo trutta) in the Adriatic drainage including the putative S. letnica endemic to Lake Ohrid

We explore the historical demography of the Adriatic lineage of brown trout and more explicitly the colonization and phylogenetic placement of Ohrid trout, based on variation at 12 microsatellite loci and the mtDNA control region. All Adriatic basin haplotypes reside in derived positions in a network that represents the entire lineage. The central presumably most ancestral haplotype in this network is restricted to the Iberian Peninsula, where it is very common, supporting a Western Mediterranean origin for the lineage. The expansion statistic R2, Bayesian based estimates of demographic parameters, and star-like genealogies support expansions on several geographic scales, whereas application of pairwise mismatch analysis was somewhat ambiguous. The estimated time since expansion (155,000 years ago) for the Adriatic lineage was supported by a narrow confidence interval compared to previous studies. Based on microsatellite and mtDNA sequence variation, the endemic Ohrid trout represents a monophyletic lineage isolated from other Adriatic basin populations, but nonetheless most likely evolving from within the Adriatic lineage of brown trout. Our results do not support the existence of population structuring within Lake Ohrid, even though samples included two putative intra-lacustrine forms. In the interests of protecting the unique biodiversity of this ancient ecosystem, we recommend retaining the taxonomic epithet Salmo letnica for the endemic Ohrid trout.     

Mollusc biodiversity in a European ancient lake system: lakes Prespa and Mikri Prespa in the Balkans

The spatial distribution of (endemic) biodiversity in ancient and potentially ancient lakes in Europe is poorly understood. Examples include Lakes Prespa and Mikri Prespa in the Central Balkans. Utilizing information of the most species-rich taxon in these lakes, the Mollusca, we therefore attempt to statistically assess and visualize the spatial distribution of biodiversity, to analyse biogeographical patterns, and to carry out a conservation assessment. We estimate that at least 40 (sub)species (29 gastropod and 11 bivalve taxa) occur in the lakes. For both lakes combined, 37.5% of the mollusc taxa are endemic. In general, the mollusc richness in Lake Mikri Prespa is lower than in Lake Prespa and less heterogeneously distributed. The highest species richness can be seen on the western and south eastern shores of Lake Prespa. Based on the presence/absence of genera, a minimum spanning tree analysis supports the sister lake relationship of both lakes, which, in turn, are most closely related to lakes in the western Balkans and not to nearby Lake Ohrid. The IUCN red list assessment revealed (A) a tendency towards mollusc faunal change, (B) a contemporary decline and potential loss of mollusc diversity, and (C) that all endemic species are of conservation concern.     

Phylogeny and historical demography of endemic fishes in Lake Biwa: the ancient lake as a promoter of evolution and diversification of freshwater fishes in western Japan

To elucidate the origins of the endemic fish of Lake Biwa, an ancient lake in Japan, and the role of the lake in the diversification of freshwater fish in western Japan, we established a molecular phylogenetic framework with an absolute time scale and inferred the historical demography of a large set of fish species in and around the lake. We used mtDNA sequences obtained from a total of 190 specimens, including 11 endemic species of Lake Biwa and their related species, for phylogenetic analyses with divergence time estimations and from a total of 2319 specimens of 42 species (including 14 endemics) occurring in the lake for population genetic analyses. Phylogenetic analysis suggested that some of the endemic species diverged from their closest relatives earlier (1.3–13.0 Ma) than the period in which the present environmental characteristics of the lake started to develop (ca. 0.4 Ma), whereas others diverged more recently (after 0.4 Ma). In contrast, historical demographic parameters suggested that almost all species, including endemic and nonendemic ones, expanded their populations after the development of the present lake environment. In phylogeographic analyses, common or very close haplotypes of some species were obtained from Lake Biwa and other regions of western Japan. The phylogenetic and historical demographic evidence suggests that there was a time lag between phylogenetic divergence and population establishment and that phenotypic adaptation of some endemic species to the limnetic environment occurred much later than the divergences of those endemic lineages. Population structure and phylogeographic patterns suggest that Lake Biwa has functioned not only as the center of adaptive evolution but also as a reservoir for fish diversity in western Japan.     

Differences in rotifer communities in two freshwater bodies of different trophic degree (Lake Ohrid and Lake Dojran,Macedonia)

We investigated monogonont rotifers in two natural Macedonian lakes that greatly differ in age, size and trophic state: Lake Ohrid and Lake Dojran. A main characteristic of Lake Ohrid is the scarcity of nutrients and consequently a low level of primary production. Lake Dojran represents a typical eutrophic lake. Results clearly indicate that species numbers are negatively correlated with trophic degree. Qualitative analyses of rotifer compositions in Lakes Ohrid and Dojran showed the presence of 70 and 55 taxa, respectively. Rotifer assemblages differed in their community structure, population densities, and the occurrence pattern of dominant species. The density of rotifers increased with increasing nutrient concentration, varying from min. 0.67 ind. L⁻¹ in June, 2006 to max. 8.2 ind. L⁻¹ in July, 2004 in Lake Ohrid, whereas min. 28.8 ind. L⁻¹ (in December, 2005) and max. 442.5 ind. L⁻¹ (in September, 2005) were recorded in Lake Dojran. Gastropus stylifer and Keratella cochlearis were the most abundant species in the pelagic zone of Lake Ohrid, averaging monthly densities of 1.2 ind. L⁻¹ and 0.6 ind. L⁻¹, respectively, thereby contributing 29% and 15% to rotifer abundance. In contrast, Lake Dojran rotifers were dominated by Brachionus spp. Brachionus diversicornis and Brachionus calyciflorus f. amphiceros were most abundant, comprising 40% and 25% of the total rotifer density. These results corroborate our idea, that the trophic state is an important factor in determining the composition of rotifer communities.     

Microsporidian Parasites Found in the Hemolymph of Four Baikalian Endemic Amphipods

At present, approximately 187 genera and over 1300 species of Microsporidia have been described, among which almost half infect aquatic species and approximately 50 genera potentially infect aquatic arthropods. Lake Baikal is the deepest and one of the oldest lakes in the world, and it has a rich endemic fauna with a predominance of arthropods. Among the arthropods living in this lake, amphipods (Crustacea) are the most dominant group and are represented by more than 350 endemic species. Baikalian amphipods inhabit almost all depths and all types of substrates. The age and geographical isolation of this group creates excellent opportunities for studying the diversity, evolution and genetics of host-parasite relationships. However, despite more than 150 years of study, data investigating the microsporidia of Lake Baikal remain incomplete. In this study, we used molecular genetic analyses to detect microsporidia in the hemolymph of several endemic species of amphipods from Lake Baikal. We provide the first evidence that microsporidian species belonging to three genera (Microsporidium, Dictyocoela and Nosema) are present in the hemolymph of Baikalian endemic amphipods. In the hemolymph of Eulimnogammarus verrucosus, we detected SSU rDNA of microsporidia belonging to the genus Nozema. In the hemolymph of Pallasea cancellous, we found the DNA of Microsporidium sp. similar to that in other Baikalian endemic amphipods; Dictyocoela sp. was found in the hemolymph of Eulimnogammarus marituji and Acanthogammarus lappaceus longispinus.     

鄱阳湖区泥鳅遗传多样性研究

采用同工酶电泳、RAPD引物PCR扩增和线粒体DNA细胞色素b基因全序列3种方法,对鄱阳湖3种斑纹泥鳅的遗传结构进行分析。同工酶电泳显示3种斑纹泥鳅在LDH、MDH和EST这3种同工酶之间存在群体和组织差异;RAPD试验结果显示6条RAPD引物在18尾鄱阳湖泥鳅个体中共扩增出了169条不同分子量的RAPD标记,其中多态性带101条,多态性带的比例为59.56%,平均每个引物产生9.39条带和5.61条多态性带,不同斑纹泥鳅的遗传多样性指数显示3种斑纹泥鳅的遗传水平处于较高水平。以特异性引物进行PCR扩增,获得了1140bp的细胞色素b基因全序列。结果显示,3种不同斑纹泥鳅中检测出了1个共有单倍型和3个特有单倍型。与共有单倍型相比,大花斑泥鳅特有单倍型有91个变异位点,而小花斑泥鳅与无花斑泥鳅特有单倍型的变异位点为20和24个。基于单倍型构建的NJ树和MP树,显示大花斑泥鳅的特有单倍型EU145023处于独立的分支上,这反映出鄱阳湖地区不同斑纹的泥鳅之间在线粒体DNACytb基因中出现了一定距离的分化。