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.     

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.     

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     

On the diversity of fishes in mountain lakes of the Amur basin

Results of the study of 11 lakes situated in the mountainous part of the Amur River basin are reported. Six fish species, namely, blunt-snouted lenok Brachymystax tumensis, Baikal-Lena grayling Thymallus baicalolenensis, Siberian stone loach Barbatula toni, Lagowski’s minnow Rhynchocypris lagowskii, Czekanowski’s minnow Rh. czekanowskii, and Amur sculpin Cottus szanaga, were found in five lakes. Four species were found in one lake (Lake Pereval’noe, basin of the Amgun’ River), and two species were found in each of the remaining lakes. Lenok and Amur sculpin were the most common species. The fauna of Lake Okonon (basin of the river Zeya) was the most specific and did not have analogs among the lake faunas; Czekanowski’s minnow and Baikal black grayling were found in this lake. The lenok ecotype that formed in the mountain lakes differed from the river lenok in some regards. The potential pathways of formation and protection of mountain lake ichthyocenoses of the Amur basin are discussed.     

Phytoplankton successions under ice cover in four lakes located in North-Eastern Sweden: Effects of liming

Phytoplankton successions under ice cover (January–March) were determined in four oligotrophic lakes (Burtjärn, Aspen, Vialamptjärn and Storkorstjärn) located in North-Eastern Sweden. The total phosphorus concentration in the lakes was less than 10 µg/L. Lake Burtjärn (reference lake) had a similar hydrology as Lake Aspen. Storkorstjärn and Vialamptjärn were of similar hydrology and had heavily colored water (>100 mg_Pt/L). Aspen as well as Vialamptjärn became continuously limed with calcium carbonate annually during the last decades. Biodiversity was considerably higher in the limed lakes (Aspen and Vialamptjärn) than in the untreated lakes (Burtjärn and Storkorstjärn). In Lake Burtjärn the most frequent species wereRhodomonas lacustris, Tabellaria flocculosa andBotryococcus braunii. Cryptophyceae (R. lacustris andCryptomonas marssonii) andDinophyceae (especiallyGymnodinium lantzschii) were common phytoplankton groups in Lake Aspen.Tabellaria flocculosa was also the most common organism in both humic lakes Storkorstjärn and Vialamptjärn, other phytoplankton groups were in the humic lakes scarce. Liming was found to have profound effects on phytoplankton communities studied.     

New host species and new distribution areas of <Emphasis Type="Italic">Salmincola lavaretus</Emphasis> (Copepoda: Lernaeopodidae): A parasite of coregonid fishes of Siberia

Data on the infestation of certain fish species by the parasitic copepod Salmincola lavaretus are presented for the first time. The infested fish species included Bauntovsky whitefish (Coregonus baunti), an endemic species from the Maloye and Bolshoye Kapylyushi lakes; Siberian whitefish (C. pidschian) and Siberian cisco (C. sardinella) from Bolshoye Kapylyushi Lake and Baunt Lake, which belong to the Tsypo-Tsypikan lake system (the Lena River basin, Transbaikalia); and Teletsky whitefish (C. lavaretus natio smitti) from Teletskoe Lake (the Ob River basin). Previously, S. lavaretus had been described as a parasite of Baikal omul (C. migratorius) and Baikal whitefish (C. baicalensis) from Baikal. The taxons of S. longimanus complex (S. longimanus, S. l. sibirica, S. svetlanovi, and S. lavaretus)—parasites of nasal fossae of grayling and coregonid fishes—were registered in the lakes of the Lake Baikal basin (Lake Baikal and Khovsgol Lake), the Lena River, the Yenisei and the Ob rivers (the largest rivers of the Arctic zoogeographic province), and the Kobdo River (the Western Mongolian province). It was assumed that S. longimanus complex is more widely distributed in the water bodies of the Arctic zoogeographic province of the Palearctic.     

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.     

Development of subfossil <Emphasis Type="Italic">Daphnia</Emphasis> and <Emphasis Type="Italic">Chaoborus</Emphasis> assemblages in relation to progressive acidification and fish community alterations in SW Sweden

Dated sediment cores from acidified and fishless Lake Gaffeln and Lake Härsvatten, SW Sweden, were analyzed for Daphnia ephippia and Chaoborus mandibles to test whether acidification history and fish extirpations could be reconstructed in a paleo-study using these easily identifiable animal remains. According to monitoring data fish were lost in both lakes from the 1950s to the 1970s. Progressive acidification prior to monitoring was confirmed by a gradual decrease and eventual loss of Daphnia ephippia in both study lakes during the first half of the twentieth century. In Lake Gaffeln mandibles of C. obscuripes appeared immediately after fish loss in 1973, and the regular presence of this species confirmed the succeeding fishless state of this lake. In Lake Härsvatten sediments C. obscuripes appeared only recently, i.e. three decades after fish extirpation, showing that the absence of C. obscuripes mandibles is not a trustworthy indicator of fish presence. Hence, the appearance of C. obscuripes was not temporally related to fish loss but confirmed the present fishless condition. Known historical presence of cyprinid fish in Lake Gaffeln was confirmed by a significantly higher proportion of fragmented mandibles of C. flavicans compared to the historically cyprinid-free Lake Härsvatten. In addition, both lake profiles displayed zero-proportions of fragmented mandibles during fishless periods. We conclude that acidification history and fish extirpations can be inferred by integrated studies on subfossil Daphnia ephippia and Chaoborus mandibles. However, during extreme ultra-oligotrophic conditions in acidified clear-water lakes subfossil Chaoborus mandibles may be too scarce to infer fish absence/presence.     

Diversity of Prokaryotes in Planktonic Communities of Saline Sol-Iletsk lakes (Orenburg Oblast,Russia)

Prokaryotic diversity was studied in the planktonic communities of six Sol-Iletsk lakes (Orenburg oblast, Russia) varying in salinity level using the Illumina technology of high-throughput sequencing. The extremely halophilic archaea of the phyla Euryarchaeota and Nanohaloarchaeota, as well as the bacterial phylum Bacteroidetes predominated in the communities of lakes with salinity of 285–300‰. Representatives of the phyla Bacteroidetes and Actinobacteria, as well as of the class Gammaproteobacteria were predominant in the lakes with salinity 110?180‰. A bloom of Cyanobacteria was observed in Bol’shoe Gorodskoe Lake (10‰ salinity). The dominant OTUs in the lakes with high salinity were represented by archaea Halonotius sp., uncultured Nanohaloarchaea, and bacteria Salinibacter sp. In the lakes with medium salinity level the dominants included gammaproteobacteria Spiribacter sp., alphaproteobacteria Roseovarius sp., flavobacteria Psychroflexus sp., unidentified archaea of the family Haloferacaceae, actinobacteria Pontimonas sp. and Rhodoluna sp. In the lake with low salinity level cyanobacteria of the genus Planktothrix were predominant. Effect of salinity on prokaryotic taxonomic richness, composition, and diversity in planktonic communities of the studied lakes was demonstrated.     

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.     

The plankton community of Lake Matano: factors regulating plankton composition and relative abundance in an ancient,tropical lake of Indonesia

Recent evidence reveals that food webs within the Malili Lakes, Sulawesi, Indonesia, support community assemblages that are made up primarily of endemic species. It has been suggested that many of the species radiations, as well as the paucity of cosmopolitan species in the lakes, are related to resource limitation. In order to substantiate the possibility that resource limitation is playing such an important role, a study of the phytoplankton and zooplankton communities of Lake Matano was implemented between 2000 and 2004. We determined species diversity, relative abundances, size ranges, and total biomass for the phytoplankton and zooplankton, including the distribution of ovigerous individuals throughout the epilimnion of Lake Matano in three field seasons. The phytoplankton community exhibited very low biomass (<15 μg l^?1) and species richness was depressed. The zooplankton assemblage was also limited in biomass (2.5 mg l^?1) and consisted only of three taxa including the endemic calanoid Eodiaptomus wolterecki var. matanensis, the endemic cyclopoid, Tropocyclops matanensis and the rotifer Horaella brehmi. Zooplankton were very small (<600 μm body length), and spatial habitat partitioning was observed, with Tropocylops being confined to below 80 m, while rotifer and calanoid species were consistently observed above 80 m. Less than 0.1% of the calanoid copepods in each year were egg-bearing, suggesting very low population turnover rates. It was concluded that chemical factors as opposed to physical or biological processes were regulating the observed very low standing crops of phytoplankton which in turn supports a very minimal zooplankton community restricted in both species composition and abundance. As chemical factors are a function of the catchment basin of Lake Matano, it is predicted that resource limitation has long played an important role in shaping the unique endemic assemblages currently observed in the food web of the lake.     

Diversity and community patterns of testate amoebae in Bau Sen and Bau Trang lakes in Binh Thuan Province,Vietnam

The species composition and distribution of testate amoebae in Bau Sen and Bau Trang lakes (Binh Thuan Province, Vietnam) were investigated. Fifty-three species, varieties and forms have been identified from the water column and sediments. Twenty-seven species, varieties and forms from the genera Arcella, Centropyxis, Cyclopyxis, Difflugia and Pyxidicula were found in Vietnam for the first time. The updated list of testate amoebae in Vietnam now includes 286 taxa. The genera Arcella, Centropyxis and Difflugia are characterized by the highest frequency of occurrence. The species diversity of Arcella, Centropyxis, Difflugia was the highest. Centropyxis aculeata, Arcella discoides, Difflugia schurmanni, Netzelia oviformis and Difflugia limnetica were the most common species. Many species have shown restricted distribution, some species have been found only in one lake and not been found in another lake. The number of observed species, varieties, and forms in each lake is 34. The average number of the species per sample on the shores of the lakes subjected to human agricultural activity was significantly higher than in the samples from the shores of the lakes with less anthropogenic disturbance. The curves of “cumulative species number vs. sampling effort” are well fitted by equations S = 7.60 N ^0.69 for Bau Sen lake and S = 12.52 N ^0.46 for Bau Trang. The curves are unsaturated, which indicates that more intensive investigations of testate amoebae should be expected to reveal more species.     

The Afromontane Cladocera (Crustacea: Branchiopoda) of the Rwenzori (Uganda–D. R. Congo): taxonomy,ecology and biogeography

The timely characterization of high-altitude freshwater habitats allows an assessment of the diversity of its biota and provides the basis for monitoring community change. In this study, we investigate the Cladocera fauna of 29 water bodies (pools, freshwater lakes, and surrounding swamps sampled at various occasions between 2005 and 2009) in the Rwenzori Mountains (Uganda, D. R. Congo), which are part of the East African Sky Island Complex. All sites except one are located above 3700 m altitude. We include notes on the morphology, taxonomy, distribution, and ecology of each recorded taxon and describe a new species of the Alona rustica-group (Alona sphagnophila n.sp.; Chydoridae). We found 11 species of which seven are restricted to Lake Mahoma, the lowest lake in our study area (2990 m) (Alona affinis barbata, A. intermedia, Alonella exisa, Alonella nana, Daphnia cf. obtusa, Pleuroxus aduncus) and/or Lake Bujuku (Daphnia cf. curvirostris, P. aduncus) (3900 m). Two taxa (Ilyocryptus cf. gouldeni, A. sphagnophila n.sp.) are restricted to Carex/Sphagnum bogs surrounding lakes in the afroalpine zone. Pigmented populations of Chydorus cf. sphaericus occur in all the sites. It is the only cladoceran species surviving the extreme alpine and nival conditions in the Rwenzori. The species is joined by A. guttata at locations at lower altitudes (ca. 3000–4000 m), present in about half of the sites. The Rwenzori Cladocera fauna is characterized by a strong extratropical temperate component and a low level of speciation/endemism. Harboring an impoverished boreal cladoceran community, Lake Mahoma is given closer attention. At 2990 m, the lake is a cold-temperate aquatic island in the tropics and may function as a stepping stone for Palaearctic taxa. We introduce a new term for high-altitude, cold-water habitats in the tropics, which act as climatic islands for extratropical freshwater faunas, Löffler Islands, in honor of Dr Heinz Löffler. In comparison to surveys in 1961, we list five new records in Lake Mahoma, which could indicate cladoceran community changes over the past few decades at ca. 3000 m in the Rwenzori. Since the species distributions correlate to temperature and catchment properties of the lakes, the Rwenzori cladoceran fauna can be expected as sensitive indicators for local changes.     

Preimaginal stages of Diptera (excluding Chironomidae and Simuliidae) of mountain lakes,their inlets and outlets in the Tatra Mountains (Slovakia)

The occurrence of preimaginal stages of Diptera (excluding Chironomidae and Simuliidae) in the Tatra Mountain lakes, their inlets and outlets, and the influence of acidification on the presence of these taxa were assessed. At 21 sites, 18 Diptera taxa were recorded in 2000 and 2001. The most frequent were Dicranota sp. (73%), Pedicia (C.)rivosa (50%) and Wiedemannia sp. (14%). Diptera were ordinated by RDA; relationships between taxa and environmental variables were significantly determined by temperature and habitat characteristics (altitude, habitat, alkalinity, Ca²⁺, pH, temperature, conductivity). The RDA diagram confirmed the existence of four Diptera assemblage groups corresponding with four habitat types. Group 1 — alpine and subalpine oligotrophic lakes at higher altitudes threatened by acidification, with the acid tolerant taxa Tipula rufina, T. alpium, Dicranota sp. and Rhypholophus sp. Group 2 — subalpine oligotrophic lakes not threatened by acidification with the acid sensitive taxa Wiedemannia sp., Hemerodromia sp., Tipula (S.) benesignata, Orimarga sp., Chionea sp., Pedicia (C.) rivosa and Bazarella subneglecta which prefer higher temperature. Group 3 — subalpine oligotrophic lakes, with the acid sensitive species Berdeniella illiesi, which prefers cold water. Group 4 — outlets of submontane oligotrophic lakes with characteristic taxa Tricyphona sp. and Tipula goriziensis. Acidified alpine and subalpine Tatra lakes are situated mostly at higher altitudes (1705–2145 m. a.s.l.) and have specific taxonomic composition.     

The impact of the invasive Ponto-Caspian amphipod <Emphasis Type="Italic">Pontogammarus robustoides</Emphasis> on littoral communities in Lithuanian lakes

The intentionally introduced Pontogammarus robustoides is the most successful amphipod invader of Lithuanian inland waters and has become established in large lakes. Its impact on littoral invertebrate communities was studied by comparing similar habitats across lakes that harbour or are devoid of the invader. In habitats where P. robustoides is well established and numerous, it significantly reduces species richness and community diversity. Moderate pontogammarid density in habitats that can sustain the native gammarid Gammarus lacustris, however, revealed no negative impact on diversity metrics. Among the lakes studied, the benthic biomass did not differ in invaded and uninvaded habitats. The biomass of indigenous invertebrates (excluding chironomids, which exhibited high lake-specific biomass variation) was lower in the places with well-established P. robustoides. A detrimental impact was observed upon the native isopod Asellus aquaticus and a negative correlation with most of the higher taxa of native invertebrates. In the invaded lake habitats that favour P. robustoides, a change in community structure and a decrease in diversity up to twofold or more are to be expected.     

Long-term change of the littoral Cladocera in the Tatra Mountain lakes through a major acidification event

In our study, we focused on littoral Cladocera living and feeding in shallow shore parts of 46 mountain lakes in the Tatra Mountains (Slovakia and Poland). The studied lakes underwent a major acidification event in the 1980s and are now in the process of recovery. Lakes were divided into three categories based on their sensitivity to acidification: 5 extremely sensitive (ES), 11 acid sensitive (AS), and 30 non-sensitive (NS) lakes. In our study, we included historical data from the literature, and data from sediment core and littoral samples, which together represent the evolution of the littoral communities from a pre-industrial period up to the present. In total, 11 littoral species were found belonging to three cladoceran families. Most of the species were members of the family Chydoridae: Alona affinis, A. quadrangularis, A. rectangula, A. guttata, Acroperus harpae, Alonella excisa, A. nana, Chydorus sphaericus, and Eurycercus lamellatus. One species belonged to each family Daphniidae (Ceriodaphnia quadrangula) and Polyphemidae (Polyphemus pediculus). The most numerous littoral taxa were Alona affinis, Acroperus harpae, and Chydorus sphaericus. All species reacted to decreased pH levels during peak acidification in the 1980s by disappearing from most of the lakes of all categories; the only persisting species was Chydorus sphaericus. Most species returned to the lakes when pH started to increase in the 1990s, although their return was noticeably slower in AS lakes. Alona quadrangularis decreased its distribution range over the studied period; Polyphemus pediculus was mostly detected in the 1910s only. The number of species was highest in all lake categories when dwarf pine was present in the lake catchment. On the whole, the littoral community was richest in NS lakes.     

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.     

Adaptations to the hyporheic in Aloninae (Crustacea: Cladocera): allocation of <Emphasis Type="Italic">Alona protzi</Emphasis> Hartwig, 1900 and related species to <Emphasis Type="Italic">Phreatalona</Emphasis> gen. nov.

Morphological study of Alona protzi Hartwig, 1900, Alona phreatica Dumont, 1983 and Alona smirnovi Petkovski &; Flößner, 1972 reveals close affinities with Alona labrosa Vasiljeva &; Smirnov, 1969. We separate these four species from the polyphyletic Alona Baird, 1843 (Anomopoda: Chydoridae). United under Phreatalona gen. nov., these taxa share primitive features on the limbs, together with specializations to a rheic life mode. Phreatalona contains some of the only true hyporheic taxa within the Cladocera. Endemism in two ancient lakes (P. smirnovi and P. labrosa) and a preference for river sediments in Europe (P. phreatica and P. protzi) suggest a long isolation from typical littoral/benthic biotopes. We discuss close links with southern vicariant Nicsmirnovius, the position of these (hypo)rheic chydorids within the subfamily and their affinities with Acroperus. We remark an independent evolution of external (habitus, postabdomen) vs. internal (limb) morphology in the protzi-complex. Phreatalona is likely tertiary in origin, evolving from a littoral alonine adapting to rheic and finally hyporheic environments. Baikal endemic P. labrosa is likely the most primitive species of the genus. We discuss adaptations and evolution in the hyporheic and the effect on dispersal and biogeography of Phreatalona.     

Management of <Emphasis Type="Italic">Nymphoides peltatum</Emphasis> using water level fluctuations in freshwater lakes of Kashmir Himalaya

Lakes have an esthetic significance that is particularly important for attracting tourism. In this context, it is often preferable for lakes to have clear water, so many lake managers attempt to achieve clear lake water by various means. However, the lakes of Kashmir Himalaya are undergoing several complex ecological changes due to, for example, increasing tourism, overfishing, and intensive agriculture, which are making these lakes less clear. One such change is the vigorous growth and development of aquatic weeds in the shallow-water areas of Kashmir Himalayan lakes. We thus, investigated the response of Nymphoides peltatum, a rapidly multiplying clonal species, to water depth, in order to determine whether water depth can be used to control the spread of this proliferating macrophyte. Different traits of the given plant species, such as the mean number of ramets, were significantly higher (F = 55.412, p = 0.000) at depth zone D1 (0–100 cm) than at depth zones D2 (101–200 cm) and D3 (201–300 cm). In all of the lakes, mean spacer length—a tool for facilitating plant spread—was observed to be significantly higher (F = 45.890, p = 0.000) at lower water levels (0–100 cm). Also, the reproductive structures (flowers) of N. peltatum showed significant variation with depth (F = 51.909, p = 0.000) and with the lake examined (F = 9.909, p = 0.001). Thus, the results obtained during the present study indicate the importance of water depth in the management of N. peltatum in various Kashmir Himalayan lakes.