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The quality of spatial analyses of biodiversity is improved by (i) utilizing study areas with well defined physiogeographical boundaries, (ii) limiting the impact of widespread species, and (iii) using taxa with heterogeneous distributions. These conditions are typically met by ecosystems such as oceanic islands or ancient lakes and their biota. While research on ancient lakes has contributed significantly to our understanding of evolutionary processes, statistically sound studies of spatial variation of extant biodiversity have been hampered by the frequently vast size of ancient lakes, their limited accessibility, and the lack of scientific infrastructure. The European ancient Lake Ohrid provides a rare opportunity for such a reliable spatial study. The comprehensive horizontal and vertical sampling of a species-rich taxon, the Gastropoda, presented here, revealed interesting patterns of biodiversity, which, in part, have not been shown before for other ancient lakes. In a total of 284 samples from 224 different locations throughout the Ohrid Basin, 68 gastropod species, with 50 of them (= 73.5%) being endemic, could be reported. The spatial distribution of these species shows the following characteristics: (i) within Lake Ohrid, the most frequent species are endemic taxa with a wide depth range, (ii) widespread species (i.e. those occurring throughout the Balkans or beyond) are rare and mainly occur in the upper layer of the lake, (iii) while the total number of species decreases with water depth, the proportion of endemics increases, and (iv) the deeper layers of Lake Ohrid appear to have a higher spatial homogeneity of biodiversity. Moreover, gastropod communities of Lake Ohrid and its feeder springs are both distinct from each other and from the surrounding waters. The analysis also shows that community similarity of Lake Ohrid is mainly driven by niche processes (e.g. environmental factors), but also by neutral processes (e.g. dispersal limitation and evolutionary histories of species). For niche-based mechanisms it is shown that large scale effects such as type of water body or water depth are mainly responsible for the similarity of gastropod communities, whereas small scale effects like environmental gradients affect gastropod compositions only marginally. In fact, neutral processes appear to be more important than the small scale environmental factors, thus emphasizing the importance of dispersal capacities and evolutionary histories of species. [ABSTRACT FROM AUTHOR]

Freshwater habitats and species living in freshwater are generally more prone to extinction than terrestrial or marine ones. Immediate conservation measures for world-wide freshwater resources are thus of eminent importance. This is particularly true for so called ancient lakes. While these lakes are famous for being evolutionary theatres, often displaying an extraordinarily high degree of biodiversity and endemism, in many cases these biota are also experiencing extreme anthropogenic impact. Lake Ohrid, the European biodiversity hotspot, is a prime example for a lake with a magnitude of narrow range endemic taxa that are under increasing anthropogenicpressure. Unfortunately, evidence for a "creeping biodiversity crisis" has accumulated over the last decades, and major socio-political changes have gone along with human-mediated environmental changes. Based on field surveys, monitoring data, published records, and expert interviews, we aimed to (1) assess threats to Lake Ohrids' (endemic) biodiversity, (2) summarize existing conservation activities and strategies, and (3) outline future conservation needs for Lake Ohrid. We compiled threats to both specific taxa (and in cases to particular species) as well as to the lake ecosystems itself. Major conservation concerns identified for Lake Ohrid are: (1) watershed impacts, (2) agriculture and forestry, (3) tourism and population growth, (4) non-indigenous species, (5) habitat alteration or loss, (6) unsustainable exploitation of fisheries, and (7) global climate change. Of the 11 IUCN (International Union for Conservation of Nature and Natural Resources) threat classes scored, seven have moderate and three severe impacts. These latter threat classes are energy production and mining, biological resource use, and pollution. We review and discuss institutional responsibilities, environmental monitoring and ecosystem management, existing parks and reserves, biodiversity and species measures, international conservation activities, and ongoing research on conservation and raising of public awareness. Following this summary, we evaluate the status quo and future of Lake Ohrid and its biota. Given the number of identified threats, it is clear that only concerted international action can stop or at least slow down further degradation of Lake Ohrid and the creeping biodiversity crisis already evident. A comprehensive conservation strategy should include measures that result in an immediate reduction of pollution, particularly with phosphorous, in order to slow down the ongoingeutrophication process. The existing watershed management should become more effective. Implementation and particularly with a view to the enforcement of national laws should be enhanced. Increased research on the lakes' limnology, biodiversity, and conservation management practices are necessary. The latter research should identify conservation priorities. Public awareness should be enhanced. Facing these parallel needs to protect the unique biodiversity of Lake Ohrid, we suggest urging (a) implementation and enforcement of the General Management Plan that would ensure long-term integrated and sustainable use of the lake and its watershed, (b) scientific studies on ecology, biodiversity and effects of human impact, (c) the establishment of Core Conservation areas (CCA), including underwater reserves, and (d) Coastal Zone Management (CZM) areas that would constitute buffer zones for the CCA around the lake. These activities should, among others, ultimately lead to a trans-boundary major conservation area of the Ohrid-Prespa region that would allow long-term integration of both humans and nature. [ABSTRACT FROM AUTHOR]

Spatial heterogeneity of biodiversity arises from evolutionary processes, constraints of environmental factors and the interaction of communities. The quality of such spatial analyses of biodiversity is improved by (i) utilizing study areas with well defined physiogeographical boundaries, (ii) limiting the impact of widespread species, and (iii) using taxa with heterogeneous distributions. These conditions are typically met by ecosystems such as oceanic islands or ancient lakes and their biota. While research on ancient lakes has contributed significantly to our understanding of evolutionary processes, statistically sound studies of spatial variation of extant biodiversity have been hampered by the frequently vast size of ancient lakes, their limited accessibility, and the lack of infrastructure around them. The small European ancient Lake Ohrid provides a rare opportunity for such a reliable spatial study. The comprehensive horizontal and vertical sampling of a species-rich taxon, the Gastropoda, presented here, revealed interesting patterns of biodiversity, which, in part, have not been shown before for other ancient lakes. In a total of 224 locations throughout the Ohrid Basin, representatives of 68 gastropod species with 50 of them being endemic (=73.5%) could be reported. The spatial distribution of these species shows the following characteristics: (i) within Lake Ohrid, the most frequent species are endemic taxa with a wide depth range, (ii) widespread species (i.e. those occurring throughout the Balkans or beyond) are rare and mainly occur in the upper layer of the lake, (iii) while the total number of species decreases with water depth, the share of endemics increases, (iv) the deeper layers of Lake Ohrid appear to have a higher spatial homogeneity of biodiversity and related environmental factors, (v) biotic interaction due to possible spillover effects may contribute to the establishment of hotspots, and (vi) eco-insularity within the Ohrid Basin occurs at two levels, at the level of the lake proper and at the level of the feedersprings. It is also shown that large scale effects such as type of water body or water depth are mainly responsible for the distribution of biodiversity. In addition, small scale effects like environmental gradients or biotic interaction affect gastropod composition within a particular depth zone. [ABSTRACT FROM AUTHOR]

Ancient Lake Ohrid on the Balkan Peninsula is considered to be the oldest ancient lake in Europe with a suggested Plio-Pleistocene age. Its exact geological age, however, remains unknown. Therefore, molecular clock data of Lake Ohrid biota may serve as an independent constraint of available geological data, and may thus also help to refine age estimates. Such evolutionary data may also help unravel potential biotic and abiotic factors that promote speciation events. Here, mitochondrial sequencing data of one of the largest groups of endemic taxa in Lake Ohrid, the leech genus Dina, is used to test whether it represents an ancient lake species flock, to study the role of horizontal and vertical barriers in Lake Ohrid for evolutionary events, to estimate the onset of intralacustrine diversification in this group based on molecular clock analyses, and to compare this data with data from other endemic species for providing an approximate time frame for the origin of Lake Ohrid. Based on the criteria speciosity, monophyly and endemicity, it can be concluded that Lake Ohrid Dina, indeed, represents an ancient lake species flock. Lineage sorting of its species, however, does not seem to be complete. Analyses of population structures of Dina spp. in the Ohrid watershed indicate a horizontal zonation of haplotypes from spring and lake populations, corroborating the role of lake-side springs, particularly the southern feeder springs, for evolutionary processes in endemic Ohrid taxa. Vertical differentiation of lake taxa, however, appears to be limited, though differences between populations from the littoral and the profundal are apparent. Molecular clock analyses indicate that the most recent common ancestor of extant species of this flock is approximately 1.99 ± 0.83 Ma old, whereas the split of the Lake Ohrid Dina flock from a potential sister taxon outside the lake is estimated at 8.30 ± 3.60 Ma. Comparisons with other groups of endemic Ohrid species indicated that in all cases, intralacustrine diversification started ≤2 Ma ago. Thus, this estimate may provide information on a minimum age for the origin of Lake Ohrid. Maximum ages are less consistent and generally less reliable. But cautiously, a maximum age of 3 Ma is suggested. Interestingly, this time frame of approximately 2-3 Ma for the origin of Lake Ohrid, generated based solely on evolutionary data, well fits the time frame most often used in the literature by geologists. Future studies must show whether this concurrence holds true. [ABSTRACT FROM AUTHOR]