Biogeography of freshwater fishes of the Balkan Peninsula

Delineating biogeographical regions is a critical step towards the establishment and evaluation of conservation priorities. In the present study, we analysed the distribution patterns of the freshwater fish of an understudied European biodiversity hotspot, the Balkan Peninsula. Based on the most extensive available database of native freshwater fish species distributions, we performed a hierarchical clustering analysis to identify the major biogeographical regions of the Balkan Peninsula. We also highlighted the ‘hottest hotspots’ of freshwater fish diversity across the delimited biogeographical regions by describing the patterns of species richness, endemic and vulnerable species; indicator species were also determined. The bioregionalisation scheme consisted of eight groups of drainage basins that correspond to distinct regions of the Balkan Peninsula. Overall, the delineated biogeographical regions varied in terms of species richness, endemism, vulnerability (i.e. extinction threats) and indicator species composition. From a conservation perspective, this study emphasises the prioritisation of areas characterised by high levels of irreplaceability (endemism) and vulnerability (i.e. the Attikobeotia region, Ionian Sea and Prespa Lakes) and stresses the necessity of implementing a network of protected freshwater areas across the Balkan Peninsula.     

Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types

Aim   We analysed the variation of species richness in the European freshwater fauna across latitude. In particular, we compared latitudinal patterns in species richness and β-diversity among species adapted to different habitat types.
Location   Europe.
Methods   We compiled data on occurrence for 14,020 animal species across 25 pre-defined biogeographical regions of European freshwaters from the Limnofauna Europaea . Furthermore, we extracted information on the habitat preferences of species. We assigned species to three habitat types: species adapted to groundwater, lotic (running water) and lentic (standing water) habitats. We analysed latitudinal patterns of species richness, the proportion of lentic species and β-diversity.
Results   Only lentic species showed a significant species–area relationship. We found a monotonic decline of species richness with latitude for groundwater and lotic habitats, but a hump-shaped relationship for lentic habitats. The proportion of lentic species increased from southern to northern latitudes. β-Diversity declined from groundwater to lentic habitats and from southern to northern latitudes.
Main conclusions   The differences in the latitudinal variation of species richness among species adapted to different habitat types are in part due to differences in the propensity for dispersal. Since lentic habitats are less persistent than lotic or groundwater habitats, lentic species evolved more efficient strategies for dispersal. The dispersal propensity of lentic species facilitated the recolonization of central Europe after the last glaciation. Overall, we stress the importance of considering the history of regions and lineages as well as the ecological traits of species for understanding patterns of biodiversity.     

Evidence of history in explaining diversity patterns in tropical riverine fish

Aim Documentation of the ongoing effect of rain forest refuges at the last glacial maximum (LGM) on patterns of tropical freshwater fish diversity. Location Tropical South and Central America, and West Africa. Methods LGM rain forest regions and species richness by drainage were compiled from published data. GIS mapping was applied to compile drainage area and contemporary primary productivity. We used multiple regression analyses, applied separately for Tropical South America, Central America and West Africa, to assess differences in species richness between drainages that were connected and disconnected to rain forest refuge zones during the LGM. Spatial autocorrelation of the residuals was tested using Moran's I statistic. We added an intercontinental comparison to our analyses to see if a historical signal would persist even when a regional historical effect (climate at the LGM) had already been accounted for. Results Both area and history (contact with LGM rain forest refuge) explained the greatest proportions of variance in the geographical pattern of riverine species richness. In the three examined regions, we found highest richness in drainages that were connected to the rain forest refuges. No significant residual spatial autocorrelation was detected after considering area, primary productivity and LGM rain forest refuges. These results show that past climatic events still affect West African and Latin American regional and continental freshwater fish richness. At the continental scale, we found South American rivers more species‐rich than expected on the basis of their area, productivity and connectedness to rain forest refuge. Conversely, Central American rivers were less species‐diverse than expected by the grouped model. African rivers were intermediate. Therefore, a historical signal persists even when a regional historical effect (climate at the LGM) had already been accounted for. Main conclusions It has been hypothesized that past climatic events have limited impact on species richness because species have tracked environmental changes through range shifts. However, when considering organisms with physically constrained dispersal (such as freshwater fish), past events leave a perceptible imprint on present species diversity. Furthermore, we considered regions that have comparable contemporary climatic and environmental characteristics, explaining the absence of a productivity effect. From the LGM to the present day (a time scale of 18,000 years), extinction processes should have played a predominant role in shaping the current diversity pattern. By contrast, the continental effects could reflect historical contingencies explained by differences in speciation and extinction rates between continents at higher time scales (millions of years).     

Contrasting phylogeography of two Western Palaearctic fish parasites despite similar life cycles

Aim

We used comparative phylogeography of two intestinal parasites of freshwater fish to test whether similarity in life cycle translates into concordant phylogeographical history. The thorny‐headed worms Pomphorhynchus laevis and P. tereticollis (Acanthocephala) were formerly considered as a single species with a broad geographical and host range within the Western Palaearctic.

Location

Central and eastern parts of Northern Mediterranean area, Western and Central Europe, Ponto‐Caspian Europe.

Methods

A mitochondrial marker (COI) was sequenced for 111 P. laevis and 50 P. tereticollis individuals and nuclear ITS1 and ITS2 sequences were obtained for 37 P. laevis and 21 P. tereticollis. Genetic divergence, phylogenetic relationships and divergence time were estimated for various lineages within each species, and their phylogeographical patterns were compared to known palaeogeographical events in Western Palaearctic. Biogeographical histories of each species were inferred.

Results

The two species show very different phylogeographical patterns. Five lineages were identified in P. laevis, partially matching several major biogeographical regions defined in the European riverine fish fauna. The early stages of P. laevis diversification occurred in the peri‐Mediterranean area, during the Late Miocene. Subsequent expansion across Western Europe and Russia was shaped by dispersal and vicariant events, from Middle Pliocene to Middle Pleistocene. By contrast, P. tereticollis has differentiated more recently within the Western and Central parts of Europe, and shows weak geographical and genetic structuring.

Conclusion

Our study highlights weak to moderate similarity in the phylogeographical pattern of these acanthocephalan parasites compared to their amphipod and fish hosts. The observed differences in the timing of dispersion and migration routes taken may reflect the use of a range of final hosts with different ecologies and dispersal capabilities. By using a group underrepresented in phylogeographical studies, our study is a valuable contribution to revealing the biogeography of host–parasite interactions in continental freshwaters.     

How unique is the tiger beetle fauna (Coleoptera,Cicindelidae) of the Balkan Peninsula?

The tiger beetle fauna of the Balkan Peninsula is one of the richest in Europe and includes 19 species or 41% of the European tiger beetle fauna. Assembled by their biogeographical origins, the Balkan tiger beetle species fall into 14 different groups that include, Mediterranean, Middle Oriental, Central Asiatic, Euro-Siberian, South and East European, Pannonian-Sarmatian, West Palaearctic, Turano-European and Afrotropico Indo-Mediterranean species. The Mediterranean Sclerophyl and the Pontian Steppe are the Balkan biogeographical provinces with the highest species richness, while the Balkan Highlands has the lowest Cicindelidae diversity. Most species are restricted to single habitat types in lowland areas of the Balkan Peninsula and only Calomera aulica aulica and Calomera littoralis nemoralis occur in respectively 3 and 4 different types of habitat. About 60% of all Balkan Cicindelidae species are found in habitats potentially endangered by human activity.     

Late‐glacial recolonization and phylogeography of European red deer (Cervus elaphus L.)

The Pleistocene was an epoch of extreme climatic and environmental changes. How individual species responded to the repeated cycles of warm and cold stages is a major topic of debate. For the European fauna and flora, an expansion–contraction model has been suggested, whereby temperate species were restricted to southern refugia during glacial times and expanded northwards during interglacials, including the present interglacial (Holocene). Here, we test this model on the red deer (Cervus elaphus) a large and highly mobile herbivore, using both modern and ancient mitochondrial DNA from the entire European range of the species over the last c. 40 000 years. Our results indicate that this species was sensitive to the effects of climate change. Prior to the Last Glacial Maximum (LGM) haplogroups restricted today to South‐East Europe and Western Asia reached as far west as the UK. During the LGM, red deer was mainly restricted to southern refugia, in Iberia, the Balkans and possibly in Italy and South‐Western Asia. At the end of the LGM, red deer expanded from the Iberian refugium, to Central and Northern Europe, including the UK, Belgium, Scandinavia, Germany, Poland and Belarus. Ancient DNA data cannot rule out refugial survival of red deer in North‐West Europe through the LGM. Had such deer survived, though, they were replaced by deer migrating from Iberia at the end of the glacial. The Balkans served as a separate LGM refugium and were probably connected to Western Asia with genetic exchange between the two areas.     

Regional biogeography of shallow reef fish and macro-invertebrate communities in the Galapagos archipelago

Aim To delineate biogeographical patterns in Galapagos shallow‐water reef fauna at regional scales. Location Galapagos Islands. Methods Fishes and macro‐invertebrates were quantitatively censused using underwater visual techniques along more than 500 transects at defined depth strata across the Galapagos archipelago. Data were analysed using multivariate techniques to define regional patterns and identify species typical of different regions. Results Subtidal communities of fishes and macro‐invertebrates on shallow reefs differed consistently in species composition across the Galapagos archipelago, with three major biogeographical groupings: (1) the ‘far‐northern area’ containing the islands of Darwin and Wolf, (2) the ‘central/south‐eastern area’, including the east coast of Isabela, and (3) the ‘western area’, encompassing Fernandina and western Isabela. In addition, the northern islands of Pinta, Marchena and Genovesa form a separate region in the central/south‐eastern area, and Bahia Elizabeth and Canal Bolivar separate from other parts of the western area. The far‐northern bioregion is characterized by high fish species richness overall, including a high proportion of species of Indo‐Pacific origin. However, very few endemic fishes or species with distributions extending south from Ecuador (‘Peruvian’ species) are present, and the bioregion also possesses relatively low species richness of mobile macro‐invertebrate taxa. By contrast, the ‘western’ bioregion possesses disproportionately high numbers of endemic fish taxa, high numbers of cool‐temperate Peruvian fish species, and high invertebrate species richness, but very few species of Indo‐Pacific origin. The Bahia Elizabeth/Canal Bolivar bioregion possesses more endemic species and fewer species with Peruvian affinities than coasts within the western bioregion. The northern bioregion of Pinta, Marchena and Genovesa represents an overlap zone with affinities to both the far‐northern and south‐eastern islands. The south‐eastern bioregion includes species from a variety of different sources, particularly ‘Panamic’ species with distributions extending north to Central America. Main conclusions On the basis of congruent divisions for reef fish and macro‐invertebrate communities, the Galapagos archipelago can be separated into three major biogeographical areas, two of which can be further subdivided into two regions. Each of these five bioregions possesses communities characterized by a distinctive mix of species derived from Indo‐Pacific, Panamic, Peruvian and endemic source areas. The conservation significance of different regions is not reflected in counts of total species richness. The regions with the lowest overall fish species richness possess a temperate rather than tropical climate and highest levels of endemism.     

Passalidae (Coleoptera: Scarabaeoidea) of the Biogeographical Province of Chocó and the West Andean Region of Colombia,with the Description of Two New Species

We present a synopsis of the Passalidae of the Chocó biogeographical province and the western slopes of the Western Andean range of Colombia as a result of field collections, examination of entomological collections, and review of the literature. We record a total of 41 species, provide an identification key, and, for the 39 species for which we were able to examine specimens, include a diagnosis and collecting data. Two new species of Passalus (Pertinax) are described and illustrated. The species of this region compose 42% of the passalid species known for Colombia. The richness of species and the high degree of endemism (34%) indicate the faunistic importance of this area, which is closely related to the fauna of lowland Central America.     

Effects of historical and contemporary factors on global patterns in avian species richness

Aim  The aim of this study was to determine how regional and historical factors influence global patterns in avian species richness.
Location  Global.
Methods  Using a comprehensive data set including 710 World Wildlife Fund terrestrial ecoregions covering nearly all the land surface of the Earth, avian species richness was compared among six biogeographical regions after accounting for sample area, elevational range and climate. Analysis of variance and multiple regressions were used. Spatial autocorrelation in model residuals was accounted for.
Results  Significant effects of region on avian species richness were found in nearly all comparisons between biogeographical regions.
Main conclusions  Regional and historical processes have played a role in regulating large-scale avian species richness patterns across the globe. Avian species richness in different regions of the world cannot be accurately predicted by a single global model. Avian species richness in areas of similar environments may differ substantially between regions, and thus avian species richness in one biogeographical region cannot be predicted using the richness–environment relationship derived from the data of another biogeographical region, even one with similar environments.     

Invasion phylogeography of the Ponto-Caspian crustacean Limnomysis benedeni dispersing across Europe

Aim  Limnomysis benedeni Czerniavsky, 1882 is a mysid crustacean native to the Ponto-Caspian (Black and Caspian Sea) rivers and estuaries, and has recently spread across Europe through intentional and unintentional introductions. We explored the structuring of genetic variation in native and non-native populations with an aim to trace the sources of the invasions, and to infer whether the spread has occurred through a single or multiple invasion waves.
Location  Native estuaries in the Ponto-Caspian basin (Volga, Don, Dnieper, Dniester, Danube) and the recently colonized range along the Danube–Rhine river systems and Lithuania.
Methods  A fragment of the mitochondrial COI gene was sequenced to assess genetic affinities and diversity in native and recently established populations.
Results  The genetic diversity in the native regions is organized into several strongly diverged haplotype groups or lineages, partly allopatric, partly sympatric. All these lineages have also spread beyond the native range. Even the recent rapid dispersal across Europe along the Danube–Rhine system towards the North Sea basin involved several lineages from the Danube delta sector. The structuring of genetic diversity among invaded sites suggests multiple invasion events to the Danube–Rhine drainage. This contrasts with data from some other Ponto-Caspian species, where a single haplotype seems to have occupied most invaded areas. There is no evidence that intentionally stocked reservoirs in the Baltic Sea basin would have contributed to further unintentional spread of L. benedeni.
Main conclusions  Limnomysis benedeni is spreading across Europe using the southern invasion corridor. The invasion most likely involved several waves from differentiated sources in the native Danube delta area.