With the biodiversity crisis continuing unchecked, we need to establish levels and drivers of extinction risk to effectively allocate conservation resources and develop targeted actions. Given that threat appears particularly high in freshwaters, we assessed the extinction risk of 1,500 randomly selected freshwater molluscs using the IUCN Red List Categories and Criteria, as part of the Sampled Red List Index project. We show that close to one third of species in our sample are estimated to be threatened with extinction, with highest levels of threat in the Palearctic, Australasia and Nearctic and among gastropods. Threat levels were higher in lotic than lentic systems. Twenty-seven species were classified as Extinct (eight bivalves and 19 gastropods), mostly from the Nearctic realm and lotic systems. Pollution and the modification of natural systems (e.g. through damming and water abstraction) were the most frequently reported threats to freshwater molluscs, with some regional variation. Given that we found little spatial congruence between species richness patterns of freshwater molluscs and other freshwater taxa, new additional conservation priority areas emerge from our study. We discuss the implications of our findings for freshwater mollusc conservation and important next steps to estimate trends in freshwater mollusc extinction risk over time.
The genus Pseudamnicola comprises a group of tiny springsnails inhabiting several continental and insular regions of the Mediterranean basin. Given the limited dispersal capabilities of these animals, it is difficult to explain the wide distribution range of the genus and, more specifically, its presence in isolated habitats, such as on islands. Thus, to investigate the process(es) that may explain these distribution patterns, we morphologically re-described and genetically analysed the six Pseudamnicola (Pseudamnicola) species occurring in the Iberian Peninsula and the nearby Balearic Islands. Genetic relationships were explored by sequencing two mitochondrial (cytochrome c oxidase subunit I and 16S rRNA) and one nuclear (28S rRNA) gene in 19 populations. Our morphological study confirmed the presence of previously described species, whereas our phylogenetic results revealed three lineages within the subgenus: one clade grouping the species from Minorca Island with an Iberian Peninsula species, a second clade grouping the three species from Majorca Island, and a third clade that consists of a single species, which occurs in both the Iberian Peninsula and Ibiza Island. Calculated speciation times show that the cladogenetic events involving the insular species seem to have occurred after the current conformation of the Balearic Islands (c. 20 Mya). Therefore, the speciation process may have been related to subsequent transmarine colonizations, probably during the Messinian Salinity Crisis, and the Pleistocene glaciations when landmass corridors connected the islands with the continent.
Valvatiform gastropods of the family Hydrobiidae are frequently found in European spring-fed systems. Their simplified and sometimes convergent morphological structures and restricted geographical ranges make the description and identification of valvatiform species challenging. In the Iberian Peninsula, to date, 20 valvatiform species have been described morphologically. To test morphological species delimitations and to evaluate the role of physical geography on their evolutionary history, we analysed two mitochondrial (COI and 16S rRNA) and two nuclear (28S rRNA and 18S rRNA) gene fragments of 16 valvatiform species from the Iberian Peninsula. Molecular species delimitation methods consistently inferred 11 of the nominal species. Our phylogenetic analyses recovered the species Islamia azarum as an independent lineage, distantly related to other genera, which we place in the new genus Deganta. Furthermore, Iberian valvatiform hydrobiids do not form a monophyletic subunit, but they are included in two distinct clades of Hydrobiidae. Divergence times indicate that speciation in these clades is likely to have coincided with the formation of major Iberian mountain ranges and river basins. Similarities of geographical subdivision and divergent times between this assemblage and the non-valvatiform genus Corrosella provide strong evidence for the role of the Iberian geographical barriers in shaping the biogeography of these springsnails.
Efforts to describe spatial patterns of freshwater diversity and to evaluate their underlying factors have traditionally been focused on some animal groups (e.g. amphibians, fish or dragonflies). Despite being a dominant component in continental aquatic ecosystems and crucial for determining priority areas for conservation, broad‐scale data on gastropod total species, endemic and threatened species richness are limited. Based on these biodiversity indices, we identify global hotspots, extinction risk along the elevational gradient and the drivers of species richness patterns in the largest group of freshwater gastropods, the family Hydrobiidae. Given the strong dependency to a nonmarine aquatic environment, the observed richness patterns of extant hydrobiid species could be significantly influenced by large‐scale geography and dispersal processes as well as climatic conditions affecting continental ecosystems. Therefore, we tested several predictions for species richness derived from ecological and evolutionary hypotheses postulated for other freshwater groups. Based on a comprehensive literature and biodiversity database review, we compiled the number of total, endemic and threatened species per freshwater ecoregion. We classified ecoregions as hotspots if each biodiversity index was in the top 25% of its range and assessed the effect of 13 environmental and evolutionary factors on species richness using generalised linear models. We identified 906 species and 157 genera of Hydrobiidae showing mainly a Nearctic–Palearctic distribution and 19 biodiversity hotspots, most located across the Mediterranean Basin. In our data set, 83% of the species were endemic to a single ecoregion. Of the 43% non‐data deficient species, we found almost three times more threatened than non‐threatened species, and extinction risk peaked at 1,500 m a.s.l. Species richness was unequally distributed over biogeographic realms, increased with higher connectivity among ecoregions, and was negatively related with annual temperature range. Latitude and precipitation seasonality explained part of the richness variation by a nonlinear relationship. The identified hotspots correspond with those of other freshwater taxa. The hump‐shaped relationship of extinction risk with elevation is likely the consequence of decreasing natural and anthropogenic perturbations at higher elevations. Global hotspots of Hydrobiidae richness represent areas of climatic stability with medium precipitation and temperature seasonality that are well connected with other hydrological basins. Our results illustrate that both evolutionary and environmental factors determine these global patterns and that future changes of the latter factors may affect hydrobiid richness.
Delicado, D., Machordom, A., Ramos, M.A. (2015). Effects of habitat transition on the evolutionary patterns of the microgastropod genus Pseudamnicola (Mollusca, Hydrobiidae). -Zoologica Scripta, 44, 403-417. Molecular phylogenies of extant species are considered effective tools to infer mechanisms of speciation. Here, we benefit from this utility to investigate the evolutionary history of an organismal group linked to different aquatic ecosystems, the microgastropod genus Pseudamnicola (family Hydrobiidae). Previous studies have found around 45 species of the nominal subgenus P. (Pseudamnicola), most of them in coastal stream localities of several Mediterranean islands and mainland territories, whereas only 12 species of the other subgenus, P. (Corrosella), have been collected from springs and headwaters of mountainous regions of the Iberian Peninsula and south of France. As springs often act as isolated habitats affecting dispersion and constraining gene flow, we supposed that the temporal history and mode of diversification of species from both subgenera should differ and therefore be reflected in their phylogenetic patterns. To assess this hypothesis, we performed a molecular phylogeny based on mitochondrial and nuclear DNA sequences and later conducted an independent analysis to examine the potential effect of certain geographic and ecological variables in the genetic divergences of the subgenera. Additionally, we estimated the ancestral area of diversification of both groups. Published anatomical revisions and our molecular analyses suggest that the genus Pseudamnicola should be divided into three genera: the two previous subgenera plus a new one described here. As postulated, the evolution of the spring organisms was strongly related to habitat fragmentation and isolation, whereas dispersal followed by divergence seem to have been the most common speciation processes for euryhaline species inhabiting coastal streams and low river stages in which waters remain connected. On the contrary, rather than habitat fragmentation or dispersion, environmental conditions have played a larger role during the deep divergent split leading to the three genera.
A deep systematic study of the Spanish hydrobiid fauna is unravelling a more complex group of species than was previously described. The use of diOE erent taxonomical and statistical methodologies helps to clarify this old classi® cation. Based on anatomical studies made for the ® rst time on specimens from the type locality of Paludina sturmi (Rosenhauer, 1856), the species is redescribed and transferred to a new genus Boetersiella. The study demonstrates that Horatia (?) sturmi (sensu Boeters, 1988 ) is a combination of two species belonging to two new, diOE erent but closely related genera, B. sturmi and Chondrobasi s levantina gen. and sp. nov. A second species for the ® rst genus, B. davisi, is also described. Morphometrical analysis (MANOVA and DFA) combining species of both genera showed clear diOE erences in some shell variables although the most valuable discriminant characters are related to the female and male genital systems and to the size and shape of the osphradium. Boetersiella mainly diOE ers from Chondrobasis in having a simple penis while a small basal penial lobe is present in the second; in addition, the distal seminal receptacle is bigger in Boetersiella and leans over the bursa copulatrix instead of over the renal oviduct as is the case in Chondrobasis. The two new genera diOE er from Horatia Bourguignat, 1887 in three main characters: the lack of a ctenidium, the presence in the female genitalia of only one seminal receptacle in a distal position and a penis simple (in Boetersiella) or with a small lobe close to its basis (in Chondrobasis). None of these characters are present in the type species Horatia klecakiana (Bourguignat, 1887). Both sets of character / character-states also diOE er from the other two valvatiform genera described for the Iberian Peninsula, Neohoratia Schu È tt, 1961 and Tarraconia Ramos and Arconada (in Ramos et al., in press). Comparisons with other European valvatiform genera were made although the lack of accurate anatomical data for most of them prevented any cladistical approach.
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