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Multilocus phylogenies have shown that soft-part anatomy alone is insufficient for the systematics of valvatiform- and trochiform-shelled springsnails of the subfamily Islamiinae (family Hydrobiidae) due to morphological similarities between taxa that are not closely related. However, a solid knowledge of their systematics and species boundaries is crucial to conserve this highly imperilled group. Here, we inferred molecular phylogenies based on two mitochondrial and two nuclear gene fragments from representatives of three morphological species of the Iberian genus Iberhoratia and performed multispecies coalescent and shell variation analyses (geometric morphometrics) to reevaluate their taxonomic status. The resulting phylogenetic analyses, which included homologous sequences of 15 other valvatiform-shelled hydrobiid species endemic to the Iberian Peninsula, depicted the topotypical specimens of I. aurorae and I. gatoa elisai distantly related to one another and the type species I. morenoi. Considering their high genetic divergence and morphological differences from other hydrobiid taxa, we have placed the former two species in the new genera Beatrix n. gen. and Actenidia n. gen. The multispecies coalescent delimitation method DELINEATE, a novel approach to the taxonomy of the Hydrobiidae, and shell morphometric data suggested two species within the 'I. gatoa elisai' clade. Based on morphological and molecular evidence, we recognized this clade as Actenidia , a new genus comprising the species A. elisai n. gen. n. comb. and A. baetica n. gen. n. sp. Our results underscore the importance of using multiple lines of evidence to delimitate species of Islamiinae and interpret their systematics. We also discuss the implications of our findings for conserving I. gatoa. [ABSTRACT FROM AUTHOR]

The fifth mass extinction event (MEE) at the Cretaceous–Palaeogene (K–Pg) boundary 66 million years ago (Ma) led to massive species loss but also triggered the diversification of higher taxa. Five models have been proposed depending on whether this diversification occurred before, during or after the K–Pg boundary and the rate of species accumulation. While the effects of the K–Pg MEE on vertebrate evolution are relatively well understood, the impact on invertebrates, particularly in freshwater ecosystems, remains controversial. One example is the hyperdiverse Hydrobiidae—the most species‐rich family of freshwater gastropods. Whereas some studies place its origin in the Jurassic or even Carboniferous, most fossil records postdate the K–Pg event. We therefore used robustly time‐calibrated multi‐locus phylogenies of >400 species representing >100 hydrobiid genera to unravel its evolutionary history and patterns of diversification. We found that the family started diversifying shortly after the K–Pg boundary (∼60 Ma; 95% highest posterior density 52–69 Ma). Lineage richness gradually increased to the present and phylogenetic diversity until ∼25 Ma. These findings suggest that diversification was not initially driven by ecological opportunity. Combining the two criteria of timing and rate of diversification, a soft‐explosive diversification model of aquatic vertebrates best fits the patterns observed. We also show that most higher hydrobiid taxa (i.e. subfamilies) diversified from the Middle Oligocene to Middle Miocene (i.e. 12–28 Ma). Two of the 15 major clades delimited are described here as new subfamilies (i.e. Bullaregiinae n. subfam. and Pontobelgrandiellinae n. subfam.), whose members are restricted to subterranean waters. Our results are an important contribution to understanding how the fifth MEE has shaped evolution and patterns of biodiversity in continental aquatic systems. Given the high extinction risks faced by many hydrobiids today, they also emphasise the need to study the biodiversity of vulnerable ecosystems. [ABSTRACT FROM AUTHOR]

Shell features are key factors for systematics and evolutionary biology studies of freshwater gastropods. Based mostly on shell morphology, 19 species of the springsnail genus Radomaniola (family Hydrobiidae) have been described. Although the scarce differentiation of various shell dimensions suggests morphological stasis, the evolutionary dynamics of shell and other anatomical structures have not yet been explored fully. By inferring a phylogeny and a time-calibrated species tree from mitochondrial (COI and 16S) and nuclear (28S) sequences of 15 recognized species, we initially examine the species diversity of the Radomaniola dataset and then, through phylogenetic comparative methods, assess the evolutionary correlation, pace and mode of 40 continuous shell and anatomical characters. By synthesizing the results of four species delimitation methods and through morphological examinations, we recognize 21 groups, for which taxonomy is discussed here. Seven new species are described. We reveal a high degree of correlated evolution between characters of the shell, which are constrained by a single morphological optimum, consistent with the morphological stasis model. Anatomical traits diverged rapidly in an unconstrained manner or toward multiple optima. These findings indicate that in order to understand the morphological evolution of springsnails, it is essential to examine different organs in detail. [ABSTRACT FROM AUTHOR]

Unravelling the drivers of species diversification through geological time is of crucial importance for our understanding of long-term evolutionary processes. Numerous studies have proposed different sets of biotic and abiotic controls of speciation and extinction rates, but typically they were inferred for a single, long geological time frame. However, whether the impact of biotic and abiotic controls on diversification changes over time is poorly understood. Here, we use a large fossil dataset, a multivariate birth–death model and a comprehensive set of biotic and abiotic predictors, including a new index to quantify tectonic complexity, to estimate the drivers of diversification for European freshwater gastropods over the past 100 Myr. The effects of these factors on origination and extinction are estimated across the entire time frame as well as within sequential time windows of 20 Myr each. Our results find support for temporal heterogeneity in the factors associated with changes in diversification rates. While the factors impacting speciation and extinction rates vary considerably over time, diversitydependence and topography are consistently important. Our study highlights that a high level of heterogeneity in diversification rates is best captured by incorporating time-varying effects of biotic and abiotic factors. [ABSTRACT FROM AUTHOR]

Aim: Rates of colonization, speciation and extinction determine species richness and endemism in insular systems. The general dynamic model of island biogeography (GDM) predicts that speciation and extinction rates depend on island area and elevation via their control on ecological limits to diversification and therefore covary with an island's geological history. Additionally, the colonization rate may increase with area and elevation through the 'target effect', which can be mediated by reduced 'environmental filtering'. Here we test whether the area and depth expansion of an island‐like ecosystem, a lake, caused a shift in diversity dynamics. Location: Lake Biwa, Japan, whose geological history and biota are well documented. Taxon: Fishes. Methods: We extended the phylogenetic island biogeography framework DAISIE (Dynamic Assembly of Island biota through Speciation, Immigration and Extinction) to accommodate time‐shifts in macroevolutionary rates and in carrying capacity. Using phylogenetic information on colonization and speciation times for the complete Lake Biwa fish community (70 taxa), we tested for a shift in macroevolutionary assembly rates and reconstructed the temporal diversity trajectory in the lake. We assessed the power to identify a shift through a wide range of scenarios and benchmarked against simulated fossil records. Results: We detected an increase in colonization rate of fishes at 0.2 million years ago (Ma), with limited support for the existence of ecological limits. The reconstructed diversity trajectory was close to a source‐sink equilibrium diversity prior to the shift and remained well below a new shift‐driven elevated equilibrium thereafter. We found sufficient power to identify an increase in colonization rate up to 1.5 Ma, whereas extinction concealed the signal of earlier shifts. Main conclusions: The fish diversity dynamics of Lake Biwa show a response to changes in area and depth and phylogenies carry a signature of these changes. The detected increase in colonization rate following Lake Biwa's expansion, elevating the fish diversity, is unlikely due to a predicted increase in ecological limits feeding back on colonization rate. We therefore call for (additional) explanations: the target effect, whereby larger islands attract more species, and reduced environmental filtering due to higher habitat diversity associated with increased lake area/depth. [ABSTRACT FROM AUTHOR]

Until recently, the evolutionary origin of aquatic organisms in the Balearic archipelago (western Mediterranean) had been scarcely addressed. The freshwater gastropod genus Pseudamnicola Paulucci, 1878 (Hydrobiidae), consisting of ∼60 species in the Mediterranean area, is widely distributed across the major islands of Majorca and Minorca. From phylogenetic datasets of limited geographic coverage, a recent well-supported split has been inferred between the Minorca and Iberian Pseudamnicola species, but the phylogenetic position of the Majorca clade composed of three species remained uncertain. In this study, we extend published sequence data for the COI and 16S rRNA mitochondrial genes and the 28S rRNA nuclear gene in western Mediterranean populations of Pseudamnicola with homologous sequences from individuals of the non-surveyed region of Morocco to assess the origin of the Balearic species from a wider geographic perspective. Our maximum likelihood and Bayesian inference analyses recovered a strongly supported clade comprising the Majorcan species and four phylogenetic lineages in Morocco, two of which are new to science (described here as P. ramosae sp. nov. and P. ouarzazatensis sp. nov.). Dating the nodes using an external molecular clock rate, younger ages were estimated for the Majorcan species (∼1 Mya) than for the Moroccan ones (∼5 Mya), which were still younger than the formation of the Balearic Islands (∼30 Mya). An independent well-supported clade grouped the Minorca and Iberian species diverging at ∼4 Mya. Thus, we found two independent evolutionary origins within the Balearic archipelago likely attributable to long-distance dispersal events from the African and Eurasian continents. :pub:24D17C1D-4A42-4722-9867-D8D84465F55A :act:85EC4DF1-9BFA-4B28-B3F2-C091212AFCF1 :act:C790EAC6-F22B-489A-93EB-CABF9EEAF282 [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Abstract: 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. [ABSTRACT FROM AUTHOR]

The western Palearctic freshwater snail genus Mercuria (Caenogastropoda: Hydrobiidae) comprises 26 species primarily distributed in lowland localities of Western Europe and North Africa. Although this genus in North Africa has received considerable attention in terms of species discoveries through morphological descriptions, its distribution and phylogenetic patterns remain unknown. Based on morphological and mitochondrial DNA (mtCOI) evidence, this study examines the three Mercuria species (M. bakeri, M. tingitana, and M. targouasensis) from Morocco identified so far. Besides expanding on information regarding the anatomy of these species, two new species (M. midarensis sp. n. and M. tensiftensis sp. n.) are described for this region and phylogenetic relationships inferred between these species and the European M. emiliana and M. similis. All Moroccan and European species were recovered as independent entities according to these phylogenetic inferences (uncorrected p-distances 2.8-8.5%) and DNA barcode data. Moroccan Mercuria species clustered with M. emiliana from Spain, although basal relationships within this clade were not well supported. Given that factors such as the season when specimens are collected, habitat type, and parasites could be responsible for the remarkable intraspecific variation observed in shell and penis morphology, it is proposed that the most efficient approach to delimit and identify Mercuria species is to combine morphological descriptions with genetic data. [ABSTRACT FROM AUTHOR]

Rapids, falls, and cascades might act as barriers for freshwater species, determining the species community up- and downstream of barriers. However, they affect community composition not only by acting as barriers but also by their influence on environmental gradients. Moreover, the directional dispersal of species along the watercourse might determine community composition. A suitable system to study these differential effects is the Congo River, the world's second largest river by discharge. The small 'Upper Congo Rapids' ecoregion features several rapids known as barrier for fish. The Wagenia Cataract at the town of Kisangani constitutes the strongest drop of the Congo River and several studies have emphasized its role as barrier for fish distribution. Alternative explanations for this pattern, however, are rarely evaluated. Though mollusks represent a vital component of the macrozoobenthos, with distribution patterns and underlying drivers often distinct from that of fishes, virtually no field surveys of the Congo River have been reported for decades. We collected and determined mollusks of 51 stations, recorded environmental conditions, and generated proxies for directional species dispersal and an indirect barrier effect. Those variables were subjected to distance-based redundancy analyses and variation partitioning in order to test whether the mollusk community compositions are better explained by an individual or combined influence of the direct and indirect effect of the cataract barrier, environmental conditions, and downstreamdirected dispersal. Our survey showed an exclusive upstream/downstream distribution for just four out of the 19 species, suggesting a limited barrier effect. We revealed no direct influence of the barrier itself on community composition but of substrate type. However, we found an indirect effect of the barrier through replacing spatially structured communities upstream of the cataract with more uniform ones downstream. Downstream-directed dispersal explained the highest fraction of variation in mollusk communities. Thus, environmental factors, the indirect cataract effect, and downstream-directed spatial proxies model mollusk community composition in concert. These results support previous studies showing a multi-factorial imprint on communities. However, a large fraction of variation community composition remained unexplained, potentially due to flood plain dynamics that (re-)shape mollusk communities constantly and a high temporal turnover, evidenced by the comparison with historical surveys. This is likely caused by the growth of Kisangani and resulting human activities. A monitoring system could allow better assessments of these impacts on communities and the conservation status of endemic species in the Wagenia Cataract. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. © 2014 The Linnean Society of London [ABSTRACT FROM AUTHOR]

Highlights: [•] We analyze modes of speciation and causes of diversification in the hydrobiid subgenus Pseudamnicola (Corrosella). [•] Coalescent methods reveal three main speciation events during the upper Miocene to Pleistocene. [•] Diversification mechanisms are a result of habitat fragmentation (non-adaptive radiation). [•] Formation of Iberian mountain ranges and configuration of the hydrographic system played important roles in speciation. [ABSTRACT FROM AUTHOR]

Several Pseudamnicola (Corrosella) populations of the central and eastern Iberian Peninsula have been ascribed to P. (C.) astieri (Dupuy, 1851), though recent evidence demonstrates the species could be endemic to the departments of Var and Alpes-Maritimes in France. Through the identification of cryptic species using a combined morphological and phylogenetic approach, this paper provides a detailed morphological description of P. (C.) astieri, clarifying its taxonomic boundaries and confirming it as a French endemic. In parallel, by comparing Pseudamnicola (Corrosella) populations from the provinces of Castellón and Valencia in Eastern Spain, it was observed that rather than P. (C.) astieri they represented a new species here described as P. (C.) hauffei sp. n. Among other characters, the two species show marked differences in shell shape, male and female genital systems, radular formula and concentration of the nervous system. P. (C.) hauffei sp. n. was also compared morphologically to another two Pseudamnicola (Corrosella) species living in nearby areas [P. (C.) hinzi Boeters, 1986 and P. (C.) navasiana (Fagot, 1907)], molecularly to P. (C.) falkneri (Boeters, 1970), the type species of the subgenus, and to the rest of the Pseudamnicola (Corrosella) species described so far. Morphological differentiation between the species is supported by a genetic divergence of 7.4% inferred from a partial sequence (658 bp) of the mitochondrial gene cytochrome c oxidase subunit I (COI). On the basis of an average 8% (5.39 to 11.15%) divergence estimated for the COI gene in other P. (Corrosella) species reported in GenBank, the existence of two specific entities is here proposed, which will have impact on conservation policies both in France and in Spain. [ABSTRACT FROM AUTHOR]

The freshwater gastropod genus Pseudamnicola Paulucci, 1878 belongs to the family Hydrobiidae Stimpson, 1865 and has been cited in Europe, Africa and Asia. The Ibero-Balearic region is considered one of the areas showing the greatest diversity of hydrobiids although few works have provided complete morphological descriptions. Seven Pseudamnicola species have been described based only on shells and some features of the male and female genitalia. This paper provides the largest set of anatomical and morphological data currently available for hydrobiids and describes five new species of Pseudamnicola (Corrosella) from the Iberian Peninsula: P. (C.) manueli sp. nov., P. (C.) bareai sp. nov., P. (C.) marisolae, sp. nov., P. (C.) iruritai sp. nov. and P. (C.) andalusica sp. nov. Two further species, P. (C.) luisi Boeters, 1984 and P. (C.) falkneri (Boeters, 1970), whose distribution areas overlap, are re-described for comparison. The new species are morphologically well differentiated, the most satisfactory characters being female and male genitalia, radula and nervous system, and are well supported by molecular data, with genetic distances between 5.39 and 11.15% for the mitochondrial DNA cytochrome c oxidase subunit I sequence. All the species inhabit a mountain region whose tectonic episodes are here invoked to explain the creation of an area of endemisms. [ABSTRACT FROM AUTHOR]

Two new species of valvatoid hydrobiids are reported from southern and eastern Spain that showed notable similarities with Horatia Bourguignat, 1887 species. However, a conchological redescription of Horatia klecakiana Bourguignat, 1887, the type species of this Balkan genus, and the study of new characters for the Iberian species suggest differences between the Balkan and the Iberian groups of species. These differences and the large geographical separation between the Balkan and the Iberian populations for species with poor dispersal capacity lead us to introduce a new generic taxon, Iberhoratia Arconada and Ramos (this paper), to include the two new species. In the Iberian Peninsula, another two species have been described in Horatia, H. (?) sturmi (Rosenhauer, 1856) and H. gatoa Boeters, 1980. The first was recently moved to a new genus, Boetersiella Arconada and Ramos, 2001, while the second is here assigned to Iberhoratia. The introduction of a new genus for the Iberian Peninsula is another indication of the richness and endemic diversity of the Hydrobiidae in this region. [ABSTRACT FROM AUTHOR]

Unravelling the drivers of species diversification through geological time is of crucial importance for our understanding of long-term evolutionary processes. Numerous studies have proposed different sets of biotic and abiotic controls of speciation and extinction rates, but typically they were inferred for a single, long geological time frame. However, whether the impact of biotic and abiotic controls on diversification changes over time is poorly understood. Here, we use a large fossil dataset, a multivariate birth-death model and a comprehensive set of biotic and abiotic predictors, including a new index to quantify tectonic complexity, to estimate the drivers of diversification for European freshwater gastropods over the past 100 Myr. The effects of these factors on origination and extinction are estimated across the entire time frame as well as within sequential time windows of 20 Myr each. Our results find support for temporal heterogeneity in the factors associated with changes in diversification rates. While the factors impacting speciation and extinction rates vary considerably over time, diversity-dependence and topography are consistently important. Our study highlights that a high level of heterogeneity in diversification rates is best captured by incorporating time-varying effects of biotic and abiotic factors.