Ecophenotypic plasticity leads to extraordinary gastropod shells found on the "Roof of the World"

The often extraordinary shell forms and shapes of gastropods found in palaeolakes, such as the highly diverse Gyraulus fauna of the famous Steinheim Basin, have been puzzling evolutionary biologists for centuries, and there is an ongoing debate whether these aberrant shell forms are indicative of true species (or subspecies) or ecophenotypic morphs. Interestingly, one of the Steinheim Gyraulus morphs – a corkscrew-like open-coiled shell – has a recent analogue in the Lake Bangong drainage system on the western Tibetan Plateau. Therefore, a combination of morphological, molecular, palaeolimnological, and ecological analyses was used in this study to assess whether the extraordinary shell shape in Gyraulus sp. from this drainage system represents a (young) ecophenotypic phenomenon or if it has been genetically fixed over an extended period of time. Our morphological, ecological, and palaeolimnological data suggest that the corkscrew-like specimens remain restricted to a small pond near Lake Bangong with an elevated pH value and that the colonization may have occurred recently. The phylogenetic reconstruction based on two gene fragments shows that these nonplanispiral specimens cluster within the previous described Tibetan Plateau Gyraulus clade N2. A network analysis indicates that some haplotypes are even shared by planispiral and nonplanispiral specimens. Given the ephemerality of the phenomenon, the compact network patterns inferred, the likely young phylogenetic age of the aberrant Gyraulus shells studied, and the ecological peculiarities of the study site, we suggest that the evolution of the aberrant shell forms on the Tibetan Plateau could likely be considered as a rapid ecophenotypic response, possibly induced by ecological stress. This finding may thus have implications for the ongoing debate about the processes that have caused the extraordinary shell diversity in palaeolakes such as the Steinheim Basin

Recurrent camouflaged invasions and dispersal of an Asian freshwater gastropod in tropical Africa

Background: Non-indigenous taxa currently represent a large fraction of the species and biomass of freshwater ecosystems. The accumulation of invasive taxa in combination with other stressors in these ecosystems may alter the habitats to which native taxa are adapted, which could elicit evolutionary changes in native populations and their ecological interactions. Assessing ecological and evolutionary consequences of invasions simultaneously may therefore be the most effective approach to study taxa with complex invasion histories. Here we apply such an integrated approach to the cerithioid gastropod Melanoides tuberculata, a model system in invasion biology. Results: Molecular phylogenetics and ancestral range reconstructions allowed us to identify several independent Asian invasions in Lakes Malawi and Tanganyika, the Congo River, Nigeria and Cameroon. Some invasive M. tuberculata populations display much variation in shell morphology, and overlap in morphospace with M. tuberculata populations native to Africa. Experiments confirmed great ecophenotyic plasticity in some invasive populations, which, in combination with the overlap in disparity with native populations, masks invaders and their dispersal through Africa. Finally, the results of geographic modeling indicate that cryptic M. tuberculata invasions occurred primarily in densely populated areas. Conclusions: We reveal the continental nature of invasions of Asian M. tuberculata to Africa. Several of the affected ecosystems have high endemicity in Cerithioidea: Lake Tanganyika has an unparalleled diversity in freshwater cerithioids (> 10 endemic genera) and the Congo Basin and Lake Malawi are home to the two largest endemic species clusters of Melanoides in Africa (similar to 12 and similar to 8 species, respectively). Cerithioids perform ecologically important functions in the benthic ecosystems of African freshwaters, but invaders and ecosystem change pose risks to their native diversity. We draw suggestions for more effective conservation strategies from our integrated approach

Cryptic intermediate snail host of the liver fluke Fasciola hepatica in Africa

Background: Snails such as Galba truncatula are hosts for trematode flukes causing fascioliasis, a zoonosis that is a major public health problem. Galba truncatula has recently been shown to be a cryptic species complex. African populations of Galba spp. are not yet studied using molecular assessments and is imperative to do so and reconstruct the centre of origin of Galba and to understand when and by what means it may have colonized the highlands of Africa and to what extent humans might have been involved in that process. Methods: Samples from all known sub-ranges throughout Africa and new samples from Europe and Asia were obtained. We used a combination of two mitochondrial (cox1 and 16S) and one nuclear (ITS2) markers and phylogenetic, divergence time estimates and phylogeographical methods to determine the identity and biogeographical affinities. We also reconstructed the colonization history including the likely mode of dispersal and tested for the presence of cryptic Galba species in Africa. Results: Galba truncatula is restricted to the Palaearctic region of the continent, namely Morocco. All sub-Saharan populations proved to be a distinct species according to the phylogenetic analyses and genetic distance. We propose to use the existing name Galba mweruensis (Connolly, 1929) for this species which is morphologically indistinguishable from the other two species hitherto known to occur in northern Africa, i.e. G. truncatula and G. schirazensis. Sub-tropical Africa has been colonized only once in either the Pliocene and possibly Miocene. Diversification within G. mweruensis is dated to the Plio-Pleistocene and thus human-mediated dispersal can be ruled out for the initial colonization of the isolated mountain ranges. There are potentially even more cryptic species in high altitude areas of Africa as outlined by the distinctness of the population found at the top of Mt. Elgon, Uganda. Conclusions: From a novel genetic inspection of available African material, a hitherto neglected distinct species, G. mweruensis, now appears a major host of F. hepatica throughout sub-Saharan Africa. A closer examination of trematode parasites hosted by this species is needed in order to understand transmission patterns in highlands throughout eastern and southern Africa. We encourage future studies to inspect other high altitudes areas in Africa in light of parasites of either veterinary or medical importance

A Complex System of Glacial Sub-Refugia Drives Endemic Freshwater Biodiversity on the Tibetan Plateau.

Although only relatively few freshwater invertebrate families are reported from the Tibetan Plateau, the degree of endemism may be high. Many endemic lineages occur within permafrost areas, raising questions about the existence of isolated intra-plateau glacial refugia. Moreover, if such refugia existed, it might be instructive to learn whether they were associated with lakes or with more dynamic ecosystems such as ponds, wetlands, or springs. To study these hypotheses, we used pulmonate snails of the plateau-wide distributed genus Radix as model group and the Lake Donggi Cona drainage system, located in the north-eastern part of the plateau, as model site. First, we performed plateau-wide phylogenetic analyses using mtDNA data to assess the overall relationships of Radix populations inhabiting the Lake Donggi Cona system for revealing refugial lineages. We then conducted regional phylogeographical analyses applying a combination of mtDNA and nuclear AFLP markers to infer the local structure and demographic history of the most abundant endemic Radix clade for identifying location and type of (sub-)refugia within the drainage system. Our phylogenetic analysis showed a high diversity of Radix lineages in the Lake Donggi Cona system. Subsequent phylogeographical analyses of the most abundant endemic clade indicated a habitat-related clustering of genotypes and several Late Pleistocene spatial/demographic expansion events. The most parsimonious explanation for these patterns would be a scenario of an intra-plateau glacial refugium in the Lake Donggi Cona drainage system, which might have consisted of isolated sub-refugia. Though the underlying processes remain unknown, an initial separation of lake and watershed populations could have been triggered by lake-level fluctuations before and during the Last Glacial Maximum. This study inferred the first intra-plateau refugium for freshwater animals on the Tibetan Plateau. It thus sheds new light on the evolutionary history of its endemic taxa and provides important insights into the complex refugial history of a high-altitude ecosystem

NeighborNet of Lake Donggi Cona <i>Radix</i> specimens based on AFLP data.

<p>Lake specimens are colored in red; drainage specimens in orange. For specimen and locality details see the Supplement (<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0160286#pone.0160286.s002" target="_blank">S1 Table</a>).</p

Results of mismatch distribution analyses for Lake Donggi Cona <i>Radix</i> specimens based on mtDNA.

<p>SSD = sum of squared deviations from the respective model; RI = raggedness index; τ = population parameter tau with 95% confidence interval; <i>P</i> = significance of the respective parameter (level 0.05).</p

STRUCTURE output (K = 8) for Lake Donggi Cona <i>Radix</i> specimens based on AFLP data.

<p>Each group assignment is represented by a colored horizontal bar. (A) Accessions ordered according to their cluster membership (K = 8: A–H). (B) Accessions ordered according to their sampling site (lake: DC04, 05, 11; drainage system: DC07, 08, 10, 12, 18, 21). (C) Geographical presentation of the structure results. Red circles: sampling sites within the lake, orange circles: sampling sites within the drainage system. Lake map redrawn from Dietze et al. [<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0160286#pone.0160286.ref021" target="_blank">21</a>].</p

Sampling sites and phylogenetic relationships of <i>Radix</i> specimens sampled in the Lake Donggi Cona drainage system.

<p>(A) Lake Donggi Cona drainage system showing the lake, the perennial eastern and northeastern inflows (thick lines), some seasonal inflows, and the gauge-controlled outflow. Lake map redrawn from Dietze et al. [<a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0160286#pone.0160286.ref021" target="_blank">21</a>]. Satellite image (upper right corner) taken from Natural Earth (free vector and raster map data at naturalearthdata.com). (B) Collapsed Bayesian-inference tree based on mtDNA (COI) data. The uncollapsed tree is given in the Supplement (see <a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0160286#pone.0160286.s001" target="_blank">S1 Fig</a>).</p

The fingernail clams (Bivalvia: Veneroida: Sphaeriidae) of Morocco: Diversity, distribution and conservation status

In Morocco, many uncertainties surround the current diversity and distribution of the bivalve family Sphaeriidae. Such information, including taxonomy and conservation status, are vital for further studies to improve the knowledge of this family in Morocco and represents the first step towards the development of a national conservation plan for all freshwater bivalves.Between 2016 and 2019, several investigations were carried out to assess the diversity and distribution of Sphaeriidae in the different basins of Morocco, covering different types of habitat (lakes, springs, rivers and small ponds). The identification of specimens and their morpho-ecological features was based on morphological and morphometric analyses. The data on the general distribution of the collected species allowed the evaluation of their conservation status as well.The shell morphology and morphometric analyses revealed the existence of five species belonging to the genus Pisidium [P. casertanum (Poli, 1791), P. (cf.) personatum Malm, 1855, P. subtruncatum Malm, 1855, P. amnicum (O. F. Müller, 1774) and Pisidium sp.] and one species to the genus of Musculium [M. lacustre (O. F. Müller, 1774)]. Sphaeriidae were found in all Moroccan basins, except Bouregreg and Sakia El Hamra-Oued Eddahab Basins. The results showed that Sebou Basin was the species richest with the occurrence of the five species, while Loukkos and Sous-Massa Basins had the lowest-species richness with one species only. The conservation status of sphaeriids in Morocco was evaluated for the first time and resulted in P. (cf.) personatum and P. subtruncatum being proposed as Least Concern and Vulnerable, respectively, while the status of Regionally Extinct is suggested for both species P. amnicum and M. lacustre.First evaluation of the diversity and species richness of the family Sphaeriidae in Morocco with an assignment of an updated conservation status of the recorded species

Data from: Conquest of the deep, old and cold: an exceptional limpet radiation in Lake Baikal

Lake Baikal is the deepest, oldest and most speciose ancient lake in the world. The lake is characterized by high levels of molluscan species richness and endemicity, including the limpet family Acroloxidae with 25 endemic species. Members of this group generally inhabit the littoral zone, but have been recently found in the abyssal zone at hydrothermal vents and oil-seeps. Here, we use mitochondrial and nuclear data to provide a first molecular phylogeny of the Lake Baikal limpet radiation, and to date the beginning of intra-lacustrine diversification. Divergence time estimates suggest a considerably younger age for the species flock compared with lake age estimates, and the beginning of extensive diversification is possibly related to rapid deepening and cooling during rifting. Phylogenetic relationships and divergence time estimates do not clearly indicate when exactly the abyssal was colonized but suggest a timeframe coincident with the formation of the abyssal in the northern basin (Middle to Late Pleistocene)