Hybridization between genetically and morphologically divergent forms of Rhagada (Gastropoda: Camaenidae) snails at a zone of secondary contact

Congeneric sympatry is rare in Australian camaenid land snails, and the reasons are poorly understood. As the first example of contact between species of the Western Australian genus Rhagada, we examined the parapatric zone between the coastally distributed Rhagada convicta and an undescribed inland species, which differ in size, shape, and banding of their shells and in their reproductive systems. Phylogenetic analysis of sequences of the mitochondrial 16S RNA gene indicated secondary contact between two distinct clades, with an average sequence divergence of 6.5%. Despite these substantial differences, the two forms hybridize in the narrow contact zone, resulting in intermediate phenotypes for shells and reproductive anatomy, and incongruence of the mitochondrial DNA (mtDNA) lineage and morphology. Hybridization appears to be asymmetric, however, as all morphological intermediates possessed the R. convicta mtDNA. Interbreeding between these deeply divergent forms challenges the use of reproductive structures as indicators of reproductive isolation in snail taxonomy. This is the first contact zone detected between mainland species of Rhagada and indicates that the widespread pattern of geographical replacement is not simply a result of allopatric divergence without subsequent contact. Instead, the perspective for understanding the radiation and geographical distributions must include interactions between the species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 348–362.     

Forest snail faunas from Crimea (Ukraine), an isolated and incomplete Pleistocene refugium

The land snail faunas of 26 forest sites and two open rocky sites in the Crimean Mountains were sampled in 2011. Of the 40 species found within the forests (about half the known fauna of Crimea as a whole), 28 were species with wide western Palaearctic distributions, and only eight were endemic to Crimea. While there were significant differences in the faunas of different sampling areas, these seemed to be a consequence of ecological differences among them rather than a product of geographical isolation and differentiation. Endemic species were large, and not entirely restricted to forest; known endemics not found in these forests are mainly typical of more open habitats. There is no local radiation of small species living in damp forest litter, as with Leiostyla species in the Transcaucasian forest refugium, and families such as the Clausiliidae with many endemic forest species in both Transcaucasia and the Carpathians are sparsely represented. The one endemic clausiliid genus, Mentissa, occurs in open as well as in wooded habitats. The present faunas are rather poor considering the soil conditions and climate, and the forests hold widespread species often associated with open habitats elsewhere. While there is evidence that these mountains provided a refuge for many animals and plants during glacial episodes further north, the forest snail fauna suggests that full forest cover did not survive throughout the Pleistocene. Rather, the present fauna contains endemics that survived in other habitats and widespread species with good powers of passive dispersal. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 424–433.     

Forest snail faunas from Georgian Transcaucasia: patterns of diversity in a Pleistocene refugium

Thirty forest sites in Georgia were sampled to obtain inventories of their land snail faunas. The sites, in nine sampling areas, ranged from Batumi near the Turkish border to Lagodekhi near the Azerbaijan border, and included parts of both the Lesser and Greater Caucasus mountains and the Colchic lowlands. Abundance was generally low, and most sites held less than 20 species. Ninety species were found overall. Although sites within some sampling areas had very similar faunas, the overall levels of similarity among sites and sampling areas was low, especially when compared with similar forests from northern central Europe. Both climate and geographical position affected similarity: although these were correlated, independent effects of each were detected, and the effect of position alone was strongest. Despite this effect, most species have ranges extending into Turkey or Ciscaucasia, a few have very limited ranges, and there is no evidence of patterns of allopatric replacement among congeners. These patterns resemble those seen in faunas from forests in lower latitudes, and suggest a slow build‐up of regional diversity by differentiation in situ, without the repeated restrictions to a multiplicity of small refugia that are characteristic of areas where fluctuations in aridity are important. The refugium has remained isolated, and has contributed little to the Holocene recolonization of areas further north. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 239–250.     

Calcareous forest seepages acting as biodiversity hotspots and refugia for woodland snail faunas

Land-snail species richness has repeatedly been found to increase with the increasing site calcium content and humidity. These two factors, reported as the main drivers of land-snail assemblage diversity, are also among the main habitat characteristics of calcareous seepages. Here we explore local species richness and compositional variation of forest spring-fed patches (i.e. seepages), to test the hypothesis that these habitats might act as biodiversity hotspots and refugia of regional snail faunas. In contrast to treeless spring fens, only little is known about land snail faunas inhabiting forest seepages. Studying 25 isolated calcareous forest seepages, evenly distributed across the White Carpathians Protected Landscape Area (SE Czech Republic), we found that these sites, albeit spatially very limited, can harbour up to 66% of the shelled land-snail species known to occur in this well-explored protected area (in total 83 species). By comparing land snail assemblages of the studied seepages with those occurring in the woodland surroundings of each site as well as those previously sampled in 28 preserved forest sites within the study area, we found the seepages to be among the most species rich sites. Although the numbers of species did not statistically differ among these three systems, we found highly significant differences in species composition. Seepage faunas were composed of many species significantly associated with spring sites, in contrast to the assemblages of both surrounding and preserved forest sites. Our results highly support the hypothesis that calcareous forest seepages might serve as refugia and biodiversity hotspots of regional land snail faunas. Protection of these unique habitats challenges both conservation plans and forest management guidelines as they might act as sources for the recolonization and restoration of forest snail assemblages particularly in areas impoverished by harvesting and clearcutting.     

A random process may control the number of endemic species

The richness of endemic species is often recognized as an indication of the distinctiveness of certain local faunas and is used for the definition of conservation hotspots as well. Faunas of different animal taxa were considered in sets of contiguous geographical units. Comparing the faunas of different units in one set, we found an exponential increase in the number of endemics when plotted against the number of non-endemics. A model of independent stochastic population dynamics under the control of environmental oscillations produces random fluctuations in the ranges of species. Ranges of endemic species are supposedly narrower than ranges of co-occurring non-endemic species. In such a case, the flow of a random process leads to an exponential relationship between numbers of co-occurring endemic and non-endemic species. This process also produces an apparent positive correlation between total species number and the percentage of endemics.     

Land snail diversity in the monsoon tropics of Northern Australia: revision of the genus Exiligada Iredale, 1939 (Mollusca: Pulmonata: Camaenidae), with description of 13 new species

Vast parts of the monsoon tropics of Australia feature semi‐arid habitats, which are generally thought to harbour a depauperate land snail fauna as compared to the mesic continental fringes, in particular the Australian wet tropics. However, our knowledge of land snails inhabiting these often remote environments is still very patchy. In order to improve the understanding of land snail diversity in the monsoon tropics, we revised the camaenid land snail genus Exiligada based on comprehensive collections, undertaken by use of helicopters on remote limestone outcrops in the Northern Territory and in Western Australia. Based on comparative analyses of shell and genital morphology and patterns of molecular differentiation, we recognize 15 species within Exiligada, 13 of which are newly described. In addition, we suggest a revised delimitation of the type species Exiligada negriensis, as compared to the latest available revision, by removing Exiligada qualis from its synonymy and recognizing it as a distinct species. A key for species identification is also provided. Molecular phylogenetic analyses strongly supported the monophyly of Exiligada with respect to other confamilial genera known to occur in the same geographical region. Most Exiligada species are narrowly endemic to restricted limestone outcrops that cover areas with a diameter of about 20 to 150 km. Within the distributional range of Exiligada, the ranges of up to seven species overlap but we never found more than three species to occur in sympatry at a given sampling site. We propose that species originated by allopatric differentiation, followed by secondary range expansion, leading to sympatric distributions. Our study confirms that less complex rock habitats in more xeric environments support no more than three sympatric species, this being likely to be a result of ecological exclusion. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 689–722.     

Drivers of Central European urban land snail faunas: the role of climate and local species pool in the representation of native and non-native species

The importance of macroclimate and dispersal limitation in the broad-scale variation of European urban land snail assemblages is likely to differ between native and non-native species because of the southern origin of many non-native snails, often spread by humans. We sampled land snails in each of 32 European cities and compiled from the literature a list of land snail species reported from the surroundings of each city. To quantify the predictive power of climate and local species pools, beta-sim dissimilarity matrices of both native and non-native species were explored using MDS and RDA ordination methods, Mantel tests with bootstrapping of each dataset, and multivariate homogeneity analysis of group variances. We observed no significant relation between the numbers of non-native species found in the cities and their surroundings (p > 0.133), while the percentage of native species in the cities derived from their local species pools decreased significantly with the increasing species richness of local faunas (r_S = ?0.75, p < 0.001). Assemblage variation of urban native species was explained mostly by the difference between mean January and July temperatures (21.3 %), with the major role of July temperature (18.0 %). In contrast, variation of non-native species assemblages was mainly explained by January temperature (19.9 %). The congruence in faunal similarities between the cities and the surrounding areas was higher in native (r = 0.46, p < 0.001) than in non-native species (r = 0.36, p < 0.001). Overall native faunas were significantly more homogeneous than the non-native faunas. Our results suggest that recent climate warming may foster geographical expansions of many non-native land snail species as their distributions are controlled mainly by January temperature.     

The biogeography of land snails in the islands of the Gulf of Guinea

Data on land snail diversity in the Gulf of Guinea islands is presented and the biogeography of the Gulf of Guinea fauna is discussed with reference to island snail faunas in the other areas of the world. Although the land snail faunas of the four islands clearly have west and central African affinities, speciation events following rare colonizations have led to high rates of endemism at the species and generic levels. The influence of island size, altitude, isolation and other factors on the development of the land snail faunas is discussed. The effect of land use changes on the land snail fauna over the last 150 years is evaluated. Current threats include recent changes in agricultural practice leading to the destruction of snail habitats in both forest and plantation areas. The potential threat posed by introduced species is also discussed. The Gulf of Guinea snail faunas appear to be in relatively good health in comparison to many other island snail faunas. However, the uniqueness of the faunas makes it imperative that further ecological and taxonomic research is undertaken both to understand the processes by which they developed and to evaluate the current status of many species in terms of distribution and threats.     

Patterns of land snail diversity over a gradient of habitat degradation: a comparison of three Czech cities

Terrestrial snail faunas were studied at 273 sites in three large Czech cities (Prague, Pilsen and Hradec Králové). The aim was to explore the effect of habitat degradation on land snail diversity in urban environments. Of the 81 species recorded, 12 were considered synanthropic and 29 anthropophobic. We assessed the power of habitat degradation and distance from the city centre as predictors of species richness for: (1) all species combined; (2) synanthropic species alone; and (3) anthropophobic alone. The highest correlation was observed between richness of athropophobic species and habitat degradation (average r = −0.69, P < 0.001), which explained on average 48% of the variation, whereas distance from the city centre explained a negligible amount of independent variation. A similar pattern was found for all species combined; however, the amount of explained variation was lower (10% on average). We found a contrasting pattern for the richness of synanthropic species whose occurrence was mainly and negatively driven by the distance from the city centre. However, the amount of explained variation was very unequal among the cities, being very low in Prague and Pilsen (8%) and quite high in Hradec Králové (36%). In conclusion, the response of local snail assemblages along a gradient of habitat degradation can be generalised as a gradual decline in species richness, especially of rare and anthropophobic species. This reduction was independent of the sites’ position within the city, which highlights the importance of natural habitats for the maintenance of snail diversity in urban environments.     

Forest snail diversity and its environmental predictors along a sharp climatic gradient in southern Siberia

Diversity patterns of forest snail assemblages have been studied mainly in Europe. Siberian snail faunas have different evolutionary history and colonization dynamics than European faunas, but studies of forest snail diversity are almost missing from Siberia. Therefore, we collected snails at 173 forest sites in the Russian Altai and adjacent areas, encompassing broad variation in climate and forest types. We found 51 species, with a maximum of 15 and an average of seven species per site. The main gradient in species composition was related to soil pH, a variable that also positively correlates with snail abundances. The second gradient was associated with climate characteristics of winter. We observed significant differences in both species richness and composition among six forest types defined based on vegetation classification. Hemiboreal continental forests were the poorest of these types but hosted several species characteristic of European full-glacial stages of the Late Pleistocene. A high snow cover in Temperate coniferous and mixed forests, protecting the soil from freezing, allowed the frost-sensitive large-bodied (>10 mm) species to inhabit this forest type. In contrast to most of the European snail assemblages studied so far we found that the factors responsible for the variation in species richness differed from those driving species composition. This may be attributed to the sharp climatic gradient and the presence of the cold-adapted species typical of the Pleistocene cold stages. We suggest that southern Siberian forests hosting these species can serve as modern analogues of full-glacial forests in periglacial Central and Eastern Europe.     

Genus age,provincial area and the taxonomic structure of marine faunas

Species are unevenly distributed among genera within clades and regions, with most genera species-poor and few species-rich. At regional scales, this structure to taxonomic diversity is generated via speciation, extinction and geographical range dynamics. Here, we use a global database of extant marine bivalves to characterize the taxonomic structure of climate zones and provinces. Our analyses reveal a general, Zipf–Mandelbrot form to the distribution of species among genera, with faunas from similar climate zones exhibiting similar taxonomic structure. Provinces that contain older taxa and/or encompass larger areas are expected to be more species-rich. Although both median genus age and provincial area correlate with measures of taxonomic structure, these relationships are interdependent, nonlinear and driven primarily by contrasts between tropical and extra-tropical faunas. Provincial area and taxonomic structure are largely decoupled within climate zones. Counter to the expectation that genus age and species richness should positively covary, diverse and highly structured provincial faunas are dominated by young genera. The marked differences between tropical and temperate faunas suggest strong spatial variation in evolutionary rates and invasion frequencies. Such variation contradicts biogeographic models that scale taxonomic diversity to geographical area.     

The influence of recent geography, palaeogeography and climate on the composition of the fauna of the central Aegean Islands

Partial Mantel tests and structural equation models were used to investigate the influence of recent geography, palaeogeography and climate on the composition of the fauna of the central Aegean Islands. The composition of land snail and isopod island faunas was significantly influenced by recent and by Pliocene geography. Only Pleistocene palaeogeography had a significant influence on the composition of tenebrionid beetle island faunas. The composition of butterfly island faunas was influenced by recent and by Miocene geographical distances. The composition of reptile island faunas was correlated with recent and Pliocene geography as well as with Pleistocene and/or Miocene geographical distances. Island area influenced only the composition of the island faunas of the volant butterflies, and not that of the less mobile land snails, land isopods, tenebrionid beetles and reptiles. This might indicate that butterflies are able to colonize large islands with suitable habitats even if such islands are comparatively far from source areas more frequently than can the nonvolant groups. Influence of a climatic parameter, namely annual precipitation, on faunal composition was found only for reptiles.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 785–795.     

Miocene land snails from Belchatow (Central Poland), I. Cyclophoridae,Pomatiasidae (Gastropoda Prosobranchia)

Three calcareous intercalations within lignite sequences in the open-cast mine Belchatów (51°15’N, 19°20’E) yield rich land snail faunas of over 40 species. Fission-track ages of these faunas (Be?-A and Bel-B) range from 18.1 ± 1.7 Ma to 16.5 ± 1.3 Ma, though the Be?-C level lies below the dated tuffitic intercalations and seems to be older. Five species of Cyclophoridae and Pomatiasidae are described including two new species:Procydotella?glazeki n. sp. andCaspicyclotus belchatoviensis n. sp.     

Gastropod associations within Malham Tarn, Yorkshire

A survey of gastropod species associations was undertaken in one locus, i.e. Malham Tarn, Yorkshire. This ranged through both the major weed beds and the rocky shores. A semiquantitative analysis, in terms of both presence-absence and relative abundance indices, suggested a clear distinction between the snail fauna present in the weed beds and that on the rocky shores, and also indicated a separation of faunas within these two major habitat types. The latter was closely correlated with degree of exposure. From this, and other evidence, it was suggested that wind, via wave action, may be a critical determinant of both the qualitative and quantitative aspects of biocoenotic structure in large bodies of fresh water.     

Molecular phylogeny and taxonomic revision of the genera Baudinella Thiele, 1931, Retroterra Solem, 1985 and Molema Köhler, 2011 endemic to the coastal Kimberley,Western Australia (Gastropoda,Camaenidae)

Baudinella Thiele, 1931, Retroterra Solem, 1985 and Molema Köhler, 2011 are three genera of camaenid land snail endemic to coastal regions of the Western Australian Kimberley. Each of these genera has fairly distinct shells, but all exhibit a rather similar configuration of the reproductive system, which is characterized by lack of a penial sheath and presence of an elongated and coiled bursa copulatrix. By combining comparative morphology of shell and penial anatomy with analyses of the mitochondrial DNA fragments 16S rRNA (16S) and cytochrome c oxidase subunit 1 (COI), the phylogenetic relationships amongst representative species of these genera are addressed to test the monophyly of taxa and to identify new species. Our results show that all three genera are members of a single camaenid radiation. Five new species are described: two new species of Baudinella, B. magna n. sp. from the Institute and Montesquieu Archipelagos and B. margaritata n. sp. from Pitta Gorge in the Prince Regent Reserve, one new species of Molema, Molema tenuicostata n. sp., from near Talbot Bay, and two new species of Retroterra, R. dichroma, and R. nana, from the Prince Regent Reserve.     

Distribution,movement, and microhabitat use of the introduced predatory snail Euglandina rosea in Hawaii: implications for management

The purposeful introduction of the land snail Euglandina rosea, which feeds exclusively on snails and slugs, has been implicated as a major factor in the decline of diverse Pacific island land snail faunas. We report on the distribution, movement patterns, and microhabitat preferences of E. rosea in a gulch in the Waianae Mountains, Oahu, Hawaii, because such data will help focus management actions at a local scale to protect native snail populations in areas where E. rosea is established. The Waianae Mountains harbor many endangered or threatened snails, most currently found in isolated habitat patches near the ridges. Conversely, most living individuals (28/29) and shells (46/56) of E. rosea were collected within the gulch, which supported higher densities of other native and non‐native snails, and was cooler and more moist than the ridges. Thirteen individuals of E. rosea were tracked (eight directly using a bobbin and thread method, and five indirectly by mark–recapture); most (10/13) moved on average <2.5 m per week (range 0.1–25.21 m), and all stayed within the gulch. Members of E. rosea preferred leaf litter over open, fern/shrub, or wood microhabitats. There were large differences in the population density of E. rosea over small spatial scales, indicating that there may be places where native snail populations could persist even in areas where populations of E. rosea are established. Identifying areas with differing population densities of E. rosea is critical for not only understanding why some native snail species may be more vulnerable to extinction, but also to locate areas where predation pressure is low and conservation efforts will be most likely to succeed.     

Forest snail faunas from Transylvania (Romania) and their relationship to the faunas of Central and Northern Europe

Forty‐three forest sites in seven sampling areas in mountainous parts of Transylvania were sampled to obtain inventories of their snail faunas, and to make comparisons between these and the faunas of similar sites further north along the Carpathian chain. Sampling areas ran from close to the Ukrainian border in the north to Retezat in the south‐west. Altogether, 83 species were found, as well as between 19 and 40 species at individual sites. Sites within the same sampling area had more species in common than in among‐area comparisons, although differences between areas did not relate to the distance between them. Such differences appear to be related more to ecological factors than to geography. Faunas of all areas were very similar to those recorded from the Polish and Ukrainian Carpathians; distance from these northern faunas had little effect on similarity. More broadly, the fauna of these forests shows a much greater affinity to those of countries to the north than to those of countries to the south. Despite the survival of forests through the last glacial period of the Pleistocene, and the greater proximity to potential Mediterranean refugia, these faunas appear to be mainly a subset of those found further north. They represent a rather small proportion of the known Romanian fauna, in which there are many endemics restricted to other habitats. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 471–479.     

Biogeographical tests of the vicariance model in Mediterranean land snails

Aim To investigate the importance of vicariance in shaping land snail faunas. Location Three data sets of Mediterranean land snails were analysed: Helicoidea of the Iberian Peninsula and the complete land snail faunas of the central and eastern Aegean Islands and of Israel and Palestine. Methods The vicariance model predicts a clustering of species ranges. We tested for clustering of species ranges with a Monte Carlo simulation. For this simulation we used a null model that generates range data sets in such a way that their range size distribution, their species richness distribution and the spatial autocorrelation of the ranges approximate the parameters in the real data set. Biotic elements (clusters of species ranges) were delimited with model‐based Gaussian clustering. A second prediction of the vicariance model is that closely related species belong to different biotic elements. This was tested with a chi‐squared test. Results The distribution areas of the Iberian Helicoidea and the Israeli/Palestinian land snails are significantly clustered. The same is true for Israeli/Palestinian land snail species belonging to groups with geographically restricted species. However, the clustering is not significant in the complete central and eastern Aegean land snail data set, the Iberian Helicoidea species, and the central and eastern Aegean land snails belonging to groups with geographically restricted species. Contrary to the prediction of the vicariance model, closely related Iberian Helicoidea species and Israeli/Palestinian land snails belong significantly more often to the same biotic element than expected by chance. The null hypothesis that closely related species are homogeneously distributed across biotic elements cannot be rejected only for the data set that includes the Israeli/Palestinian land snails belonging to groups with geographically restricted species. Main conclusions The patterns found in the central and eastern Aegean land snail and the Iberian Helicoidea data sets do not correspond with the predictions of the vicariance model. This indicates that speciation modes other than vicariance were frequent, or that the distribution areas of many species in these faunas were largely modified by extensive post‐speciation dispersal and/or regional extinction. Hardly any possible vicariance events that might have contributed to the origin of the observed biotic elements in the Israeli/Palestinian land snail fauna could be identified. On the contrary, the biotic elements of the Israeli/Palestinian land snail fauna correspond with ecological factors. Vicariance has had only a limited influence on current biogeographical patterns in Mediterranean land snails.     

Altitudinal species turnover in southern Tanzania shown by anurans: Some zoogeographical considerations

A transect was used to investigate the pattern of altitudinal species turnover shown by the rich fauna of frogs and toads in southeastern Tanzania, and to compare the pattern with those shown by transects taken further south. The transect, taken from the Udzungwa range to the southeastern coast, was divided into 50 m intervals. As with the southern transects, there appears to be a major grouping into lowland and highland sets, with opposing and overlapping subtraction margins that form a replacement‐transition zone. A distributionally complex assemblage of species occurs in this zone. When a similarity value (the Dice‐S?rensen index, expressed as a percentage) taken between the montane ridge and lowland faunas was compared with those of the southern transects, a markedly higher species turnover (low similarity value) is shown in Tanzania. Both lowland and highland faunas have more species than equivalent faunas to the south; the highland fauna is particularly rich in genera not known to occur further south. Yet overall, the pattern of altitudinal turnover shown in the Tanzanian transect is in agreement with the pattern shown in the southern transects. The transects discussed in this paper, combined with other evidence, suggest a need for re‐evaluating the common practice of including all sub‐Saharan Africa in a single so‐called Afrotropical Region.     

Biogeographic patterns in the Australian chondrichthyan fauna

The major biogeographic structure and affinities of the Australian chondrichthyan fauna were investigated at both interregional and intraregional scales and comparisons made with adjacent bioregions. Faunal lists were compiled from six geographical regions with species from these regions assigned to distributional classes and broad habitat categories. Australian species were further classified on provincial and bathomic structure following bioregionalization outputs from regional marine planning. About 40% of the world's chondrichthyan fauna occurs in Indo-Australasia (482 species) of which 323 species are found in Australian seas. The tropical Australian component, of which c. 46% of taxa are regional endemics, is most similar to faunas of Indonesia, New Guinea and New Caledonia. The temperate Australian component is most similar to New Zealand and Antarctica with about half of its species endemic. Highest levels of Australian endemism exist in bathomes of the outer continental shelf and upper slope. A relatively high proportion of regional endemism (57% of species) on the slope in the poorly surveyed but species-rich Solanderian unit is probably due to high levels of large-scale habitat complexity in the Coral Sea. The richness of demersal assemblages on the continental shelf and slope appears to be largely related to the spatial complexity of the region and the level of exploration. Much lower diversity off Antarctica is consistent with the pattern in teleosts. The complex chondrichthyan fauna of Australia is confirmed as being amongst the richest of the mega-diverse Indo-West Pacific Ocean. Species-level compositions of regional faunas across Indo-Australasia differ markedly because of moderate to high levels of intraregional speciation. Faunal assemblages in Australian marine provinces and bathomes differ from each other, supporting a broader pattern for fishes that underpins a marine planning framework for the region.