Diversity of Central European urban biota: effects of human‐made habitat types on plants and land snails

Aim Urbanization is associated with strong changes in biodiversity, but the diversity of plant and animal assemblages varies among urban habitats. We studied effects of urban habitats on the diversity of vascular plants and land snails in 32 large cities. Location Central Europe, Belgium and the Netherlands. Methods The species composition of all vascular plants that had not been planted by humans, and all land snails, was recorded in seven 1‐ha plots within each city. Each plot contained one urban habitat type representing a different disturbance regime: historical city square, boulevard, residential area with compact building pattern, residential area with open building pattern, park, early successional and mid‐successional site. For each plot, we obtained temperature and precipitation data. The effects of climate and habitat types on species composition were quantified using ordination methods with an adjusted variation partitioning algorithm. Differences in species composition among urban habitats were described using statistically determined diagnostic species, and differences in alpha, beta and gamma diversity were quantified. Results A total of 1196 plant and 87 snail species were recorded. Habitat type explained higher proportions of the total variation in both plant and snail species composition (11.2 and 8.2%, respectively) than did climate (4.6 and 6.3%). For both taxa, the main differences in species composition were observed between strongly urbanized sites in city centres and early successional and mid‐successional sites. For vascular plants, the number of species was lowest in city squares and boulevards, and highest at successional sites and in residential areas with compact building patterns. Beta diversity of vascular plants calculated for the same habitat types among cities was highest for squares and successional sites. The number of snail species was lowest in city squares and at early successional sites, and highest at mid‐successional sites. The highest beta diversity of snail assemblages among cities was observed within the city square and early successional habitat types, and the lowest within residential area habitat types. Main conclusions Urban habitats differ notably in the diversity of their vascular plant flora and land snail fauna. Understanding the habitat‐related biodiversity patterns in urbanized landscapes will allow projections of future impacts of urban land‐use changes on the biota.     

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.     

High diversity and regional endemism in land snails of eastern Tanzania

In February/March 1995 we collected land snails (including slugs) at 12 stations in eastern Tanzania. A total of 571 person-hours yielded 9174 snails assigned to 159 morpho-species. The richest two sites each (<4ha of uniform forest) had 50 species (Amboni Cave) and 48 species (near Amani, Usambaras), nearly as great as the most species-rich sites known in the world; sieving of litter and soil would probably yield more species. In lowland (coastal) forests, both diversity and endemism seemed to decrease from north to south. Most snail species were found within only one of four coastal or one montane geographic regions, indicating substantial regional endemism. Only one species (Achatina fulica) appeared in all five regions, and 84% of all other species were found in only one (61%) or two regions (23%). The predatory streptaxids comprised about half the species and a third of the individuals at the Usambara site, an extremely high ratio of carnivores. Small snails (< 5mm greatest adult shell dimension) – many of which are probably undescribed species – comprise a substantial proportion of Tanzanian molluscan diversity; more surveys are needed, especially because of human pressures on the few forest patches remaining.     

The distribution of pond snail communities across a landscape: separating out the influence of spatial position from local habitat quality for ponds in south-east Northumberland,UK

Ponds support a rich biodiversity because the heterogeneity of individual ponds creates, at the landscape scale, a diversity of habitats for wildlife. The distribution of pond animals and plants will be influenced by both the local conditions within a pond and the spatial distribution of ponds across the landscape. Separating out the local from the spatial is difficult because the two are often linked. Pond snails are likely to be affected by both local conditions, e.g. water hardness, and spatial patterns, e.g. distance between ponds, but studies of snail communities struggle distinguishing between the two. In this study, communities of snails were recorded from 52 ponds in a biogeographically coherent landscape in north-east England. The distribution of snail communities was compared to local environments characterised by the macrophyte communities within each pond and to the spatial pattern of ponds throughout the landscape. Mantel tests were used to partial out the local versus the landscape respective influences. Snail communities became more similar in ponds that were closer together and in ponds with similar macrophyte communities as both the local and the landscape scale were important for this group of animals. Data were collected from several types of ponds, including those created on nature reserves specifically for wildlife, old field ponds (at least 150 years old) primarily created for watering livestock and subsidence ponds outside protected areas or amongst coastal dunes. No one pond type supported all the species. Larger, deeper ponds on nature reserves had the highest numbers of species within individual ponds but shallow, temporary sites on farm land supported a distinct temporary water fauna. The conservation of pond snails in this region requires a diversity of pond types rather than one idealised type and ponds scattered throughout the area at a variety of sites, not just concentrated on nature reserves. Handling editor: B. Oertli     

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.     

Habitats of relict terrestrial snails in southern Siberia: lessons for the reconstruction of palaeoenvironments of full‐glacial Europe

Aim Shells of fossil molluscs are important for palaeoecological reconstructions. However, the habitat requirements of snail species typical of central European full‐glacial loess sediments are poorly known because most of them became very rare or extinct in Europe. The recent discovery of an almost complete extant assemblage of such species in mountainous regions of central Asia enables more precise characterization of their habitats, which may significantly improve reconstructions of Pleistocene environments. Location Altai Mountains, Russia. Methods Terrestrial snail assemblages, vegetation composition and selected environmental variables were recorded at 118 sites along a gradient of climatic continentality in the Russian Altai. Habitat characteristics of sites where species typical of the full‐glacial period occurred were described using a classification tree. Results Seven of the eight species that are typical of central European full‐glacial loess sediments were found in the study area. They were confined to cool areas with January mean temperatures below ?17 °C, but occurred mainly in sheltered habitats with a warmer microclimate, such as scrub or open woodland. Pupilla loessica and Vallonia tenuilabris had the broadest habitat range, occurring from woodland to dry steppe. Unexpectedly, Columella columella, Pupilla alpicola, Vertigo genesii, V. parcedentata and V. pseudosubstriata were found mainly in wooded fens and shrubby tundra rather than in open steppe. Most of these seven species were recorded in base‐rich wooded fens. Very dry open steppe habitats usually supported no snails. Main conclusions Habitat ranges of the studied snails in the Altai indicate that the full‐glacial landscapes of central European lowlands that harboured these species were not completely dominated by open and dry loess steppe. Most probably they contained a significant component of shrubby vegetation, patches of wet habitats, and probably also areas of woodland at sites with a favourable mesoclimate.     

Patterns of diversity and habitat relationships in terrestrial mollusc communities of the Pukeamaru Ecological District, northeastern New Zealand

Aim The New Zealand terrestrial mollusc fauna is among the most speciose in the world, with often remarkably high richness at lowland forest sites. We sought to elucidate general explanations for patterns of richness in terrestrial mollusc communities by analysis of species coexistence and habitat relationships within a New Zealand district fauna. Location Pukeamaru Ecological District, eastern North Island, New Zealand. Methods We sampled molluscs using qualitative methods at twenty-three sites and quantitatively by frame sampling of scrubland-forest floor litter at sixteen of these sites and analysed patterns of species richness and turnover in relation to regional species pools and local habitat attributes. We then tested for nonrandom assemblage of taxa along diversity and habitat gradients. Results Ninety-four indigenous mollusc species were recorded from a district fauna estimated at 102 indigenous species: only two species were endemic. From the presumptive geological history of the district, the low endemism, and Brooks parsimony and indicator species analyses of faunal relationships, the communities were indicated to have resulted by accumulation of colonists from other New Zealand districts since the Miocene. Richness ranged from two or three indigenous species in dune habitats to fifty-nine species in a floristically rich forest. Beta diversity was high and site occupancy per species was low, indicating communities structured by successive replacement of ecological equivalents. Sites differing in vegetation had characteristic species assemblages, indicating a degree of habitat specialization. Canonical correspondence analysis indicated that canopy tree species, canopy height, floristic diversity, altitude, litter mass, and litter pH were important determinants of species assemblage in scrubland and forest. Richness was strongly associated with site floristic diversity and, for litter-dwelling species, the pH of litter substrate. High richness occurred at those sites supporting molluscs in high abundance. Shell-shape distributions were essentially Cainian unimodal, with communities dominated by snail species with subglobose to discoidal shells. Mean and variance of shell size increased with mollusc species richness and floristic diversity at sites, indicating dominance of communities by small-shelled species at early successional or floristically poor sites, and increased richness resulting from addition of larger snails into vacant niches. Shifts in shell form were associated with sympatry in several congeneric taxa. Main conclusions The underdispersion of shell shape, relative to faunas elsewhere in the world, indicates that community structure in New Zealand land snail faunas has been constrained by limited phylogenetic diversity and/or by convergence upon successful adaptations. The remarkably high richness that characterizes these communities indicates special conditions allow coexistence of numerous species. The relationship between floristic diversity at sites and the richness, diversity, and shell-size distributions of the molluscs suggests assemblages structured around niche partitioning among competing species. While there is an element of congruence between vegetation and mollusc pattern, this study indicates that assembly rules will be defined, and spatial pattern predicted, only through a better understanding of the linkage between regional species pool, organism traits, environment, and local community assemblage.     

Diversity of the Western Carpathian flysch grasslands: Do extremely species-rich plant communities coincide with a high diversity of snails?

Upland fringes of the White Carpathians (Czech Republic) are known to support grasslands with the world’s highest local plant species richness. We investigated whether this unusually high plant richness has a parallel in snail communities, whether patterns of species composition of snail and plant communities in grasslands co-vary and how they are affected by local environment and landscape history. We compared plant and snail communities of dry to mesic grasslands in three neigh bouring regions: (1) hilly lowland of the Central Moravian Carpathians, (2) upland fringes and (3) upland of the White Carpathians. Both snail and plant communities exhibited a strong gradient in species composition associated with altitude, annual temperature and precipitation, soil calcium and pH. However, there was no correlation between local species richness of plants and snails in individual plots. The upland fringes of the White Carpathians were richest in snail species, probably due to intermediate environmental conditions, supporting the occurrence of species with contrasting environmental requirements. The highest local numbers of plant species were also recorded there, although differences among regions were not significant. The regional species richness of plants was also highest in the upland fringes, whereas that of snails was highest in the hilly lowland. Similarities in the diversity patterns of plants and snails among regions suggest the importance of regional factors for local richness, although local abiotic factors, which are partly correlated with the three regions, also influence local species composition and richness.     

鄱阳湖沿岸带螺类的物种组成及其生境

2011年5月和6月在鄱阳湖沿岸带的62个样点采集了淡水螺类样本并调查了生境类型,分析了淡水螺类的物种组成及其生境分布特征。结果表明:共采集到淡水螺类19种,分属田螺科（8种）、觿螺科（8种）、椎实螺科（2种）和黑螺科（1种）。与前人的研究相比,本次调查的螺类物种数有较大的下降,但田螺科和觿螺科的物种仍是整个螺类群落的主要组成部分。淤泥生境的螺类物种组成与其他生境的差异较大,淤泥生境的螺类物种丰富度显著高于沙土、砂石和水草3种生境（P0.05）,表明螺类对生境具有一定的栖息偏好性。最后详细讨论了淡水螺类种数变化、物种组成及其生境分布的影响因素。       

Bellwether of the Canaries: anthropogenic effects on the land snail fauna of the Canary Islands

Natural areas near human-modified landscapes experience factors that may affect local biodiversity at levels commensurate with natural environmental factors. The land snails of the Canary Islands provide excellent opportunities to evaluate the importance of anthropogenic agents in mediating the diversity and distribution of species. Land snails are particularly sensitive to disturbance and are an integral part of terrestrial ecosystems. This study analyzed the distributions and abundances of terrestrial macrosnail shell assemblages at 60 localities throughout the coastal scrub biome of the Canary Islands. This was accomplished using data on natural and anthropogenic variables to assess their relative importance in governing land snail diversity. A total of 34,801 dead shells represented a diverse malacofauna with highly localized endemism. Due to uncertain species identifications, samples from the 18 sites from the two easternmost islands are described, but excluded from statistical analyses. Regression tree analysis indicated that proximity to agricultural sites was the most important predictor of species diversity. Sites with no or very little agricultural area (≤ 0.167 km²) within a 1 km radius had significantly higher richness and diversity. These results have implications for Canary Islands conservation. Protected areas that are patchworks of natural and agricultural landscapes are still subject to native biodiversity loss because of anthropogenic impacts even when the footprint of agriculture is small.     

On the flexibility of adaptation processes in cold-hardy insects

A shift in the adaptation strategy depending on the temperature regime within the species range has been revealed in three species of cold-hardy insects inhabiting central Yakutia: Pieris rapae, Acanthocinus aedilis, and Cossus cossus. In warmer climates of northern Europe, these species adopt the strategy of freezing avoidance, whereas in the extremely cold climate of central Yakutia they become freezing-tolerant. This shift probably provides for the ability of these species for unlimited migration and settling in habitats with diverse climatic conditions.     

Estimating and conserving patterns of invertebrate diversity: a test case of New Zealand land snails

Aim Using New Zealand land snails as a case study, we evaluated recent spatial modelling approaches for the analysis of diversity in species‐rich invertebrate groups. Applications and prospects for improved conservation assessment were investigated. Location New Zealand. Methods The study used a spatially extensive and taxonomically comprehensive, plot‐based dataset on community structure in New Zealand land snails. Generalized regression analysis and spatial prediction (GRASP) was used to model and predict species richness as a function of environmental variables (including aspects of climate, soils and vegetation). Generalized dissimilarity modelling (GDM) was used to model turnover in species composition in relation to environmental and geographical distances, and to assess community similarity and the representativeness of the reserve network. Results Observed land snail richness in 20 × 20 m plots ranged from 1 to 74 (mean 17.5). The GRASP model explained a modest 27% of the variation in richness. The GDM model explained 57% of the variation in species turnover and indicated approximately equal amounts related to environmental (Cody’s beta diversity) and geographical distance (Cody’s gamma diversity). Temperature and moisture were the most important environmental variables. Results indicate that snail distributions are not only sorted by environment but are also strongly influenced by historical effects consistent with those expected of poorly dispersing taxa that have persisted in refugia during past climatic change. The GDM model enabled spatial classifications of snail communities, highlighting diverse communities in heterogeneous regions, such as the South Island mountains, and also enabled continuous depictions of community similarity and adequacy of New Zealand’s protected natural areas network. Main conclusions The GRASP and GDM analyses allowed us to model and depict spatial patterns of diversity in land snail communities involving 845 species, and produce community classifications and estimates of community similarity. These tools advance conservation assessment in species‐rich groups, but require further conceptual and methodological development.     

Habitat and integration within indigenous communities of Xeropicta derbentina (Gastropoda: Hygromiidae) a recently introduced land snail in south-eastern France

Xeropicta derbentina is a recently introduced land snail in Provence, France. The present work aims at describing the habitats and the land snail communities that it has invaded. Also, its impact on the species richness and diversity of indigenous communities is assessed. Two areas have been studied in order to cover the range of conditions that X. derbentina is encountering within its current French distribution. Sixty‐five samples were analysed from the most Mediterranean site, Auriol, and 140 from a northern site, Craus de St‐Michel‐l’Observatoire. From these samples, X. derbentina is found to be a thermophilous open‐land species mostly inhabiting human‐impacted habitats. It occurs in large populations (more than 100 individuals collected in 15 min) in habitat characterizing the first stages of secondary successions, from cultivated crops to dense fallow lands. However, within a heterogeneous landscape and if enough time has allowed its dispersal, X. derbentina can be found in almost any kinds of habitat, such as garrigues and woodlands. Consequently, X. derbentina occurs in diverse land snail communities. At the local scale, its large populations could result in a significant decrease in species richness and diversity. At the regional scale, however, the introduction of X. derbentina seems to have increased land snail species richness.     

High resilience of Mediterranean land snail communities to wildfires

In the Mediterranean region, wildfires have devastating effects on animals with limited mobility. With their poor dispersal abilities, their habitats on vegetation and in litter, and their sensitivity to humidity and shade, we expected land snails to be an interesting model to assess short, medium and long-term impact of fires on fauna biodiversity and their resilience. Stratified sampling was carried out on 12 sampling sites in garrigues and forests of Provence (southeastern France), according to fire regime (number of fires, fire intervals and age of the last fire) over the past 30 years. Data were investigated using diversity indexes, Kruskal–Wallis test, dendrogram of affinities and Correspondence Analysis (CA). We found, however, that Mediterranean land snail communities are particularly resilient to fires. Although abundance is drastically reduced in the short-term, species richness and community diversity are preserved provided that the time lapse between two successive fires is longer than the time required for recovery (i.e. around 5 years). This high community resilience in the short-term may be partly due to ecological and ethological aptitudes of land snails. However, these astonishing results, which have implications for conservation biology, are mainly due to the presence, within burned areas, of cryptic refuges that allow initial land snail survival, malacofauna persistence after successive fires and consistent biogeographical patterns in the long-term.     

Priority sites for conservation of land snails in Gabon: testing the umbrella species concept

In order to assess the potential for molluscan conservation of a protected area considered representative of regional megafauna, we sampled molluscs inside and outside Lopé National Park in Gabon. In the northern part of Lopé National Park, 116 stations were surveyed and 71 species collected. Outside the park, 37 stations yielded 96 species, including 71 in Lastoursville, a small limestone area where molluscs are significantly more abundant than in other collecting sites. Lastoursville is among the richest sites known for molluscs in Africa. Overlap between sampling areas was limited, with 20.0% of the species found only in Lopé National Park, and 40.8% of the species found only outside. This suggests that Lopé National Park does not protect the whole molluscan diversity of central Gabon. Given the high levels of allopatric diversity of tropical land snails, conservation strategies cannot be the same for snails and for wide-ranging vertebrates. Protecting small areas with a high abundance and diversity of molluscs would be less expensive and as efficient for molluscan conservation as protecting large tracts of rainforest. Despite limited general knowledge of central African molluscs, robust estimates of site-specific diversity can be produced. Limestone areas harbour a remarkable biodiversity: sites such as Lastoursville would be ideal candidates for small protected areas dedicated to the conservation of land snails, and would complement the role of large protected areas.     

Agriculture erases climate‐driven β‐diversity in Neotropical bird communities

Earth is experiencing multiple global changes that will, together, determine the fate of many species. Yet, how biological communities respond to concurrent stressors at local‐to‐regional scales remains largely unknown. In particular, understanding how local habitat conversion interacts with regional climate change to shape patterns in β‐diversity—differences among sites in their species compositions—is critical to forecast communities in the Anthropocene. Here, we study patterns in bird β‐diversity across land‐use and precipitation gradients in Costa Rica. We mapped forest cover, modeled regional precipitation, and collected data on bird community composition, vegetation structure, and tree diversity across 120 sites on 20 farms to answer three questions. First, do bird communities respond more strongly to changes in land use or climate in northwest Costa Rica? Second, does habitat conversion eliminate β‐diversity across climate gradients? Third, does regional climate control how communities respond to habitat conversion and, if so, how? After correcting for imperfect detection, we found that local land‐use determined community shifts along the climate gradient. In forests, bird communities were distinct between sites that differed in vegetation structure or precipitation. In agriculture, however, vegetation structure was more uniform, contributing to 7%–11% less bird turnover than in forests. In addition, bird responses to agriculture and climate were linked: agricultural communities across the precipitation gradient shared more species with dry than wet forest communities. These findings suggest that habitat conversion and anticipated climate drying will act together to exacerbate biotic homogenization.     

Evaluating the role of regional and local processes in structuring a larval trematode metacommunity of Helisoma trivolvis

Metacommunity theory has advanced our understanding of how local and regional processes affect the structure of ecological communities. While parasites have largely been omitted from metacommunity research, parasite communities can provide the large sample sizes and discrete boundaries often required for evaluating metacommunity patterns. Here, we used assemblages of flatworm parasites that infect freshwater snails (Helisoma trivolvis) to evaluate three questions: 1) what factors affect individual host infections within ponds? 2) Is the parasite metacommunity structured among ponds? And 3) what is the relative role of local versus regional processes in determining metacommunity structure and species richness among ponds? We examined 10 821 snails from 96 sites in five park complexes in the San Francisco Bay area, California, and found 953 infections from six parasite groups. At the within‐pond level, infection status of host snails correlated positively with individual snail size and pond infection prevalence for all six parasite groups. Using an ordination method to test for metacommunity structure, we found that the parasite metacommunity was organized in a non‐random pattern with species responding individually along an environmental gradient. Based on a model selection approach involving local and regional predictors, parasite species richness and metacommunity structure correlated with both local abiotic (pH and total dissolved nitrogen) and biotic (non‐host mollusk density, and H. trivolvis biomass) factors, with little support for regional predictors. Overall, this trematode metacommunity most closely followed the predictions from the species sorting or mass effects metacommunity paradigm, in which community diversity is filtered by local site characteristics.     

Evaluating the efficacy of land snail survey techniques in Hawaii: implications for conservation throughout the Pacific

Terrestrial micromolluscs (snails with an adult maximum shell dimension <5 mm) constitute a considerable proportion of the land snail fauna of the Pacific. However, micromolluscs are often underestimated in biological surveys because of size bias. It has been argued that visual searches are preferable on Pacific islands because: (1) size biases are limited based on the understanding that most native Pacific island land snails are very small, and (2) amount of labor is less than other methods such as soil surveys and adequate for inventory purposes (though not for abundance assessments). To test whether visual surveys and soil surveys were accurately recording all taxa, land snail inventories were completed in three forest reserves (5 sampling sites in each) on the island of Oahu, Hawaii. Visual surveys involved 30-min visual search in a 10 m² site; soil surveys involved sieving leaf litter and topsoil from four 0.3 m² quadrats and extracting snails with the aid of a microscope. The data indicate a size and microhabitat bias associated with both techniques. Visual surveys consistently collected large arboreal and litter-dwelling species but missed a significant portion of micromolluscs, while soil surveys collected micromolluscs but missed larger snails. Because of such biases, employing both methods is critical for collecting all taxa at a survey location. As such, we recommend that future land snail surveys on Pacific Islands incorporate both survey techniques. Obtaining a complete inventory is critical if we are to understand species distributions and patterns of diversity and make well-informed conservation recommendations.     

Diversity patterns of the terrestrial snail fauna of Nyungwe Forest National Park (Rwanda), a Pleistocene refugium in the heart of Africa

We investigated the land snail fauna of Nyungwe Forest National Park in south‐western Rwanda. Fifty plots at altitudes between 1718 and 2573 m were studied. In total, 3461 specimens were collected and were assigned to 102 land snail species. With respect to land snail species, Nyungwe Forest is the richest forest known in Africa. A comparison with other forests in the northern Albertine Rift indicates that land snail species richness in this region is significantly correlated with distance from Pleistocene forest refugia. The high beta diversity in Nyungwe is the result of a high species turnover between sites, which has biogeographical and ecological origins. Nyungwe Forest is situated on the Congo–Nile divide where species of different geographical origin may meet. Moreover, Nyungwe Forest offers a high diversity of habitats because it extends across a wide range of altitudinal zones. Species richness decreased with increasing altitude. It was also correlated with the presence of bare rocks that offer additional microhabitats and shelter. Although the occurrences of different land snail species in Nyungwe Forest were significantly clustered, only a minority of the species could be assigned to a group of species with similar occurrences. The majority of the species respond individualistically to environmental variables. The significant nestedness of the occurrences of the land snail species in Nyungwe was mainly correlated with altitude. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 363–375.