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.     

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.     

The importance of littoral forest remnants for indigenous bird conservation in southeastern Madagascar

The littoral forests of Madagascar are relatively unexplored ecosystems that are considered seriously threatened by deforestation and habitat fragmentation. We set out to describe the bird communities inhabiting the littoral forest remnants in three different sub-regions of southeastern Madagascar to determine the national importance of these forests for bird conservation. In total, 77 bird species were found inhabiting 14 littoral forest remnants. Of these species, 40 are endemic to Madagascar and a further 21 are endemic to the Indian Ocean sub-region, consisting of Madagascar, the Comoros and the Mascarenes. The matrix habitats (Melaleuca forests, marécage swamp forest, Eucalyptus plantations and Erica grassland) that immediately surround the littoral forests were depauperate of bird species and contained few species that were found within the littoral forests. The geographic location of littoral forest remnants had an important role in determining what bird species occurred within them, with the northern remnants having similar bird communities to nearby humid forest whilst the most southern remnant had a bird community that resembled those of nearby spiny forest habitats. Eleven bird species that have been previously described as being habitat-restricted endemics to either spiny forests or humid forests, were found in littoral forest remnants. These results suggest that these littoral forests may play an important transitional role between the two other major natural habitats (spiny forest and humid forest) of southeastern Madagascar. On this basis we advocate that the littoral forest remnants of southeastern Madagascar should be afforded continuing conservation priority.     

Chance and necessity: land-snail faunas of Sao Miguel, Azores, compared with those of Madeira

The land-snail faunas of 23 sites from forest, disturbed andopen habitats were sampled on São Miguel, Azores archipelago.Collectively, the samples contained nearly all the known faunaof the island. Forest sites were the richest; while the numberof non-endemic species differed little between habitats, forestsheld more endemic species, many of which were widely distributed.Open habitats have no unique endemic species. These resultsare compared with similar data from Madeira. Although the twoislands are of the same size, and share the same Laurisilvaforest cover, their faunas differ in important aspects of diversity.While many non-endemic species are common to both islands, andoccur in similar numbers, endemic species are more numerouson Madeira, even though the numbers at individual forest sitesare, on average, fewer than on São Miguel. Many endemicsare confined to open, coastal habitats. These differences arediscussed in terms of the islands' histories, the range of habitatsavailable and the ease of dispersal from continental sources.Age, accidents of colonization and the range of natural habitatsavailable must have major effects on the faunas, but presentendemic faunal diversity is more a product of speciation andextinction than of immigration. (Received 13 May 2006; accepted 7 July 2006)     

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.     

Biogeographic patterns of the East African coastal forest vertebrate fauna

The archipelago-like coastal forest of East Africa is one of the highest priority ecosystems for biodiversity conservation worldwide. Here we investigate patterns of species richness and biogeographic distribution among birds, mammals and reptiles of these forests, using distribution data obtained from recently published reviews and information collated by the WWF Eastern Africa Coastal Forest Ecoregion Programme. Birds and mammals species were divided into forest specialists and generalists, and forest specialist reptiles into ‘coastal’ and ‘forest’ endemics. The species richness of birds and generalist mammals increased with area, and is probably a result of area-dependent extinction. Only in birds, however, species richness increased with decreasing isolation, suggesting possible isolation-dependent colonization. Forest diversity, associated to altitudinal range, is important for specialist birds and mammals, whose species richness increased with wider altitudinal range. The number of relict coastal endemic and forest endemic reptiles was higher in forests with wider altitudinal ranges and on relatively higher altitude, respectively. Such forests have probably provided a suitable (and perhaps stable) environment for these species through time, thus increasing their persistence. Parsimony analysis of distributions (PAD) and cluster analyses showed geographical distance and general ecological similarity among forests as a determinant factor in bird distribution patterns, with compositional similarity decreasing with increasing inter-forest distance. Compositional similarity patterns of mammals among the forests did not show a strong geographical correspondence or a significant correlation with inter-forest distance, and those of reptiles were not resolved, with very low similarity levels among forest faunas. Our results suggest that the relative importance (and causal relationship) of forest attributes affecting the distribution of the East African coastal forest vertebrate fauna varies depending on life history traits such as dispersal ability and forest specialization. The groupings in PAD are partly congruent with some of the previous classifications of areas of endemism for this region, supporting the ‘naturalness’ of these regions.     

Pattern and process in the distribution of North American freshwater fish

Published species lists were analysed to determine the contributions of dispersal, habitat preference, river channel size, body size, and glacial history to large‐scale patterns in freshwater fish species richness in North America, north of central Mexico. Total species richness declines to the north and west but the pattern for endemics differs from that of widespread species. Mississippi Basin regions are more species rich than more isolated, coastal, regions. Richness declines more rapidly with increasing latitude in riverine specialist than in habitat generalist species. Levels of endemism are greatest in species found in small‐ to medium‐sized river channels. The strong Rapoport effect, more marked in migratory than resident species, is correlated with habitat preference, channel size, and glacial history. Body size increases with latitude, largely as a result of a trend from small resident to large migrant species. In unglaciated regions, ancestral species survived in large habitats because these are longer‐lived, more extensive, less isolated and more stable than headwaters, permitting larger populations and lower extinction levels. Reduced levels of gene flow in small, peripheral, channels isolated by larger downstream habitats have resulted in the production of many, small range, small‐bodied species. The latitudinal richness gradient is a consequence of speciation and extinction events in unglaciated faunas and an increasing domination of faunas by generalist, large bodied, large channel, recolonizing species in more northern regions. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 46–61.     

Shell size and shape in Madeiran land snails: do niches remain unfilled?

The distribution of shell heights and diameters in the mainly endemic Madeiran land snail fauna shows the bimodal pattern of high- and low-spired shells found in many other faunas. Field and laboratory studies show that shell shape is associated with the angle of substrate on which the snails crawl; as elsewhere, tall spired species use vertical surfaces or burrow in soft material. Flattened species predominate on horizontal surfaces, while globular species are less specific in their preferences. Detailed comparisons with the fauna of N.W. Europe show that the proportion of high-spired species in the Madeiran fauna is low, and large high-spired species associated with vertical surfaces are very few in number despite an apparent abundance of suitable habitats. Amongst low-spired species, one family, the Helicidae, dominates the Madeiran fauna. While the overall distribution of size in these species is much as in Europe, Madeiran helicids extend into smaller size classes than do those in Europe, and they appear to fill a gap in the scatter created by the absence of other families.
Non-endemic species, other than those strictly associated with man-made environments, are generally small in size. In the upper scatter, their size distribution parallels that of endemics, but in the lower scatter they constitute the whole of the smallest size classes.
The role of interspecific competition in determining these distributions is discussed. The range of helicid sizes is compatible with a relaxation of competition or predator pressure relative to other areas, but in the upper scatter there appear to be gaps in the range of size and shape expected despite a long period in which the fauna could evolve. This could indicate the existence of adaptive troughs blocking, or delaying, radiation over the full spectrum of size and shape.     

Does a ruderal strategy dominate the endemic flora of the West African forests?

Aim  To understand the distribution pattern of endemic plant species in West African rain forests, one of the global priority areas for biodiversity conservation.
Location  Upper Guinean forests, West Africa (Senegal, Guinea-Bissau, Guinea, Sierra Leone, Liberia, Côte d'Ivoire, Ghana and Togo).
Methods  We used herbarium collections from the whole Upper Guinean region ( sensu White 1981) to analyse the distribution patterns of 216 vascular plant species (approximately one-third of the endemic flora ranging from herbs to trees) that are restricted to these rain forests. We related species distribution ranges and species commonness to the main environmental variables and species life-history traits.
Results  We found that most endemic species of West African forests have large distribution ranges and ruderal strategies. Among all plant life-forms, species with small ranges were restricted to very moist places whereas more widespread species were less dependent on the availability of water. These more common species seem able to tolerate drier conditions, indicated by the minimum rainfall conditions where they are found and the positive correlation with local habitat openness. Light-demanding endemics were more common and widespread than shade-tolerant species, and there was a trend towards wind-dispersed species having a higher proportion of records in open places. This suggests that how widespread and common an endemic species is depends on being able to tolerate open and drier conditions.
Main conclusions  A ruderal strategy seems to be key for understanding the success of endemics in the Upper Guinean forests, and indicates the important role of disturbances in shaping the composition of these tropical forests. Our results have large implications for predicting the potential effect of increasing drought on rare endemic plant species of West African forests.     

Conservation value of secondary forest habitats for endemic birds,a perspective from two widely separated tropical ecosystems

Tropical secondary forests are increasingly widespread, but their potential for conserving endemic birds remains unclear. Previous studies report different results; however all have been restricted to geographically discreet locations. This is important as different ecosystems are influenced by different external factors, possibly influencing conservation potential. Here we use consistent survey methods to examine how endemic bird richness varies between primary and secondary forest habitats in two widely separated tropical ecosystems, providing a more global context for evaluating the conservation value of secondary forests. Research was completed in Lambusango Forest Reserve (LFR) on Buton Island, Sulawesi, and Cusuco National Park (CNP), a Honduran cloud forest reserve. Bird communities in both forests were surveyed using 50 m radius point counts. Vulnerability assessments based on ecological theory on avifaunal assemblages were then conducted, which suggested endemics in LFR to be more susceptible to disturbance than those in CNP. Contrary to the results from our vulnerability assessments, endemics in CNP were less tolerant of moderate habitat modification than those in LFR. Richness of Mesoamerican endemics per study site declined significantly between core zone forest (6.34 ± 0.81) and more degraded forest in the boundary zone (3.86 ± 0.69). Richness of Wallacean endemics was similar in primary (4.89 ± 1.68) and disturbed secondary forest (4.52 ± 1.62). We recommend considering local and regional biogeographical and ecological factors when determining the conservation value of secondary forests, and suggest examples of potential importance, including differential community richness, influence of figs and human settlement patterns.     

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.     

Survey of the terrestrial and freshwater molluscan fauna of the Pilliga forest area in northern inland New South Wales,Australia

The Pilliga forest in northern inland New South Wales, Australia, is one of the largest surviving remnants of native forest on the western slopes of the Great Dividing Range. The Pilliga landscape is a challenging environment for molluscs, dominated by dry sclerophyll forest and with limited and largely ephemeral aquatic habitats. A field survey of the area in 2006–2012 identified a surprisingly rich and relatively intact aquatic native molluscan fauna with five species of bivalves in three families and nine species of freshwater gastropods (four families), including some rare species and range extensions. The native land snail fauna comprised 18 species (six families), including an unusually rich pupillid fauna with nine species. Some range extensions are recorded and some species are narrow-range endemics. The distributions of many aquatic and terrestrial species were correlated with geology or soil type. Introduced molluscs were predominantly found in anthropogenic habitats and include two freshwater gastropods (two families) and nine terrestrial snails and slugs (eight families). This study provides insight into the original molluscan fauna of the western slopes prior to landscape-scale agricultural development and provides a benchmark for future reference.     

Diversity and endemism of ciliates inhabiting Neotropical phytotelmata

While the diversity and distribution of macro-organisms living in phytotelmata (plant-container habitats) is well known, detailed taxonomic work on micro-organisms living in the same environments is limited. As a model clade of microbial eukaryotes, sampling of ciliates in Neotropical bromeliad tanks increased, and Neotropical phytotelmata such as bamboo stumps and tree holes were newly sampled. Thirty-three isolates from Brazil, Costa Rica, Dominican Republic, Jamaica and Mexico were sequenced for small subunit rDNA, and placed into a phylogenetic context using non-phytotelmata GenBank accessions. This and the morphological investigations discovered 45 undescribed, possibly endemic ciliate species. The potential endemics are from throughout most clades of the ciliate tree of life, and there is evidence of speciation within the Neotropical phytotelmata habitat. Our data show the number of potential Neotropical phytotelmata-endemic ciliate species increasing as more phytotelmata are sampled. While the new data show that the supposed endemics are mainly recruited from moss and ephemeral limnetic habitats, the bromeliad ciliate fauna is quite distinct from those of other limnetic habitats, lacking many typical and common freshwater genera, such as Coleps, Colpidium, Frontonia, Paramecium, Glaucoma, Nassula, Stylonychia and Trithigmostoma. There is no indication that specific ciliates are confined to specific bromeliads.     

Habitat selection and population trends in terrestrial bird species of Robinson Crusoe Island: habitat generalists versus forest specialists

Habitat loss and degradation on oceanic islands are key processes leading to population decline of endemic birds and facilitating the establishment of invasive bird species. In this study, carried out in the Robinson Crusoe Island, we assessed density and habitat selection of terrestrial bird species, including juan fernandez firecrown and juan fernandez tit-tyrant, two endemics, as well as green-backed firecrown and austral thrush, which apparently originate from the mainland. Results show that perturbed habitats contained a low density of the endemic species whereas the mainland species were significantly more abundant in perturbed scrub habitats. Bird species show different habitat selection patterns, with endemics selecting for native forest and mainland species selecting for perturbed habitats, or using them at random. Bird species experienced temporal trends in their overall population sizes, with the endemic tit-tyrant suffering a significant decline in its population size of about 63% between 1994 and 2009. Only mainland species exhibited temporal changes in habitat use, significantly reducing their densities in the preferred scrub habitats, possibly as a response to decreased habitat quality. Thrushes apparently were able to compensate the population decrease in one non native habitat type by using native forests, a habitat giving them the opportunity of preying on nests of endemic species. We conclude that endemic bird species behave as specialists whereas the mainland species must be treated as invasive generalists on Robinson Crusoe Island.     

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.     

Tree invasion in managed tropical forests facilitates endemic species

Aim To determine whether different abundances of introduced species of Cinchona (Rubiaceae) affect species composition and facilitate species richness in managed tropical forests, to test whether any facilitative effects on understorey species depend on forest type, and to investigate whether facilitative effects can be attributed to the ‘substitutive facilitation model’. Location Makawao Forest Reserve on Maui, Hawai’i, USA. Methods Cinchona species (Cinchona pubescens and Cinchona calisaya) were mapped within various forest types. In three forest types (ageing Eucalyptus and Pinus plantations, and near‐natural Acacia koa forests), we analysed environmental parameters (e.g. canopy cover, litter cover, pH value and soil depth) and the species composition of Cinchona‐invaded and non‐invaded plots; data were compared based on Cinchona cover and forest types. Habitat modelling for several endemic species and tree ferns was carried out to test whether Cinchona cover is an important variable for the probability of occurrence of these endemics. Results Cinchona species have naturalized mainly in Eucalyptus and Pinus plantations and Acacia koa forests and here add an additional shrub layer. In contrast to other studies, we revealed facilitative effects of Cinchona on native species within all forest types. Species richness is about 20% higher in invaded plots than in non‐invaded plots, and these show a nearly 50% higher proportion of endemic species, including tree ferns. The proportion of endemics even increases with increasing Cinchona cover. For several endemics, Cinchona is found to be an important variable for the probability of occurrence, and the removal of Cinchona cover as an explanatory variable lowers the model fit. In addition to Cinchona, variables delineating vegetation structure and light availability have a strong effect on the model fit. Main conclusions In the structurally simplified Hawaiian forests studied, Cinchona facilitated endemic species in accordance with the ‘substitutive facilitation model’. This contrasts with the results of an earlier study in the naturally treeless Galápagos highlands, which revealed a sharp decrease in the abundance of endemics under Cinchona canopy. These results illustrate that, through the same structural change (addition of a vegetation layer), an invasive species may exert divergent effects across different ecosystem types. The facilitation of endemic understorey species by invasive tree species in managed forests leads to a dilemma in conservation but also to new perspectives for ecosystem restoration.     

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.     

Terrestrial small mammal diversity and abundance in central Benin: comparison between habitats,with conservation implications

We performed a terrestrial small mammal species inventory in the Agoua and Wari‐Maro forest reserves (Benin). Four localities were sampled, and in each locality, three habitats were surveyed: dense forest, open forest or woodland savannah and shrub savannah. This is the first comprehensive inventory for small mammals in central Benin. We captured 794 small mammals representing twenty species (six shrew species, fourteen rodent species). Three new species that need to be described were recorded. We observed a mixture of both true forest species and of species adapted to a wider range of habitats ranging from savannah to forest clearings. Species with either Sudanian or Guinea–Congolian affinities were recorded, as well as a new species endemic to Togo and Benin. This rich biodiversity underlines the urgent need for an effective protection of these forests. The Sudanian species Crocidura cf. foxi was more abundant in Wari‐Maro than in Agoua forest, while the Guineo–Congolian species Praomys misonnei and Hylomyscus sp were only captured in Agoua forest. These results are in agreement with the fact that these two forests belong to two distinct chorological zones.     

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.     

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.