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.     

Sea‐slug invasion of the land

Invasion of the land has been a considerable challenge for aquatic organismal lineages, primarily because of desiccation and lack of physical support in air. Gastropod molluscs are among the most successful animals in this habitat shift because of their hard exoskeleton, whereas land slugs, regarded as descendants of land snails, gradually lost their shells in sheltered, moist environments. We present solid evidence of terrestrial invasion by a lineage of sea slugs; this invasion is represented by a newly discovered species, Aiteng marefugitus (Panpulmonata: Acochlidia), from a humid tropical rainforest on a small island in the Republic of Palau, western Pacific. The slug lineage seems to have invaded the forest habitat directly from the upper littoral zone in the Cenozoic.     

Morphological parallelism of sympatric cave-dwelling microsnails of the genus Georissa at Mount Silabur,Borneo (Gastropoda,Neritimorpha, Hydrocenidae)

Parallel evolution in phenotype may result when closely related taxa are adapting in the face of similar ecological pressures. Here, we discuss possible parallelism in shell morphology in the context of the microgeographic phylogeography of two conchologically distinct sympatric hydrocenid snails inhabiting a limestone outcrop and its cave system, Georissa pyrrhoderma and Georissa silaburensis, respectively, at Mount Silabur in Sarawak, Malaysian Borneo. Our results show a certain degree of morphological parallelism of a third, possibly new, cryptic Georissa species within the same cave that strongly diverged from its above-ground sister species, G. pyrrhoderma. We found that both sympatric cave species have shifted from a more sculptured, conical shell toward a broader, less sculptured form.     

Divergence in the shell morphology of the land snail genus Aegista (Pulmonata: Bradybaenidae) under phylogenetic constraints

The role of natural selection in phenotypic evolution is central to evolutionary biology. Phenotypic evolution is affected by various factors other than adaptation, and recent focus has been placed on the effects of phylogenetic constraints and niche conservatism on phenotypic evolution. Here, we investigate the relationship between the shell morphology and habitat use of bradybaenid land snails of the genus Aegista and clarify the causes of the divergence in shell morphology among phylogenetically related species. The results of ancestral state reconstruction showed that arboreal species have evolved independently from ground‐dwelling species at least four times. A significant association was found between shell shape and habitat use, despite the existence of a certain degree of phylogenetic constraint between these traits. A principal component analysis showed that arboreal species tend to have a relatively high‐spired shell with a narrow umbilicus. By contrast, ground‐dwelling species have a low‐spired shell with a wide umbilicus. Although the latitude and elevation of the sampling locations showed no relationship with shell morphology, the geology of the sampling locations affected the shell size of arboreal species. The development of a well‐balanced shell shape is one effective method for reducing the cost of locomotion under the force of gravity in each life habitat, resulting in the divergence in shell morphology and the independent evolution of morphologically similar species among different lineages. The present study suggests that ecological divergence is probably the cause of shell morphology divergence in land snails. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 229–241.     

Patterns of ecomorphological convergence among mainland and island Anolis lizards

Differing selective pressures on islands versus the mainland may produce alternative evolutionary outcomes among closely related lineages. Conversely, lineages may be constrained to produce similar outcomes in different mainland and island environments, or mainland and island environments may not differ significantly. Among the best‐studied island radiations are Caribbean Anolis lizards. Distinct morphotypes, or ‘ecomorphs’, have been described, and the same ecomorphs have evolved independently on each Greater Antillean island. The mainland Anolis radiation has received much less attention. We use a large morphological data set and a novel phylogenetic hypothesis to show that mainland Anolis did not evolve the same morphotypes as island Anolis, despite some island species being more closely related to mainland species than to island species that share their morphotype. A maximum of four of the six Caribbean ecomorphs were found to exist on the mainland, and just 15 of 123 mainland species are assignable to a Caribbean ecomorph. This result was insensitive to differing taxon samples and alternative phylogenetic hypotheses. Mainland convergence to a Caribbean ecomorph occurs only among species assigned to the grass‐bush ecomorph. Thus, the ecomorphs that have evolved convergently multiple times in the Caribbean have not evolved in parallel on the mainland. These results are consistent with the hypothesis that mainland and island environments offer different selective pressures. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 852–859.     

The founding of Mauritian endemic coffee trees by a synchronous long‐distance dispersal event

The stochastic process of long‐distance dispersal is the exclusive means by which plants colonize oceanic islands. Baker's rule posits that self‐incompatible plant lineages are unlikely to successfully colonize oceanic islands because they must achieve a coordinated long‐distance dispersal of sufficiently numerous individuals to establish an outcrossing founder population. Here, we show for the first time that Mauritian Coffea species are self‐incompatible and thus represent an exception to Baker's rule. The genus Coffea (Rubiaceae) is composed of approximately 124 species with a paleotropical distribution. Phylogenetic evidence strongly supports a single colonization of the oceanic island of Mauritius from either Madagascar or Africa. We employ Bayesian divergence time analyses to show that the colonization of Mauritius was not a recent event. We genotype S‐RNase alleles from Mauritian endemic Coffea, and using S‐allele gene genealogies, we show that the Mauritian allelic diversity is confined to just seven deeply divergent Coffea S‐RNase allelic lineages. Based on these data, we developed an individual‐based model and performed a simulation study to estimate the most likely number of founding individuals involved in the colonization of Mauritius. Our simulations show that to explain the observed S‐RNase allelic diversity, the founding population was likely composed of fewer than 31 seeds that were likely synchronously dispersed from an ancestral mainland species.     

Small‐scale genetic structuring in a tropical cave snail and admixture with its above‐ground sister species†

We analyse the phylogeographical structure in the cave snail Georissa filiasaulae Haase & Schilthuizen, 2007 (Gastropoda: Hydrocenidae) and its above‐ground sister species G. saulae (van Benthem‐Jutting, 1966) at limestone outcrops in Sabah, Malaysian Borneo. Morphometric and 16S mitochondrial DNA data for some 220 individuals reveal strong morphological differentiation, despite ongoing unidirectional gene flow from the epigean into the hypogean environment, strong, small‐scale genetic structuring within the cave and underground dispersal between caves that were previously thought to be isolated. We discuss these results – which constitute the first phylogeographical analysis of a terrestrial cave snail – in the light of speciation in cave organisms and across ecotones in general. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 727–740.     

Comparative phylogeography of mainland and insular species of Neotropical molossid bats (Molossus)

Historical events, habitat preferences, and geographic barriers might result in distinct genetic patterns in insular versus mainland populations. Comparison between these two biogeographic systems provides an opportunity to investigate the relative role of isolation in phylogeographic patterns and to elucidate the importance of evolution and demographic history in population structure. Herein, we use a genotype‐by‐sequencing approach (GBS) to explore population structure within three species of mastiff bats (Molossus molossus, M. coibensis, and M. milleri), which represent different ecological histories and geographical distributions in the genus. We tested the hypotheses that oceanic straits serve as barriers to dispersal in Caribbean bats and that isolated island populations are more likely to experience genetic drift and bottlenecks in comparison with highly connected ones, thus leading to different phylogeographic patterns. We show that population structures vary according to general habitat preferences, levels of population isolation, and historical fluctuations in climate. In our dataset, mainland geographic barriers played only a small role in isolation of lineages. However, oceanic straits posed a partial barrier to the dispersal for some populations within some species (M. milleri), but do not seem to disrupt gene flow in others (M. molossus). Lineages on distant islands undergo genetic bottlenecks more frequently than island lineages closer to the mainland, which have a greater exchange of haplotypes.     

Phylogeny and historical biogeography of the cave-adapted shrimp genus Typhlatya (Atyidae) in the Caribbean Sea and western Atlantic

Aim To infer phylogenetic relationships among five species of the cave‐adapted shrimp genus Typhlatya in order to test competing hypotheses of dispersal and colonization of the disjunct cave localities occupied by these five species. Location Typhlatya species are found in caves and anchialine ponds across the northern margin of the Caribbean Sea, along the Mediterranean and Adriatic coasts and on oceanic islands in the Atlantic and eastern Pacific oceans. This study focuses on five species, one from Bermuda, one from the Caicos Islands and three from the Yucatan Peninsula of Mexico. Methods Partial sequences (c. 1400 bp) from the mitochondrial cytochrome b, 16S rDNA and COI genes were obtained from representative samples of the five species. Phylogenetic inference was carried out with maximum parsimony and maximum likelihood analyses. Parsimony networks were constructed for the Bermudian species Typhlatya iliffei and one Yucatan species Typhlatya mitchelli, to determine the degree of connectivity among populations inhabiting different cave systems. Results All three land masses were recovered as monophyletic. The two insular marine species from Bermuda and the Caicos Islands formed a clade, while the three continental freshwater species from the Yucatan Peninsula formed another. Within both Bermuda and the Yucatan, shared haplotypes were found in different cave systems, suggesting recent or ongoing gene flow among populations in both locales. Main conclusions The two insular marine Typhlatya species originated from an ancestral marine population, possibly already cave‐adapted, that is suggested to have colonized the Caicos Islands and subsequently dispersed to Bermuda via the Gulf Stream. Divergence estimates suggest that colonization occurred before the formation of present‐day anchialine cave habitat, which did not form on either island until the late Pliocene to early Pleistocene. Divergence estimates also indicate that the Yucatan freshwater species split before the formation of freshwater cave habitat in the Yucatan. These species could have inhabited crevicular marine habitats before the late Pliocene/early Pleistocene in the Yucatan or elsewhere in the Caribbean, and subsequently migrated to freshwater caves once they formed.     

Winter bird communities in the Åland archipelago: an island biogeographic point of view

The distribution of wintering land birds was examined in the archipelago of Åland (60°N, 20°E) along two ecological dimensions: (1) the island continuum from small skerries ((0.5 ha) to the mainland of Åland (97000 ha), and (2) the habitat range of the mainland of Åland. In the species-area relationship, both the exponential and power function models fitted well. The increase of the number of species with island size was interpreted as a consequence of increasing habitat diversity on larger islands. The smallest islands supported only few seed-eating species utilizing the islands in a fine-grained fashion; none of these birds belonged to the breeding fauna of the islands. With increasing island size and habitat diversity, insectivores and species tied to the human culture were added. On the mainland of Åland, more than half of the birds were observed near human habitations: bird communities wintering in pine forests were richer than in deciduous forests. Proportion of wintering species was greatest in the breeding communities of pine forests. The proportion of wintering species was presumably regulated by the degree of structural change in the habitat between summer and winter, this change being most drastic in deciduous habitats The species observed could be broadly classified into three feeding categories: opportunists ., omnivores and scavengers, dependent on the diverse food items provided by man; food specialists , species éating seeds and berries, ranging widely in the islands and habitats; and insectivorous habitat specialists , inhabiting coniferous habitats, and islands with coniferous (pine) forests. Opportunists, in particular, gel increasing wintering opportunities by the impact of man.     

Enhanced colour polymorphisms in island populations of the land snail Euhadra peliomphala

Variations in visible genetic polymorphisms are assumed to decrease in populations on small islands because of intense founder effects, genetic drift and inbreeding. However, we have found evidence of a marked enhancement of colour polymorphisms within populations on small oceanic islands that were colonized from the mainland. The source populations on the mainland of the land snail Euhadra peliomphala in four oceanic islands were estimated by phylogenetic analysis of mitochondrial DNA sequences. Diversity of shell colour was higher in the island populations than in the source populations on the mainland. In addition, the shell colour morphs differed not only among populations from different islands but also between the island populations and the source populations on the mainland. By contrast, no mtDNA variations were found in any of the island populations, even though the source populations possessed high mtDNA diversity. Thus, components of colour morphs changed in the island populations after their colonization, and colour polymorphisms are enhanced in these islands despite the loss of genetic variation. The above findings suggest that ecological mechanisms such as morphological release owing to a release from competition may overcome the tendency toward reduced genetic variation in islands to enhance the colour polymorphism.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 417–425.     

Even the Smallest Non-Crop Habitat Islands Could Be Beneficial: Distribution of Carabid Beetles and Spiders in Agricultural Landscape

Carabid beetles and ground-dwelling spiders inhabiting agroecosystems are beneficial organisms with a potential to control pest species. Intensification of agricultural management and reduction of areas covered by non-crop vegetation during recent decades in some areas has led to many potentially serious environmental problems including a decline in the diversity and abundance of beneficial arthropods in agricultural landscapes. This study investigated carabid beetle and spider assemblages in non-crop habitat islands of various sizes (50 to 18,000 square metres) within one large field, as well as the arable land within the field, using pitfall traps in two consecutive sampling periods (spring to early summer and peak summer). The non-crop habitat islands situated inside arable land hosted many unique ground-dwelling arthropod species that were not present within the surrounding arable land. Even the smallest non-crop habitat islands with areas of tens of square metres were inhabited by assemblages substantially different from these inhabiting arable land and thus enhanced the biodiversity of agricultural landscapes. The non-crop habitat area substantially affected the activity density, recorded species richness and recorded species composition of carabid and ground-dwelling spider assemblages; however, the effects were weakened when species specialised to non-crop habitats species were analysed separately. Interestingly, recorded species richness of spiders increased with non-crop habitat area, whereas recorded species richness of carabid beetles exhibited an opposite trend. There was substantial temporal variation in the spatial distribution of ground-dwelling arthropods, and contrasting patterns were observed for particular taxa (carabid beetles and spiders). In general, local environmental conditions (i.e., non-crop habitat island tree cover, shrub cover, grass cover and litter depth) were better determinants of arthropod assemblages than non-crop habitat island size, indicating that the creation of quite small but diversified (e.g., differing in vegetation cover) non-crop habitat islands could be the most efficient tool for the maintenance and enhancement of diversity of ground-dwelling carabids and spiders in agricultural landscapes.     

Evolution of cave living in Hawaiian Schrankia (Lepidoptera: Noctuidae) with description of a remarkable new cave species

Although temperate cave‐adapted fauna may evolve as a result of climatic change, tropical cave dwellers probably colonize caves through adaptive shifts to exploit new resources. The founding populations may have traits that make colonization of underground spaces even more likely. To investigate the process of cave adaptation and the number of times that flightlessness has evolved in a group of reportedly flightless Hawaiian cave moths, we tested the flight ability of 54 Schrankia individuals from seven caves on two islands. Several caves on one island were sampled because separate caves could have been colonized by underground connections after flightlessness had already evolved. A phylogeny based on approximately 1500 bp of mtDNA and nDNA showed that Schrankia howarthi sp. nov. invaded caves on two islands, Maui and Hawaii. Cave‐adapted adults are not consistently flightless but instead are polymorphic for flight ability. Although the new species appears well suited to underground living, some individuals were found living above ground as well. These individuals, which are capable of flight, suggest that this normally cave‐limited species is able to colonize other, geographically separated caves via above‐ground dispersal. This is the first example of an apparently cave‐adapted species that occurs in caves on two separate Hawaiian islands. A revision of the other Hawaiian Schrankia is presented, revealing that Schrankia simplex, Schrankia oxygramma, Schrankia sarothrura, and Schrankia arrhecta are all junior synonyms of Schrankia altivolans. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 114–139.     

Habitat constraints limit the distribution and population density of redwing francolin, Francolinus levaillantii, in the highland grasslands of Mpumalanga province, South Africa

We investigated a variety of habitat characteristics along a land‐use gradient in an attempt to determine which factors may have influenced the population decline of redwing francolin, Francolinus levaillantii, in commercial pasture farms in the highland grasslands of Mpumalanga province, South Africa. Within three microhabitats (wetlands, rocky outcrops and grassland), number and diversity of food plants, height of the sward and percentage cover were significantly lower in habitat exposed to commercial grazing with sheep. Increased cover and height of the sward were positively correlated with higher diversity and availability of francolin food plants. Crop analyses revealed a lower intake in the number of food plant species, but higher incidence of invertebrates, ingested by birds collected in heavily grazed and frequently burned grasslands. Intestinal caecae were significantly (P < 0.001) longer in birds collected from grazed/burned grasslands – possibly associated with a low quality, more fibrous diet. Both the quantity and quality of habitat in terms of food abundance, diversity and cover are important factors influencing the distribution of redwing francolin in these grasslands.     

Seed plants of Fiji: an ecological analysis

An annotated list of indigenous Fijian seed plant genera is presented and comprises 484 genera and 1315 species in 137 families. The relative diversity of the largest families and genera in Fiji is indicated and compared with floras in New Caledonia and the Upper Watut Valley, Papua New Guinea. Differences and similarities appear to be due to biogeographical/phylogenetic factors rather than ecological differences or means of dispersal. Generic diversity for the seed plants as a whole is greatest between 0–100 m and decreases monotonically with altitude. However, in the largest family, Orchidaceae, maximum diversity occurs between 200–400 m. Fifty percent of the families are recorded from shore habitat. Twenty‐seven percent of the families and 80 species occur in or around mangrove, where the most diverse families are Orchidaceae, Rubiaceae, and the legumes. Some of the mangrove‐associate species are pantropical or Indo‐Pacific but most are locally or regionally endemic. Fifty‐six percent of the Fijian families are recorded on limestone. Twenty‐nine species are restricted to limestone and 12 species usually occur on limestone. The importance of calcium in reducing the effects of salinity is emphasized and 39 species are recorded from both mangrove and limestone. A plagiotropic habit occurs in 38 species which occur on limestone or around beaches, and 20 of these are Pacific endemics. Genera restricted to higher altitudes include many present elsewhere in Melanesia but absent from Australia despite suitable habitat there, again indicating the importance of biogeographical and historical factors. Altitudinal anomalies in Fiji taxa are cited and include 7 anomalously high records from northern Viti Levu, a site of major uplift, and 22 anomalously low altitudinal records in the Lau Group, a site of subsidence. It is suggested that the Fijian flora has not been derived from immigrants from Asia, but has evolved more or less in situ. Taxa would have survived as metapopulations on the individually ephemeral volcanic islands always found at oceanic subduction zones and hot spots, and the atolls which characterize areas of subsidence. The complex geology of Fiji is determined by its position between two subduction zones of opposite polarity, the Vanuatu and Tonga Trenches, in what is currently a region of transform faulting. The large islands comprise fragments of island arcs that have amalgamated and welded together. There has been considerable uplift as well as subsidence in the islands and it is suggested that both these processes have had drastic effects on the altitudinal range of the taxa. Limestone and mangrove floras could have provided a widespread, diverse ancestral species pool from which freshwater swamp forest, lowland rainforest, dry forest, secondary forest, thickets, and montane forest have been derived during phases of uplift. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 89 , 407–431.     

Are island plant communities more invaded than their mainland counterparts?

Questions: Are island vegetation communities more invaded than their mainland counterparts? Is this pattern consistent among community types? Location: The coastal provinces of Catalonia and the para‐oceanic Balearic Islands, both in NE Spain. These islands were connected to the continent more than 5.35 million years ago and are now located <200 km from the coast. Methods: We compiled a database of almost 3000 phytosociological relevés from the Balearic Islands and Catalonia and compared the level of invasion by alien plants in island versus mainland communities. Twenty distinct plant community types were compared between island and mainland counterparts. Results: The percentage of plots with alien species, number, percentage and cover percentage of alien species per plot was greater in Catalonia than in the Balearic Islands in most communities. Overall, across communities, more alien species were found in the mainland (53) compared to the islands (only nine). Despite these differences, patterns of the level of invasion in communities were highly consistent between the islands and mainland. The most invaded communities were ruderal and riparian. Main conclusion: Our results indicate that para‐oceanic island communities such as the Balearic Islands are less invaded than their mainland counterparts. This difference reflects a smaller regional alien species pool in the Balearic Islands than in the adjacent mainland, probably due to differences in landscape heterogeneity and propagule pressure.     

Managing rock outcrops to improve biodiversity conservation in Australian agricultural landscapes

Summary  Rocky outcrops are prominent geological features in agricultural landscapes worldwide. Reptiles are a major component of these habitats and some species are restricted to, and more abundant on, rocky outcrops than in remnant vegetation. Rock outcrops are important to reptiles because they provide resources that are often limited in the surrounding landscape (e.g. micro-gradients in climatic conditions, basking- and retreat-sites). However, there is a knowledge gap in the literature addressing the conservation value of small, rocky outcrops. Management may be necessary to reverse habitat degradation in these systems. We identify four key areas of management that need to be addressed to improve outcrop habitat values and enhance biodiversity conservation in agricultural landscapes. Specific actions involve: (i) protecting outcrops from processes that cause damage to rock microhabitat, (ii) monitoring and managing changes in vegetation structure to maintain thermally suitable environments, (iii) applying integrated pest animal control and (iv) improving matrix management to enhance inselberg function and landscape connectivity. Further research is required to evaluate the efficacy of different management regimes on outcrop biota. We hope this paper will provide the stimulus for land managers to incorporate rocky outcrops in future biodiversity conservation programmes.     

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.     

Effects of habitat history and extinction selectivity on species-richness patterns of an island land snail fauna

Aim Local‐scale diversity patterns are not necessarily regulated by contemporary processes, but may be the result of historical events such as habitat changes and selective extinctions that occurred in the past. We test this hypothesis by examining species‐richness patterns of the land snail fauna on an oceanic island where forest was once destroyed but subsequently recovered. Location Hahajima Island of the Ogasawara Islands in the western Pacific. Methods Species richness of land snails was examined in 217 0.25 × 0.25 km squares during 1990–91 and 2005–07. Associations of species richness with elevation, current habitat quality (proportion of habitat composed of indigenous trees and uncultivated areas), number of alien snail species, and proportion of forest loss before 1945 in each area were examined using a randomization test and simultaneous autoregressive (SAR) models. Extinctions in each area and on the entire island were detected by comparing 2005–07 records with 1990–91 records and previously published records from surveys in 1987–91 and 1901–07. The association of species extinction with snail ecotype and the above environmental factors was examined using a spatial generalized linear mixed model (GLMM). Results The level of habitat loss before 1945 explained the greatest proportion of variation in the geographical patterns of species richness. Current species richness was positively correlated with elevation in the arboreal species, whereas it was negatively correlated with elevation in the ground‐dwelling species. However, no or a positive correlation was found between elevation and richness of the ground‐dwelling species in 1987–91. The change of the association with elevation in the ground‐dwelling species was caused by greater recent extinction at higher elevation, possibly as a result of predation by malacophagous flatworms. In contrast, very minor extinction levels have occurred in arboreal species since 1987–91, and their original patterns have remained unaltered, mainly because flatworms do not climb trees. Main conclusions The species‐richness patterns of the land snails on Hahajima Island are mosaics shaped by extinction resulting from habitat loss more than 60 years ago, recent selective extinction, and original faunal patterns. The effects of habitat destruction have remained long after habitat recovery. Different factors have operated during different periods and at different time‐scales. These findings suggest that historical processes should be taken into account when considering local‐scale diversity patterns.