Ecological and morphological patterns in communities of land snails of the genus Mandarina from the Bonin Islands

The land snail genus Mandarina has undergone extensive radiation within the Bonin Islands in the west Pacific. The preferred above-ground vegetation heights of sympatric species were clearly different. They separated into arboreal, semi-arboreal, exposed ground and sheltered ground ecotypes. Shells of species with different ecotypes differ markedly, but shells of species with the same ecotype are very similar to each other. Shell morphologies of some phylogenetically distantly related species with the same ecotype were indistinguishable. Character evolution estimated parsimoniously using a phylogenetic tree suggests that the speciation among sympatric species is accompanied by ecological and morphological diversification. In addition, species coexistence of Mandarina is related to niche differentiation. The above findings suggest that ecological interactions among species contribute to the ecological and morphological diversification and radiation of these land snails in this depauperate environment.     

Ecological diversity and speciation in land snails of the genus Mandarina from the Bonin Islands

 Endemic land snails of the genus Mandarina of the oceanic Bonin Islands offer an example of habitat and character divergence among closely related species. The molecular phylogenies of Mandarina show that a divergence of arboreal, semiarboreal, and ground-dwelling species has occurred repeatedly in different times, areas, and lineages. Ecological diversification is suggested to be important for the coexistence of Mandarina species based on the facts that sympatric species are typically highly differentiated ecologically and morphologically, and that species occupying similar habitats do not coexist. The ecological diversification of Mandarina has occurred without much genetic divergence compared with that of its mainland relatives. This suggests that morphological and ecological diversifications are accelerated in depauperate environments where there are fewer competitors and predators. Although the details of the reproductive isolation mechanisms are not understood and further examination is needed, the rapid evolution of prezygotic isolation is the main cause of speciation in Mandarina. In particular, ecological diversification may be an effective barrier to gene exchange between two species. Because of incomplete postmating isolation and the lower genetic divergence among species of Mandarina, breakdowns of reproductive isolation have frequently occurred as a result of habitat change. It is important to estimate the effect of hybridization on species diversification in future studies. Received: February 7, 2002 / Accepted: October 22, 2002 Acknowledgments I express my sincere thanks to A. Davison, B.C. Clarke, A. Guiller, D. Thomaz, K. Tomiyama, I. Hayami, and K. Tanabe for helpful advice and assistance. This study was supported by grants from the Japan Society for the Promotion of Science and the Nippon Life Insurance Foundation.     

Ecological character displacement caused by reproductive interference

We carried out a theoretical investigation of whether ecological character displacement can be caused by reproductive interference. Our model assumes that a quantitative character is associated with both resource use and species recognition, and that heterospecific mating incurs costs. The model shows that ecological character displacement can occur as a consequence of evolution of premating isolation; this conclusion is based on the premise that resource competition is less intense between species than within species and that the ecological character also contributes to premating isolation. When resource competition between species is intense, extinction of either species may occur by competitive exclusion before ecological character divergence. Some observational studies have shown that character displacement in body size is associated with not only resources use but also species recognition. We propose that body size displacement can occur as a consequence of evolution of premating isolation. Our results suggest that ecological character displacement results from reproductive character displacement.     

Ecological and community-wide character displacement: the next generation

Ecological character displacement, mostly seen as increased differences of size in sympatry between closely‐related or similar species, is a focal hypothesis assuming that species too similar to one another could not coexist without diverging, owing to interspecific competition. Thus, ecological character displacement and community‐wide character displacement (overdispersion in size of potential competitors within ecological guilds) were at the heart of the debate regarding the role of competition in structuring ecological communities. The debate has focused on the evidence presented in earlier studies and generated a new generation of rigorous, critical studies of communities. Character displacement research in the past two decades provides sound statistical support for the hypothesis in a wide variety of taxa, albeit with a phylogenetically skewed representation. A growing number of studies are strongly based in functional morphology, and some also demonstrate actual morphologically related resource partitioning. Phylogenetic models and experimental work have added to the scope and depth of earlier research, as have theoretical studies. However, many challenging ecological and evolutionary issues, regarding both selective forces (at the inter‐ and intraspecific level) and resultant patterns, remain to be addressed. Ecological character displacement and community‐wide character displacement are here to stay as the focus of much exciting research.     

Simultaneous character convergence and divergence in Western Australian land snails

Three distantly related genera of land snails, Amplirhagada Iredale, 1933, Quistrachia Iredale, 1939, and Westraltrachia Iredale, 1933, overlap in distributions only in the Napier and Oscar Ranges, east of Derby, Western Australia. The first two remain allopatric, Quistrachia having a more eastern range, while Westraltrachia overlaps both ranges. In the zone of sympatry, with changes proceeding from east to west, Westraltrachia diverges in feeding, and there is massive, gradual convergence in shell size, shape and colour between Amplirhagada and Westraltrachia. Shells of both genera depart markedly from their distinctive allopatric appearances to form a new morphotype in the north-west Napier Ranges. The species there are so similar in appearance that the two genera cannot be identified at a distance of 0.5 m. Thirteen species-level taxa are involved in these changes, four allopatric Amplirhagada , seven allopatric Westraltrachia, Quistrachia monogramma and Rhagada basedowana.
Westraltrachia species, the only large land snails in their main area of distribution (south-east Kimberlcy), are pre-adapted to graze on an occasionally present food resource, algal-fungal blooms on limestone seepage faces. They entered the Oscar and Napier Ranges, where this food resource is more abundant, and large land snails that are generalized feeders, Amplirhagada or Quistrachia , were already present and abundant. Hence feeding divergence occurred. Such specialization by Westraltrachia to minimize competition is readily understandable.
The selective pressures that led to the striking convergence in shell features in north-west Napier taxa are unknown.     

Characterization of 17 microsatellite loci in the Japanese land snail genera Mandarina,Ainohelix, and Euhadra (Mollusca,Gastropoda, Pulmonata)

Land snails of the genera Mandarina, Euhadra and Ainohelix are useful for understanding the ecology and evolution of speciation and adaptation, so we have developed 17 microsatellite loci for these species. As in other land snails, most of the loci are highly polymorphic compound repeats, with a great size range between alleles. The loci should be useful in understanding gene‐flow, genetic structure and speciation in these species.     

Character displacement, frequency-dependent selection, and divergence of shell colour in land snails Mandarina (Pulmonata)

Endemic land snails of the genus Mandarina of the oceanic Bonin Islands offer an exceptional example of habitat and character divergence among closely related species. In this study, microhabitat differences between sympatric ground-dwelling species were studied by distinguishing habitats on the basis of vegetation and types of litter. In all sites where two ground species coexisted, segregation occurred with each species showing preference for the microhabitat in which they were found. When they were in sympatry, one species was predominant in relatively wet and sheltered sites and the other in relatively dry and exposed sites. Although most species can live in both types of habitat, occupation by one species is inhibited by occupation by another. This suggests that competitive interaction between sympatric species caused segregation. Except for populations that have undergone interspecific hybridization, no examples were found of sympatric populations of two ground species sharing a similar shell colour. Species that were predominant in relatively wet and sheltered sites possessed shells with dark coloration and their colour patterns were mostly of one type. Species that were predominant in relatively dry and exposed sites possessed shells with bright coloration and their color patterns were polymorphic. Most populations from areas in which single species were distributed had shells with medium coloration. Microhabitat differentiation between sympatric species possibly caused diversification of shell colour, because bright shells are advantageous in sites where snails are largely exposed, and dark shells are advantageous in sites in where they are mostly sheltered from sunlight. In addition, frequency-dependent selection by predators hunting by sight may have operated to maintain colour polymorphism in the populations which are restricted to exposed habitats by competition with other sympatric species. This reveals the importance of interaction among closely related species as a cause of diversification in ecological and morphological traits.     

Ecological and morphological determinants of evolutionary diversification in Darwin's finches and their relatives

Darwin''s finches are a classic example of adaptive radiation, a process by which multiple ecologically distinct species rapidly evolve from a single ancestor. Such evolutionary diversification is typically explained by adaptation to new ecological opportunities. However, the ecological diversification of Darwin''s finches following their dispersal to Galápagos was not matched on the same archipelago by other lineages of colonizing land birds, which diversified very little in terms of both species number and morphology. To better understand the causes underlying the extraordinary variation in Darwin''s finches, we analyze the evolutionary dynamics of speciation and trait diversification in Thraupidae, including Coerebinae (Darwin''s finches and relatives) and, their closely related clade, Sporophilinae. For all traits, we observe an early pulse of speciation and morphological diversification followed by prolonged periods of slower steady‐state rates of change. The primary exception is the apparent recent increase in diversification rate in Darwin''s finches coupled with highly variable beak morphology, a potential key factor explaining this adaptive radiation. Our observations illustrate how the exploitation of ecological opportunity by contrasting means can produce clades with similarly high diversification rate yet strikingly different degrees of ecological and morphological differentiation.     

Ecological opportunity and the adaptive diversification of lineages

The tenet that ecological opportunity drives adaptive diversification has been central to theories of speciation since Darwin, yet no widely accepted definition or mechanistic framework for the concept currently exists. We propose a definition for ecological opportunity that provides an explicit mechanism for its action. In our formulation, ecological opportunity refers to environmental conditions that both permit the persistence of a lineage within a community, as well as generate divergent natural selection within that lineage. Thus, ecological opportunity arises from two fundamental elements: (1) niche availability, the ability of a population with a phenotype previously absent from a community to persist within that community and (2) niche discordance, the diversifying selection generated by the adaptive mismatch between a population's niche‐related traits and the newly encountered ecological conditions. Evolutionary response to ecological opportunity is primarily governed by (1) spatiotemporal structure of ecological opportunity, which influences dynamics of selection and development of reproductive isolation and (2) diversification potential, the biological properties of a lineage that determine its capacity to diversify. Diversification under ecological opportunity proceeds as an increase in niche breadth, development of intraspecific ecotypes, speciation, and additional cycles of diversification that may themselves be triggered by speciation. Extensive ecological opportunity may exist in depauperate communities, but it is unclear whether ecological opportunity abates in species‐rich communities. Because ecological opportunity should generally increase during times of rapid and multifarious environmental change, human activities may currently be generating elevated ecological opportunity – but so far little work has directly addressed this topic. Our framework highlights the need for greater synthesis of community ecology and evolutionary biology, unifying the four major components of the concept of ecological opportunity.     

Shell shape and habitat use in the North‐west Pacific land snail Mandarina polita from Hahajima,Ogasawara: current adaptation or ghost of species past?

The endemic land snail genus Mandarina of the Ogasawara Islands provides an excellent model system to investigate adaptive radiation. Previously, it has been shown that coexisting species of the islands segregate by microhabitat, so that they are either predominantly found on the ground in relatively wet and sheltered sites, dry and exposed sites, or else are arboreal. Moreover, shell morphology correlates with microhabitat, so that species in wet and sheltered sites tend to have high-spired shells with a high aperture, and those in dry and exposed sites tend to have relatively low-spired shells with a wide aperture. We have now found that on Hahajima, Mandarina polita have variable shell morphology, and there is a correlation between morphology and the depth of leaf litter, as well as the presence/absence of other terrestrial species. Specifically, when high-spired terrestrial Mandarina ponderosa is present, M. polita tend to be low-spired and have a large aperture, indicative of character displacement. When M. ponderosa is absent, the shell shape of M. polita is much more variable, the overall spire is higher, individuals are found in deeper litter, and there is a strong correlation between litter depth and spire height. We argue that these patterns are due to local adaptation, but it remains possible that they are an artefact due to the 'ghost of species past'.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 149–159.     

Labile ecotypes accompany rapid cladogenesis in an adaptive radiation of Mandarina (Bradybaenidae) land snails

The endemic land snail genus Mandarina of the oceanic Bonin Islands shows exceptionally diverse morphological and ecological traits. Previous studies have already provided evidence that speciation on different islands of the three main archipelagos was such that similar ecotypes evolved independently in different lineages and islands. Here we present data to show that the same species can have different ecotypes. As most of the characters involved are inherited, then variation between ecotypes must represent genetic differences between populations. We then show that the radiation on the Bonin Islands is derived from a single colonization event, and use a mitochondrial phylogeny to provide evidence for a burst of cladogenesis soon after colonization. As divergent selection has previously been implicated in causing differences between Mandarina species, and theory predicts that most of the speciation should have taken place early in their history, then the study adds to the evidence for an adaptive radiation by ecological speciation in Mandarina . However, while the diversity of ecotypes present at each site is dependent on the regime of natural selection and competition, geography still must have an important role.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 269–282.     

Origin and diversification of Hibiscus glaber, species endemic to the oceanic Bonin Islands, revealed by chloroplast DNA polymorphism

Abstract Two woody Hibiscus species co-occur in the Bonin Islands of the northwestern Pacific Ocean: Hibiscus glaber Matsum. is endemic to the islands, and its putative ancestral species, Hibiscus tiliaceus L., is widely distributed in coastal areas of the tropics and subtropics. To infer isolating mechanisms that led to speciation of H. glaber and the processes that resulted in co-occurrence of the two closely related species on the Bonin Islands, we conducted molecular phylogenetic analyses on chloroplast DNA (cpDNA) sequences. Materials collected from a wide area of the Pacific and Indian Oceans were used, and two closely related species, Hibiscus hamabo Siebold Zucc. and Hibiscus macrophyllus Roxb., were also included in the analyses. The constructed tree suggested that H. glaber has been derived from H. tiliaceus, and that most of the modern Bonin populations of H. tiliaceus did not share most recent ancestry with H. glaber. Geographic isolation appears to be the most important mechanism in the speciation of H. glaber. The co-occurrence of the two species can be attributed to multiple migrations of different lineages into the islands. While a wide and overlapping geographical distribution of haplotypes was found in H. tiliaceus, localized geographical distribution of haplotypes was detected in H. glaber. It is hypothesized that a shift to inland habitats may have affected the mode of seed dispersal from ocean currents to gravity and hence resulted in geographical structuring of H. glaber haplotypes.     

Molecular biogeography and diversification of the endemic terrestrial fauna of the Hawaiian Islands

Oceanic islands have played a central role in biogeography and evolutionary biology. Here, we review molecular studies of the endemic terrestrial fauna of the Hawaiian archipelago. For some groups, monophyly and presumed single origin of the Hawaiian radiations have been confirmed (achatinelline tree snails, drepanidine honeycreepers, drosophilid flies, Havaika spiders, Hylaeus bees, Laupala crickets). Other radiations are derived from multiple colonizations (Tetragnatha and Theridion spiders, succineid snails, possibly Dicranomyia crane flies, Porzana rails). The geographic origins of many invertebrate groups remain obscure, largely because of inadequate sampling of possible source regions. Those of vertebrates are better known, probably because few lineages have radiated, diversity is far lower and morphological taxonomy permits identification of probable source regions. Most birds, and the bat, have New World origins. Within the archipelago, most radiations follow, to some degree, a progression rule pattern, speciating as they colonize newer from older islands sequentially, although speciation often also occurs within islands. Most invertebrates are single-island endemics. However, among multi-island species studied, complex patterns of diversification are exhibited, reflecting heightened dispersal potential (succineids, Dicranomyia). Instances of Hawaiian taxa colonizing other regions are being discovered (Scaptomyza flies, succineids). Taxonomy has also been elucidated by molecular studies (Achatinella snails, drosophilids). While molecular studies on Hawaiian fauna have burgeoned since the mid-1990s, much remains unknown. Yet the Hawaiian fauna is in peril: more than 70 per cent of the birds and possibly 90 per cent of the snails are extinct. Conservation is imperative if this unique fauna is to continue shedding light on profound evolutionary and biogeographic questions.     

Phylogenetic analysis of ecological and morphological diversification in Hispaniolan trunk-ground anoles (Anolis cybotes group)

Abstract Anolis lizards in the Greater Antilles partition the structural microhabitats available at a given site into four to six distinct categories. Most microhabitat specialists, or ecomorphs, have evolved only once on each island, yet closely related species of the same ecomorph occur in different geographic macrohabitats across the island. The extent to which closely related species of the same ecomorph have diverged to adapt to different geographic macro-habitats is largely undocumented. On the island of Hispaniola, members of the Anolis cybotes species group belong to the trunk-ground ecomorph category. Despite evolutionary stability of their trunk-ground microhabitat, populations of the A. cybotes group have undergone an evolutionary radiation associated with geographically distinct macrohabitats. A combined phylogeographic and morphometric study of this group reveals a strong association between macrohabitat type and morphology independent of phylogeny. This association results from long-term morphological evolutionary stasis in populations associated with mesic-forest environments ( A. c. cybotes and A. marcanoi ) and predictable morphometric changes associated with entry into new macrohabitat types (i.e., xeric forests, high-altitude pine forest, rock outcrops). Phylogeographic analysis of 73 new mitochondrial DNA sequences (1921 aligned sites) sampled from 68 geographic populations representing 12 recognized species and subspecies diagnoses 16 allopatric or parapatric groupings of populations differing from each other by 5–18% sequence divergence. At least some of these groupings appear to have attained species-level divergence from others. Evolutionary specialization to different macrohabitat types may be a major factor in the evolutionary diversification of Greater Antillean anoles.     

The recent history and population structure of five Mandarina snail species from subtropical Ogasawara (Bonin Islands, Japan)

The effect of Pleistocene climate change on the organisms of tropical and subtropical regions is rather poorly understood. We therefore studied the land snail genus Mandarina (Bradybaenidae) of oceanic Ogasawara (Bonin Islands, Japan), with the aim of using population genetic data to understand their recent history. Our analysis of a mitochondrial 16S ribosomal RNA region from more than 600 snails in five ground-living species suggests that populations on the small islands of Mukoujima, Anejima, Imotojima and Meijima, as well as on the low-lying southern and central parts of Hahajima, have probably undergone recent bottlenecks followed by subsequent expansions. Except between the main island of Hahajima and Mukouijima, there is almost no evidence for gene flow among islands even though the islands were connected repeatedly by land bridges through the Pleistocene. Within islands the population structure is severe, suggestive of a long-term, low level of gene flow (F(ST) is frequently greater than 0.5 among geographically close populations). Finally, there is a marked genetic patchiness, meaning that genetically close populations are sometimes separated by genetically distant populations. These patterns could be a consequence of expansion from bottlenecks, low active dispersal and founder effects caused by rare long-distance migrants. Unfortunately, the exact nature of the refugia and bottlenecks remains unknown because the palaeoclimate of this region is poorly understood. Dating the population size changes is also challenging because the molecular clock is uncertain. We suggest, however, that arid conditions or deforestation induced by decreased atmospheric CO(2) may have been the main factor in determining population size.     

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.     

Origin and evolution of endemic plants of the Bonin (Ogasawara) Islands

The Bonin Islands are typical oceanic islands, located at the western part of the North Pacific Ocean and approximately 1,000 km south of mainland Japan. This archipelago consists of about 20 small islands. Although floristic diversity is low due to the small area and limited environmental diversity, the Bonin Islands harbor unique endemic flora as in other well-known oceanic islands. This paper presents a brief summary of the results obtained from recent studies on the endemic flora of the Bonin Islands. The results are reviewed in relation to the four stages of the evolution of endemic flora in the oceanic islands; migration, establishment, enlargement and diversification. The ancestors of the flora originated mostly from tropical and subtropical Southeast Asia or mainland Japan by rare events of long distance dispersal. The proportion of bird-dispersed species is relatively high as for other oceanic islands. Genetic data sets obtained from allozyme variation in some endemic species suggest that migration occurred several million years ago and genetic diversity is correlated with current population size. At the time of establishment, self-compatible plants are expected to have an advantage. However, the percentage of dioecious plants is relatively high. This is partly due to evolutionary changes from hermaphroditic ancestors to dioecy which occurred in two genera in the Bonin Islands. In addition, there are some examples of evolutionary changes from herbaceous ancestors to woody endemics. Adaptive radiation is found in some genera, although the number of congeneric endemic species is less than five. Studies of allozyme variation inPittosporum, Symplocos andCrepidiastrum showed that genetic identity is generally very high between congeneric species in spite of their distinct morphologies. This result suggests that divergence of these species occurred rather recently and distinct morphological differences are based on a limited number of genetic changes.     

Rates of morphological evolution are correlated with species richness in salamanders

The tempo and mode of species diversification and phenotypic evolution vary widely across the tree of life, yet the relationship between these processes is poorly known. Previous tests of the relationship between rates of phenotypic evolution and rates of species diversification have assumed that species richness increases continuously through time. If this assumption is violated, simple phylogenetic estimates of net diversification rate may bear no relationship to processes that influence the distribution of species richness among clades. Here, we demonstrate that the variation in species richness among plethodontid salamander clades is unlikely to have resulted from simple time-dependent processes, leading to fundamentally different conclusions about the relationship between rates of phenotypic evolution and species diversification. Morphological evolutionary rates of both size and shape evolution are correlated with clade species richness, but are uncorrelated with simple estimators of net diversification that assume constancy of rates through time. This coupling between species diversification and phenotypic evolution is consistent with the hypothesis that clades with high rates of morphological trait evolution may diversify more than clades with low rates. Our results indicate that assumptions about underlying processes of diversity regulation have important consequences for interpreting macroevolutionary patterns.     

Distribution of land snails (Mollusca, Gastropoda, Pulmonata) on the island of Gran Canaria (Canary Islands) in relation to protected natural areas

The oceanic island of Gran Canaria, part of the Macaronesian Archipelagos, is highly deteriorated, a situation which has had negative repercussions for endemic species. A recent law, however, established 32 Protected Natural Areas in an attempt to remedy the problem. In the present work we show the distribution of common (non-endemic) land snails on the island. The snails were largely introduced by man over the past five centuries. A comparison between the distribution of the endemic land snails and the protected areas shows the latter to be generally adequate. We also suggest some modifications to the law in two Areas and the addition to the list of two further Areas so as to guarantee protection of several endemic species currently under threat (according to the criteria of the IUCN): 2 critically endangered, Hemicycla saulcyi saulcyi and Napaeus isletae, 2 vulnerable, Theba arinagae and Obelus despreauxii, and 1 lower risk, Hemicycla berkeleyi.     

Species richness in Madeiran land snails, and its causes

Aim  To unravel the reasons why a group of islands of 800 km² area should support over 200 extant land snail species.
Location  The temperate Atlantic Madeiran archipelago.
Methods  Distributional surveys and model testing.
Results  Repeated volcanic events have changed the topography of the islands and created periodic further isolates. There has been climatic fluctuation and varying sea level. Individual locations are not species-rich, but there is replacement from one subregion to another. Evidence of competitive interactions is lacking.
Main conclusions  High diversity results from the fortuitous coincidence of rates of geological and climatic change and isolation on the one hand, and migration and genetic divergence on the other. Hubbell's neutral model of biodiversity explains species richness if a factor is added describing structural instability or periodicity, here called the geodetic rate. This geodetic, biological interaction can explain why some archipelagos are species-rich.