Allopatric Speciation Model Research Articles

The genetic structure and demographic history of Zabelia tyaihyonii, endemic to Korean limestone karst forests, based on genome‐wide SNP markers

AbstractSimilar to the global phenomenon, many plant species endemic to Korean limestone karst forests are at risk of extinction due to human intervention. Zabelia tyaihyonii is a familiar shrub, called “Hardy abelia” and “Fragrant abelia” growing in the karst forests of Korea, where it is one of the most threatened species. We investigated the genetic structure and demographic history of Z. tyaihyonii, which allow us to develop appropriate conservation and management strategies. The genetic structure was evaluated using a total of 187 samples from 14 populations, covering the entire distribution of Z. tyaihyonii in South Korea. We utilized 254 and 1753 SNP loci obtained via MIG‐seq (Multiplexed ISSR Genotyping by sequencing) for structure and demographic analyses, respectively. The population demographic modeling was performed with site frequency spectrum. To gain further historical insights, we also employed ENM (Ecological Niche Modeling). We found two distinct clusters (CLI and CLII) of ancient origin (ca. 490 ka). Despite CLII experiencing a more severe bottleneck, both clusters showed similar levels of genetic diversity, indicating mutual historical gene flow. Their historical distribution range seems to have changed very little. We proposed a historical distribution scenario for Z. tyaihyonii, taking into account its intrinsic factors, and emphasized a more complex response to Quaternary climate change beyond simple allopatric speciation models. These findings provide valuable insights for conservation and management strategies for Z. tyaihyonii.

Plumage assisted divergence in a vocally complex island endemic: The Dicrurus paradiseus species complex in Sri Lanka

Models of allopatric speciation within an island biogeographic framework suggest that the division of ancestral mainland populations leads to one or more allopatric island species predominantly through natural and sexual selection or genetic drift. Here we studied phenotypic divergence in a phylogenetic framework in the Dicrurus paradiseus allospecies complex in Sri Lanka, a continental island located in the Indian plate, to understand the complexity of phenotypic divergence on an island. Members of the genus Dicrurus are known as drongos and are conserved in morphology and plumage, but highly variable in vocalization due to vocal learning and mimicry. Two closely related drongos are found in Sri Lanka: the endemic D. lophorinus (or D. paradiseus lophorinus to many authors) found in the wet zone of the island and the widespread continental species D. paradiseus, which inhabits the dry zone. Sampling from all major populations, and voucher specimens from museums across their range in Sri Lanka, we examined phenotypic and genetic variation in this group. The phenotype showed two clusters: birds with a fish-like tail and erect crest (D. lophorinus), and birds with elongated tail streamers with backwardly curved crest (D. paradiseus). There was no significant difference in the vocal traits compared. The genetic variation was examined using two nuclear (Myo 2 and c-mos) and two mitochondrial (ND2 and Cytb) loci and the phylogenetic relationship was analyzed using the Bayesian inference coalescent-based species tree estimation method. The quantitative criteria for species delimitation provided a score sufficient to consider these two taxa as distinct species by considering measurements of body and plumage, acoustics, behaviour and distribution. The phylogeny supports distinct species status for the Sri Lanka Drongo (Dicrurus lophorinus) and that the D. lophorinus and D. paradiseus sister pair diverged since 1.35 mya. The variation in the crest and the tail plumage (components of phenotype) were the main contributors of the divergence, despite the similarity in general appearance and vocalization of the allopatric species.

Multi-queen breeding is associated with the origin of inquiline social parasitism in ants

Social parasites exploit the brood care behavior of their hosts to raise their own offspring. Social parasites are common among eusocial Hymenoptera and exhibit a wide range of distinct life history traits in ants, bees, and wasps. In ants, obligate inquiline social parasites are workerless (or nearly-so) species that engage in lifelong interactions with their hosts, taking advantage of the existing host worker forces to reproduce and exploit host colonies’ resources. Inquiline social parasites are phylogenetically diverse with approximately 100 known species that evolved at least 40 times independently in ants. Importantly, ant inquilines tend to be closely related to their hosts, an observation referred to as ‘Emery’s Rule’. Polygyny, the presence of multiple egg-laying queens, was repeatedly suggested to be associated with the origin of inquiline social parasitism, either by providing the opportunity for reproductive cheating, thereby facilitating the origin of social parasite species, and/or by making polygynous species more vulnerable to social parasitism via the acceptance of additional egg-laying queens in their colonies. Although the association between host polygyny and the evolution of social parasitism has been repeatedly discussed in the literature, it has not been statistically tested in a phylogenetic framework across the ants. Here, we conduct a meta-analysis of ant social structure and social parasitism, testing for an association between polygyny and inquiline social parasitism with a phylogenetic correction for independent evolutionary events. We find an imperfect but significant over-representation of polygynous species among hosts of inquiline social parasites, suggesting that while polygyny is not required for the maintenance of inquiline social parasitism, it (or factors associated with it) may favor the origin of socially parasitic behavior. Our results are consistent with an intra-specific origin model for the evolution of inquiline social parasites by sympatric speciation but cannot exclude the alternative, inter-specific allopatric speciation model. The diversity of social parasite behaviors and host colony structures further supports the notion that inquiline social parasites evolved in parallel across unrelated ant genera in the formicoid clade via independent evolutionary pathways.

Phylogeography of the endangered orchids Cypripedium japonicum and Cypripedium formosanum in East Asia: Deep divergence at infra‐ and interspecific levels

AbstractTo date, little is known about the past evolutionary trajectories of rare and endangered orchids native to mainland China, Japan, and Korea (the CJK region). In this study, we focus on two endangered orchids, Cypripedium japonicum (present in the three countries) and C. formosanum (endemic to Taiwan), to understand the divergence/speciation models that would have been operating in this group, including genetic diversity, geographic structure, and colonization pathways across the region. Using a combination of five cpDNA regions, we reconstructed phylogenetic trees and investigated the genetic diversity/structure of 20 populations. Ecological niche modeling was used to gain insight into the paleodistribution and dispersal corridors at the Last Glacial Maximum and to survey climatic niche differences. Populations from mainland China + Korea, Japan, and Taiwan formed three distinct monophyletic lineages and were placed into separate genetic clusters, agreeing with geographic barriers and species boundaries. Populations of C. japonicum in mainland China harbored the highest diversity, suggesting the presence of multiple glacial refugia. The Korean populations would have originated from either western/central or eastern China, probably using a dispersal corridor across the East China Sea shelf. The divergence of C. formosanum is proposed under an allopatric speciation model, also highly influenced by a climate niche shift. In the context of previous studies, a deep divergence in cpDNA sequences between Chinese + Korean and Japanese populations of C. japonicum may be taken as an example of the speciation events of the CJK flora since the late Neogene that have led to its current species richness.

Multilocus phylogeography and ecological niche modeling suggest speciation with gene flow between the two Bamboo Partridges

BackgroundUnderstanding how species diversify is a long-standing question in biology. The allopatric speciation model is a classic hypothesis to explain the speciation process. This model supposes that there is no gene flow during the divergence process of geographically isolated populations. On the contrary, the speciation with gene flow model supposes that gene flow does occur during the speciation process. Whether allopatric species have gene flow during the speciation process is still an open question.MethodsWe used the genetic information from 31 loci of 24 Chinese Bamboo Partridges (Bambusicola thoracicus) and 23 Taiwan Bamboo Partridges (B. sonorivox) to infer the gene flow model of the two species, using the approximate Bayesian computation (ABC) model. The ecological niche model was used to infer the paleo-distribution during the glacial period. We also tested whether the two species had a conserved ecological niche by means of a background similarity test.ResultsThe genetic data suggested that the post-divergence gene flow between the two species was terminated before the mid-Pleistocene. Furthermore, our ecological niche modeling suggested that their ecological niches were highly conserved, and that they shared an overlapping potential distribution range in the last glacial maximum.ConclusionsThe allopatric speciation model cannot explain the speciation process of the two Bamboo Partridges. The results of this study supported a scenario in which speciation with gene flow occurring between the allopatric species and have contributed to our understanding of the speciation process.

Gulf Coast vicariance shapes phylogeographic history of a North American freshwater mussel species complex

AbstractAimFreshwater mussels share habitat and are parasites of freshwater fishes during the larval life stage. Therefore, models of fish biogeography may also explain the historical biogeography of freshwater mussels. We tested this assumption using predictions of three biogeographic models constructed for northern Gulf of Mexico drainages on a freshwater mussel species complex. Specifically, we tested (1) if speciation was due to vicariant events of fluctuating sea levels that separated lineages east‐west of the Mobile Basin (Central Gulf Coast speciation hypothesis), (2) if the timing of divergences occurred 8.5–3.5 MYA (Gulf Coast allopatric speciation model) and (3) if diversification in Mississippi River populations was recent and for evidence of population increase consistent with range expansion into northern deglaciated regions (Pleistocene glaciation model).LocationEastern North America.TaxonFreshwater mussels (Bivalvia: Unionidae), Lampsilis teres and L. floridensis.MethodsWe collected 249 specimens from 73 localities across the group's distribution. We used three molecular markers (COI, NDI & ITSI) to conduct time calibrated Bayesian phylogenetic analyses, phylogeographic analyses (AMOVA & SAMOVA) and demographic analyses including Bayesian skyline plots.ResultsLampsilis teres and L. floridensis are allopatric species whose distributions meet at the eastern edge of the Mobile Basin. Speciation was estimated to occur in the late Miocene. Populations from isolated river systems surrounding the Gulf of Mexico are almost all monophyletic. Mississippi drainage samples formed a shallow clade with recent diversification and showed evidence of recent population expansion.Main conclusionsThe historical biogeography of the L. teres species complex is broadly consistent with tested ichthyofaunal models. The timing of speciation and intraspecific divergences correspond to low sea‐level events suggesting that Gulf Coast sea‐level fluctuations are responsible for dispersal (sea‐level recession) and subsequent cladogenesis (sea‐level inundation). Consistent with numerous other freshwater studies, we found the Mobile Basin to be a suture zone, which may be due to the narrow, offshore continental shelf.

Morphological and molecular evidence support the taxonomic separation of the medically important Neotropical spiders Phoneutria depilata (Strand, 1909) and P. boliviensis (F.O. Pickard-Cambridge, 1897) (Araneae, Ctenidae).

The species of the genus Phoneutria (Ctenidae), also called banana spiders, are considered amongst the most venomous spiders in the world. In this study we revalidate P. depilata (Strand, 1909), which had been synonymized with P. boliviensisis (F.O. Pickard-Cambridge, 1897), using morphological and nucleotide sequence data (COI and ITS-2) together with species delimitation methods. We synonymized Ctenus peregrinoides, Strand, 1910 and Phoneutria colombiana Schmidt, 1956 with P. depilata. Furthermore, we designated Ctenus signativenter Strand, 1910 as a nomen dubium because the exact identity of this species cannot be ascertained with immature specimens, but we note that the type locality suggests that the C. signativenter syntypes belong to P. depilata. We also provide species distribution models for both species of Phoneutria and test hypotheses of niche conservatism under an allopatric speciation model. Our phylogenetic analyses support the monophyly of the genus Phoneutria and recover P. boliviensis and P. depilata as sister species, although with low nodal support. In addition, the tree-based species delimitation methods also supported the separate identities of these two species. Phoneutria boliviensis and P. depilata present allopatric distributions separated by the Andean mountain system. Species distribution models indicate lowland tropical rain forest ecosystems as the most suitable habitat for these two Phoneutria species. In addition, we demonstrate the value of citizen science platforms like iNaturalist in improving species distribution knowledge based on occurrence records. Phoneutria depilata and P. boliviensis present niche conservatism following the expected neutral model of allopatric speciation. The compiled occurrence records and distribution maps for these two species, together with the morphological diagnosis of both species, will help to identify risk areas of accidental bites and assist health professionals to determine the identity of the species involved in bites, especially for P. depilata.

Progressing diversification and biogeography of the mesopelagic Nematoscelis (Crustacea: Euphausiacea) in the Atlantic

Abstract Evolutionary mechanisms driving the diversification of pelagic animals remain poorly understood, partly due to the high levels of gene flow in the open ocean. We use molecular phylogenetics, morphological, and phylogeographic approaches to test the allopatric speciation model in respect to the Atlantic krill genus Nematoscelis. Based on our observations, we hypothesize that diversification in genus Nematoscelis may occur through three progressing stages: (i) populations separated geographically and divergence occurred in the mitochondrial COI gene only (Nematoscelis megalops, one clade of Nematoscelis tenella), (ii) morphology diverged (clades of Nematoscelis microps and Nematoscelis atlantica), and (iii) the nuclear H3 gene diverged (clades of N. tenella). Our results confirm allopatric expectations and the dispersal-limiting speciation model. We propose that the primary driver of diversification is geographic isolation coupled with hydrology-linked gene barriers at ∼14–22°N (new) and ∼30°S. The second driver preventing hybridization of diverged populations is linked to external morphology, i.e. enlarged photophores and chitin saddles on the pleon of males, which facilitate sexual selection by female choice. Same-male forms, even belonging to different species, rarely co-occur, which makes the selection effective and not biased. Our results implicate a significant role of non-copulatory characters in Nematoscelis speciation.

Reproduction and life span characterization of Artemia urmiana in Lake Urmia, Iran (Branchiopoda: Anostraca)

Urmia Lake is one of the largest permanent hypersaline lakes in the world. The lake can be characterized as an oligotrophic lake concerning phytoplankton production. Due to low precipitation in recent years in Urmia Lake’s catchment area, the water level has declined as much as 4 m compared to 15 years ago. Irrespective of its recent evolution, Urmia Lake has always shown fluctuations in historical and geological times which may cause some ecological partition. Regarding allopatric speciation model of Artemia we assume that several populations of Artemia are generated during last years and living in Urmia Lake. To investigate Artemia population diversity in Urmia Lake, Artemia urmiana Gunther 1890 cysts were collected from five sites in Lake Urmia, Iran. These sites in part represent areas with different physicochemical conditions. To detect possible differences between stations, discriminant analysis (DA) was performed on data of survival, growth, and reproduction at two salinities (75 and 150 g.l-1) in laboratory culture tests. The performed DA implied that for the reproductive and life span characters, in both salinities, the stations of Golmankhaneh can be determined as the separated group among the studied stations which can be considered for industrial projects and evolutionary studies.

Secondary contact after allopatric divergence explains avian speciation and high species diversity in the Himalayan-Hengduan Mountains

The geographical context of speciation is important for understanding speciation and community assembly. However, the predominant mode of speciation in the Himalayan-Hengduan Mountains (HHMs), a global biodiversity hotspot, remains unknown. Here, we examined the role of geography in speciation using four pairs of sister or closely related avian species that currently co-occur in the HHMs. While multilocus network analyses based on nine to eleven genes revealed deep splits between these species, several allelic networks based on individual loci suggested phylogenetic paraphyly implying a recent history of divergence. Following extensive sampling in the contact zones of these species pairs, the coalescence-based approximate Bayesian computation approach supported no gene flow during their divergence and was consistent with an allopatric speciation model. We further estimated the divergence times of the four species pairs during the middle and late Pleistocene, which were characterized by increased amplitudes of glacial variability. We found a positive relationship between their divergence times and current sympatry levels, supporting a scenario of secondary contact following allopatric speciation. The Pleistocene glacial-interglacial cycles may have led to the initial geographic population isolation; ecological divergence or mate choice might further accelerate their differentiation during secondary contact, facilitating their speciation and species accumulation in the mountainous landscape. Our findings reveal the critical role of geographic isolation in speciation in the HHMs and shed light on how this biodiversity hotspot aggregates numerous species.

Oh, what a tangled web we weave

Oh, what a tangled web we weave

Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species.

Pristimantis is the most diverse genus of tetrapods comprising 532 described species. It contains a large number of morphologically cryptic species that are being discovered with the assistance of genetic evidence. We use molecular, morphological, bioacoustic, and environmental data to assess the phylogenetic relationships and determine the species within an Andean clade of Pristimantis, which is distributed from central Ecuador to northern Peru. We assign to this clade the name Huicundomantis and propose it as a subgenus. Our results show that Huicundomantis is composed of two large clades which we name as the P. phoxocephalus species group and the P. cryptomelas species group. Huicundomantis is composed of 28 species of which 12 have been described and 16 are new. We describe 11 of these undescribed species. The most effective characters to discriminate among species are DNA sequences, qualitative morphology, and advertisement calls. Morphometric and environmental characters are not very useful to define species limits. We clarify the identity of P. riveti and show that populations from southern Ecuador traditionally ascribed to P. riveti are a new species, P. lutzaesp. nov. We also show that P. prometeii is a junior synonym of P. hampatusami. The current diversity and geographic distribution of Huicundomantis are consistent with a model of allopatric speciation. All species have a restricted distribution range (less than 4330 km2) and are assigned to the Red List categories Data Deficient or threatened with extinction. We provide new reasons to increase conservation efforts for these species and their habitat. Taking our results into account, Pristimantis species richness in Ecuador increases from 211 to 221 species, and the number of species endemic to Ecuador from 119 to 129.

Ancient habitat shifts and organismal diversification are decoupled in the African viper genus Bitis (Serpentes: Viperidae)

AbstractAimThe expansion of open habitats during the mid‐Miocene has been hypothesized as a driver of allopatric speciation for many African taxa. This habitat‐dependent mode of diversification has been implicated in the shift from C3 (e.g. forest/woodland) to C4 dominated systems (i.e. open savanna, grasslands) in a number of African squamates. We examined this hypothesis using a genus of African viperid snakes (Bitis) with both open habitat and forest‐dwelling representatives.LocationAfrica.MethodsA comprehensive multilocus dataset was used to generate a calibrated species tree using a multispecies coalescent model. Individual gene trees and patterns of nuclear allele sharing were used to assess species monophyly and isolation. To test the habitat‐dependent evolution hypothesis, we generated an ancestral character state reconstruction for open and closed habitats using the dated phylogeny. This was related to the timing of open habitat expansion and forest/woodland contraction in Africa.ResultsThe genus Bitis originated in the Oligocene, with species level diversification in the late Miocene/Pliocene. Four well‐supported clades correspond to the recognized subgenera Bitis, Keniabitis, Macrocerastes and Calechidna. Several previously unrecognized lineages potentially represent cryptic species.Main conclusionsHabitat‐dependent evolution does not appear to have been a main driver for generic level viperine diversification: the ancestral state for Bitis was open habitat and at least one clade moved into forest in the Miocene, long after forest had contracted and fragmented. Forest‐dependent species diversified only in the late Miocene, presumably as forest became further reduced in extent, fitting an allopatric model of speciation. Although our results do not favour a general pattern of habitat‐dependent diversification in Bitis, cladogenesis within the subgenus Calechidna for “arenicolous” species (Bitis caudalis complex) and “rupicolous” species (B. atropos‐cornuta complex), corresponds to the aridification of southwest Africa. This suggests there are subtleties not captured in the broad open habitat category, which are relevant for understanding the role of habitat‐dependent evolution.

Climate variability impacts on diversification processes in a biodiversity hotspot: a phylogeography of ancient pseudoscorpions in south-western Australia

Abstract The south-western division of Australia is the only biodiversity hotspot in Australia and is well-known for extreme levels of local endemism. Climate change has been identified as a key threat for flora and fauna, but very few data are presently available to evaluate its impact on invertebrate fauna. Here, we derive a molecular phylogeography for pseudoscorpions of the genus Pseudotyrannochthonius that in the south-west are restricted to regions with the highest rainfall. A dated molecular phylogeny derived from six gene fragments is used for biogeographic reconstruction analyses, spatial mapping, environmental niche-modelling, and to infer putative species. Phylogenetic analyses uncover nine clades with mostly allopatric distributions and often small linear ranges between 0.5 and 130 km. Molecular dating suggests that the origins of contemporary diversity fall into a period of warm/humid Palaeogene climates, but splits in the phylogeny coincide with major environmental shifts, such as significant global cooling during the Middle Miocene. By testing several models of historical biogeography available for the south-west, we determine that Pseudotyrannochthonius is an ancient relict lineage that principally follows a model of allopatric speciation in mesic zone refugia, although there are derivations from this model in that some species are older and distribution patterns more complex than expected. Ecological niche models indicate that drier and warmer future climates will lead to range contraction towards refugia of highest rainfall, probably mimicking past variations that have generated high diversity in these areas. Their conservation management will be crucial for preserving the unique biodiversity heritage of the south-west.

Dispersal and the transition to sympatry in vertebrates.

Under allopatric speciation models, a key step in the build-up of species richness is population dispersal leading to the co-occurrence of previously geographically isolated forms. Despite its central importance for community assembly, the extent to which the transition from spatial segregation (allopatry or parapatry) to coexistence (sympatry) is a predictable process, or alternatively one governed by chance and the vagaries of biogeographic history, remains poorly understood. Here, we use estimated divergence times and current patterns of geographical range overlap among sister species to explore the evolution of sympatry in vertebrates. We show that rates of transition to sympatry vary predictably according to ecology, being faster in marine or strongly dispersive terrestrial clades. This association with organism vagility is robust to the relative frequency of geographical speciation modes and consistent across taxonomic scales and metrics of dispersal ability. These findings reject neutral models of dispersal assembly based simply on evolutionary age and are not predicted by the main alternative view that range overlap is primarily constrained by biotic interactions. We conclude that species differences in dispersal limitation are fundamental in organizing the assembly of ecological communities and shaping broad-scale patterns of biodiversity over space and time.

A new dynamic null model for phylogenetic community structure.

Phylogenies are increasingly applied to identify the mechanisms structuring ecological communities but progress has been hindered by a reliance on statistical null models that ignore the historical process of community assembly. Here, we address this, and develop a dynamic null model of assembly by allopatric speciation, colonisation and local extinction. Incorporating these processes fundamentally alters the structure of communities expected due to chance, with speciation leading to phylogenetic overdispersion compared to a classical statistical null model assuming equal probabilities of community membership. Applying this method to bird and primate communities in South America we show that patterns of phylogenetic overdispersion – often attributed to negative biotic interactions – are instead consistent with a species neutral model of allopatric speciation, colonisation and local extinction. Our findings provide a new null expectation for phylogenetic community patterns and highlight the importance of explicitly accounting for the dynamic history of assembly when testing the mechanisms governing community structure.

Baja California disjunctions and phylogeographic patterns in sympatric California blennies

Baja California disjuncts, which are marine species that are present in the northern Sea of Cortez and along the Pacific Coast of California and Baja California, but absent from the south, have been shown to be interesting models of allopatric speciation. While at least 19 species of fish are Baja California disjuncts, no study has ever evaluated the genetic patterns of closely related species. Here, we used three sympatric California blennies (genus Hypsoblennius), where two species have disjunct populations and one does not. Based on one mitochondrial and one nuclear molecular marker, we found that the two disjunct species exhibit different genetic patterns, one species showing disjunct populations as reciprocally monophyletic assemblages, while the other species showed evidence of gene flow between disjunct populations. In addition, the non-disjunct species, H. gilberti, exhibits high gene flow along the California and Baja California coasts. I hypothesized that adult habitat and historical vicariant events, more so than pelagic larval duration, played major roles in shaping the currently observed distribution and genetic patterns for these species.

Ecological divergence and speciation between lemur (Eulemur) sister species in Madagascar

Understanding ecological niche evolution over evolutionary timescales is crucial to elucidating the biogeographic history of organisms. Here, we used, for the first time, climate-based ecological niche models (ENMs) to test hypotheses about ecological divergence and speciation processes between sister species pairs of lemurs (genus Eulemur) in Madagascar. We produced ENMs for eight species, all of which had significant validation support. Among the four sister species pairs, we found nonequivalent niches between sisters, varying degrees of niche overlap in ecological and geographic space, and support for multiple divergence processes. Specifically, three sister-pair comparisons supported the null model that niches are no more divergent than the available background region. These findings are consistent with an allopatric speciation model, and for two sister pairs (E.collaris-E.cinereiceps and E.rufus-E.rufifrons), a riverine barrier has been previously proposed for driving allopatric speciation. However, for the fourth sister pair E.flavifrons-E.macaco, we found support for significant niche divergence, and consistent with their parapatric distribution on an ecotone and the lack of obvious geographic barriers, these findings most strongly support a parapatric model of speciation. These analyses thus suggest that various speciation processes have led to diversification among closely related Eulemur species.

Phylogeography of the sand dollar genus Mellita: Cryptic speciation along the coasts of the Americas

Sand dollars of the genus Mellita are members of the sandy shallow-water fauna. The genus ranges in tropical and subtropical regions on the two coasts of the Americas. To reconstruct the phylogeography of the genus we sequenced parts of the mitochondrial cytochrome oxidase I and of 16S rRNA as well as part of the nuclear 28S rRNA gene from a total of 185 specimens of all ten described morphospecies from 31 localities. Our analyses revealed the presence of eleven species, including six cryptic species. Sequences of five morphospecies do not constitute monophyletic molecular units and thus probably represent ecophenotypic variants. The fossil-calibrated phylogeny showed that the ancestor of Mellita diverged into a Pacific lineage and an Atlantic+Pacific lineage close to the Miocene/Pliocene boundary. Atlantic M. tenuis, M. quinquiesperforata and two undescribed species of Mellita have non-overlapping distributions. Pacific Mellita consist of two highly divergent lineages that became established at different times, resulting in sympatric M. longifissa and M. notabilis. Judged by modern day ranges, not all divergence in this genus conforms to an allopatric speciation model. Only the separation of M. quinquiesperforata from M. notabilis is clearly due to vicariance as the result of the completion of the Isthmus of Panama. The molecular phylogeny calibrated on fossil evidence estimated this event as having occurred ∼3Ma, thus providing evidence that, contrary to a recent proposal, the central American Isthmus was not completed until this date.

The Diversity of Land Molluscs—Questions Unanswered and Questions Unasked*

Studies on the diversity of land molluscs raise a number of crucial questions that remain unanswered, or in some cases have not been asked. At the most fundamental level, we need to question the comparability of species concepts used in different studies, which influence our assessment of overall diversity and of regional differentiation. Current estimates of global diversity are likely to be too low. Beyond that, the assumption that all species can be represented by equivalent digits needs to be challenged: the range of size, habits and trophic levels needs to be taken into account. While higher taxonomic categories can be used as rough proxies for similar ecology, we have very little detailed information other than for shell size and shape, where the huge range suggests some radical differences within and among faunas. The range of niches occupied may differ among faunas: are we dealing with comparable taxonomic entities? Our knowledge of microhabitat requirements is severely deficient.Regional species richness varies much more than maximum local (site) richness across faunas. The poor regional faunas of areas subject to glacial or periglacial Pleistocene conditions are not always impoverished at local level. While this might suggest the presence of competitive interactions, the evidence for competition is scarce. There are many cases where closely-related species live in sympatry at the smallest scales. Nevertheless, there are cases of parapatric or microallopatric distributions of congeners, and metapopulation dynamics can facilitate the co-existence of potential competitors. There may be no general rule; cases of both character displacement and convergence in sympatry are reported.While most studies accept the traditional model of allopatric speciation to account for the development of diversity, the balance between vicariance and dispersal as driving forces remains undetermined, and clearly varies from case to case. Poor powers of active dispersal may permit genetic differentiation over very short distances, but there are a number of spectacular cases of long-distance passive dispersal that suggest that our direct estimates of dispersal are questionable. Better knowledge of actual dispersal and of the ecological requirements of land mollusc species should be priorities for the future.