BackgroundReconstructing the higher relationships of pulmonate gastropods has been difficult. The use of morphology is problematic due to high homoplasy. Molecular studies have suffered from low taxon sampling. Forty-eight complete mitochondrial genomes are available for gastropods, ten of which are pulmonates. Here are presented the new complete mitochondrial genomes of the ten following species of pulmonates: Salinator rhamphidia (Amphiboloidea); Auriculinella bidentata, Myosotella myosotis, Ovatella vulcani, and Pedipes pedipes (Ellobiidae); Peronia peronii (Onchidiidae); Siphonaria gigas (Siphonariidae); Succinea putris (Stylommatophora); Trimusculus reticulatus (Trimusculidae); and Rhopalocaulis grandidieri (Veronicellidae). Also, 94 new pulmonate-specific primers across the entire mitochondrial genome are provided, which were designed for amplifying entire mitochondrial genomes through short reactions and closing gaps after shotgun sequencing.ResultsThe structural features of the 10 new mitochondrial genomes are provided. All genomes share similar gene orders. Phylogenetic analyses were performed including the 10 new genomes and 17 genomes from Genbank (outgroups, opisthobranchs, and other pulmonates). Bayesian Inference and Maximum Likelihood analyses, based on the concatenated amino-acid sequences of the 13 protein-coding genes, produced the same topology. The pulmonates are paraphyletic and basal to the opisthobranchs that are monophyletic at the tip of the tree. Siphonaria, traditionally regarded as a basal pulmonate, is nested within opisthobranchs. Pyramidella, traditionally regarded as a basal (non-euthyneuran) heterobranch, is nested within pulmonates. Several hypotheses are rejected, such as the Systellommatophora, Geophila, and Eupulmonata. The Ellobiidae is polyphyletic, but the false limpet Trimusculus reticulatus is closely related to some ellobiids.ConclusionsDespite recent efforts for increasing the taxon sampling in euthyneuran (opisthobranchs and pulmonates) molecular phylogenies, several of the deeper nodes are still uncertain, because of low support values as well as some incongruence between analyses based on complete mitochondrial genomes and those based on individual genes (18S, 28S, 16S, CO1). Additional complete genomes are needed for pulmonates (especially for Williamia, Otina, and Smeagol), as well as basal heterobranchs closely related to euthyneurans. Increasing the number of markers for gastropod (and more broadly mollusk) phylogenetics also is necessary in order to resolve some of the deeper nodes -although clearly not an easy task. Step by step, however, new relationships are being unveiled, such as the close relationships between the false limpet Trimusculus and ellobiids, the nesting of pyramidelloids within pulmonates, and the close relationships of Siphonaria to sacoglossan opisthobranchs. The additional genomes presented here show that some species share an identical mitochondrial gene order due to convergence.
Aim We collate and analyse data for land snail diversity and endemism, as a means of testing the explanatory power of the general dynamic model of oceanic island biogeography (GDM): a theoretical model linking trends in species immigration, speciation and extinction to a generalized island ontogeny. Location Eight oceanic archipelagos: Azores, Canaries, Hawaii, Galápagos, Madeira, Samoa, Society, Tristan da Cunha. Methods Using data obtained from literature sources we examined the power of the GDM through its derivative ATT2 model (i.e. diversity metric = b1 + b2Area + b3Time + b4Time2), in comparison with all the possible simpler models, e.g. including only area or time. The diversity metrics considered were the number of (1) native species, (2) archipelagic endemic species, and (3) single‐island endemic species. Models were evaluated using both log‐transformed and untransformed diversity data by means of linear mixed effect models. For Hawaii and the Canaries, responses of different major taxonomic groups were also analysed separately. Results The ATT2 model was always included within the group of best models and, in many cases, was the single‐best model and was particularly successful in fitting the log‐transformed diversity metrics. In four archipelagos, a hump‐shaped relationship with time (island age) is apparent, while the other four archipelagos show a general increase of species richness with island age. In Hawaii and the Canaries outcomes vary between different taxonomic groups. Main conclusions The GDM is an intentionally simplified representation of environmental and diversity dynamics on oceanic islands, which predicts a simple positive relationship between diversity and island area combined with a humped response to time. We find broad support for the applicability of this model, especially when a full range of island developmental stages is present. However, our results also show that the varied mechanisms of island origins and the differing responses of major taxa should be taken into consideration when interpreting diversity metrics in terms of the GDM. This heterogeneity is reflected in the fact that no single model outperforms all the other models for all datasets analysed.
Aim To quantify the influence of past archipelago configuration on present‐day insular biodiversity patterns, and to compare the role of long‐lasting archipelago configurations over the Pleistocene to configurations of short duration such as at the Last Glacial Maximum (LGM) and the present‐day. Location 53 volcanic oceanic islands from 12 archipelagos worldwide—Azores, Canary Islands, Cook Islands, Galápagos, Gulf of Guinea, Hawaii, Madeira, Mascarenes, Pitcairn, Revillagigedo, Samoan Islands and Tristan da Cunha. Time period The last 800 kyr, representing the nine most recent glacial–interglacial cycles. Major taxa studied Land snails and angiosperms. Methods Species richness data for land snails and angiosperms were compiled from existing literature and species checklists. We reconstructed archipelago configurations at the following sea levels: the present‐day high interglacial sea level, the intermediate sea levels that are representative of the Pleistocene and the low sea levels of the LGM. We fitted two alternative linear mixed models for each archipelago configuration using the number of single‐island endemic, multiple‐island endemic and (non‐endemic) native species as a response. Model performance was assessed based on the goodness‐of‐fit of the full model, the variance explained by archipelago configuration and model parsimony. Results Single‐island endemic richness in both taxonomic groups was best explained by intermediate palaeo‐configuration (positively by area change, and negatively by palaeo‐connectedness), whereas non‐endemic native species richness was poorly explained by palaeo‐configuration. Single‐island endemic richness was better explained by intermediate archipelago configurations than by the archipelago configurations of the LGM or present‐day. Main conclusions Archipelago configurations at intermediate sea levels—which are representative of the Pleistocene—have left a stronger imprint on single‐island endemic richness patterns on volcanic oceanic islands than extreme archipelago configurations that persisted for only a few thousand years (such as the LGM). In understanding ecological and evolutionary dynamics of insular biota it is essential to consider longer‐lasting environmental conditions, rather than extreme situations alone.
Mitochondrial DNA hyperdiversity is primarily caused by high mutation rates (µ) and has potential implications for mitogenome architecture and evolution. In the hyperdiverse mtDNA of Melarhaphe neritoides (Gastropoda: Littorinidae), high mutational pressure generates unusually large amounts of synonymous variation, which is expected to (1) promote changes in synonymous codon usage, (2) reflect selection at synonymous sites, (3) increase mtDNA recombination and gene rearrangement, and (4) be correlated with high mtDNA substitution rates. The mitogenome of M. neritoides was sequenced, compared to closely related littorinids and put in the phylogenetic context of Caenogastropoda, to assess the influence of mtDNA hyperdiversity and high µ on gene content and gene order. Most mitogenome features are in line with the trend in Mollusca, except for the atypical secondary structure of the methionine transfer RNA lacking the TΨC-loop. Therefore, mtDNA hyperdiversity and high µ in M. neritoides do not seem to affect its mitogenome architecture. Synonymous sites are under positive selection, which adds to the growing evidence of non-neutral evolution at synonymous sites. Under such non-neutrality, substitution rate involves neutral and non-neutral substitutions, and high µ is not necessarily associated with high substitution rate, thus explaining that, unlike high µ, a high substitution rate is associated with gene order rearrangement.
Oceanic islands have been the grand stage of documented extinctions. In view of limited resources, efficient prioritization is crucial to avoid the extinction of taxa. This work lists the top 100 management priority species for the European archipelagos of the Macaronesian region (Azores, Madeira and the Canary Islands), taking into account both their protection priority and their management feasibility. Bryophytes, vascular plants, molluscs, arthropods and vertebrates were scored by species experts following two sets of criteria: (i) protection priority, including ecological value, singularity, public institutions' management responsibilities and social value; (ii) management feasibility, including threats knowledge and control feasibility, external socio-economical support for management and biological recovery potential. Environmental managers weighted the same criteria according to their management importance. Final species scores were determined by the combination of both species valuation and criteria weighting. Vascular plants dominate the Top 100 list, followed by arthropods and vertebrates. The majority of listed taxa are endemic to one archipelago or even to a single island. The management feasibility criteria did not dictate that all taxa must be eminently endangered, as for most of the species it should be relatively easy to control threats. The main advantages of this process are the independent participation of scientists and conservation managers, the inclusion of criteria on both protection priority and management feasibility and the taxonomically unbiased nature of the process. This study provides a potentially useful biodiversity conservation tool for the Macaronesian archipelagos that could be readily implemented by the respective regional governments in future legislation.
The Pleistocene (Eemian) outcrops of Lagoinhas and Prainha, located at Santa Maria Island (Azores), were investigated and their fossil mollusc content reported. These studies revealed that the last glaciation affected two groups of molluscs: the ‘warm‐guest’ gastropods with West African or Caribbean affinities (e.g. Conus spp., Cantharus variegatus, Bulla amygdala, Trachypollia nodulosa) and shallow bivalve species mainly associated with sandy habitats (Ensis minor, Lucinella divaricata, and probably Laevicardium crassum). In this paper we focus on this group of bivalves, which has since locally disappeared from the Azores. We relate the local disappearance of these bivalves in the Azores with the lack of sand in the shelf. The specific characteristics of the Santa Maria shelf combined with the sea‐level drop during the Weichselian prevented deposition of the lowstand deposits and permitted erosion of the previous ones, leaving the shelf without a sediment cover. Copyright © 2008 John Wiley & Sons, Ltd.
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