Abstract. To re-evaluate the relationships of the major bivalve lineages, we amassed detailed morpho-anatomical, ultrastructural and molecular sequence data for a targeted selection of exemplar bivalves spanning the phylogenetic diversity of the class. We included molecular data for 103 bivalve species (up to five markers) and also analysed a subset of taxa with four additional nuclear protein-encoding genes. Novel as well as historically employed morphological characters were explored, and we systematically disassembled widely used descriptors such as gill and stomach 'types'. Phylogenetic analyses, conducted using parsimony direct optimisation and probabilistic methods on static alignments (maximum likelihood and Bayesian inference) of the molecular data, both alone and in combination with morphological characters, offer a robust test of bivalve relationships. A calibrated phylogeny also provided insights into the tempo of bivalve evolution. Finally, an analysis of the informativeness of morphological characters showed that sperm ultrastructure characters are among the best morphological features to diagnose bivalve clades, followed by characters of the shell, including its microstructure. Our study found support for monophyly of most broadly recognised higher bivalve taxa, although support was not uniform for Protobranchia. However, monophyly of the bivalves with protobranchiate gills was the best-supported hypothesis with incremental morphological and/or molecular sequence data. Autobranchia,
The systematics of the molluscan class Bivalvia are explored using a 5-gene Sanger-based approach including the largest taxon sampling to date, encompassing 219 ingroup species spanning 93 (or 82%) of the 113 currently accepted bivalve families. This study was designed to populate the bivalve Tree of Life at the family level and to place many genera into a clear phylogenetic context, but also pointing to several major clades where taxonomic work is sorely needed. Despite not recovering monophyly of Bivalvia or Protobranchia-as in most previous Sanger-based approaches to bivalve phylogeny-our study provides increased resolution in many higher-level clades, and supports the monophyly of Autobranchia, Pteriomorphia, Heteroconchia, Palaeoheterodonta, Heterodonta, Archiheterodonta, Euheterodonta, Anomalodesmata, Imparidentia, and Neoheterodontei, in addition to many other lower clades. However, deep nodes within some of these clades, especially Pteriomorphia and Imparidentia, could not be resolved with confidence. In addition, many families are not supported, and several are supported as non-monophyletic, including Malletiidae, Nuculanidae, Yoldiidae, Malleidae, Pteriidae, Arcidae, Propeamussiidae, Iridinidae, Carditidae, Myochamidae, Lyonsiidae, Pandoridae, Montacutidae, Galeommatidae, Tellinidae, Semelidae, Psammobiidae, Donacidae, Mactridae, and Cyrenidae; Veneridae is paraphyletic with respect to Chamidae, although this result appears to be an artifact. The denser sampling however allowed testing specific placement of species, showing, for example, that the unusual Australian Plebidonax deltoides is not a member of Donacidae and instead nests within Psammobiidae, suggesting that major revision of Tellinoidea may be required. We also showed that Cleidothaerus is sister group to the cementing member of Myochamidae, suggesting that Cleidothaeridae may not be a valid family and that cementation in Cleidothaerus and Myochama may have had a single origin. These results highlight the need for an integrative approach including as many genera as possible, and that the monophyly and relationships of many families require detailed reassessment. NGS approaches may be able to resolve the most recalcitrant nodes in the near future.
Although several studies have evaluated the genetic structure and phylogeographic patterns in many species of marine invertebrates, a general model that applies to all of them remains elusive. For example, some species present an admixture of populations with high gene flow, whereas others exhibit more complex patterns characterized by small‐scale unstructured genetic heterogeneity, even at a local scale. These differences are thought to be due to clear biological aspects such as direct versus indirect development, or the presence of lecithotrophic versus planktotrophic larvae, but few studies compare animals with similar distributions and life modes. Here, we explore the phylogeographic and genetic structure patterns in two chiton (Chiton olivaceus and Lepidopleurus cajetanus) and one abalone (Haliotis tuberculata) species co‐occurring in the same habitat. Samples were obtained from shallow rocky bottoms along the Iberian Peninsula (Atlantic and Mediterranean coasts), Italy, Croatia and Greece, and the mitochondrial markers COI and 16S rRNA gene were sequenced. Our data show evidence of admixture and population expansion in C. olivaceus and H. tuberculata, whereas L. cajetanus exhibited a ‘chaotic patchiness’ pattern defined by a high genetic variability with locality‐exclusive haplotypes, high genetic divergence, and a lack of geographic structure. Shared haplotypes were sampled in both coasts of Iberia (for H. tuberculata) and in the Western and Eastern Mediterranean (for C. olivaceus), potentially indicating high dispersal ability and a recent expansion. The processes underlying the fine‐scale structuring in L. cajetanus remain a mystery. These results are especially interesting because the reproductive mode of the two chitons is similar but differs from that of the abalone, with a veliger larva, while instead the genetic structure of C. olivaceus and H. tuberculata are similar, thus contrasting with predictions based on the life history of the three molluscs and showing that the genetic patterns of marine species may be shaped by many factors, including historical ones.
Palpigradi are a poorly understood group of delicate arachnids, often found in caves or other subterranean habitats. Concomitantly, they have been neglected from a phylogenetic point of view. Here we present the first molecular phylogeny of palpigrades based on specimens collected in different subterranean habitats, both endogean (soil) and hypogean (caves), from Australia, Africa, Europe, South America and North America. Analyses of two nuclear ribosomal genes and COI under an array of methods and homology schemes found monophyly of Palpigradi, Eukoeneniidae and a division of Eukoeneniidae into four main clades, three of which include samples from multiple continents. This supports either ancient vicariance or long-range dispersal, two alternatives we cannot distinguish with the data at hand. In addition, we show that our results are robust to homology scheme and analytical method, encouraging further use of the markers employed in this study to continue drawing a broader picture of palpigrade relationships.
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