title: Speciation in the dark: diversification and biogeography of the deep-sea gastropod genusScaphanderin the Atlantic Ocean.
abstract: OBJECTIVE
  The aim of this work was to improve understanding about the mode, geography and tempo of diversification in deep-sea organisms, using a time-calibrated molecular phylogeny of the heterobranch gastropod genusScaphander.
  
  
  METHODS
  Atlantic and Indo-West Pacific (IWP) oceans.
  
  
  METHODS
  Two mitochondrial gene markers (COIand16S) and one nuclear ribosomal gene (28S) from six Atlantic species ofScaphander, and four IWP species were used to generate a multilocus phylogenetic hypothesis using uncorrelated relaxed-clock Bayesian methods implemented in beast and calibrated with the first occurrence ofScaphanderin the fossil record (58.7-55.8 Ma).
  
  
  RESULTS
  Two main clades were supported: clade A, with sister relationships between species and subclades from the Atlantic and IWP; and clade B, with two western Atlantic sister species. Our estimates indicate that the two earliest divergences in clade A occurred between the middle Eocene and late Miocene and the most recent speciation occurred within the middle Miocene to Pleistocene. The divergence between the two western Atlantic species in clade B was estimated at late Oligocene-Pliocene.
  
  
  CONCLUSIONS
  The prevailing mode of speciation inScaphanderwas allopatric, but one possible case of sympatric speciation was detected between two western Atlantic species. Sister relationships between IWP and Atlantic lineages suggest the occurrence both of vicariance events caused by the closure of the Tethyan Seaway and of dispersal between the two ocean basins, probably around South Africa during episodic disruptions of the deep-sea regional current system caused by glacial-interglacial cycles. Cladogenetic estimates do not support comparatively older diversification of deep-sea faunas, but corroborate the hypothesis of a pulse of diversification centred in the Oligocene and Miocene epochs. Amphi-Atlantic species were found to occur at deeper depths (bathyal-abyssal) and we hypothesize that trans-Atlantic connectivity is maintained by dispersal between neighbouring reproductive populations inhabiting the abyssal sea floor and by dispersal across the shelf and slope of Arctic and sub-Arctic regions.
authors.affiliation: Marine Biodiversity Research Group Department of Biology University of Bergen 5006 Bergen Norway.
authors.forename: Mari H
authors.initials: MH
authors.lastname: Eilertsen
authors.affiliation: Phylogenetic Systematics and Evolution Research Group Department of Natural History University Museum of Bergen University of Bergen 5020 Bergen Norway.
authors.forename: Manuel António E
authors.initials: MA
authors.lastname: Malaquias
keywords: Allopatric speciation; Cephalaspidea; Heterobranchia; Mollusca; Tethyan vicariance; amphi‐Atlantic; dispersal; diversification pulse; marine biogeography