Geographic clines and stepping-stone patterns detected along the East Pacific Rise in the vetigastropod Lepetodrilus elevatus reflect species crypticism

Three different molecular markers (i.e. seven allozyme loci, two nuclear gene loci and, mtCOI DNA sequences) were used to assess the genetic structure of the vent gastropod Lepetodrilus elevatus collected from three vent fields along the East Pacific Rise (13°N, 9°50′N and 17°S). While allozymes and nuclear loci suggested a strong stepping-stone pattern, a multivariate analysis performed on allozymic frequencies showed the presence of two distinct evolutionary lineages: the first situated in the north from 13°N to 9°50′N and the second in the south from 9°50′N to 17°S. The analysis of mitochondrial DNA sequences confirmed the separation of L. elevatus into two distinct clades with a divergence of 6.5%, which is consistent with the interspecific level of sequence variation in other vent species. A divergence time of 6–14 Mya was estimated between the two clades from previous clock calibrations. Our results suggest that these taxa followed an allopatric speciation between the northern and southern parts of the EPR with a recent demographic expansion of the southern clade to the north and a subsequent secondary contact (clade hybridisation). This speciation was probably reinforced by a habitat specialisation of the two cryptic species because the southern clade was mainly found associated with mussel-dominated communities and the northern clade with tubeworm-dominated communities. However, the analysis of shell morphology failed to separate the two cryptic species based on this sole criterion although they differed from Lepetodrilus elevatus galriftensis (Galapagos population) by a higher shell elevation. Within each clade, genetic differentiation was not related to the distance across populations and could be within vent field as important as between fields. While both clades appear to be in expansion since their speciation, significant excesses in heterozygotes suggest a very recent and local bottleneck at 17°S, probably due to massive site extinction in this region.