Mitochondrial gene variation in Mercenaria clam sibling species reveals a relict secondary contact zone in the western Gulf of Mexico

We investigated phylogeographic relationships among American Mercenaria taxa by assessing variation in a 444 nucleotide fragment of the mitochondrial 16S ribosomal gene in clams sampled from four representative sites in January to November 1994. Three of these sites were in the Gulf of Mexico, one was on the Atlantic coast in South Carolina. Direct sequencing of this amplified gene fragment in 85 individuals revealed 21 haplotypes. Phylogenetic analyses consistently resolved this variation into three well supported clades, and within-clade genetic divergence levels were markedly lower than among-clade values. One of the clades, A, was taxon-specific, in that it solely and exclusively contained specimens of M. mercenaria (Linnaeus, 1758) sampled in South Carolina. The other two clades, B and C, were the most divergent and both encompassed specimens of M. campechiensis (Gmelin, 1791) and of M. campechiensis texana (Dall, 1902), sampled from the three Gulf of Mexico sites. Clade B was found at high frequencies at all three Gulf sites, whereas Clade C occurred at low frequencies at two western Gulf sites. We interpret this pattern as resulting from the secondary contact and introgression of two allopatrically differentiated Mercenaria taxa in the western Gulf of Mexico. Clade C haplotypes may represent relict mitochondrial lineages from original Gulf Mercenaria spp. populations that predate massive mitochondrial introgression by M. campechiensis. We further propose that the M. campechiensis texana nuclear genome is a mosaic, heavily weighted toward M. campechiensis, but containing some relict alleles inherited from the precontact population, especially those governing shell characteristics, which may be adaptive in cohesive sediments of bays and estuaries in the northwestern Gulf of Mexico.