Phylogenetic relationships, genetic diversity and biogeography of menhadens, genus Brevoortia (Clupeiformes, Clupeidae).

Brevoortia Gill 1861 is a genus of the Clupeidae (Teleostei) that includes six species of fishes commonly known as menhadens in eastern North America and "savelhas" or "saracas" in southeastern South America. Species of Brevoortia are important components of the marine food web of coastal ecosystems in the Atlantic and contribute significantly to fisheries. In this study, the first phylogenetic and biogeographic hypotheses including all species of Brevoortia are presented. A total of 113 specimens were analyzed using three molecular markers (two mitochondrial: COI and 16s; and one nuclear: RAG2). Maximum Parsimony, Maximum Likelihood and Bayesian Inference were employed to estimate phylogenetic relationships. A Bayesian multispecies coalescent approach was used to estimate a dated phylogeny, which supported biogeographic analyses of ancestral geographic ranges. Results corroborate previous hypotheses that the four North Atlantic species are grouped in two clades, one composed of B. tyrannus (Latrobe, 1802) and B. patronus Goode, 1878, and the second including B. smithi Hildebrand, 1941 and B. gunteri Hildebrand, 1948. The South Atlantic B. aurea (Spix and Agassiz, 1829) and B. pectinata (Jenyns, 1842) form a third clade, which is sister to the clade composed of B. smithi and B. gunteri. The monophyly and validity of the six nominal species of Brevoortia were not supported. Results also indicate that Brevoortia originated in the North Atlantic during the middle Miocene (about 15 Mya). A cooling event of the tropical Atlantic at around 10 Mya likely facilitated the range expansion of the genus to the South Atlantic, whereas a significant warming of the tropical Atlantic waters during the late Miocene at 6-7 Mya possibly promoted the isolation between the northern and southern counterparts of that ancestral lineage. The relevance of the Florida Peninsula in association with sea level fluctuations for the diversification within Brevoortia is also discussed.