Evolutionary affinity of billfishes (Xiphiidae and Istiophoridae) and flatfishes (Plueronectiformes): Independent and trans-subordinal origins of endothermy in teleost fishes

Billfishes (Scombroidei) and tunas (Scombridae), both considered part of the suborder Scombroidei, have long been studied by biologists largely because of their remarkable physiological and anatomical muscular adaptations associated with regional endothermy and continuous swimming. These attributes, combined with analyses of other morphological and molecular data, have led to a general perception that tunas and billfishes are close relatives, though this hypothesis has been vigorously debated. Using Bayesian phylogenetic analysis of nine mitochondrial and three nuclear loci (>7000 bp), we show that billfishes are only distantly related to tunas, but rather share strong evolutionary affinities with flatfishes (Pleuronectiformes) and jacks (Carangidae). This phylogenetic relationship is striking because of the marked variation in phenotype and niche across these trans-ordinal groups of fishes. Billfishes and flatfishes have each evolved radically divergent morphological and physiological features: elongated bills and extraocular heater organs in billfishes, and cranial asymmetry with complete eye migration during ontogenetic development in flatfishes. Despite this divergence, we identify synapomorphies consistent with the hypothesis of a common billfish/flatfish/jack ancestor.