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Morphology of the clasper musculature in rays (Chondrichthyes; Elasmobranchii: Batoidea), with comments on their phylogenetic interrelationships

The subclass Batoidea comprise skates, electric rays, stingrays, guitarfishes, and sawfishes, and their interrelationships are still problematical despite recent morphological and molecular phylogenetic studies. The most recent morphological phylogeny indicates that guitarfishes are a polyphyletic group, and that the phylogenetic placement of Platyrhina and Platyrhinoidis is still unclear. Several molecular studies suggest that guitarfishes (except Zanobatus) and sawfishes comprise the monophyletic order Rhinopristiformes, and that thornback rays (Platyrhinidae, Platyrhina, and Platyrhinoidis) are more closely related to the electric rays (Torpediniformes); rhinopristiforms have recently been supported by morphological data as well. The clasper musculature of batoids suggests an alternative pattern of interrelationships for thornback rays, with the m. dilatator attached to the dorsal terminal 1 cartilage by a series of tendons found only in Rhinopristiformes and Platyrhinidae, suggesting that they are closely related. Furthermore, Rajiformes, Rhinopristiformes, and Platyrhinidae exclusively share a reduced m. extensor lateralis, suggesting that these taxa form a monophyletic group. This study identifies new synapomorphies that corroborate the separate monophyly of Rajiformes, Torpediniformes, and Myliobatiformes: the m. dilatator divided into dorsal and ventral bundles and the presence of a single m. flexor are found only in Rajiformes; the extensor lateralis with a laminar shape is a derived character of Torpediniformes; and the exclusive m. flexor medialis arising on the puboischiadic bar is derived for Myliobatiformes.