Chirocentrids as engrauloids: evidence from suspensorium, branchial arches, and infraorbital bones (Clupeomorpha, Teleostei)

The Chirocentridae is a family of highly specialized large predatory clupeomorphs composed of two species from coastal waters of the Indian and western Pacific Oceans. Peculiarities of the anatomy of these fishes have puzzled ichthyologists who attempted to resolve their phylogenetic relationships. Despite controversy, it is currently accepted that the Chirocentridae is a family of Clupeiformes, included with the Clupeidae in the superfamily Clupeoidea. New data support an alternative hypothesis. Seven previously unreported derived character states from the suspensorium, branchial arches, and infraorbitals strongly indicate a hitherto unsuspected sister group relationship between the Chirocentridae and Engrauloidea, which comprises approximately 140 species of the commercially important fishes known as anchovies. These are character states: (1) the anterior margin of metapterygoid located anterior to the quadrate; (2) the ventral limb of hyomandibula and quadrate not separated by the metapterygoid; (3) the posterodorsal margin of metapterygoid in line with the condyle of articulation of the hyomandibula with the opercle; (4) the presence of a laminar outgrowth of the anterior margin of the quadrate; (5) the endochondral portion of the quadrate in the shape of an isosceles triangle; (6) the presence and arrangement of autogenous tooth plates on ceratobranchials 1 to 3; and (7) posterior region of infraorbital 1 well developed and extending along the ventral margin of infraorbital 2. Three of those character states are further modified and hypothesized as synapomorphies of the Engrauloidea: (1') a substantial portion of the metapterygoid situated anterodorsal to the quadrate, (2') articulation between the ventral limb of the hyomandibula and the quadrate, and (7') infraorbitals 1 and 3 articulating by means of a well-developed laminar process of the posterior region of infraorbital 1. The separation of the dorsal, paired elements of the branchial arches of the Chirocentridae and representative Engrauloidea is apomorphic within the Clupeoidei, and constitutes circumstantial evidence for the sister group relationship between those clades. Microphagy within the Engrauloidea is secondary, homoplastic to the same condition present in other clades of the Clupeiformes. The decomposition of character complexes into discrete morphological characters and its use in phylogenetic inference is discussed. The sister group relationship between the Chirocentridae and Engrauloidea renders the Clupeoidea paraphyletic. A new classification of the Clupeoidei, with the inclusion of the Chirocentridae in the Engrauloidea, is proposed. (C) 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156, 363-383. (AU)