First multilocus and densely sampled timetree of trevallies, pompanos and allies (Carangoidei, Percomorpha) suggests a Cretaceous origin and Eocene radiation of a major clade of piscivores Francesco Santini ⇑ , Giorgio Carnevale Dipartimento di Scienze della Terra, Università degli Studi di Torino, Torino 10125, Italy article info Article history: Received 21 August 2014 Revised 21 October 2014 Accepted 23 October 2014 Available online 1 November 2014 Keywords: Carangidae Echeneidae Coryphaenidae Rachycentridae Coral reefs Molecular clock abstract Carangoid fishes (trevallies, pompanos, jacks, dolphinfishes, cobias and remoras) include about 159 spe- cies of marine fishes found in tropical and temperate waters worldwide (Froese and Pauly, 2014). Many carangoids are powerful swimmers and active piscivores in and around coral-reef ecosystems. Some carangoid lineages, such as dolphinfishes, have evolved a pelagic lifestyle, while remoras spend their adult life attached to cetaceans, sharks, manta rays and large teleosts, feeding off skin parasites or left- overs from their host’s meals. In spite of their taxonomic diversity, ecological dominance, economic importance to humans, and a rich fossil record dating to the Paleogene, relatively little is currently known about the tempo of evolution of this group. Here we present the results of the first time calibration study of carangoid fishes. Using a fossil-calibrated molecular timetree that includes 133 species of carangoids (85% of extant species), we show that this group originated in the Late Cretaceous and that several major lineages were already present before the K-Pg extinction. All major clades were in existence by the end of the Eocene, even though significant diversification has continued to occur throughout the his- tory of this group. Ó 2014 Elsevier Inc. All rights reserved. 1. Introduction Carangoid fishes represent a highly successful group of 159 species (Nelson, 2006; Froese and Pauly, 2014) that includes tre- vallies, pompanos, jacks and allies (family Carangidae, 148 spe- cies), remoras (Echeneidae, eight species), dolphinfishes (Coryphaenidae, two species) and the cobia (Rachycentridae, one species). All species in this group are primarily or exclusively mar- ine, even though several species of carangids are known to enter brackish waters, especially as juveniles (Froese and Pauly, 2014). While carangids are predominantly coastal fishes, found in tropical to temperate water worldwides, both coryphaenids and the cobia are found in open waters, and several species of remoras live as commensals on large pelagic vertebrates, such as sharks, turtles, billfishes and whales. Carangoids show a great range of body size, varying from less than 20 cm in total length in Alepes kleinii to the 2 m of Lichia amia or Coryphaena. This group is also very variable in shape, with deep-bodied forms such as Parastromateus niger, as well as more streamlined, oblong taxa, such as the species of Seri- ola and Trachurus. The remoras are well known for the dramatic transformation of the spiny dorsal fin, which in adults is modified into a sucking disk that allows adhesion to a host species (O’Toole, 2002; Britz and Johnson, 2012). Many species of carangids feed predominantly on zooplankton, while remoras are opportunist feeders that can eat ectoparasites or shedding skin off their hosts, as well as scraps of food from the meals of their symbiont, in addition to plankton. The majority of carangids, however, as well as the two species of dolphinfishes and the cobia, are active predators of both large invertebrates, such as cephalopods, and other teleost fishes. Carangoids are generally powerful swimmers, and are highly sought after by sportfishermen worldwide; they are also very important for both artisanal fisheries in many tropical countries, as well as for larger scale fisheries in European and American waters (Smith-Vaniz, 1999). In spite of their economic importance to humans and ecological role as top predators in many coastal ecosystems worldwide, cur- rently very little is known about the evolutionary history of this group. Carangid phylogenetic relationships have been investigated in a number of comparative morphological studies focusing on both adult and larval characters (Suzuki, 1962; Smith-Vaniz and Staiger, 1973; Smith-Vaniz, 1984, 1986, 1999; Gushiken, 1988; Hilton and Johnson, 2007; Springer and Smith-Vaniz, 2008; http://dx.doi.org/10.1016/j.ympev.2014.10.018 1055-7903/Ó 2014 Elsevier Inc. All rights reserved. ⇑ Corresponding author at: Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, Torino 10125, Italy. E-mail address: francesco.santini@alumni.utoronto.ca (F. Santini). Molecular Phylogenetics and Evolution 83 (2015) 33–39 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev