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Fig. 2. (A) Cretaceous–Pleistocene (C–Pl) time scale showing relative cooling in benthic ocean temperature based on stable oxygen isotope ratios. Warmest times were late Paleocene (Pa), while coolest were Pleistocene. Numbers in circles mark the approximate timing of five major paleoceanographic events: (1) Central American land bridge formation (3 mya); (2) beginning of thermohaline circulation (22 mya); (3) closure of the Tethys seaway (25 mya); (4) Antarctic glaciation and polar cooling (30 mya); (5) Northward tectonic drift of Australia (36 mya). Data summarized from Macdougall (1996); Fordyce and de Muizon (2001); Graham and Dickson (2004). Other Tertiary epochs abbreviated on the left axis are: Eocene, E; Oligocene, O; Miocene, M; Pliocene, P. (B) Phylogeny for the Sardini and Thunnini based on the earliest known occurrences of fossil remains (orange circles) of two extant Sardini genera (G, Gymnosarda; S, Sarda) and extant tuna genera (A, Auxis; E/K, Euthynnus or Katsuwonus; T, Thunnus). (Note that Thunnus fossils extend from the late Paleocene to the Pliocene.) The extinct Paleothunnus (P) is considered to be closely related to the tuna–bonito common ancestor. Data from Bannikov (1985); Carroll (1988); Monsch (2000; personal communication). Bonito genera abbreviations: S and G, as above; C, Cybiosarda; O, Orcynopsis. Tuna abbreviations as in Fig. 1; the two Thunnus subgenera (Neothunnus and Thunnus), each with four species, are indicated. (Note: terminal lines indicate number of extant species in each tuna genus.)





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