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Osteology and Systematic Position of the Eocene Salmonid Eosalmo

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Osteology and Systematic Position of the Eocene Salmonid Eosalmo ZoologicalJournal ofthe Linnean Sociep (1999), 125: 279-31 I. With 9 figures Article ID: zjls.1997.0166, available online at http://www.idealibrary.com on 10 Ehl Osteology and systematic position of the Eocene salmonid TEosalmo dr@woodensis Wilson &om western North America MARK V.H. WILSON1*AND GUO-QING LI1" I Laboratory for Ertebrate Paleontology, Department of Biological Sciences, Universig of Alberta, Edmonton, Alberta T6G 2E9, Canada Institute of Vertebrate Paleontology and Paleoanthropology, Academia Sinica, rl 0. Box 643, Beging 100044, China Received May 1996; acceptedfor Publication April 1997 The fossil salmonid tEosulmo dnzoodensis was originally described from fragmentary speci- mens. Study of new material of this fossil species confirms that it is a stem-group salmonine, with a mixture of primitive and derived salmonine features in its skull, but with its postcranial skeleton essentially of modern salmonine construction. Two autapomophies define the genus TEosalrno: a long anterodorsal process of the subopercle meeting the dorsal edge of the bone at an angle of about 60°, and a thin dermal basihyal plate apparently lacking teeth. Its salmonine relationship is supported by eight derived features: (1) posterior part of frontal widely expanded above autosphenotic, (2) hyomandibular fossa on pterotic long, (3) posterior part of endopterygoid extending posteriorly and broadly overlapped by both metapterygoid and quadrate, (4)premaxillary process of maxilla extending dorsally at an angle larger than loo, (5)infraorbitals 3 to 5 narrow and covering less than anterior half of hyomandibula, (6) presence of suprapreopercle, (7) anterior end of preopercular canal on horizontal arm distinctly turning to anteroventral corner of preopercle, (8) first uroneural amplified into large fan-shaped stegural, and (9) scales small, with more than two lateral line scales per vertebral centrum. Salmonidae are a monophyletic family defined by at least three synapomorphies: posterior surface of epiotic with sulcus, peg-and-socket connection in caudal skeleton, and tetraploid karyotype. Within the Salmonidae, Thymallinae and Salmoninae form a clade based on features from premaxilla, supramaxilla, anguloarticular, and supraorbital. 0 1999 The Linnean Society of London ADDITIONAL KEY WORDS:-Eocene ~ tEosulmo ~ North America ~ osteology phylogeny - stem-group salmonine. CONTENTS Introduction ....................... 280 Methods and material ................... 281 Methods ...................... 281 Materials ...................... 282 Abbreviations ..................... 282 * Corresponding author: E-mail: [email protected] 279 0024-4082/99/030279 + 33 $30.00/0 0 1999 The Linnean Society of London 280 M. V. H. WILSON AND C.-Q. IJ Systematic description ................... 283 Genus TEosalmo Wilson, 1977 ......... ...... 283 TEosalmo dn@ooodensis Wilson, 1977 ........ ...... 2 84 Description ..................... 2 84 Discussion ................. ...... 2 99 New evidence on the ostcology of tEosalmo ........... 303 Autapomorphies of tEosalmo .......... ...... 303 Synapomorphies of Salmonidae ............... 304 Interrelationships among three subfamilies of Salmonidae ...... 304 Phylogeny ofthe subfamily Salmoninae ...... ...... 305 Conclusions ................. ...... 307 Acknowledgements .............. ...... 307 References ....................... 307 Appendix .................. ...... 310 INTRODUC‘I’ION Salmonids are euteleostean fishes that have high value in commercial and sport fisheries (Wu & Wu, 1992; Nelson, 1994). They have long been interesting to ichthyologists not only because of their great economic significance but also because of their problematic phylogenetic relationships (see Sanford, 1990), their ecological adaptations including anadromous habits (Stearley, 1992), and their geographic distributions. Numerous systematic studies have been done on the family since 1960, including treatments of morphology (e.g. Norden, 196 1 ; Vladykov, 1962, 1963; Kolyushev, 197 1; Shaposhnikova, 1968, 197 1, 1975; Gorshkov et al., 1979; Cavender, 1980; Behnke, 1980; Williams, 1987; HolGk et al., 1988; Smith & Stearley, 1989; Sanford, 1990; Stearley, 1990; Stearley & Smith, 1993; Burnham-Curtis & Smith, 1994), karyology (e.g. Booke, 1968; Hartley, 1987; Phillips, Pleyte & Ihssen, 1989; Phillips, Van Ert & Pleyte, 1989), allozymes (e.g. Tsuyuki & Roberts, 1966; Utter, Allendorf & Hodgins, 1973; Johnson, 1980), ontogeny (e.g. Balon, 1980; Pavlov, 1980; Kendall & Behnke, 1984), and DNA sequences (e.g. Berg & Ferris, 1984; Skurikhina, Tugarina & Mednikov, 1986; Thomas, Withler & Beckenbach, 1986; Grewe, Billington, & Hebert, 1990; Partti-Pellinen et al., 1991; Phillips & Pleyte, 1991). However, these studies have not yet removed all the uncertainties about the phylogeny of this group, particularly those related to its origin and early evolution. Wilson (1974, 1977) described what is still regarded as the oldest known fossil salmonid, in the genus TEosalmo, based on specimens from Eocene lacustrine rocks at a locality near Smithers, British Columbia, Canada. f.Eosalmo has since been found in the Eocene near Princeton, British Columbia (Wilson, 199610) and near Republic, Washington, USA (Wilson, 1996a). This fossil salmonid was originally thought to be morphologically intermediate between extant thymallines and sal- monines (Wilson, 1977). Recently, Stearley & Smith (1993) reconstructed the phylogeny of Salmonidae based on a cladistic analyses of 1 19 characters, presenting a cladogram suggesting that TEosalmo is the sister group to the Salmoninae. This suggestion, though generally in agreement with the ideas of Wilson (1977), was vulnerable to criticism because 41 of the 1 19 characters used in Stearley and Smith’s analyses were not available (i.e. ‘missing data’) for TEosalmo. This paper summarizes new information on the osteology of TEosalmo based on a review of the previously described specimens together with new materials from western North America. It also documents several synapomorphies for the genus OSTEOLOGY AND PHYLOGENY OF tEOSAW.IO 28 1 fEosalmo, and uses these data to place the fossil species within the phylogeny of the Salmonidae. METHODS AND MATERIAL Methods Terms used for general description follow Wilson (1977), Rojo (199 l), and Stearley & Smith (1993). For caudal skeleton, terms mainly follow Arratia (199 l), and Arratia & Schultze (1992). Numerical phylogenetics programs (PAUP 3.1.1, Swofford, 1993; and MacClade 3.04, Maddison & Maddison, 1992) were used in carrying out cladistic analyses of character states for reconstruction of the phylogeny. Characters used for cladistic analysis are mainly those from the skeletal system that are observable in both fossil and Recent specimens. All of the characters are unordered and equally weighted in our analyses. Each of the states of a given character is placed in square brackets following the number representing the character (e.g. 10[1] is character state 1 of character 10). Missing data are coded by question marks in the data matrix. There remains considerable controversy over the question of the phylogenetic position of the Salmonidae within the Teleostei. Williams (1 987) presented evidence favouring a sister group relationship of salmonids with esocoids; however, Johnson and Patterson (1 996) reviewed recent studies and advocated a sister group relationship of salmonoids (coregonids + salmonids) with osmeroids (including galaxioids); salmonoids + osmeroids in turn were held to be the sister group of argentinoids. Wilson and Williams (1992) and Johnson and Patterson (1996) agree that Hypomesus is the most primitive extant osmerid. In this paper, we have used data for Hypomesus, the esocoid Novumbra, and the argentinoid Algentina as out-group taxa to root the cladogram and thereby establish the polarity of character-state transformations within the Salmonidae. In addition to tEosalmo dnjhooodensis, five other fossil species have been included in the subfamily Salmoninae. They are (1) tRhabdofario lacustris (Cope, 1870) from the Miocene to Pliocene of Idaho and Oregon (see Kimmel, 1975; Smith, 1975, 1981; Smith et al., 1982), (2) tSmilodonichthys rastrosus (Cavender & Miller, 1972) from the Miocene to Pliocene of Oregon and California (also see Barnes, McLeod & Raschke, 1985), (3) tSalmo cyniclope (La Rivers, 1964) from the Miocene of Nevada, (4) tSalmo australis (Cavender & Miller, 1982) from the Pliocene of southwestern Mexico, and (5) a fossil from eastern Asia referred by Sytchevskaya (1986) to tEosalmo. Too little is known about the fifth fossil for inclusion in this study. The other four fossil species were recently referred to the genus Oncorhynchus (see Kendall, 1988; Stearley & Smith, 1993; Nelson, 1994). Therefore, the generic names f Rhabdofario and tSmilodonichthys are considered to be synonymous with Oncorhynchus. Comparative Recent fishes consist of dried skeletons, specimens fixed in formalin and preserved in alcohol or cleared and stained with alizarin. Data for measurements and counts made following Hubbs and Lagler (1964) are from the more complete fossil specimens and from the Recent fishes. Taxa marked with daggers ‘7’ preceding their names are exclusively known as fossils. 282 M. V. H. WILSON AND G.-Q LI Materials In addition to those tEosalmo specimens listed in the systematic section, the following materials were used in this study for comparison and for checking information in the literature. Alizarin prepared extant specimens: Novumbra hubbsi, specimens
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