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Phylogenetic Relationships Within the South American Fish Family

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Phylogenetic Relationships Within the South American Fish Family Zoological Journal of the Linnean Society, 2008, 154, 70–210. With 53 figures Phylogenetic relationships within the South American fish family Anostomidae (Teleostei, Ostariophysi, Characiformes) BRIAN L. SIDLAUSKAS1–4* and RICHARD P. VARI FLS4 1Committee on Evolutionary Biology & University of Chicago, 1025 E. 57th St., Culver Hall 402, Chicago, IL 60637, USA 2Division of Fishes, The Field Museum, 1400 S. Lake Shore Drive, Chicago, IL 60605, USA 3National Evolutionary Synthesis Center, 2024 W. Main St. A200, Durham, NC 27705, USA 4Department of Vertebrate Zoology, MRC-159, National Museum of Natural History, PO Box 37012, Smithsonian Institution, Washington, DC 20013-7012, USA Received 17 September 2007; accepted for publication 17 September 2007 Analysis of a morphological dataset containing 152 parsimony-informative characters yielded the first phylogenetic reconstruction spanning the South American characiform family Anostomidae. The reconstruction included 46 ingroup species representing all anostomid genera and subgenera. Outgroup comparisons included members of the sister group to the Anostomidae (the Chilodontidae) as well as members of the families Curimatidae, Characidae, Citharinidae, Distichodontidae, Hemiodontidae, Parodontidae and Prochilodontidae. The results supported a clade containing Anostomus, Gnathodolus, Pseudanos, Sartor and Synaptolaemus (the subfamily Anostominae sensu Winterbottom) albeit with a somewhat different set of relationships among the species within these genera. Anostomus as previously recognized was found to be paraphyletic and is split herein into two monophyletic components, a restricted Anostomus and the new genus Petulanos gen. nov., described herein. Laemolyta appeared as sister to the clade containing Anostomus, Gnathodolus, Petulanos, Pseudanos, Sartor and Synapto- laemus. Rhytiodus and Schizodon together formed a well-supported clade that was, in turn, sister to the clade containing Anostomus, Gnathodolus, Laemolyta, Petulanos, Pseudanos, Sartor and Synaptolaemus. Anostomoides was sister to the clade formed by these nine genera. Leporinus as currently defined was not found to be monophyletic, although certain clades within that genus were supported, including the species with subterminal mouths in the former subgenus Hypomasticus which we recognize herein as a genus. Abramites nested in Leporinus, and Leporellus was found to be the most basal anostomid genus. The presence of cis- and trans-Andean species in Abramites, Leporellus, Leporinus and Schizodon, all relatively basal genera, suggests that much of the diversification of anostomid species pre-dates the uplift of the Andean Cordilleras circa 11.8 million years ago. Several important morphological shifts in anostomid evolution are illustrated and discussed, including instances of convergence and reversal. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 70–210. ADDITIONAL KEYWORDS: Anostominae – biogeography – cladistics – headstanders – morphology – Neotropics – new genus – osteology – South America. INTRODUCTION western Colombia south to the central portions of Argentina. The greatest diversity at both the specific Members of the Anostomidae inhabit myriad fresh- and the generic levels occurs east of the Andes Cor- water environments in tropical and subtropical South dilleras where representatives of the family inhabit America that range from the Río Atrato of north- all major drainages from the Río Orinoco basin (Lasso et al., 2004b) to the Río de La Plata system (Garavello *Corresponding author. E-mail: [email protected] & Britski, 2003). The vast reaches of the Amazon and 70 No claim to original US Government works. Journal compilation © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 70–210 PHYLOGENY OF ANOSTOMID FISHES 71 Figure 1. Rhytiodus microlepis, USNM 229095, 121.5 mm Figure 3. Synaptolaemus cingulatus, AMNH 91095, SL; Brazil, Amazonas, Ilha da Marchantaria; lateral 78 mm SL; Venezuela, Bolivar, Río Paraguay, at Carapo view. Rapids, just upriver of mouth of Río Carapo; lateral view. Figure 4. Leporinus friderici, FMNH 116829, 180.5 mm Figure 2. Abramites hypselonotus, MCZ 59240, 98.6 mm SL; Suriname, Rechter Coppename River; lateral view. SL; Ecuador, Río Punino, above Coca; lateral view. Sartor (Myers & Carvalho, 1959: figs 1–3; Sidlauskas the adjoining Río Orinoco basin are inhabited by the 2007: fig. 2D). Although some of the highly modified largest number of genera and species, including some genera such as Sartor and Synaptolaemus (Fig. 3) of the most modified members of the family. The have limited geographical distributions, small body Atlantic versant of the Guayana shield in Guyana, sizes (c. 0.1 m standard length (SL)) and are appar- Suriname and French Guiana also harbours many ently rare in nature based on museum holdings, other species (Planquette, Keith & Le Bail, 1996), while genera [e.g. Anostomoides, Laemolyta, Leporinus a less diverse fauna inhabits the hydrographically (Fig. 4), Rhytiodus and Schizodon] are geographically highly-dissected trans-Andean portions of South widespread and have large body sizes up to approxi- America. West of the Andean Cordilleras, anostomids mately 0.3 m SL. The larger species are commonly are present in the arch from Lago Maracaibo of the exploited in commercial and artesanal food fisheries Caribbean Sea versant as far south as the Guayas (Goulding, 1980, 1981). Many anostomids feed largely basin of Ecuador, which drains into the Pacific Ocean. on aquatic vegetation (Goulding, Leal Carvalho & Although few species inhabit the trans-Andean Ferreira, 1988); other species have more diverse regions, most that do occur there are endemic to very diets including larval and adult insects, nematodes, small regions. For example, Abramites eques and Lep- sponges, larval fishes and detritus (Knöeppel, 1972; orinus muyscorum are known only from trans-Andean Winterbottom, 1974b; Saul, 1975; Taphorn, 1992; Colombia (Steindachner, 1900, 1902; Vari & Williams, Santos & Rosa, 1998; Balassa et al., 2004). 1987) and Schizodon corti only from Lago Maracaibo The first anostomids known to western science were in Venezuela (Vari & Raredon, 1991). described by Linnaeus (1758) who based his species Overall body form in the Anostomidae ranges from descriptions on specimens previously reported on by the elongate, fusiform species of Rhytiodus (Fig. 1; Gronovius (1756). Günther (1864) first united various Géry, 1977: 180; Santos, 1980: fig. 12) to the deep- anostomid genera in his ‘Anostomina’, but the next bodied, laterally compressed species of Abramites century brought little agreement about relationships (Fig. 2; Vari & Williams, 1987: fig. 3). Most anosto- of what is now recognized as the Anostomidae to other mids, however, are moderately elongate species that taxa. Boulenger’s Anostominae (1904) significantly are rounded in cross-section. Despite their similarity expanded the Anostomina of Günther (1864) to in overall body form, members of the Anostomidae include a number of taxa now assigned to other fami- demonstrate a wide range of jaw, tooth, neurocranium lies (see summary in Vari, 1998), but none of those and suspensorium morphologies (Sidlauskas, 2007). taxa is currently considered to be closely related to Mouth positions range from distinctly subterminal the Anostomidae. Regan (1911) proposed a family within Hypomasticus, particularly in Hypomasticus Anostomidae that encompassed the Anostomidae, mormyrops (Steindachner, 1875: pl. 6) and Hypomas- Chilodontidae, Curimatidae and Prochilodontidae of ticus pachycheilus (Britski, 1976; Sidlauskas, 2007: present concepts, an insightful grouping under fig. 2B) to supraterminal and posteriorly directed in current hypotheses of relationships (Vari, 1983). No claim to original US Government works. Journal compilation © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 70–210 72 B. L. SIDLAUSKAS and R. P. VARI Gregory & Conrad (1938) recognized a broader Anos- sively. The partial revisionary studies cited above tomidae consisting of Regan’s (1911) family of that have yielded many previously undescribed species, name plus taxa now included by Langeani (1998) in and numerous questions remain as to which of the the Hemiodontidae (sensu Greenwood et al., 1966). approximately 140 nominal species should be recog- Géry (1961a: 108), reiterating the concept of Regan nized and how they can be discriminated (Garavello (1911), united the present Anostomidae (his Anos- & Britski, 2003). These issues and others preclude an tominae) with the Chilodontidae, Prochilodontidae attempt to develop a species-level phylogeny in the and Curimatidae (his Chilodinae, Prochilodontinae Anostomidae at this time. and Curimatinae), without providing explicit evidence This study had four goals: in support of that grouping. Roberts (1973: 221) 1. to propose a hypothesis of higher level phyloge- hypothesized an explicit relationship between the netic relationships within the Anostomidae; Anostomidae and Prochilodontidae. Subsequently, 2. to define the nominal genera in the Anostomidae Vari (1983: 50) detailed numerous synapomorphies in on the basis of derived features; support of the hypothesis that the Anostomidae was 3. to determine the number of times that major tran- the sister group to the Chilodontidae rather than the sitions in jaw orientation have evolved within the Prochilodontidae as proposed by Roberts (1973). Anostomidae; and Hypotheses of intrafamilial phylogenetic
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