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      External Morphology of Lophiosilurus alexandri Steindachner, 1876 during Early Stages of Development, and Its Implications for the Evolution of Pseudopimelodidae (Siluriformes)

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          Abstract

          Pseudopimelodidae are Neotropical catfishes characterized by having slightly to strongly depressed body in fully developed specimens. The largest species of the family with 500 mm SL, Lophiosilurus alexandri, experiences impressive changes in body shape during development, becoming extremely depressed when fully developed. Accordingly, Lophiosilurus alexandri is an ideal species to observe the morphological changes during ontogeny, and to seek solid interpretations on the polarity of characters. Specimens of distinct larval periods (yolk sac, flexion and postflexion; n = 186 specimens) and juvenile stages (n = 20) were analyzed. Changes in body shape, position of mouth and eye, morphology of fins and pigmentation were observed during the development of Lophiosilurus. Larvae (5.7–11.2 mm standard length) had pigmentation concentrated on the head and parts of body, eyes small and pigmented, short barbels, and well-developed finfold. Juveniles (15.9–28.1 mm standard length) had body shape similar to adult, with head depressed and bearing bony ridges, large mouth, dorsally-oriented eyes, small barbels and well-developed shoulder bulges (cleithral width). The greatest morphological changes in the development of L. alexandri occurred during the postflexion larval stage. Relative to standard length, measurements of snout length, head depth and body depth are smaller in juveniles than in larvae, but body width is larger. New interpretations on the phylogenetic characters related to these changes are provided in view of the two alternative hypotheses of the evolution of Pseudopimelodidae.

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          Most cited references 3

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          A phylogenetic analysis of the major groups of catfishes (Teleostei: Siluriformes) using rag1 and rag2 nuclear gene sequences.

          Higher-level relationships among catfishes were investigated by parsimony, maximum likelihood and Bayesian analyses of two nuclear genes across 110 catfish species representing 36 of 37 families and Conorhynchos (family incertae sedis). Analysis of 3660 aligned base pairs from the rag1 and rag2 genes confirms monophyly of Siluriformes, of most siluriform families and of a number of multifamily groups, some recognized, some novel. South American Loricarioidei are recovered as the sistergroup to other catfishes which are divided into Diplomystidae and Siluroidei. This result contrasts with the prevailing hypothesis that Diplomystidae is the sister to all other catfishes. Monophyly of Siluroidei is supported by rag data including a unique three-codon deletion from rag1. Deep within Siluroidei are 12 large, strongly supported groups with poorly resolved interrelationships. Five are single families: Cetopsidae, Plotosidae, Chacidae, Siluridae and Pangasiidae. Four others are monophyletic taxa ranked here as superfamilies: Clarioidea (Clariidae, Heteropneustidae), Arioidea (Ariidae, Anchariidae), Pimelodoidea (Pimelodidae, Pseudopimelodidae, Heptapteridae, Conorhynchos), Ictaluroidea (Ictaluridae, Cranoglanididae). South American Doradoidea (Doradidae, Auchenipteridae) and Aspredinidae are a sistergroup pair. Sisoroidea (without Aspredinidae), Ailia+Laides, Horabagridae, and Bagridae (without Rita) form a large, predominantly Asian clade, "Big Asia." Mochokidae, Malapteruridae, Amphiliidae, Claroteidae, and African schilbids are united as a species-rich African clade, "Big Africa." The three large continental clades, "Big Asia," "Big Africa" and Neotropical Loricarioidei suggest a prevalence of intracontinental diversification of catfishes. South America is the home of the Gymnotiformes, putative sistergroup of catfishes, plus two of the deepest siluriform clades, Loricarioidei and Diplomystidae, thus suggesting an ancient siluriform presence if not origin there. The rag phylogeny does not identify any African-South American catfish clade. The well-known African-Asian relationships within families Clariidae and Bagridae are confirmed, as is the recently found North American-Asian relationship between Ictaluridae and Cranoglanididae.
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            Phylogeny and jaw ontogeny of beloniform fishes.

            To investigate jaw evolution in beloniform fishes, we reconstructed the phylogeny of 54 species using fragments of two nuclear (RAG2 and Tmo-4C4) and two mitochondrial (cytochrome b and 16S rRNA) genes. Our total molecular evidence topology refutes the monophyly of needlefishes (Belonidae) and halfbeaks (Hemiramphidae), but supports the monophyly of flyingfishes (Exocoetidae) and sauries (Scomberesocidae). Flyingfishes are nested within halfbeaks, and sauries are nested within needlefishes. Optimization of jaw characters on the tree reveals a diverse array of evolutionary changes in ontogeny. During their development, needlefishes pass through a "halfbeak" stage that closely resembles the adult condition in the hemiramphid halfbeaks. The reconstruction of jaw transitions falsifies the hypothesis that halfbeaks are paedomorphic derivatives of needlefishes. Instead, halfbeaks make up a basal paraphyletic grade within beloniforms, and the needlefish jaw morphology is relatively derived. The parallel between needlefish ontogeny and beloniform phylogeny is discussed, and clades amenable to future morphological analysis are proposed.
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              Morphology of the gas bladder in bumblebee catfishes (Siluriformes, Pseudopimelodidae).

              The gross morphology of the gas bladder is described and compared for representatives of all valid genera of Pseudopimelodidae (Siluriformes). Cephalosilurus albomarginatus and species of Batrochoglanis, and Microglanis have the most basic form: a large, cordiform gas bladder with a simple internal T-shaped septum. Cephalosilurus apurensis, C. fowleri, and C. nigricauda also have a large, cordiform gas bladder, but they have well-developed trabeculae associated with the internal T-shaped septum, and a pair of well-developed constrictor muscles inserted on the external wall; the latter feature is present in most species of Pimelodidae, but absent in all other catfishes. The monotypic Lophiosilurus alexandri also has well-developed constrictor muscles, and its gas bladder is moderately sized. The species of Pseudopimelodus and Cruciglanis have a diminutive gas bladder partially divided into two lateral sacs without internal communication, and lack constrictor muscles. The parapophysis of the fourth vertebra is a wide and long shelf connected to the dorsal surface of the gas bladder in most pseudopimelodid genera. However, in the species of Pseudopimelodus and Cruciglanis the parapophysis of the fourth vertebra is shorter and has its anterior ramus folded back, partially covering the gas bladder anteroventrally; and the tympanic opening is smaller than in species of the other genera. Five phylogenetic characters are proposed based on the morphology of the gas bladder and associated structures in species of Pseudopimelodidae, and the evolution of those characters in the family is discussed. J. Morphol., 2011. © 2011 Wiley-Liss, Inc. Copyright © 2011 Wiley-Liss, Inc.
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                Author and article information

                Affiliations
                [1 ]Programa de Pós-Graduação em Ciências Biológicas, Departamento de Biologia Animal e Vegetal, Centro de Ciências Biológicas, Universidade Estadual de Londrina, 86051–980, Londrina, PR, Brazil
                [2 ]Programa de Pós-Graduação em Ecologia de Ambientes Aquáticos Continentais, Departamento de Biologia, Universidade Estadual de Maringá, Maringá, PR, Brazil
                James Cook University, AUSTRALIA
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: FMA OAS. Performed the experiments: FMA OAS. Analyzed the data: FMA JLOB AB OAS. Contributed reagents/materials/analysis tools: FMA AB OAS. Wrote the paper: FMA JLOB AB OAS. Drawings: OAS. Photo: FMA.

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                15 April 2016
                2016
                : 11
                : 4
                27082430 4833337 10.1371/journal.pone.0153123 PONE-D-15-34845
                © 2016 Assega et al

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

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                Figures: 8, Tables: 1, Pages: 20
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                Funding
                Funded by: Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (Capes)
                Award Recipient :
                The work was supported by CNPq (to AB), Fundação Araucária (to OAS and JLOB) and fellowship grants from CAPES (to FMA). This is part of author FMA’s Msc thesis, developed under the Programa de Pós-Graduação em Ciências Biológicas. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
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