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BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Development and variation of the Suspensorium of primitive Catfishes (Teleostei: Ostariophysi) and their phylogenetic relationships by G. ARRATIA BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 32 1992 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at BONNER ZOOLOGISCHE MONOGRAPHIEN Die Serie wird vom Zoologischen Forschungsinstitut und Museum Alexander Koenig herausgegeben und bringt Originalarbeiten, die für eine Unterbringung in den „Bonner zoologischen Beiträgen" zu lang sind und eine Veröffenthchung als Monographie rechtfertigen. Anfragen bezüglich der Vorlage von Manuskripten sind an die Schriftleitung zu richten; Bestellungen und Tauschangebote bitte an die Bibliothek des Instituts. This series of monographs, published by the Zoological Research Institute and Museum Alexander Koenig, has been estabhshed for original contributions too long for inclu- sion in „Bonner zoologische Beiträge". Correspondence concerning manuscripts for publication should be addressed to the editor. Purchase orders and requests for exchange please address to the library of the institute. L'Institut de Recherches Zoologiques et Museum Alexander Koenig a etabh cette serie de monographies pour pouvoir publier des travaux zoologiques trop longs pour etre inclus dans les „Bonner zoologische Beiträge". Toute correspondance concernante des manuscrits pour cette serie doit etre adressee ä I'editeur. Commandes et demandes pour echanges adresser ä la bibliotheque de I'insti- tut, s. V. p. BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 32, 1992 Preis: 35,— DM Schriftleitung/Editor: G. Rheinwald Zoologisches Forschungsinstitut und Museum Alexander Koenig Adenauerallee 150—164, D-5300 Bonn 1, Germany Druck: Rheinischer Landwirtschafts-Verlag G.m.b.H., 5300 Bonn 1 ISBN 3-925382-34-8 ISSN 0302-671X © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Development and variation of the Suspensorium of primitive Catfishes (Teleostei: Ostariophysi) and their phylogenetic relationships by G. ARRATIA BONNER ZOOLOGISCHE MONOGRAPHIEN, Nr. 32 1992 Herausgeber: ZOOLOGISCHES FORSCHUNGSINSTITUT UND MUSEUM ALEXANDER KOENIG BONN © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at CIP-Kurztitelaufnahme der Deutschen Bibliothek Arratia, G.: Development and variation of the Suspensorium of primitive Catfishes (Tele- ostei: Ostariophysi) and their phylogenetic relationships / by G. Arratia. Hrsg.: Zoolog. Forschungsinstitut und Museum Alexander Koenig, Bonn. — Bonn: Zoolog. Forschungsinst. und Museum Alexander Koenig, 1992. (Bonner zoologische Monographien ; Nr. 32) ISBN 3-925382-34-8 NE: GT © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at CONTENTS Page Introduction 5 Methods 6 General methodology 6 Cladistic methodology 6 Character determination 7 Outgroup comparison 7 Materials examined 7 Terminology 11 Autopalatine 12 Pterygoid bones 12 Metapterygoid 12 Entopterygoid 12 Ectopterygoid 13 Dermo + metapterygoid 14 Ectopterygoid + subpalatine toothplate 14 Sesamoid pterygoids 14 'Entopterygoid' type 1 14 'Entopterygoid' type 2 14 'Entopterygoid' type 3 15 'Entopterygoid' type 4 15 'Entopterygoid' type 5 15 'Entopterygoid' type 6 16 'Entopterygoid' type 7 16 'Ectopterygoid' type 1 16 Additional pterygoids 16 Pterygoid type 1 16 Origin and ossification of bones of Suspensorium 16 Suspensorium of ostariophysans other than catfishes 17 Gonorynchiforms 17 Cypriniforms 26 Characiforms 28 Gymnotoids 31 Suspensorium of catfishes 33 Siluroids with a simple quadrate 37 Diplomystids 37 tHypsidorids 47 Ictalurids 48 Nematogenyids 55 Ariids 58 Siluroids with a complex quadrate 58 Ictalurids 58 'Pimelodids' 61 'Bagrids' 66 Schilbeids 66 Trichomycterids 68 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Page Comparison among certain siluriforms 68 Autopalatine 69 Subautopalatine toothplate 72 Antorbital 73 Metapterygoid 74 Dermo + metapterygoid 77 Quadrate-metapterygoid-maxillary ligament 77 Quadrate and symplectic 78 Hyomandibula 79 Dermal pterygoids 82 Sesamoid 'entopterygoid' 83 Sesamoid 'ectopterygoid' 84 Opercle and preopercle 85 Comparison among ostariophysans and their relationships 86 First cladistic analysis 105 Second cladistic analysis 107 Relationships among primitive catfishes 108 First cladistic analysis 108 Results 126 Second cladistic analysis 128 Results 128 Overall Results 128 Conclusions 130 Early ontogeny 130 Homology of chondral bones of the Suspensorium 130 Homology of entopterygoid and 'entopterygoid' 131 Homology of ectopterygoid and 'ectopterygoid' 132 Homology of certain hgaments 132 Diversity of bones of the Suspensorium 132 Variation 133 The Suspensorium of ostariophysans 133 Higher categories of Siluriformes 134 Siluriformes 134 Diplomystoidei 134 Siluroidei 135 tHypsidoroidea 135 Siluroidea 135 Acknowledgements 135 Abstract 136 Zusammenfassung 138 Resumen 139 Literature cited 140 Appendices 148 Appendix 1 148 Appendix 2 149 © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 5 INTRODUCTION Catfishes are represented by over 2000 species assigned to about 32 families, largely distributed throughout South America, Africa, and Asia. Catfishes have traditionally been considered to exempHfy a generalized morphology despite the tremendous varia- bility of some structures between groups (Regan 1911, Alexander 1965, Howes 1983a, 1985, Arratia 1987a, 1990a, b). Phylogenetic investigations of most siluroid families have not been attempted, probably because our knowledge of this geographically widespread and morphologically diverse group is poor. Recently, during the study of the family Diplomystidae (Arratia 1987a), I was faced with the problem of the identification of the pterygoid bones in diplomystids. I found that in some young Diplomystes camposensis an additional bone is present between the hyomandibula, quadrate, and metapterygoid (Arratia 1987a: 54, 98, Fig. 25A). Diplo- mystidae lack the bones traditionally recognized as the ectopterygoid (or pterygoid), and entopterygoid (endopterygoid or mesopterygoid); and yet some specimens may have one or two small bones. However, these do not occupy the position of the ecto- and entopterygoids in other teleosts (Arratia 1987a: 25, 40, 54, 98, 99, Figs. 6, 16, 25 A—D; Arratia & Schultze 1991: Fig. 36). As a consequence of this particular problem in diplomystids, I decided to study ontogenetic series of other catfishes to check whether the currently recognized metapterygoid, ectopterygoid, and entopterygoid of catfishes are homologous within catfishes and homologous with those of other teleosts. I present here a detailed study of the ontogeny of the palatoquadrate and associated dermal and tendon bone pterygoids, dorsal part of the hyoid arch (hyo-symplectic), and their morphological relationships in primitive catfishes that have a small, triangular simple quadrate such as diplomystids (Diplomystidae), Ictalurus (Ictaluridae), and Nematogenys (Nematogenyidae). Further, I will compare these ontogenies to those of primitive catfishes that have a complex shaped quadrate such as Noturus (Ictaluridae) and Parapimelodus ("Pimelodidae"). However, before we can examine these ontogenies I will consider the ontogenetic studies of the Suspensorium of primitive ostariophysans. Homologization of bones will follow two criteria: 1) embryonic origin and ontogeny of the bones and 2) shape, position and relationship of bones following Remane (1952); then these results will be tested in a phylogenetic context following Wiley (1981) and Ax (1987) to determine homologous and non-homologous characters. My usage of homo- logy and non-homology follows Ax (1987: 152): "Homologous features are features in two or more evolutionary species which go back to one and the same feature of a com- mon stem species. They may have been taken over from the stem species unchanged or else with evolutionary transformation." "Non-homologous features in two or more evolutionary species are features which were not present in the common stem species; they were evolved independent to each other." © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 6 METHODS General methodology Most specimens were cleared and double stained by the author for both cartilage and bone, following the procedure described in Schultze & Arratia (1986) and Arratia & Schultze (in press). The length of the specimens refers to the standard length, both in the text and figure captions; and the measurement was taken before clearing and staining. Figures were prepared by the author using a Wild M5A stereo-dissecting microscope equipped with polarized hght and camera lucida. Young specimens were examined with high resolution Olympus and Leitz compound microscopes, equipped with phase con- trast and polarized light. Figures showing the lateral view of the left Suspensorium ex- actly portray the position of the bones in situ and in addition, the hyomandibula main- tains its precise relationship with the neurocranium. The figures were prepared while the
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