TAXONOMIC AND PHYLOGENETIC REVISION OF THE FAMILY CHIASMODONTIDAE (PERCIFORMES: ACANTHOMORPHA) Except where reference is made to the work of others, the work described in this dissertation is my own or was done in collaboration with my advisory committee. This dissertation does not include proprietary or classified information. _____________________________ Marcelo R. S. de Melo Certificate of Approval: _____________________________ ____________________________ Kenneth M. Halanych Jonathan W. Armbruster, Chair Professor Professor Biological Sciences Biological Sciences _____________________________ _____________________________ Craig Guyer Larry M. Page Professor Curator of Fishes Biological Sciences Florida Museum of Natural History University of Florida _____________________________ George T. Flowers Dean Graduate School Auburn University TAXONOMIC AND PHYLOGENETIC REVISION OF THE FAMILY CHIASMODONTIDAE (PERCIFORMES: ACANTHOMORPHA) Marcelo R. S. de Melo A Dissertation Submitted to the Graduate Faculty of Auburn University in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Auburn, Alabama August 10, 2009 TAXONOMIC AND PHYLOGENETIC REVISION OF THE FAMILY CHIASMODONTIDAE (PERCIFORMES: ACANTHOMORPHA) Marcelo R. S. de Melo Permission is granted to Auburn University to make copies of this dissertation at its discretion, upon request of individuals or institutions and at their expense. The author reserves all publication rights. _____________________________ Signature of Author _____________________________ Date of Graduation iii VITA Marcelo Roberto Souto de melo, son of Abraão Cavalcanti de Bessa and Amélia Souto de Melo, was born in Brasília, Brazil, on July 9, 1976. He received a Bachelor of Science in Biology in 1999 from Universidade Federal de Goiás (Brazil). In 2001, Marcelo earned a Master in Science degree in Biology at the Brazilian Museu Nacional, Universidade Federal do Rio de Janeiro. In 2004 he was awarded with a fellowship from the Brazilian government to join the Graduate School at Auburn University. Marcelo finished his degree requirements for a Doctor of Philosophy in Biological Sciences in August 10, 2009. iv TAXONOMIC AND PHYLOGENETIC REVISION OF THE FAMILY CHIASMODONTIDAE (PERCIFORMES: ACANTHOMORPHA) Marcelo R. S. de Melo Doctor of Philosophy, August 10, 2009 (M.Sc. Universidade Federal do Rio de Janeiro, 2001) (B. S. Universidade Federal de Goiás, 1999) 563 Typed Pages Directed by Jonathan W. Armbruster The family Chiasmodontidae, commonly known as swallowers, comprises four genera and 33 species distributed in the meso and bathypelagic regions of the Atlantic, Pacific, Indian and Southern Oceans. The species of Chiasmodon and Pseudoscopelus have disjunct distribution, and those of Dysalotus and Kali, wide distributions. The genus Chiasmodon is composed of seven species, two of them new: C. niger; C. subniger; C. braueri; C. microcephalus; C. pluriradiatus; C. asper n. sp.; and C. harteli n. sp. Pseudoscopelus has 16 species, five of them new: P. scriptus; P. sagamianus; P. altipinnis; P. cephalus; P. obtusifrons; P. scutatus; P. aphos; P. parini; P. astronesthidens; P. australis; P. pierbartus; P. bothrorrhinos n. sp.; P. lavenbergi n. sp.; P. paxtoni n. sp.; P. cordilluminatus, n. sp.; and P. odontoglossum sp. n. Dysalotus has only three species, one new: D. alcocki; D. oligoscolus; and D. acanthobrychos. Kali has v seven species, two of them new: K. indica; K. kerberti; K. macrodon; K. macrura; K. parri; K. colubrina, n. sp.; and K. falx, n. sp. The chiasmodontids have several morphologic adaptations related to life in the deep-sea and swallowing of enlarged preys. Those characteristics include the reduction of some bones and muscles, diverse dentition, enlarged mouth, a very distensible stomach and body walls, photophores, and the presence of superficial neuromasts on head and body. One of the most remarkable characteristics is the innumerous superficial neuromasts, which are present in all chiasmodontids. The superficial neuromasts are more concentrated in the head, but are also present on body along the trunk lateral-line canal, on upper and lower lobes of the caudal fin, and dorsal part, anterior and to the first- dorsal fin. The distribution of neuromasts is described, as well as their innervation. A phylogenetic analysis was made based on 161 morphological characters obtained from osteology, myology, lateral line system, dentition and miscellaneous. The monophyly of the Chiasmodontidae is corroborated by 24 synapomorphies. Within the chiasmodontids, two major clades are formed: the Chiasmodon clade is composed of Chiasmodon and Pseudoscopelus, and supported by 18 synapomorphies; and the Kali clade is composed by Dysalotus and Kali, and supported by 22 synapomorphies. The monophyly of the genus Chiasmodon is supported by 16 synapomorphies; Pseudoscopelus, by 12 synapomorphies; Dysalotus, by 18 synapomorphies; and Kali, by 23 synapomorphies. vi ACKNOWLEDGMENTS The author would like to thank to: Jonathan W. Armbruster, for his guidance and suggestions throughout during my studies. The committee members, Kenneth M. Halanych, Dave Johnson, Craig Guyer, and Larry M. Page for their critical review. The Armbruster lab and family for their friendship during the past five years: Shobnon Ferdous; Lesley de Souza; Keith Ray; Nathan Lujan; Ricardo Betancur; David Werneke; and Brian, Will, and Heather Armbruster. John Paxton (Australian Museum) and Karsten Hartel (Museum of Comparative Zoology – Harvard University) for their support and suggestions during this study. Cindy Keplado and H.J. Walker (University of California in San Diego) for their nicely collaboration on a publication. Auburn University staff, especially Jack Feminella, Mary Mendonça, Sandra Riddle and, in name of the International Education Office crew, Ken McNabb. Peter Raucci, from the Institute of International Education. Financial support was provided by a graduate studies fellowship from the Brazilian governmental agency CAPES, (process BEX 2030/03–9); Schultz fund of the Division of Fishes (USNM), and Ernst Mayr Grants (Harvard University). This work is dedicated to my mother, Amélia Melo, with love. vii Style manual or journal used: Zootaxa. Computer software used: Microsoft Word 2003 (text), Microsoft Excel 2003 (tables), Corel Draw Graphics Suite 12 (figures), Adobe Photoshop CS2 (figures), JMP version 5 (statistic analysis); PAUP* version 4.0b10 (maximum parsimony reconstructions, genetic distance calculations, hypothesis testing), MacClade 4 (dataset handling, ancestral character reconstructions), Mesquite 2.6 (ancestral character reconstructions), TreeRot version 2b2 (produce commands for performing Decay analysis). viii TABLE OF CONTENTS LIST OF TABLES........................................................................................................... xii LIST OF FIGURES .......................................................................................................... xv CHAPTER 1 – INTRODUCTION TO THE FAMILY CHIASMODONTIDAE...............1 General Introduction ........................................................................................................2 Objectives ........................................................................................................................5 References........................................................................................................................7 CHAPTER 2 – REVISION OF THE GENUS CHIASMODON JOHNSON ...................11 Abstract..........................................................................................................................12 Introduction....................................................................................................................12 Material and Methods ....................................................................................................14 List of abbreviations ................................................................................................. 16 Results............................................................................................................................17 Key to the genera of Chiasmodontidae..................................................................... 17 Key to the species of Chiasmodon............................................................................ 19 Chiasmodon Johnson 1864 ....................................................................................... 21 Chiasmodon niger Johnson 1864.............................................................................. 27 Chiasmodon subniger Garman 1899 ........................................................................ 31 Chiasmodon braueri Weber 1913............................................................................. 35 Chiasmodon microcephalus Norman 1929............................................................... 38 Chiasmodon pluriradiatus Parr 1933........................................................................ 41 Chiasmodon asper, new species ............................................................................... 44 Chiasmodon harteli, new species ............................................................................. 47 Discussion......................................................................................................................52 Identification of juveniles ......................................................................................... 53 Unidentified specimen.