nm u Ottawa L'Universite canadicnne Canada's university run FACULTE DES ETUDES SUPERIEURES l^^l FACULTY OF GRADUATE AND ET POSTOCTORALES u Ottawa POSDOCTORAL STUDIES L'UniversU6 Cdiiddienne Canada's university Holly Rae Stephens TutluFbTLA7r¥sE7Xu™ .M:§£-.i?.i9l91l).. GRADE/DEGREE Department of Biology FACULTE, ECOLE, DEPARTEMENT / FACULTY, SCHOOL, DEPARTMENT Taxonomic Revision of the Flatfish Genera Zebrias Jordan and Snyder, 1900 and Pseudaesopia Chabanaud, 1934 with Notes on Aesopia Kaup, 1858 (Pleuronectiformes: Soleidae) TITRE DE LA THESE / TITLE OF THESIS Francois Chapleau DIRECTEUR (DIRECTRICE) DE LA THESE / THESIS SUPERVISOR CO-DIRECTEUR (CO-DIRECTRICE) DE LA THESE / THESIS CO-SUPERVISOR Guy Drouin Lynn Gillespie Kathleen Conlan Gary W. Slater Le Doyen de la Faculte des etudes superieures et postdoctorales / Dean of the Faculty of Graduate and Postdoctoral Studies Taxonomic revision of the flatfish genera Zebrias Jordan and Snyder, 1900 and Pseudaesopia Chabanaud, 1934 with notes on Aesopia Kaup, 1858 (Pleuronectiformes: Soleidae) Holly R. Stephens Thesis submitted to the Faculty of Graduate and Postdoctoral Studies University of Ottawa In partial fulfillment of the requirements for the M.Sc. degree in the Ottawa-Carleton Institute of Biology These soumise a la Faculte des etudes superieures et postdoctorales, Universite d'Ottawa en vue de l'obtention de la maitrise LTnstitut de biologie d'Ottawa-Carleton © Holly R. Stephens, Ottawa, Canada, 2010 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 Ottawa ON K1A 0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-74168-9 Our file Notre reference ISBN: 978-0-494-74168-9 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extra its substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. 1*1 Canada ABSTRACT The taxonomic status of the 25 nominal species of the Indo-Pacific flatfish genus Zebrias (Pleuronectiformes: Soleidae) was revised based on a morphological study of 531 specimens, including 36 type specimens. Two distinct, monophyletic groups were identified differing in the confluence and attachment of the caudal fin to adjacent dorsal and anal fins. Species most closely associated to the type species Pleuronectes zebra were included in the genus Zebrias while the remaining species were assigned to the genus Pseudaesopia to accommodate Zebrias species having a shallowly confluent caudal fin with adjacent fins, among other characters. Ten species are now recognized and re-described in Zebrias while six species are recognized and re- described in Pseudaesopia. Comprehensive generic definitions and identification keys to species are provided. In addition, the closely related and monospecific genus Aesopia is briefly discussed. Finally, potentially useful characters for a future phylogenetic study are discussed. RESUME Le statut taxinomique des 25 especes nominales du genre indopacifique Zebrias (Pleuronectiformes: Soleidae) a ete examine suite a une etude morphologique de 531 specimens incluant 36 specimens types. Deux groupes monophyletiques ont ete identifies sur la base de la confluence et l'attachement de la nageoire caudale aux derniers rayons des nageoires anale et dorsale. Les especes etroitement associees avec l'espece type, Pleuronectes zebra ont ete placees dans le genre Zebrias. Les autres especes nominales ont ete placees dans le genre Pseudaesopia pour accommoder les especes ayant, entre autres caracteres, une nageoire caudale peu confluente avec les nageoires adjacentes. Dix especes sont reconnues comme valides dans Zebrias alors que six especes sont validees dans Pseudaesopia. Ces especes sont re-decrites. Une definition exhaustive des genres ainsi que des cles d'identification des especes sont presentees. Le statut du genre Aesopia est discute brievement. Finalement, des caracteres potentiellement informatifs au niveau phylogenetique sont discutes. TABLE OF CONTENTS Acknowledgements vi List of Figures ix List of Tables xix Abstract 1 Resume 2 General Introduction 3 Materials and Methods 12 Chapter 1: A revision of the flatfish genus Zebrias with notes on Aesopia (Pleuronectiformes: Soleidae) 20 Abstract 21 Resume 22 Introduction 23 Genus Zebrias 38 Descriptions Zebrias altipinnis 52 Zebrias annularis 59 Zebrias cancellatus 68 Zebrias captivus 77 Zebrias craticula 85 Zebrias maculosus 95 Zebrias penescalaris 100 Zebrias quagga 110 Zebrias scalaris 122 Zebrias zebra 131 Species Key 148 Genus Aesopia 151 References 159 Chapter 2: A taxonomic revision of the flatfish genus Pseudaesopia (Pleuronectiformes: Soleidae) 185 Abstract 186 Resume 187 Introduction 188 Genus Pseudaesopia 196 Descriptions Pseudaesopia callizona 208 Pseudaesopia cochinensis 215 Pseudaesopia japonica 224 Pseudaesopia lucapensis 235 Pseudaesopia regani 242 Pseudaesopia synapturoides 251 Species Key 264 References 266 General Conclusion 272 Phylogenetic Observations 274 Key to Genera 276 Appendix A 280 Appendix B 285 Appendix C 290 ACKNOWLEDGEMENTS First and foremost, I would like to thank my supervisor, Dr. Francois Chapleau. I could not have asked for a better mentor. I thank him for his limitless patience and understanding, as well as his invaluable guidance over the years. His passion for flatfish is infectious and I am grateful to have had the opportunity to work with him. I would also like to extend my sincerest thanks to the Smithsonian Institution in Washington, D.C. for granting me access to their facilities and collections, as well as the assistance I received during my visits. I am in deep gratitude to Dr. Thomas Munroe for his generous assistance throughout my thesis. His guidance and wealth of knowledge were of critical importance to my thesis and were greatly appreciated. Tom, it always brings a smile to my face when I recall your funny anecdotes. I would like to thank Martha for her wonderful hospitality; I had a wonderful time in Virginia with her and Tom and I will miss our breakfast excursions on Thursdays. I thank Cheryl Ames for leaving me with such wonderful memories of my time in Washington, particularly the Halloween spent in Old Virginia. I would also like to thank Sandra Raredon for training me in the use of x-ray equipment. To the wonderful people in the Ichthyology Department at the Australian Museum, my exceptional experience in Australia was largely attributable to the wonderful staff I had the pleasure of working alongside. I thank Mark McGrouther, Amanda Hay, Michelle Yerman, Jeff Leis, John Paxton, Sally Reader, and Sascha Schulz for making me feel so welcome, and for all their help. I'd like to extend a special thank you to Amanda and James for their assistance in taking radiographs of my specimens. In addition, I would like to thank my advisory committee members, Dr. Kathleen Conlan, Dr. Lynn Gillespie, and Dr. Keith Seifert. Their guidance, feedback, and expert advice were exceptionally helpful to me in structuring and focusing my study. There are no words to describe how thankful I am for the endless support and encouragement of my friends and family. I am eternally grateful to my labmates, Rene Duplain, Krystal Lapierre, Josiane Vachon, our "unofficial labmate" Eric Young, and Matti Emery. I consider myself very lucky to have had the opportunity to work alongside such wonderful friends who were always available to assist me with my work or share in a laugh. Some of my best memories of this thesis are of the time we spent together. I would like to thank my mom, Charis, my father, Larry, and my twin sister, Carrie, for always believing in me, even at times when I doubted myself. The pride and encouragement they regularly expressed helped me continue to push forward and I will forever be grateful for that. I am also incredibly grateful to Monique Gaudreault for her invaluable support and assistance in making it through some of the more challenging aspects of my degree. I am indebted to the institutions that provided the material