Taxonomy and Ecology of the Deep-Pelagic Fish Family Melamphaidae, with Emphasis on Interactions with a Mid- Ocean Ridge System
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TAXONOMY AND ECOLOGY OF THE DEEP-PELAGIC FISH FAMILY MELAMPHAIDAE, WITH EMPHASIS ON INTERACTIONS WITH A MID- OCEAN RIDGE SYSTEM by Kyle Allen Bartow A Dissertation Submitted to the Faculty of The Charles E. Schmidt College of Science in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Florida Atlantic University Boca Raton, FL December 2010 Copyright by Kyle Bartow 2010 ii ACKNOWLEDGEMENTS The research in this dissertation is due to funding from Tracey Sutton from the U.S. National Science Foundation Ocean Sciences Division – Biological Oceanography Program (OCE 0623568). Funding for travel and tuition were received for various parts of this project from Florida Atlantic University and Virginia Institute of Marine Science. I am grateful to the crew of the RV G.O. Sars, MAR-ECO and the staff of the Bergen Museum for the collection and curation of samples. I would also like to thank the U.S. National Museum of Natural History - Division of Fishes for allowing me into their enormous collection of melamphaid fishes and being so flexible and accommodating during the largest blizzard I've ever been in. The final entity that I would like to thank is MAR-ECO, through whose association I have been afforded many of these opportunities. I would like to thank Tracey Sutton for initially believing in me and my interest in deep-sea fish and research. Tracey not only offered me a place to fulfill my goals, but offered much support and encouragement during many of my trials and tribulations. Thanks are also due to the members of my advisory committee: Edie Widder, Jon Moore, C. Edward Proffitt and Brian Lapointe. Their edits and input into my project allowed it to mature into the final product that it is today. At Florida Atlantic University I would like to thank Geri Mayer, Jenny Govender and Becky Dixon. Without their help, it would have been impossible to get through the everyday challenges of being a graduate student and to find the answers to all my school related questions and issues. iv Finally, I would like to thank all the members of my family. My parents Allen and Kozie Bartow, Ginny Hustvedt and Dan Oberholzer not only for their love and support but also showing me that what I am doing is both important and interesting. The Knight family (Tim, Kathleen, Kristin, Kaitlin, Jared and Jordan) deserves many thanks as well. They were not only my friends, but like a second family to me growing up and played a major role in the person I am today. I also thank my wife, Jen, deserves a lot of my gratitude, not only for all the love she's shown, but for dealing with my weirdness, meltdowns and the extreme amount of input and advice she has given to this project. And I thank our friends Allyson and Johnnie Schumacher, Sara Fitzgerald and Joey Johnson for keeping me grounded, sane and keeping my life balanced and preventing grad student burnout. ABSTRACT Author: Kyle Bartow Title: Taxonomy and Ecology of the Deep-Pelagic Fish Family Melamphaidae, with Emphasis on Interactions with a Mid-Ocean Ridge System Institution: Florida Atlantic University Dissertation Advisor: Dr. Tracey Sutton Degree: Doctor of Philosophy Year: 2010 Much of the world’s oceans lie below a depth of 200 meters, but very little is known about the creatures that inhabit these deep-sea environments. The deep-sea fish family Melamphaidae (Stephanoberyciformes) is one such example of an understudied group of fishes. Samples from the MAR-ECO (www.mar-eco.no) project represent one of the largest melamphaid collections, providing an ideal opportunity to gain information on this important, but understudied, family of fishes. The key to the family presented here is the first updated, comprehensive key since those produced by Ebeling and Weed (1963) and Keene (1987). Samples from the 2004 MAR-ECO cruise and the U.S. National Museum of Natural History provided an opportunity to review two possible new species, the Scopelogadus mizolepis subspecies, and a Poromitra crassiceps species complex. Results show that Scopeloberyx americanus and Melamphaes indicoides are new species, while the two subspecies of Scopelogadus mizolepis are most likely only one species and the Poromitra crassiceps complex is actually several different species of Poromitra. vi Data collected from the MAR-ECO cruise provided an opportunity to study the distribution, reproductive characteristics and trophic ecology of the family Melamphaidae along the Mid-Atlantic Ridge (MAR). Cluster analysis showed that there are five distinct groups of melamphaid fishes along the MAR. This analysis also supported the initial observation that the melamphaid assemblage changes between the northern and southern edges of an anti-cyclonic anomaly that could be indicative of a warm-core ring. Analysis of the reproductive characteristics of the melamphaid assemblage revealed that many of the female fishes have a high gonadosomatic index (GSI) consistent with values found for other species of deep-sea fishes during their spawning seasons. This may indicate that melamphaids use this ridge as a spawning ground. Diets of the melamphaid fishes were composed primarily of ostracods, amphipods, copepods and euphausiids. Scopelogadus was the only genus shown to have a high percent of gelatinous prey in their digestive system, while Melamphaes had the highest concentration of chaetognaths. This work presents data on the ecology and taxonomy of the family Melamphaidae and provides a strong base for any future work on this biomass-dominant family of fishes. vii TAXONOMY AND ECOLOGY OF THE DEEP-PELAGIC FISH FAMILY MELAMPHAIDAE, WITH EMPHASIS ON INTERACTIONS WITH A MID- OCEAN RIDGE SYSTEM LIST OF TABLES .......................................................................................................... x LIST OF FIGURES ....................................................................................................... xiii CHAPTER 1. INTRODUCTION .................................................................................. 1 OVERVIEW ............................................................................... 1 THE MID-ATLANTIC RIDGE ................................................. 1 MAR-ECO .................................................................................. 6 THE FAMILY MELAMPHAIDAE ........................................... 7 Melamphaid Taxonomy ................................................... 8 Feeding Ecology ............................................................. 8 Reproduction and Growth............................................... 11 OBJECTIVES ............................................................................. 12 CHAPTER 2. KEY TO THE SPECIES OF MELAMPHAIDAE ................................. 15 INTRODUCTION ...................................................................... 15 MATERIALS AND METHODS ................................................ 15 KEY TO THE GENERA AND SPECIES .................................. 17 CHAPTER 3. REVIEW OF MELAMPHAID TAXONOMY AND SYSTEMATICS ..................................................................................... 28 INTRODUCTION ...................................................................... 28 MATERIALS AND METHODS ................................................ 29 Museum Samples ............................................................. 29 Species Examined............................................................ 29 Species Identification ...................................................... 30 Counts and Measures ...................................................... 31 RESULTS AND DISCUSSION ................................................. 35 CONCLUSIONS......................................................................... 65 CHAPTER 4. DISTRIBUTION OF THE DEEP-PELAGIC BIGSCALE FISHES (TELEOSTEI: MELAMPHAIDAE) ALONG THE NORTHERN MID-ATLANTIC RIDGE ............................................... 66 viii ABSTRACT ................................................................................ 66 INTRODUCTION ...................................................................... 67 MATERIALS AND METHODS ................................................ 70 RESULTS ................................................................................... 75 DISCUSSION ............................................................................. 85 CHAPTER 5. GROWTH AND REPRODUCTION OF THE FAMILY MELAMPHAIDAE ................................................................................ 92 INTRODUCTION ...................................................................... 92 MATERIALS AND METHODS ................................................ 93 Dissection ........................................................................ 95 Statistical Analyses ......................................................... 97 RESULTS ................................................................................... 100 Size and Growth .............................................................. 100 Reproduction ................................................................... 100 DISCUSSION ............................................................................. 112 CHAPTER 6. TROPHIC ECOLOGY OF THE MELAMPHAID FISHES .................. 115 INTRODUCTION ...................................................................... 115 MATERIALS AND METHODS ................................................ 117 Gelatinous Zooplankton Collection ................................ 118 Dissection .......................................................................