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Douglas G. Barton O University of Albexta Microstratigraphic variation in presnvational patterns and menstic counts of Amywn aggreganun (Teleostei: Catostomidae) bma 10 000-year intemal of the Eocene vmed lake deposits of Horsefly, British Columbia Douglas G. Barton O A thesis subrnitted to the Faculty of Graduate Snidies and Research in partial fulfdlment of the requirements for the degree of Master of Science Deparmient of Biological Sciences National Library Bibliothèque nationale du Canada Acquisitions and Acquisitions et Bibliographie Services senrices bibliographiques 395 Wellington Street 395. nie Wellington OtÉawaûN K1AW OtrawaON K1AW hnada Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence aiiowing the exclusive permettant a la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distrii.ute or sell reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/nlm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conseme la proprieté du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. ABSTRACT The 10 000-year W inmal of the Eocene Horsefly locality in British Columbia is ideal for rnicrosmtigraphic snidies. The sediments are preserved in yearly laminations, or varves, which ailow smdy of variation in ecological and morphologicai characters with an exceptionally high temporal reçolution. A taphonomic snidy of the fsh supports a lake mode1 for the H3 section in which the bottom waters are perennially anoxic (or hypoxic), but sdace waters ovemim regularly (monomitic or dimictic). Temporal trends in the preservation of fish suggest longer-terni cycles of shaUowing and deepening of the deposit Phenotypic change in the most common species in this interval, Amyzon aggregarwn (Cypriniformes: Catostomidae), is resaicted to meristic elements. Numbers of menstic elements Vary considerably through the interval, but much of this variation can likely be attributed to ecophenotypic effects. Periods of evolutionary stasis are suggested by the lack of phenotypic change which cannot be amibuteci to environmental change. Implications of these results on pdeoenvironrnents, patterns and processes of phenotypic change. and temporal averaging are discussed. ACKNOWLEDGEMENTS Firstly, special thanks an due to my supervisor, Mark V. H. Wilson. who got me started on this project, and who has pmvided support (financial and otherwise) throughout. Thanks are also due to the other membof rny supervisory comminee who have provided helpful cornments on this thesis in various stages of preparation: R C. Fox. J. S. Nelson, and N. Loveîi. Other financial support has been provided by the Department of Biological Sciences in the fom of a Graduate Teaching Assisgntship, and a Graduate Studies and Research Scholarship. Travel to the Society of Venebrate Paleontology annual mee~gin Chicago in 1997 was funded by the J. Gordin Kaplan Graduate Studies Award. I would like to thank an anonymous landowner for access to the Horsefly locaiïty. SpeRai thanks are also due to my field crew of Man Lindoe and Gavin Hanke, for putting up with weeks of standing waist-deep in the frigid Horsefly River, and for their after-work conversations (verbal and otherwise) around the fire. Assistance in the field was also provided by my mom, Jean Barton (despite her difficulties in crossing the river). Allan Lindoe has also provided invaluable assistance in the lab, preparing many of the specimens collected in previous years. and providing me with expert assistance and advice. 1 would also like to thank Ken Puph of the Fraser Center for Non-Marine Eocene Research in British Columbia. Ken provideci assistance while in the field, and 1 have benefited from numerous discussions with him on the Eocene fossils and locdities of British Columbia. Richard HeMa at the Royal British Columbia Museum (RBCM)was very helpful in planning rny field work and acquiring the necessary permissions. Joan Kerik, also at the Royal British Columbia Museum, has been very helpful in keeping me up to date on the specimens that we donated to the RBCM as part of Our coilecting agreement with the anonymous landowner. 1 would also like to thank Steve Cumbaa and Kieran Shepherd at the Museum of Nature (National Museum of Canada - MC)for access to specimens in their collections, and for information on pas collecting nips by the museum. Throughout my thesis, 1 have benefited hmthe support and fnendship of many colleagues and friends too numerous to mention individually. 1 would also like to thank my parents for theû encouragement during this thesis. Finally, a special thanks to Davis Hill, who not only has supported and encourageci me throughout, but has aiso inspired me to work diligentiy and efficiently during the final stages of this thesis. TABLE OF CONTENTS CHAPTER 1: INTRODUCTION . introduction . History of the Site . Geological Age -Y H3 Section . Varves Literature Cited CHAPTER 2: TAPHONOMlC VARlABILITY AN PALEOENVIRONMENT OF THE E Introduction . Taphonorny and Preservational Patterns of Fish . Further Research Approach - Material and Methods . Resul ts Generai Variation in Preservation Discussion . Seasonal Change/Y early Cycle Environmental Change Conclusions . Literature Cited CHAPTER 3: PHENOTYPIC CHANGE IN AMYZON THROUGH THE H3 INTERVAL Introduction . Microevolutionary Studies Meristic Variation . Materials and Methods -ty Methods Analyses and Statistics Results . Discussion . Repeatability . Summary S tatistics Correlations - Temporal Trends Conclusions . Literature Cited CHAPTER 4: CONCLUSIONS Pdeoenvironmentd Reconstructions Phenotypic Change . Concluding Rernarks - Literature Cited LIST OF TABLES Table 3- 1 : Repeatability of meristic counts . Table 3-2: Summary statistics of meristic counts of Amyzon aggregana hmthe H3 interval compared to previous studies of the species Table 3-3: Correlation rnatrix of meristic couna of Amyzon hmthe H3 interval LIST OF FIGURES CHAPTER 1: Figure 1- 1: Map of Horsefly locality . Figure 1-2: Exposure of the Horsefly locality Figure 1-3: Snatigraphic section of the Horsefiy locality - figure 1-4: Reference snatigraphic column for the H3 intemal Figure 1-5: S tylized sûatigraphic section of the H3 interval - Figure 1-6: Frequency distribution of thicknesses of vaxves . Figure 1-7: Temporal distribution of fish in the H3 section . CHAPTER 2: Figure 2-1: Buoyancy responses of fish carcasses - Figure 2-2: Taphonomic processes and preservational patterns associated with different temperatures Figure 2-3: Strarigraphic section of the H3 interval . 38 Figure 2-4: S pecirnens of Amyzon with no disarticulation - . 43 Figure 2-5: Specimens of Amyzon with patterns of disarticulation atmbuted to partial floatation . 44 Figure 2-6: Specimens of Amyzon with patterns of disarticulation amibuted to scavenging . 45 Figure 2-7: Temporal distribution of fish in the H3 section . 47 Figure 2-8: Size-fiequency dismbution of Amyzon from the H3 section . 48 Figure 2-9: Temporal trends in size of Amyzon . 50 Figure 2- 10: Temporal trends of disarticulation over the H3 interval. 5 1 Figure 2- 11 : Temporal trends of disarticulation over a 7 15-year interval . 52 Figure 2- 12: Cornparison of percentage of specimens with skuli disarticulation and intensity of skuîl disarticulation . 63 Figure 2- 13: Relative lake level changes in the X3 section through time. 68 Figure 3-1 : Cornparison of patterns of phenotypic change predicted by phyletic gradualism and punctuated equilibrium Figure 3-2: Effccts of gaps in the fossil record on the appearance of phenotypic change . Figure 3-3: Stratigraphie section of the H3 interval . Figure 3-4: Representative specimen of monaggreganvn from the H3 interval. Figure 3-5: Temporal trends in meristic counts across the H3 in terval Figure 3-6: Principal components analysis of meristic counts of dorsai fin rays, anal fin rays, dorsal vertebrae, and caudal vertebrae Figure 3-7: Principal components analysis of meristic counts of dorsal fin rays, dorsal vertebrae, predorsal vertebrae and total vertebrae . Figure 3-8: Temporal trends in meristic counts across a 715-year in tend Figure 3-9: Relative lake level changes in the H3 section through time - LIST OF ABBREVIATIONS Institutions: RBCM - Royal British Columbia Museum ROM - Royal Ontario Museum UALVP - University of Alberta Laboratory for Vertebrate Paleontology Counts and Measurements: (see p. 98- 100 for explanations): AFR - Anal fin rays AP - Anal pterygiophores BD - Body depth CFR - Total caudal fin rays CV - Caudal vertebrae DFR - Dorsal fin rays DV - Dorsal vertebrae lCFR - Lower caudal fin rays PAV - Post-anal vertebrae PCV - &-caudal vertebrae pDFR - Redorsal fin rays PDV - Predorsd vertebrae PFR - Pectoral fin rays SL - Standard length TV - Total vertebrae uCFR - Upper caudal fin rays VfR - Pelvic fin rays CHAPTER 1: INTRODUCTION Introduction Over the past several decades, paleontologists have become increasingly interested in leamhg about pattems and rates of change in different biological and paleontological features over short time intervals in the fossil record (e.g. Bell et al. 1987; McCune 1987; Carroll 1997). Microstratigraphic snidies,
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