Movement Patterns and Biology of White Sucker in a Riverine
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VARIATION IN EARLY LIFE-HISTORY CHARACTERISTICS OF SYMPATRIC RAINBOW SMELT POPULATIONS IN LAKE UTOPIA, NEW BRUNSWICK by Jennifer Lynn Shaw Bachelor of Environmental Science, University of Guelph 1996 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF Masters of Science In the Graduate Academic Unit of Biology Supervisor: R.A. Curry, Ph.D., UNB, Biology Examining Board: T.J. Benfey, Ph.D., UNB, Biology – Chair R.A. Cunjak, Ph.D., UNB, Biology – Internal Examiner M. Wiber, Ph.D., UNB, Anthropology – External Examiner This thesis is accepted by the Dean of Graduate Studies THE UNIVERSITY OF NEW BRUNSWICK January, 2006 © Jennifer L. Shaw, 2006 ABSTRACT Three sympatric morphotypes of rainbow smelt (Osmerus sp.) have been identified in Lake Utopia, New Brunswick. The ‘giant’ form is larger (20.2 ± 2.8 cm SD), has fewer gill rakers, and spawns earliest before other forms and in different streams. The ‘normal’ form is smaller (13.1 ± 1.9 cm) with increased numbers of gill rakers. The ‘dwarf’ form is the smallest (9.9 ± 0.9 cm) and has the highest gill raker count. Both normal and dwarf smelt spawn in the same streams with normal forms beginning earlier and dwarf forms extending spawning longer. If body size is the keystone biological feature among morphotypes, then differences in egg size or differential growth rates exist at some time during their ontogeny. We tested this prediction by comparing egg size, spawning date, incubation time, hatch size, and growth to determine when a divergence in size occurs. While some characteristics appeared stable, others displayed inter- annual variability. Giant larvae hatched earlier, at a larger size and consistently grew more rapidly as age 0+ fish. Divergence between normal and dwarf forms was less stable, differing between years. The forms hatched at the same size, however timing varied and divergence in growth occurred at age 0+, 1+ or 2+ fish. We suggest that genetic factors are most important for maintaining giant morphotypes and environmental factors, such as lake and stream temperatures are important in regulating the normal and dwarf morphotypes in Lake Utopia today. ii ACKNOWLEDGMENTS I would like to begin by thanking my supervisor, Dr. Allen Curry for giving me this opportunity and for providing guidance, direction and support along the way. Thank you to my supervisory committee, Dr. Stephan Peake and Mr. Steve Currie for providing direction and constructive feedback at various stages of the process. This project would not have been completed without the help of many individuals and organizations. Funding was provided by the New Brunswick Wildlife Trust Fund and Fisheries and Oceans Canada’s, Student Subvention Grant Program. Staff at the St. Andrews Biological Station, Marine Fish Division of Fisheries and Oceans Canada provided equipment and lab support for otolith microstructure procedures. The New Brunswick Department of Environment and Local Government provided water temperature data. Thank you to Marcia Chiasson, Emily Kitts, Chad Doherty, Mark Gautreau, Eric Chernoff and Jonathon Freedman who provided much needed help in the field and in the lab. I would also like to acknowledge and thank the many graduate students, summer students, staff and faculty who provided help, shared ideas and were around to talk when I needed it. Lastly, I would like to thank my mom and dad, Karen and Ken, my sisters, Lori and Michelle and my husband, Eric for always being there for me and supporting me. You are what matters most. iii TABLE OF CONTENTS ABSTRACT……………………………………………………………………………..ii ACKNOWLEDGMENTS ..................................................................................... iii TABLE OF CONTENTS .....................................................................................iv LIST OF FIGURES .............................................................................................vi 1 GENERAL INTRODUCTION ..................................................... 1 1.1 Life-History Variation ................................................................. 1 1.2 Evolutionary Significance of Size and Growth ........................... 3 1.3 Rainbow Smelt Biology.............................................................. 4 1.4 Life-history Variation in Rainbow Smelt ..................................... 5 1.5 Ecology of Morphotypes ............................................................ 6 1.6 Genetics .................................................................................... 8 1.7 Species At Risk Designation...................................................... 9 1.8 Research Objectives and Thesis Outline................................... 9 2 METHODS............................................................................... 11 2.1 Study Area............................................................................... 11 2.2 Early Life-History Characteristics............................................. 12 2.3 Growth ..................................................................................... 14 2.3.1 Field Sampling..........................................................................14 2.3.2 Otolith Microstructure Procedure..............................................15 2.4 Statistical Analysis ................................................................... 18 3 RESULTS ................................................................................ 23 3.1 Early Life-History Characteristics............................................. 23 iv 3.1.1 Spawning, Incubation, and Hatching Period.............................23 3.1.2 Water Temperature ..................................................................24 3.1.3 Egg and Hatch Size..................................................................25 3.2 Growth ..................................................................................... 26 3.2.1 Larval Growth ...........................................................................26 3.2.2 Adult Age and Growth ..............................................................27 3.2.3 Lake Temperature and Growth.................................................28 4 DISCUSSION .......................................................................... 41 4.1 Size Differences in Early Life................................................... 41 4.2 Evolutionary Origin of Morphotypes......................................... 44 4.3 Summary ................................................................................. 48 5 LITERATURE CITED............................................................... 49 CURRICULUM VITAE v LIST OF FIGURES Figure 1: Lake Utopia (45°10’, 66°47’) and its smelt spawning tributaries. The giant form spawns in Mill Lake Stream, Trout Lake Stream and Spear Brook. Normal and dwarf forms spawn in Smelt, Unnamed and Second Brooks. ...............................................................................20 Figure 2: Relationship between otolith growth and somatic growth. Larval smelt captured in Lake Utopia in 2003 ( ▲ ), r2 = 0.93 (n=142) and 2004, r2 = 0.90 ( ○ ) (n=140). Adult giant, normal and dwarf smelt captured by dip-nets during spawning in Mill Lake Stream, Smelt Brook, Second Brook and Unnamed Brook in 1999 ( ■ ), r2 = 0.58 (n=158) and 2003 ( Δ ), r2 = 0.90 (n=108). ...................................................................21 Figure 3: Spawning ( ▄▄ ), incubation ( ▄▄ ) and hatching ( ▄▄ ) period of giant smelt in Mill Lake Stream, and normal and dwarf smelt in Smelt, Unnamed and Second brooks in Lake Utopia, 2004. Hatch marks indicate approximation of dates. ......................................................29 Figure 4: Mean daily water temperature (± 1 SD) of spawning tributaries in Lake Utopia for giant smelt (Mill Lake Stream and Spear Brook) and normal and dwarf smelt (Second Brook, Unnamed Brook and Smelt Brook), 2004.................................................................................................30 Figure 5: Mean egg size (± 1 SD) of giant ( ■ ), normal ( O ) and dwarf ( ▲ ) mature female smelt captured in Trout Lake Stream, Mill Lake Stream, Unnamed Brook, Smelt Brook and Second Brook in 2004 (total sample size n=120). Mean hatching length (± 1 SD) of larval smelt captured in Mill Lake Stream, Second Brook and Smelt Brook in 2004, and Smelt Brook and Unnamed Brook in 2002 (total sample size n=360). .....................................................................................31 Figure 6: Hatching date calculated from otolith daily growth rings of normal and dwarf larvae captured by Neuston trawls in Lake Utopia in summer 2003 (n=138) and 2004 (n=96). No larvae of giant smelt were captured...........................................................................................33 vi Figure 7: Size and age of 0+ larval fish captured by Neuston trawls throughout the summer in 2003 and 2004 in Lake Utopia. ................................35 Figure 8: Back-calculated, mean length-at-age of dwarf ( ▲, -SD ) and normal ( ○ , +SD) larval smelt captured in Lake Utopia on June 25, July 2, July 25, August 26 and September 18, 2003 (n=73), and on June 1, June 28, July 18, July 29 and August 28, 2004 (n=62)....................36 Figure 9: Back-calculated mean length-at-age (± 1 SD) of adult giant ( ■ ), normal ( ○ ) and dwarf ( ▲ ) smelt captured by dip-nets during spawning in Mill Lake Stream, Smelt Brook, Second Brook and Unnamed Brook in 1999 (n=108) and in Mill Lake Stream, Trout Lake Stream, Smelt Brook, Second Brook and Unnamed