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Sexual Selection, Fertilization Dynamics and the Use of Alternative Mating Tactics in the Hermaphroditic Seabass Serranus Subligarius Mia Susan Adreani

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Sexual Selection, Fertilization Dynamics and the Use of Alternative Mating Tactics in the Hermaphroditic Seabass Serranus Subligarius Mia Susan Adreani Florida State University Libraries Electronic Theses, Treatises and Dissertations The Graduate School 2011 Sexual Selection, Fertilization Dynamics and the Use of Alternative Mating Tactics in the Hermaphroditic Seabass Serranus Subligarius Mia Susan Adreani Follow this and additional works at the FSU Digital Library. For more information, please contact [email protected] THE FLORIDA STATE UNIVERSITY COLLEGE OF ARTS AND SCIENCES SEXUAL SELECTION, FERTILIZATION DYNAMICS AND THE USE OF ALTERNATIVE MATING TACTICS IN THE HERMAPHRODITIC SEABASS SERRANUS SUBLIGARIUS By MIA S. ADREANI A dissertation submitted to the Department of Biological Science in partial fulfillment of the requirements for the degree of Doctor of Philosophy Degree Awarded: Spring Semester, 2011 The members of the committee approve the dissertation of Mia Adreani defended on October 25, 2010. ______________________________ Don Levitan Professor Directing Dissertation ______________________________ Joseph Travis Professor Directing Dissertation ______________________________ Zuoxin Wang University Representative ______________________________ David Houle Committee Member ______________________________ Christopher Koenig Committee Member Approved: _________________________________________ P. Bryant Chase, Chair, Dept of Biological Science The Graduate School has verified and approved the above-named committee members. ii ACKNOWLEDGEMENTS There are many people who have contributed, directly and indirectly, to this work, and without them, this would not have been possible. First, I thank my advisors, Joe Travis and Don Levitan, whose intelligence, guidance and humor got me through these last several years. I thank my committee members: Chris Koenig, David Houle and Zuoxin Wang for their input and patience. Many people have helped me in the field and I am forever grateful for their assistance: Shannon Chaplin, Andy David, Mark Endries, Nikki Fogarty, Meghan Kirk, Clemens Lakner, Megan Lowenberg, Katie McGhee, Chris Stallings, Brian Storz. In addition, I had a great deal of lab assistance: Shannon Chaplin, Dave Ferrell, Casey Grace, Nate Jue, Meagan Kirk, Alex Marsh, Andres Plata-Stapper, Matt Schrader. I also had a wonderful group of colleagues with whom I could discuss this research, both formally and informally: Denise Akob, Janna Fierst, Nikki Fogarty, Becca Hale, Phil Hastings, Pierson Hill, Nate Jue, Katie Lotterhos, Megan Lowenberg, Katie McGhee, Andres Plata- Stapper, Matt Schrader, Barb Shoplock, Anna Strimaitas, Casey terHorst, Bob Warner, as well as the many years of EERDG participants. This work was partially funded a Brenda Weems Bennison grant, PADI grant, and NSF grants to both Joe Travis and Don Levitan. Site support was generously provided by∗ the NOAA marine fisheries lab in Panama City, FL, St. Andrews State Park, Panama City, FL and the FSU Coastal and Marine Laboratory in St. Teresa, FL. Finally, I want to thank my family for their continued love and support. iii TABLE OF CONTENTS List of tables v List of figures vi Abstract viii 1. INTRODUCTION 1 2. DO SIZE STRUCTURE, HABITAT AND EARLY DEVELOPMENT 14 INFLUENCE THE USE OF ALTERNATIVE MATING TACTICS? 3. THE EFFECT OF POPULATION SIZE STRUCTURE AND ALTERNATIVE MATING TACTICS ON THE FERTILIZATION SUCCESS OF A HERMAPHRODITIC SEABASS 37 4. CONCLUSIONS 57 APPENDIX – ACUC PROTOCOL APPROVAL 61 REFERENCES 62 BIOGRAPHICAL SKETCH 77 iv LIST OF TABLES Table 2.1: Summary of (a) ANOVA using the total number of spawns with season and site as factors and temperature as a covariate and (b) ANOVA using the proportion of streak spawns with season and site as factors and sea-surface temperature as a covariate. 26 Table 2.2: Percentage of histologically analyzed gonads of S. subligarius at each of five stages of development: immature, mature testis only, mature inactive, mature active, and mature postspawning. The samples analyzed histologically represent a subsample of the sample size used for GSI and age analysis. For each month, the range of gonadosomatic indices (GSI = 100*(gonad mass/body mass)) is shown in percent. 27 v LIST OF FIGURES Figure 1.1: Comparisons of the Drosophila data of Bateman’s (1948) paper describing the conditions under which variation in mating may differ among males and females leading to sexual selection on traits that confer greater fitness (solid lines, modified from Bateman, 1948) to a simultaneous hermaphrodite in which selection is stronger on the male role (arbitrary data-dashed line). The gain curves are steeper for hermaphrodites and more similar for males and females than the gonochoristic flies. The gray dotted line indicates the conditions Charnov associated with hermaphroditism being favored (both male and female fitness curves are saturating). 6 Figure 1.2: Possible trade-offs between male and female function for hermaphroditism (convex line) and dioecy (concave line) as discussed by Charnov (1982). 8 Figure 2.1: Map of St. Andrews State Park in Panama City, Florida, highlighting the three locations of observation and their average fish densities; Outer Jetty, Lagoon, and Inner Jetty. 28 Figure 2.2: Number of spawns (clear diamonds) across the spawning season. Surface temperature (black squares) taken from a nearby NOAA buoy across spawning season. Water temperature correlates with spawning activity (Pearson’s correlation, r95 = 0.60, p = 0.032). 29 Figure 2.3: Fish densities taken along permanent 50-m band transects across two spawning seasons at each of the three spawning locations (inner jetty, lagoon, outer jetty). 30 Figure 2.4: Results of correlation between fish density and the proportion of streaking behaviors. Inner jetty (low density, diamonds): Pearson’s correlation, r15 = -0.147, p = 0.875; Lagoon (intermediate density, squares): Pearson’s correlation, r9 = 0.051, df = 1, p = 0.323; Outer jetty (high density, triangles): Pearson’s correlation, r13 = 0.079, df = 1, p = 0.192). 31 Figure 2.5: Incidence of pair spawning and streak spawning in S. subligarius from animals observed in the field, at each of three locations within the study site that vary naturally in density; inner jetty (low density), lagoon (intermediate density), outer jetty (high density). 32 Figure 2.6: Size structure data presented as a proportion of total density from observations along permanent density transects at each of three sites (a) Inner jetty, (b) Lagoon and (c) Outer jetty. Relative sizes were used and individuals were classified as small (<65 mm SL), medium (65-80 mm SL) or large (>80 mm SL). Black bars = small; dark gray bars = medium; light gray bars = large. 33 vi Figure 2.7: Frequency of different male mating behaviors (pair spawn and streak spawn) of each size class at the three different sites using total number of fish observed over two mating seasons. 34 Figure 2.8: Relationship between algal cover and incidence of streak spawning during two field seasons (a) 2006 and (b) 2007, where high algal cover at local spawning sites predicts a greater proportion of streak spawning than bare rock. Chi-sq; 5.20, df = 1, p = 0.023 (2006). Chi-sq; 5.97, df = 1, p = 0.015 (2007). 35 Figure 2.9: Cross section of a gonad of Serranus subligarius, taken from the area near the base (posterior) portion, indicating basic morphology and three of the gamete development stages; spermatozoa, primary oocytes and yolk globule oocytes. 36 Figure 3.1: Drawing showing typical pair spawning event, with courtship occurring over rocks and culminating in a short spawning rush into the water column, cupping behavior, and finally, the release of gametes. Black lines drawn onto to individual in cupping position indicate the body angle measurement taken at point of gamete release as a test of altered behavior using ImageJ software analysis. 47 Figure 3.2: Fertilization rates obtained from field-collected samples during the spawning seasons of 2006, 2007, 2008. Variation in fertilization rate increases with the spawning season. 48 Figure 3.3: Percent of eggs fertilized over the spawning season in 2006 and 2007. Spawns that included a streaker were removed from this analysis and a best fit regression line suggests a slightly negative, but nonsignificant result (2006 (solid line): n = 13, R2 = 0.0512; 2007 (dashed line): n = 19, R2 = 0.3016). 49 Figure 3.4: Truncated data showing the fertilization rates late in the spawning season (August-October). Pair spawning (dark circles, n = 23) and streak spawning (open circles, n = 16) fertilization rates are shown and participation of streakers results in lower overall fertilization success (t-test, df = 37, p = 0.001). 50 Figure 3.5: Total sperm output estimates (in millions) for pair spawns and pair spawns with at least one streaker participating (n = 22, t-test, p = 0.047). 51 Figure 3.6: The average number of eggs in a single parcel does not differ between spawns of a pair only and ones that include at least one streaker (n = 23, t-test, p = 0.67). 52 Figure 3.7: Proportion of spawns including a streaker from each of the four treatments of the field manipulation experiment. Streak spawning occurred at significantly higher rates in the high density, mixed size structure treatment and the presence of small fish drove the incidence of streak spawning (streak*density (hi/lo): Chi-sq=0.743, p=0.388; streak*structure (equal/mixed): Chi-sq: 3.930, p=0.047; streak*size (small/large): Chi-sq: 17.183, p<0.0001). 53 vii Figure 3.8: Proportion of eggs fertilized during the manipulation experiment, with pair spawns that included at least one streaker (black bar) and those that were only a pair of fish (white bar). Pair spawning fish had a greater proportion of eggs fertilized than those with streak spawners (t-test: n = 27, t=3.14, p=0.012). 54 Figure 3.9: Body angle of female fish during gamete release, with pair spawning only (black bar) and pair spawners with streakers (white bar). Body angle was significantly greater for females when streak spawners were present (n = 32, Watson-Williams test, p = 0.041). 55 Figure 3.10: Body angle of male fish during gamete release, with pair spawning only (black bar) and pair spawners with streakers (white bar).
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