Factors Affecting Nesting Success in Florida Bass Micropterus Floridanus

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Title:
Factors Affecting Nesting Success in Florida Bass Micropterus Floridanus
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1 online resource (34 p.)
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english
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Slagle, Zachary Joseph
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University of Florida
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Gainesville, Fla.
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Degree:
Master's ( M.S.)
Degree Grantor:
University of Florida
Degree Disciplines:
Fisheries and Aquatic Sciences, Forest Resources and Conservation
Committee Chair:
ALLEN,MICHEAL S
Committee Co-Chair:
BERKSON,JAMES
Committee Members:
TRIPPEL,NICHOLAS A

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Subjects / Keywords:
bass -- fisheries -- florida -- floridanus -- freshwater -- micropterus -- nest -- nesting -- recruitment -- reproductive -- sportfish -- success
Forest Resources and Conservation -- Dissertations, Academic -- UF
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Fisheries and Aquatic Sciences thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
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Abstract:
Bed fishing in Florida Bass Micropterus floridanus fisheries has made nest survival a topic of interest for managers, but Florida Bass nesting has rarely been studied. My objectives were to determine what habitat and temporal characteristics (substrate, depth, water temperature) Florida Bass seek for nesting, to estimate nest survival, and to examine factors that influence nest survival to the swim-up fry stage. Four lakes were snorkeled in North-central Florida each year between 2010-2013 to survey bass nests. Snorkelers found 313 nests, of which 75% were found on vegetation. Guarding male fish were only seen on 64% of active nests, indicating lower nest fidelity than has been reported in other Micropterus spp. Florida Bass daily nest survival was estimated at 0.885. The most important factors in determining nest success were presence of a male bass and depth of the nest, whereas sizes of the male bass and of the brood were less important. Stunted growth in the study populations could have decreased power of male size for statistical tests. Florida Bass have a longer available spawning period compared to other Micropterus spp. and may not need to defend their nests as tenaciously as their northern counterparts, relying on multiple spawns in a year.
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In the series University of Florida Digital Collections.
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Includes vita.
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Includes bibliographical references.
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Description based on online resource; title from PDF title page.
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This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Statement of Responsibility:
by Zachary Joseph Slagle.
Thesis:
Thesis (M.S.)--University of Florida, 2014.
Local:
Adviser: ALLEN,MICHEAL S.
Local:
Co-adviser: BERKSON,JAMES.

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1 FACTORS AFFECTING NESTING SUCCESS IN FLORIDA BASS MICROPTERUS FLORIDANUS By ZACHARY JOSEPH SLAGLE A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2014

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2 2014 Zachary Joseph Slagle

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3 ACKNOWLEDGMENTS First I would like to thank the Florida Fish and Wildlife Conservation Commission for funding of my schooling and project. I would like to thanks my advisor Mike Allen for guiding me towards better scientific thinking and practice. My committee members Jim Berkson and Nick Trippel have provided me with excellent help over the course of writing this paper. My fellow graduate students Steph Shaw and Dan Gwinn have each helped me in many ways to improve this work. Nick Cole, Mike Stansel, and many others provi ded assistance in the field. Finally, my fianc Bailey Trump and my immediate family Tim, Mary Beth, and Gabe have given me moral support.

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4 TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................ ................................ ................................ .. 3 LIST OF TABLES ................................ ................................ ................................ ............ 5 LIST OF FIGURES ................................ ................................ ................................ .......... 6 ABSTRACT ................................ ................................ ................................ ..................... 7 CHAPTER 1 INTRODUCTI ON ................................ ................................ ................................ ...... 8 2 METHODS ................................ ................................ ................................ .............. 11 Study Site ................................ ................................ ................................ ............... 11 Snorkeling Surveys ................................ ................................ ................................ 11 Analysis ................................ ................................ ................................ .................. 13 3 RESULTS ................................ ................................ ................................ ............... 15 4 DISCUSSION ................................ ................................ ................................ ......... 17 APPENDIX A TABLES ................................ ................................ ................................ .................. 21 B FIGURES ................................ ................................ ................................ ................ 25 LIST OF REFERENCES ................................ ................................ ............................... 30 BIOGRAPHIC AL SKETCH ................................ ................................ ............................ 34

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5 LIST OF TABLES Table page A 1 Summary of the top 15 nest Information Criterion, corrected for small sample size (AICc). ............................ 22 A 2 Daily nest su rvival estimates, standard errors, and upper and lower confidence intervals for model averaged parameters. ................................ ........ 23 A 3 Individua l parameters ranked by relative importance by summing the AICc model weights for each model that contains the parameter (the strongest weight a parameter can receive is 1.000). ................................ .......................... 24

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6 LIST OF FIGURES Figure page B 1 Map of the four study lakes (Big Fish, Devils Hole, Keys, and Picnic) in north central Florida. ................................ ................................ ................................ .... 25 B 2 The depth of nests versus Julian date with a linear regression model ( P < 0.01, R 2 = .266). ................................ ................................ ................................ 26 B 3 Nest count by temperature (C) on day found. ................................ ................... 27 B 4 Daily nest survival estimates and 95% confidence intervals averaged across all models for male presence, lake, nest score, and temperature. ...................... 28 B 5 Mean estimated sizes (total length in cm) for nest guarding males in three of the four study lakes (mean standard error). ................................ ..................... 29

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7 Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science FACTORS AFFECTING NESTING SUCCESS IN FLORIDA BASS MICROPTERUS FLORIDANUS By Zachary Joseph Slagle May 2014 Chair: Micheal Allen Major: Fisheries and Aquatic Sciences Bed fishing in Florida Bass Micropterus floridanus fisheries has made nest survival a topic of interest for managers, but Florida Bass nesting has rarely been studied. My objectives were to determine what habitat and temporal characteristics (substrate, depth, water temperature) Florida Bass seek for nest ing, to estimate nest survival, and to examine factors that influence nest survival to the swim up fry stage. Four lakes were snorkeled in n orth c entral Florida each year between 2010 2013 to survey bass nests. Snorkelers found 313 nests, of which 75% were found on vegetation. Guarding male fish were only seen on 64% of active nests, indicating lower nest fidelity than has been reported in other Micropterus spp. Florida Bass daily nest survival was estimated at 0.885. The most important factors in determini ng nest success were presence of a male bass and depth of the nest, whereas sizes of the male bass and of the brood were less important. Habitat and temporal characteristics had minimal effects on nest survival. Stunted growth in the study populations coul d have decreased power of male size for statistical tests. My results indicate lower nest fidelity for Florida Bass than has been reported for other black basses, but presence of the male was the most effective predictor of nest success.

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8 CHAPTER 1 INTRODUCTION Black Bass Micropterus spp. fishing in Florida is a major source of revenue for the state of Florida and a primary management and research co ncern for the Florida Fish and Wildlife Conservation Commission (FWC). In 2006, anglers in Florida spent 14 million angler days fishing for black bass, generating US$1.25 billion in economic impact and supporting 12,000 jobs (U.S. Department of Interior 20 06; FWC 2011). Florida Bass M. floridanus grow to larger sizes than Largemouth Bass M. salmoides (Bailey and Hubbs 1949), and thus the trophy Florida Bass fishery attracts anglers from across the world. Florida Bass are also widely stocked across the Unite d States due to their potential for growth to trophy size (Siepker and Casto Yerty 2008). Florida Bass nesting has rarely been studied, and there is concern from Florida anglers that fishing bass nests has negative impacts on the populations (FWC 2011). As a result, the FWC has identified Florida Bass nesting as an important research topic Male Florida Bass create shallow nests in the substrate where females then deposit their eggs (Miller 1975). The male bass guards the fertilized eggs from predators unti 1988; Cooke et al. 2002). The male often stays with the swim up fry until the fry ball disperses. Spawning in north central Florida begins in January and continues through June, starting when water temperatures are 15 17C and generally peaking in March (Chew 1974; Rogers and Allen 2009). The success of a brood is determined by a number of physical and biological factors, including size of male, timing of the nest (and associated water t emperature), and age of the male bass (Gingerich and Suski 2011; Parkos et al. 2011). Development

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9 of a brood from eggs to swim up fry is a common indicator of a successful nest (Brown 1984). Many spawning attempts are not successful, and brood abandonment is common due to storms, sudden temperature changes, and nest predation (Phillip et al. 1997; Steinhart et al. 2005, Gingerich and Suski 2011). Larger and older male Largemouth Bass create and guard nests that are more successful than smaller, younger cons pecifics (Parkos et al. 2011) Size of the male bass is also positively related to nest success in Smallmouth Bass M. dolomieu (Gingerich and Suski 2011). In both species, larger bass spawn earlier in the nesting season (Goodgame and Miranda 1993), and thu s their fry are larger than their smaller conspecifics and more likely to survive (Miranda and Hubbard 1994; Ludsin and DeVries 1997; Parkos and Wahl 2010). However, Rogers and Allen (2009) found no increase in overwinter survival for earlier spawned Flori da Bass. Size of the brood significantly influences male aggressio n (Suski and Phillip 2004); larger broods cause male bass to defend more aggressively Nest guarding bass are highly vulnerable to angling. Males guarding larger broods are more likely to s trike a lure than those defending small broods (Suski and Phillip 2004). While angling, scientists have managed to hook and land over 50% of bedding males (Phillip et al. 1997; Suski and Phillip 2004). Further, high visibility of bass nests compared to the surrounding substrate can make nests easy to locate for anglers in clear angled from the nest, the brood is much less likely to be successful owing to nest predation (Suski et al. 20 03; Hanson et al. 2008). Probability of nest failure is positively related to the amount of time the fish is held off the nest (Suski and Philipp 2004).

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10 Population level impacts of b ed fishing are unknown Largemouth B ass recruitment is relatively constant for a range of population densities (Gwinn and Allen 2010; Allen et al. 2011), which could make individual nest succ ess unimportant to recruitment. That is, fewer nests do not necessarily lead to fewer recruits the following year due to density de pendent responses in juvenile survival or growth (Walters and Martell 2004) Parkos and Wahl (2010) suggested that managers should focus on nest success in order to improve recruitment in Largemouth Bass populations, and nest success remains a potentially important consideration in a fishery. Florida Bass nesting success has not been well studied, and there is a need to understand how factors such as fish size, brood size, and habitat influence nest success. A survey of bass anglers performed by the FWC found that bed fishing for bass is a major public concern (FWC 2011). As a result, FWC has identified impacts to nesting success of bass as a priority research need. Largemouth and Smallmouth Bass nesting success has been studied (e.g. Steinhart et al. 200 5, Parkos et al. 2011), but little is known about Florida Bass nesting success and their relation to the other Micropterus spp. Daily nest survival has only been estimated for Smallmouth Bass (Steinhart et al. 2005; Suski and Ridgway 2007), and nesting fac tors for Florida Bass have gone similarly unstudied. The objectives of this study were to determine habitat characteristics of Florida Bass nests (such as temperature, substrate, and depth), to assess factors influenc ing nest success, and to estimate daily nest survival for Florida Bass.

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11 CHAPTER 2 METHODS Study Site The four lakes (Big Fish, Devils Hole, Keys, and Picnic Lakes) for this study are located on private, undeveloped land in Putnam County, Florida (Figure B 1). All lakes are relatively shall ow and represent a range of trophic statuses (eutrophic to mesotrophic; Canfield and Hoyer 1992). The study lakes range from 3 18 ha in surface area and 2.3 4.0 m mean depth (Canfield and Hoyer 1992). All four study lakes have abundant populations of Florida Bass Micropterus floridanus and Bluegill Lepomis macrochirus Other species include Lake Chubsucker Erimyzon sucetta Warmouth Lepomis gulosus in Keys, Picnic, and Devils Hole lakes; Florida Gar Lep isosteus platyrhincus in Devils Hole Lake and Seminole Killifish Fundulus seminolis in Big Fish Lake (Canfield and Hoyer 1992). Snorkeling Surveys I evaluated brood success at each study lake via nest surveys, beginning with a systematic division of each lake into feasible sampling units. The littoral area (depth 0 3 m) of each lake was mapped and divided into transects using ArcMap 9.3 software (ESRI, Redlands, California). The littoral zone of the smaller lakes (< 5 ha; Keys Pond, Big Fish) was divid ed into four transects of equal area, whereas the littoral zone of larger lakes (> 5 ha; Picnic and Devils Hole) was divided into ten transects of equal area. These transects allowed snorkelers to sample at least two randomly selected transects from each lake on each sample day. Nest surveys began once Florida Bass spawning began. Water temperature was monitored using Onset HOBO temperature loggers placed at 1 m water depth. Boat

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12 surveys to observe the littoral zone of each lake were conducted once wee kly starting before water temperatures approached the lower range of spawning temperatures reported for Florida Bass in Florida (15 17C; Chew 1974). When evidence of spawning activity was observed (i.e. nest building, increased presence of adult bass i n the littoral zone), snorkel surveys were initiated. Snorkel surveys were conducted twice weekly (every three to four days) on all four lakes for the duration of the 2010 2012 spawning seasons and once weekly for the 2013 spawning season. One half of ea ch of the smaller lakes (Keys and Big Fish) was snorkeled during each survey with a quarter of the lake being covered by each of two to three snorkelers. Thus, the entire lake was snorkeled each week (2010 2012) or every two weeks (2013) for these two lak es. In the larger lakes (Picnic and Devils Hole), two to three new transects were randomly chosen for each survey resulting in a total of two to six new transects covered in each lake each week. To cover the littoral zone represented in each transect, sn orkelers swam in a zigzag pattern perpendicular to the shoreline out to approximately 3 m depth. Each bass nest observed was marked with an individually numbered tag and the approximate location was described and marked on a map of the lake. Snorkelers collected a variety of data at each active nest they encountered. Active nests were defined as cleared areas of substrate that contained living eggs and/or larvae. Information collected for each nest included the depth of the nest (m), the developmental s tage of the offspring (egg, yolk sac fry or swim up fry), the relative number of eggs in the nest ( nest score 1 5; Kubacki 1992), and a description of the nest

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13 substrate. Presence or absence of the male was noted, and when present, the total length of the guarding male was estimated visually to the nearest centimeter. Observed nests were revisited upon successive survey days until brood success or failure was determined. Snorkelers would revisit each nest on every survey following first detection and not e whether the brood was present, whether a swim up fry ball was in the nearby water column, and whether fungus was present on the eggs. A brood was deemed successful if at least one free swimming fry was found in association with the nest, while disappeara nce of the brood or fungus on all eggs resulted in a failed nest. If researchers were unable to find the nest on successive surveys the nest was categorized as an undetermined fate. Once a fate determination was made, nest visits were ended. Analys e s I us ed the Program MARK (G. White, Colorado State University, personal communication; available at http://warnercnr.colostate.edu/~gwhite/mark/mark.htm) to conduct a nest survival analysis, which allows for unbiased estimates of daily nest survival for Micropt erus spp. (Steinhart et al. 2005, Suski and Ridgway 2007). The Program R (R Core Team, Vienna, Austria) package RMark (Laake and Rexstad 2008) was used to create models in program MARK. The MARK nest survival analysis is based on Mayfield (1961), which all ows for nests of unknown fate to be included in an analysis. Program MARK uses a maximum likelihood from the binomial distribution with the Mayfield method, providing the ability to compare individual covariates among nests. I hypothesized that nest survi val would be influenced by up to six variables ( depth, male presence, lake, nest score, average water temperature on day found, and year ), but limiting my analyses to a few combinations of these variables was deemed to

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14 be arbitrary. I used multimodel inference (Anderson 2008) in order to evaluate the model for inference. Therefore, n est survival models were built using all combinations of possible explan atory variables, including up to six parameters, for a total of 64 models. s ize (AICc; Dinsmore et al. 2002, Burnham and Anderson 2004). I did not test goodness of fit for these models, as there is no unbiased, practical method for evaluation of nest survival models (Sturdivant et al. 2007; Walker et al. 2013). I also used RMark to generate mean daily nest survivals for individual covariates. For each model, an estimat e of daily nest survival was generated for each nest. These nest survivals were then averaged across covariates of interest so nest survival was estimated for each value of each covariate Finally, the covariate specific estimates were multiplied by the m AICc probability for each model (Anderson 2008) The results are estimates of daily nest survival for each covariate value, weighted by the model probabilities. Higher ranked models contribute more to the parameter estimates. Thus, relationships bet ween those covariates and nest survival could be inferred Finally, I used RMark to compare each predictor variable based on relative importance in all models combined This approach sums the AICc weights for each model that a variable appears in, allowing a comparison of variable importance across all models evaluated (Anderson 2008). The higher ranked variables in terms of relative importance are the most related to nest survival, whereas lower ranking indicates a variable that is less related to nest sur vival

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15 CHAPTER 3 RESULTS A total of 334 nests were surveyed across four years (2010 2013) of nesting, of which 21 nests were dropped due to incomplete information. The data thus comprised 313 nests with 176 successes for an observed nesting success of 56% All lakes except Devils Hole exhibited skipped spawning during at least one year (Shaw and Allen 201 4 ). Nest numbers differed greatly among lakes (Big Fish = 17 nests; Devils Hole = 285; Keys = 8; Picnic = 3). Three quarters ( 75%) of nests incorporated m acrophytes in the nest depression, including 20% of nests on Spatterdock Nuphar advena and 10% of nests on Maidencane Panicum hemitomon Thus, use of aquatic plants was very common for spawning fish in this study. We observed a nest guarding male on 201 nests, or 64% of the surveyed nests. Depth of the nests increased with increasing Julian date (Figure B 2). Nesting began when lake temperature reached 15C and ceased above 27C, with most nesting taking place between 20 24C (Figure B 3). In 201 3, only two temperature loggers were recovered (from Big Fish and Keys Lakes), so temperatures for nests in Devils Hole Lake (n = 26) were estimated with linear regression models based on data from 2010 2012. The model was significant (F = 13500, df = 426, P < 0.001) and thus temperatures were estimated for nests found in Devils Hole during 2013. No nests were found in Picnic Lake in 2013. Male presence, lake, and depth of nest all appeared in the nest survival models that with the most AICc support, but ma ny models received substantial relative support in the AIC model selection framework (Table A 1). There were five models with This infers that no single model or single parameter had very strong support relative to o ther candidate models. However, the models with

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16 the most support shared several parameters in common, including lake, presence of a guarding male bass, and temperature of the lake when the nest was found. Nest score, depth, and year all appeared infrequen tly in the models that had the strongest support from AICc. Daily nest survival for all nests was 0.885 0.04 (median standard error). Presence of a guarding male and increasing temperature both increased daily nest survival but not substantially (Figu re B 4). Nest score did not appear to impact nest survival. The lakes showed some variability in nest survival but the small sample size in Picnic Lake created wide confidence intervals (n=3; Table A 2). When parameters were ranked by relative importanc e, male presence was the most important parameter, followed by temperature, lake, and egg score (Table A 3). Year and depth were less important. Overall, my predictor variables had relatively minor impacts on nest survival, but the predictor variables that came out as important were consistent among the most parsimonious models.

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17 CHAPTER 4 DISCUSSION In this study, I found that male bass presence, depth, nest size, and other factors have little i mpact on Florida Bass nest survival. Male presence was identified as the most important factor in determining nest survival compared to the other variables. However, the influence of male presence on nest survival was not strong (0.85 with male not observed versus 0.92 if male observed), and thus, none of my predictor variables had very strong effects on nest survival. Florida Bass nest survival and substrate use in this study was similar to previously published estimates for black basses. Daily nest survival for Florida Bass (0.88) was simil ar to Smallmouth Bass in other studies (0.86 0.96 ; Steinhart et al. 2005; Suski and Ridgway 2007). No studies on other black basses have estimated daily nest survival, and instead, researchers have relied upon nest success/failure based on presence of swim up fry. My estimates of nest survival are consistent with the values from Steinhart et al. (2005) and Suski and Ridgway (2007). Florida Bass utilized macrophytes for nest habitat, following behavior observed by Chew (1974), who reported 95% of Florida B ass nests on vegetative matter, primarily Maidencane rhizomes. Florida Bass in other studies have likewise shown a preference for both Spatterdock and Maidencane (Bruno et al. 1990). P rior studies on other basses have found that larger males have a higher nest success rate, guard more eggs, and are more efficient at guarding the nest (Weigmann and Baylis 1995; Gingerich and Suski 2011; Parkos et al. 2011) but I found no strong influence of male length on nest survival. The male bass in this study were ofte n scared off of the nest before the researcher coul d estimate the size of the fish. Active nests

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18 without a guarding bass accounted for 36% of surveyed nests. When analyses were limited to nests with a guarding male bass, male size was not an important vari able (compared to the other variables using either AICc ranking or parameter importance) The bass populations in this study did not have a large size distribution (Figure B 5) and exhibit ed slow gro wth (Shaw and Allen 2014 ) T hus, I had only moderate cont rast in male size through this study. Because so many nests lacked presence of a visible male, I did not further evaluate male size as a predic tor variable in the analyses. I found no evidence that male size influenced nest survival for fish in the size r ange of these populations but future studies should investigate whether male size is important for nest success in Florida Bass. Florida Bass in this study displayed less aggressive nest guarding behavior compared to other Micropterus spp in other studie s Prior studies of black bass have either made no mention of active nests without a guarding male (Gingerich and Suski es have similarly targeted male bass on nests or made no mention of active nests without apparent parental care (Lukas and Orth 1995; Gingerich and Suski 2011; Landsman et al. 2011). In this study, presence of a guarding male bass appeared have little effe ct on daily survival, but a lack of a guarding bass did not indicate a failed nest 81 broods with no guarding male detected (during any of the nest visits) were successfully raised to the swim up phase. The guarding males that were sighted tended to swim off immediately, which is unlike reported behavior of Micropterus spp. located in n orthern, colder climates. I believe

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19 cases where male s were not observed did not mean that a male was not present and guarding, but fish were timid and moved off the nest s as snorkelers approached How Florida Bass react to predators could be a mechanism for why fish were more timid than Largemouth Bass in other studies. American Alligator Alligator mississippi ensis i s present i n virtually every water body in Florida. Florida Bass share these waters and possibly have developed a flight response to large swimming animals This could explain why Florida Bass leave the nest when a snorkeler approaches unlike the northern black basses (where alligat ors as predators do not occur) Florida Bass in this study were timid in guarding the nest against approaching snorkelers, which may infer that they guard their nests less aggressively than largemouth bass at more northern latitudes. The longer period o f favorable spawning temperatures could influence diminished nest guarding behavior compared to Largemouth and Smallmouth Basses. Florida Bass occur at lower latitude than most Micropterus spp., increasing the time period when water temperatures are optim al for spawning. Florida Bass in these lakes spawned several times per year in some cases (Devils Hole Lake, Shaw and Allen 201 4 ), which has been found in other Florida Bass populations (Issac et al. 1998; Waters and Nobel 2004). This could potentially ma ke individual spawning attempts less important to individual fish. However, Florida Bass in other studies have shown greater nest guarding behavior and greater nest fidelity (N. Trippel, FWC, personal communication). Thus, it is not clear that the lack of aggressive nest guarding in this study results because fish have the potential to re nest, or if it is a more general pattern for Florida Bass.

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20 Florida Bass nest guarding behaviors have wider implications for freshwater fisheries management. Florida Bass nesting behaviors differ somewhat from other Micropterus spp. and patterns identified in other black bass nest survival did not hold true for Florida Bass in this study Loss of a guarding male bass has a m ajor impact on other bass brood survival (Suski e t al. 2003; Suski and Phillip 2004; Steinhart et al. 2005). However, such a loss may have a smaller effect on Florida Bass broods compared to other black bass broods. Loss of a nesting Florida Bass due to angling remains a concern for brood survival Furth er, the diminished nest guarding behavior could indicate lowered aggressiveness overall, and previous studies have found Florida Bass are more difficult to catch by angling (Zolczynski and Davies 1976; Kleinsasser et al. 1990) than Largemouth Bass This re duced vulnerability to angling could further lessen the impact of bed fishing on Florida Bass, and further studies are needed to investigate this topic. The relationship between brood survival and their eventual recruitment to the population remains unclea r, but negative impacts on bass populations may be diminished in Florida Bass populations.

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21 APPENDIX A TABLES

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22 Table A 1. Summary corrected for small sample size (AICc ). Model Parameters AICc weight Deviance S(~Lake + Male + Score + Temp) 7 591.22 0.000 0.202 577.12 S(~Lake + Male + Score + Temp + Year) 8 592.29 1.074 0.118 576.17 S(~Lake + Male + Temp) 6 592.67 1.456 0.097 580.60 S(~Lake + Male + Temp + Year) 7 593.00 1.786 0.083 578.91 S(~Depth + Lake + Male + Score + Temp) 8 593.19 1.971 0.075 577.06 S(~Depth + Lake + Male + Score + Temp + Year) 9 594.31 3.089 0.043 576.15 S(~Depth + Lake + Male + Temp) 7 594.56 3.344 0.038 580.46 S(~Male + Score + Temp) 4 594.64 3.422 0.036 586.60 S(~Lake + Male + Score) 6 594.89 3.674 0.032 582.82 S(~Depth + Lake + Male + Temp + Year) 8 594.99 3.770 0.031 578.86 S(~Male + Score + Temp + Year) 5 595.05 3.828 0.030 584.99 S(~Male + Temp + Year) 4 595.08 3.866 0.029 587.05 S(~Depth + Male + Score + Temp) 5 595.64 4.426 0.022 585.59 S(~Male + Temp) 3 595.77 4.554 0.021 589.75 S(~Depth + Lake + Male + Score) 7 596.12 4.901 0.017 582.02

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23 Table A 2. Daily nest survival estimates, standard errors, and upper and lower confidence intervals for model averaged parameters. Nests Daily Survival SE Lower Confidence Interval Upper Confidence Interval Overall 313 0.89 0.04 0.80 0.97 No Male 112 0.85 0.05 0.74 0.92 Male Present 201 0.92 0.03 0.85 0.95 Big Fish 17 0.90 0.03 0.82 0.94 Devils Hole 285 0.87 0.01 0.85 0.89 Keys 8 0.94 0.04 0.78 0.99 Picnic 3 0.19 5.89 0.00 1.00

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24 Table A 3. Individual parameters ranked by relative importance by summing the AICc model weights for each model that contains the parameter (the strongest weight a parameter can receive is 1.000) Parameter Relative Importance Weight Male Presence 1 0.996 Temp 2 0.867 Lake 3 0.789 Score 4 0.632 Year 5 0.404 Depth 6 0.298

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25 APPENDIX B FIGURES Figure B 1. Map of the four study lakes (Big Fish, Devils Hole, Keys, and Pi cnic) in north central Florida.

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26 Figure B 2. The depth of nests versus Julian date with a linea r regression model ( P < 0.01, R 2 = .266). 0 0.5 1 1.5 2 2.5 3 3.5 4 0 20 40 60 80 100 Depth of nest (m) Julian Date

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27 Figure B 3. Frequency plot of n est count ed on water temperature (C). 0 10 20 30 40 50 60 70 80 15 16 17 18 19 20 21 22 23 24 25 26 27 Number of nests Temperature C

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28 Figure B 4. Daily nest survival estimates and 95% confidence intervals averaged across all models for male presence, lak e, nest score, and temperature.

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29 Figure B 5. Mean estimated sizes (total length in cm) for nest guarding males in three of the four study lakes (mean standard error). 20.0 22.0 24.0 26.0 28.0 30.0 32.0 34.0 36.0 38.0 40.0 Big Fish Devil's Hole Keys Estimated male total length (cm)

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30 APPENDIX B LIST OF REFERENCES Allen, M. S., M. W. Rogers, M. J. Catalano, D. G. Gwinn, and S. J. Walsh. 2011. Evaluating the potential for stock size to limit recruitment in Largemouth Bass. Transactions of the American Fisheries Society 140:1093 1100. Anderson, D.R 2008. Model Based Inference in the Life Sciences: A Primer on Evidence. Springer, New York City. Bailey, R M. and C.L. Hubbs. 1949. The black basses (Micropterus) of Florida, with description of a new species. University of Michigan Occasional Papers of the Museum of Zoology 516: 1 40. Brown, J. A. 1984. Parental care and the ontogeny of predator avoidance in two species of centrarchid fish. Animal Behaviour 32:113 119. Bruno, N.A., R.W. Gregory, and H.L. Schramm. 1990. Nest Sites Used by Radio Tagged Lar gemouth Bass in Orange Lake, Florida. North American Journal of Fisheries Management 10:80 84. Burnham, K. P., and D. R. Anderson. 2004. Model selection and multi model inference: a practical information theoretic approach, 2nd edition. Springer, New York City. Canfield, D. E., and M. V. Hoyer. 1992. Aquatic macrophytes and their relation to the limnology of Florida lakes, Bureau of Aquatic Plant Management, Florida Department of Natural Resources, Gainesville, FL. Chew, R. L. 1974. Early life history of th e Florida Largemouth Bass. Dingell Johnson Project F 24 R. Florida Game and Freshwater Fish Commission, Tallahassee, FL. Conover, D. O. 1992. Seasonality and the scheduling of life history at different latitudes. Journal of Fish Biology 41:161 178. Cooke, S. J., D. P. Philipp, and P. J. Weatherhead. 2002. Parental care patterns and energetics of Smallmouth Bass (Micropterus dolomieu) and Largemouth Bass (Micropterus salmoides) monitored with activity transmitters. Canadian Journal of Zoology 80:756 770. Din smore, S. J., G. C. White, and F. L. Knopf. 2002. Advanced techniques for modeling avian nest survival. Ecology 83:3476 3488. Florida Fish and Wildlife Conservation Commission (FWC). 2011. Black bass management plan (2010 2030).

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31 Gingerich, A. J., and C. D. Suski. 2011. The role of progeny quality and male size in the nesting success of Smallmouth Bass: integrating field and laboratory studies. Aquatic Ecology 45:505 515. Goodgame, L. S., and L. E. Miranda. 1993. Early growth and survival of age 0 Largemouth Bass in relation to parental size and swim up time. Transactions of the American Fisheries Society 122:131 138. Gwinn, D. C., and M. S. Allen. 2010. Exploring population level effects of fishery closures during spawning: an example using Largemouth Bass. Transactions of the American Fisheries Society 139:626 634. Hanson, K. C., M. A. Gravel, T. Redpath, S. J. Cooke, and M. J. Siepker. 2008. Latitudinal variation in physiological and behavioral responses of nest guarding Smallmouth Bass to common recreation al angling practices. Transactions of the American Fisheries Society 137:1558 1566. Isaac, J., T. M. Kimmel, R. W. Bagley, V. H. Staats, and A. Barkoh. 1998. Spawning behavior of Florida Largemouth Bass in an indoor raceway. The Progressive Fish Culturist 60:59 62. Isely, J. J., R. L. Noble, J. B. Koppelman, and D. P. Philipp. 1987. Spawning period and first year growth of Northern, Florida, and intergrade stocks of Largemouth bass. Transactions of the American Fisheries Society 116:757 762. Kleinsasser, L. J., J. H. Williamson, and B. G. Whiteside. 1990. Growth and catchability of Northern, Florida, and F1 hybrid Largemouth bass in Texas ponds. North American Journal of Fisheries Management 10:462 468. Kubacki, M. F. 1992. The effectiveness of a closed seas on for protecting nesting University of Illinois, Urbana. Laake, J., and E. Rexstad. 2008. Appendix C RMark an alternative approach to building linear models in MARK. Landsman, S. J., A. J. Gingerich, D. P. Philipp, and C. D. Suski. 2011. The effects of temperature change on the hatching success and larval survival of Largemouth Bass Micropterus salmoides and Smallmouth Bass Micropterus dolomieu. Journal of Fish Biology 78:1200 12. Lud sin, S. A., and D. R. DeVries. 1997. First year recruitment of Largemouth Bass: The interdependency of early life stages. Ecological Applications 7:1024 1038. Lukas, J. A., and D. J. Orth. 1995. Factors affecting nesting success of Smallmouth Bass in a reg ulated Virginia stream. Transactions of the American Fisheries Society 124:726 735.

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32 Mayfield, H. 1961. Nesting success calculated for exposure. The Wilson Bulletin 73:255 261. Miller, R. J. 1975. Comparative behavior of centrarchid basses. Pages 85 94 in Black bass biology and management. Sport Fishing Institute, Washington, D.C. Miranda, L. E., and W. D. Hubbard. 1994. Winter survival of age 0 Largemouth Bass relative to size, predators, and shelter. North American Journal of Fisheries Management 14:790 796. Parkos, J. J., and D. H. Wahl. 2010. Intra and intersystem variation in Largemouth Bass recruitment: reproduction, prey availability, and the timing of year class establishment. Transactions of the American Fisheries Society 139:1372 1385. Parkos, J. J., D. H. Wahl, and D. P. Philipp. 2011. Influence of behavior and mating success on brood specific contribution to fish recruitment in ponds. Ecological Applications 21:2576 86. Philipp, D. P., C. A. Toline, M. F. Kubacki, D. B. F. Philipp, and F. J. S. Phelan. 1997. The impact of catch and release angling on the reproductive success of Smallmouth Bass and Largemouth Bass. North American Journal of Fisheries Management 17:557 567. Ridgway, M. S. 1988. Developmental stage of offspring and brood defense in Smallmouth Bass (Micropterus dolomieu). Canadian Journal of Zoology 66:1722 1728. Rogers, M. W., and M. S. Allen. 2009. Exploring the generality of recruitment hypotheses for Largemouth Bass along a latitudinal gradient of Florida lakes. Transactions of th e American Fisheries Society 138:23 37. Rogers, M. W., M. S. Allen, and W. F. Porak. 2006. Separating genetic and environmental influences on temporal spawning distributions of Largemouth Bass (Micropterus salmoides). Canadian Journal of Fisheries and Aqua tic Sciences 63:2391 2399. Shaw, S. L., and M. S. Allen. 201 4 Localized spatial and temporal variation in reproductive effort of Florida Bass. Transactions of the American Fisheries Society 143:85 96. Siepker, M. J., and M. Casto Yerty. 2008. A survey of fishery agency supplementary Largemouth Bass stocking practices in large United States reservoirs. Pages 439 449 in Balancing fisheries management and water uses for impounded river systems. American Fisheries Society, Bethesda, Maryland.

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33 Steinhart, G. B., N. J. Leonard, R. A. Stein, and E. A. Marschall. 2005. Effects of storms, angling, and nest predation during angling on Smallmouth Bass (Micropterus dolomieu) nest success. Canadian Journal of Fisheries and Aquatic Sciences 62:2649 2660. Sturdivant, R., J Rotella, and R. Russell. 2007. A smoothed residual based goodness of fit statistic for nest survival models. Studies in Avian Biology 34: 45 54. Suski, C. D., J. H. Svec, J. B. Ludden, F. J. S. Phelan, and D. P. Philipp. 2003. The effect of catch and release angling on the parental care behavior of male Smallmouth Bass. Transactions of the American Fisheries Society 132:210 218. Suski, C. D., and D. P. Philipp. 2004. Factors affecting the vulnerability to angling of nesting male Largemouth an d Smallmouth Bass. Transactions of the American Fisheries Society 133:1100 1106. Suski, C. D., and M. S. Ridgway. 2007. Climate and body size influence nest survival in a fish with parent al care. The Journal of Animal E cology 76:730 9. U.S. Department of t he Interior 2006. 2006 National survey of fishing, hunting, and wildlife associated recreation Florida report. Wagner, T., A. K. Jubar, and M. T. Bremigan. 2006. Can habitat alteration and spring angling explain Largemouth Bass nest success? Transactions of the American Fisheries Society 135:843 852. Walker, J., J.J. Rotella S.E. Stephens, M.S. Lindberg J.K. Ringelman C. H unter, and A.J. Smith. 2013. Time lagged variation in pond density and primary productivity affects duck nest survival in the Prairi e Pothole Region. Ecological applications : a publication of the Ec ological Society of America 23: 1061 74. Walters, C.J. and S.J.D. Martell. 2004. Fisheries ecology and management. Princeton University Press, Princeton, N.J. Waters, D. S., and R. L. Noble. 2004. Spawning season and nest fidelity of Largemouth Bass in a tropical reservoir. North American Journal of Fisheries Management 24:1240 1251. Wiegmann, D. D., and J. R. Baylis. 1995. Male body size and paternal behaviour in Smallmouth Bass, Micropterus dolomieu (Pisces: Centrarchidae). Animal Behaviour 50:1543 1555. Zolczynski, S. J., and W. D. Davies. 1976. Growth characteristics of the Northern and Florida subspecies of Largemouth Bass and their hybrid, and a comparison of catchability between the sub species. Transactions of the American Fisheries Society 105:240 243.

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34 BIOGRAPHICAL SKETCH Zachary Slagle received a M.S. in Fisheries and Aquatic Sciences from the B.S. came in Fisheries Science from Virginia Polytechnic Institute and State University in 2010. He has worked for the U.S. Forest Service Southern Research Station, the University of Florida School of Forest Resources and Conservation, and the U.S. Park Servi ce.