An Evaluation of Prior Residency and Habitat Effects on the Persistence of Settling Reef Fishes

<p>Both habitat complexity and competitive interactions can shape patterns of distribution and abundance of species. I evaluated the separate and joint effects of competitive interactions and habitat complexity on the survival of young fishes (Family Labridae) on coral reefs. First, I developed (in Chapter 2) a quantitative approach to evaluate potential resource (i.e., niche) overlap among groups of co-occurring species. Using appropriate transformations and probability models, I show that different types of data (e.g., categorical, continuous, count or binary data, as well as electivity scores) give rise to a standard measure of niche overlap, with the overlap statistic between two species defined as the overlapping area between the distributions for each species. Measurements derived from different types of data can be combined into a single multivariate analysis of niche overlap by averaging over multiple axes. I then describe null model permutation tests that differentiate between species occupying similar and different niches within my unified indices. I then implemented this approach (in Chapter 3) to evaluate potential habitat overlap among eight species of wrasse (Gomphosus varius, Halichoeres hortulanus, H. trimaculatus, Pseudocheilinus hexataenia, Scarus sordidus, Stethojulis bandanensis, Thalassoma hardwicke and T. quinquevittatum), and used these results to inform my subsequent field experiments. In a field assay, I identified the presence of T. quinquevittatum as having the greatest negative effect on survival of transplanted T. hardwicke from a suite of three candidate species which were most similar in habitat use to T. hardwicke (the other two candidate species were G. varius and P. hexataenia). In a subsequent field experiment, I tested how competition with T. quinquevittatum and structural refuge interact to influence the postsettlement survival of T. hardwicke. Competition with T. quinquevittatum and structural refuge both altered the survival of T. hardwicke, although their effects were not interactive, indicating that structural complexity did not mitigate the negative effects of competition. Survival of T. hardwicke was 2.3 times greater in treatments without T. quinquevittatum relative to those with T. quinquevittatum, and 2.8 times greater in treatments with structural refuge relative to treatments without structural refuge. Thalassoma hardwicke and T. quinquevittatum often enter reef communities asynchronously, resulting in competitive pressures faced by earlyarriving individuals that potentially differ from those experienced by late-arriving individuals. In a series of field experiments, I investigated whether the strength of intra-cohort competitive interactions between recent T. hardwicke and T. quinquevittatum settlers were dependent upon the sequence and temporal separation of their arrival into communities. Survival rates for both species were greatest in the absence of competitors, but when competitors were present, survival rates were maximized when competitors arrived simultaneously. Survival rates declined as each species entered the community progressively later than its competitor. Further, reversals in the sequence of arrival reversed competitive outcomes. Results provide empirical evidence for competitive lotteries in the maintenance of species diversity in demographically open marine systems, while also highlighting the importance of temporal variation in the direction and magnitude of interaction strengths. To further our understanding of how timing of arrival influences interaction strengths, I tested whether increasing the availability of complex habitat attenuates or enhances timing-of-arrival effects. Results from this field experiment indicated that aggression by early-arriving individuals towards late-arriving individuals increased as arrival times diverged. When aggression was weak, subordinate individuals were not displaced from complex habitat. Experimental increases in the availability of complex habitat resulted in increased survival of subordinates, presumably by disrupting predation pressure. However, when aggression was intense, competitive subordinates were displaced from complex habitat (regardless of the amount of complex habitat available), and this likely increased their exposure to predators. Overall, the experimental and observational components of this thesis emphasise heterogeneity in competitive environments experienced by recently settled reef fishes. These results highlight the important role that priority effects and habitat complexity play in determining the persistence of reef fish settlers, and illustrate how ecological contexts can add considerable variation to realised interaction strengths.</p>

An Evaluation of Prior Residency and Habitat Effects on the Persistence of Settling Reef Fishes

<p>Both habitat complexity and competitive interactions can shape patterns of distribution and abundance of species. I evaluated the separate and joint effects of competitive interactions and habitat complexity on the survival of young fishes (Family Labridae) on coral reefs. First, I developed (in Chapter 2) a quantitative approach to evaluate potential resource (i.e., niche) overlap among groups of co-occurring species. Using appropriate transformations and probability models, I show that different types of data (e.g., categorical, continuous, count or binary data, as well as electivity scores) give rise to a standard measure of niche overlap, with the overlap statistic between two species defined as the overlapping area between the distributions for each species. Measurements derived from different types of data can be combined into a single multivariate analysis of niche overlap by averaging over multiple axes. I then describe null model permutation tests that differentiate between species occupying similar and different niches within my unified indices. I then implemented this approach (in Chapter 3) to evaluate potential habitat overlap among eight species of wrasse (Gomphosus varius, Halichoeres hortulanus, H. trimaculatus, Pseudocheilinus hexataenia, Scarus sordidus, Stethojulis bandanensis, Thalassoma hardwicke and T. quinquevittatum), and used these results to inform my subsequent field experiments. In a field assay, I identified the presence of T. quinquevittatum as having the greatest negative effect on survival of transplanted T. hardwicke from a suite of three candidate species which were most similar in habitat use to T. hardwicke (the other two candidate species were G. varius and P. hexataenia). In a subsequent field experiment, I tested how competition with T. quinquevittatum and structural refuge interact to influence the postsettlement survival of T. hardwicke. Competition with T. quinquevittatum and structural refuge both altered the survival of T. hardwicke, although their effects were not interactive, indicating that structural complexity did not mitigate the negative effects of competition. Survival of T. hardwicke was 2.3 times greater in treatments without T. quinquevittatum relative to those with T. quinquevittatum, and 2.8 times greater in treatments with structural refuge relative to treatments without structural refuge. Thalassoma hardwicke and T. quinquevittatum often enter reef communities asynchronously, resulting in competitive pressures faced by earlyarriving individuals that potentially differ from those experienced by late-arriving individuals. In a series of field experiments, I investigated whether the strength of intra-cohort competitive interactions between recent T. hardwicke and T. quinquevittatum settlers were dependent upon the sequence and temporal separation of their arrival into communities. Survival rates for both species were greatest in the absence of competitors, but when competitors were present, survival rates were maximized when competitors arrived simultaneously. Survival rates declined as each species entered the community progressively later than its competitor. Further, reversals in the sequence of arrival reversed competitive outcomes. Results provide empirical evidence for competitive lotteries in the maintenance of species diversity in demographically open marine systems, while also highlighting the importance of temporal variation in the direction and magnitude of interaction strengths. To further our understanding of how timing of arrival influences interaction strengths, I tested whether increasing the availability of complex habitat attenuates or enhances timing-of-arrival effects. Results from this field experiment indicated that aggression by early-arriving individuals towards late-arriving individuals increased as arrival times diverged. When aggression was weak, subordinate individuals were not displaced from complex habitat. Experimental increases in the availability of complex habitat resulted in increased survival of subordinates, presumably by disrupting predation pressure. However, when aggression was intense, competitive subordinates were displaced from complex habitat (regardless of the amount of complex habitat available), and this likely increased their exposure to predators. Overall, the experimental and observational components of this thesis emphasise heterogeneity in competitive environments experienced by recently settled reef fishes. These results highlight the important role that priority effects and habitat complexity play in determining the persistence of reef fish settlers, and illustrate how ecological contexts can add considerable variation to realised interaction strengths.</p>

Lunar rhythms in growth of larval fish

Growth and survival of larval fishes is highly variable and unpredictable. Our limited understanding of this variation constrains our ability to forecast population dynamics and effectively manage fisheries. Here we show that daily growth rates of a coral reef fish (the sixbar wrasse, Thalassoma hardwicke ) are strongly lunar-periodic and predicted by the timing of nocturnal brightness: growth was maximized when the first half of the night was dark and the second half of the night was bright. Cloud cover that obscured moonlight facilitated a ‘natural experiment’, and confirmed the effect of moonlight on growth. We suggest that lunar-periodic growth may be attributable to light-mediated suppression of diel vertical migrations of predators and prey. Accounting for such effects will improve our capacity to predict the future dynamics of marine populations, especially in response to climate-driven changes in nocturnal cloud cover and intensification of artificial light, which could lead to population declines by reducing larval survival and growth.

Recruitment of a coral reef fish: roles of settlement, habitat, and postsettlement losses

Multiple processes typically influence patterns of abundance. Despite this widely accepted view, many studies continue to approach ecological questions from a single-factor, or, at most, a two-factor perspective. Here, I evaluate the consequences of considering, separately and jointly, the effects of three factors (larval settlement, reef resources, and postsettlement losses) on spatial patterns of abundance of a marine reef fish, the six bar wrasse (Thalassoma hardwicke). Using correlational methods commonly employed in single-factor studies, I show that local patterns of abundance of juvenile wrasse could be attributed entirely to either (1) patterns of abundance of settlement habitat, or (2) patterns of larval settlement. This result occurred because habitat and presumed larval delivery covaried in space. I manipulated abundance of settlement habitat in a field experiment to uncouple this covariation and found subsequent settlement to be simultaneously influenced by both factors. However, joint effects of habitat and settlement failed to account for patterns of abundance of juvenile wrasse without also considering a third factor - postsettlement losses - which were density-dependent and substantially modified patterns of settlement. These results illustrate (1) how multifactorial explanations may be falsely refuted when incomplete sets of multiple factors are considered, and (2) how single-factor explanations may misrepresent underlying multifactorial causation of ecological patterns. Uncovering the interactive role of multiple factors in determining ecological patterns of interest requires a shift from single-factor approaches to more pluralistic perspectives.

Born at the right time? A conceptual framework linking reproduction, development, and settlement in reef fish

© 2017 by the Ecological Society of America Parents are expected to make decisions about reproductive timing and investment that maximize their own fitness, even if this does not maximize the fitness of each individual offspring. When offspring survival is uncertain, selection typically favors iteroparity, which means that offspring born at some times can be disadvantaged, while others get lucky. The eventual fate of offspring may be further modified by their own decisions. Are fates of offspring set by birthdates (i.e., determined by parents), or can offspring improve upon the cards they've been dealt? If so, do we see adaptive plasticity in the developmental timing of offspring? We evaluate these questions for a coral reef fish (the sixbar wrasse, Thalassoma hardwicke) that is characterized by extreme iteroparity and flexible larval development. Specifically, we monitored larval settlement to 192 small reefs over 11 lunar months and found that most fish settled during new moons of a lunar cycle (consistent with preferential settlement on dark nights). Settlement was significantly lower than expected by chance during the full moon and last quarter of the lunar cycle (consistent with avoidance of bright nights). Survival after settlement was greatest for fish that settled during times of decreasing lunar illumination (from last quarter to new moon). Fish that settled on the last quarter of the lunar cycle were ~10% larger than fish that settled during other periods, suggesting larvae delay settlement to avoid the full moon. These results are consistent with a numerical model that predicts plasticity in larval development time that enables avoidance of settlement during bright periods. Collectively, our results suggest that fish with inauspicious birthdates may alter their developmental trajectories to settle at better times. We speculate that such interactions between parent and offspring strategies may reinforce the evolution of extreme iteroparity and drive population dynamics, by increasing the survival of offspring born at the “wrong” time by allowing them to avoid the riskiest times of settlement.

Born at the right time? A conceptual framework linking reproduction, development, and settlement in reef fish

© 2017 by the Ecological Society of America Parents are expected to make decisions about reproductive timing and investment that maximize their own fitness, even if this does not maximize the fitness of each individual offspring. When offspring survival is uncertain, selection typically favors iteroparity, which means that offspring born at some times can be disadvantaged, while others get lucky. The eventual fate of offspring may be further modified by their own decisions. Are fates of offspring set by birthdates (i.e., determined by parents), or can offspring improve upon the cards they've been dealt? If so, do we see adaptive plasticity in the developmental timing of offspring? We evaluate these questions for a coral reef fish (the sixbar wrasse, Thalassoma hardwicke) that is characterized by extreme iteroparity and flexible larval development. Specifically, we monitored larval settlement to 192 small reefs over 11 lunar months and found that most fish settled during new moons of a lunar cycle (consistent with preferential settlement on dark nights). Settlement was significantly lower than expected by chance during the full moon and last quarter of the lunar cycle (consistent with avoidance of bright nights). Survival after settlement was greatest for fish that settled during times of decreasing lunar illumination (from last quarter to new moon). Fish that settled on the last quarter of the lunar cycle were ~10% larger than fish that settled during other periods, suggesting larvae delay settlement to avoid the full moon. These results are consistent with a numerical model that predicts plasticity in larval development time that enables avoidance of settlement during bright periods. Collectively, our results suggest that fish with inauspicious birthdates may alter their developmental trajectories to settle at better times. We speculate that such interactions between parent and offspring strategies may reinforce the evolution of extreme iteroparity and drive population dynamics, by increasing the survival of offspring born at the “wrong” time by allowing them to avoid the riskiest times of settlement.

Cryptic density dependence: effects of spatio-temporal covariation between density and site quality in reef fish

The importance and strength of density dependence continues to engender debate because of its central importance to population dynamics and regulation. Here, we show how confounding effects of site quality can mask strong effects of density dependence. In particular, we explore spatiotemporal variation and covariation among (1) densities of newly settled coral reef fish (Thalassoma hardwicke), (2) environmental characteristics, and (3) the strength of density-dependent mortality. Environmental features of patch reefs were spatially and temporally variable and influenced density-dependent survival. Higher-quality sites (i.e., reefs possessing features that yield greater numbers of recruits at any given settlement level) received greater settlement, and this relationship masked the operation of density dependence when variation in quality among sites (or times) was not distinguished (a common approach in many Observational studies of density dependence). Our work illustrates how spatiotemporal covariation in settlement density and site quality can obscure patterns of density dependence at larger scales, contributing to a phenomenon we call "cryptic density dependence." Acknowledging patterns and consequences of covariance may alter the way we study population dynamics, especially of marine organisms, where the link between processes that affect settlement and post-settlement survival remains relatively poorly understood.

Recruitment of a coral reef fish: roles of settlement, habitat, and postsettlement losses

Multiple processes typically influence patterns of abundance. Despite this widely accepted view, many studies continue to approach ecological questions from a single-factor, or, at most, a two-factor perspective. Here, I evaluate the consequences of considering, separately and jointly, the effects of three factors (larval settlement, reef resources, and postsettlement losses) on spatial patterns of abundance of a marine reef fish, the six bar wrasse (Thalassoma hardwicke). Using correlational methods commonly employed in single-factor studies, I show that local patterns of abundance of juvenile wrasse could be attributed entirely to either (1) patterns of abundance of settlement habitat, or (2) patterns of larval settlement. This result occurred because habitat and presumed larval delivery covaried in space. I manipulated abundance of settlement habitat in a field experiment to uncouple this covariation and found subsequent settlement to be simultaneously influenced by both factors. However, joint effects of habitat and settlement failed to account for patterns of abundance of juvenile wrasse without also considering a third factor - postsettlement losses - which were density-dependent and substantially modified patterns of settlement. These results illustrate (1) how multifactorial explanations may be falsely refuted when incomplete sets of multiple factors are considered, and (2) how single-factor explanations may misrepresent underlying multifactorial causation of ecological patterns. Uncovering the interactive role of multiple factors in determining ecological patterns of interest requires a shift from single-factor approaches to more pluralistic perspectives.

Cryptic density dependence: effects of spatio-temporal covariation between density and site quality in reef fish

The importance and strength of density dependence continues to engender debate because of its central importance to population dynamics and regulation. Here, we show how confounding effects of site quality can mask strong effects of density dependence. In particular, we explore spatiotemporal variation and covariation among (1) densities of newly settled coral reef fish (Thalassoma hardwicke), (2) environmental characteristics, and (3) the strength of density-dependent mortality. Environmental features of patch reefs were spatially and temporally variable and influenced density-dependent survival. Higher-quality sites (i.e., reefs possessing features that yield greater numbers of recruits at any given settlement level) received greater settlement, and this relationship masked the operation of density dependence when variation in quality among sites (or times) was not distinguished (a common approach in many Observational studies of density dependence). Our work illustrates how spatiotemporal covariation in settlement density and site quality can obscure patterns of density dependence at larger scales, contributing to a phenomenon we call "cryptic density dependence." Acknowledging patterns and consequences of covariance may alter the way we study population dynamics, especially of marine organisms, where the link between processes that affect settlement and post-settlement survival remains relatively poorly understood.