Environmental and Life-history Factors Influencing Juvenile Demography of a Temperate Reef Fish

<p>Realistic population models and effective conservation strategies require a thorough understanding of the processes that drive variation in individual growth and survival, particularly within life stages that are subject to high mortality. For fragmented marine populations it is also important to consider how processes driving variation performance may vary through space and time. In this study I assess the interaction of two primary factors driving juvenile demography: benthic habitat composition and larval history traits, in a temperate reef fish, Forsterygion lapillum (the common triplefin). It is well understood that juveniles of many marine organisms are closely associated with structured nearshore habitats as they provide resources (refuge and food sources) that are critical for juvenile growth and/or survival. Nursery habitats are often assessed using measures of fitness of juveniles inhabiting them (e.g. rates of growth). However individual fitness measures may not only be indicative of conditions experienced in the benthic phase, but also an individual's prior history. Recent evidence suggests that variation in larval traits at settlement (e.g., size and age at settlement, larval growth rate) can impact on subsequent ecological performance (e.g., feeding ability and/or predator avoidance) and therefore influence subsequent fitness (i.e. rates of growth and/or probabilities of survival). I used otolith microstructure to assess separate and joint effects of habitat composition and larval traits on the growth of young F. lapillum. Both macroalgal composition of habitat patches and larval traits affected juvenile growth rates, and results suggested that habitat composition may have the potential to mediate fitness-related advantages that may accrue to certain individuals as a result of paternal effects and/or larval dispersal history. Quantifying spatio-temporal variability in the post-settlement fitness of Individuals with that differ in larval traits is essential for effective spatial management of marine populations. I further explore the joint effects of macroalgal composition and larval traits, within the context of additional spatial and temporal environmental variation. Results provide direct evidence that habitat can mediate the strength of carryover effects, but that the impact of habitat was variable between local populations and settlement events through time. In chapter 4 of my thesis, I focus on how small-scale variation in macroalgal composition within a nursery habitat (while controlling for individual variation) can affect the strength of density dependent growth and survival rates of F. lapillum. Density-dependent survival is evident during the first 30 days after settlement, and the strength of density dependence varied as a function of macroalgal composition. Resulting variation in estimates of nursery value (i.e., the number of late-stage juveniles produced per area unit of habitat) highlight the importance of incorporating local scale variation in juvenile demography into assessments of nursery habitat. Lastly, I assess a potential strategy of fishes to persist in a wide range of benthic environments. The ability to adjust traits (i.e., phenotypic plasticity) may allow organisms that encounter a range of unpredictable environmental conditions to maximise fitness within a single generation. In chapter 5 I explore patterns of variation in morphology of juvenile F. lapillum from two different subpopulations and from different macroalgal habitats. I evaluate possible evidence for constraints on morphological variation arising from variation in growth rate prior to and following settlement. Results suggest that for organisms with complex life cycles, variation in growth rates experienced during dispersal may constrain plasticity in later stages.</p>

Environmental and Life-history Factors Influencing Juvenile Demography of a Temperate Reef Fish

<p>Realistic population models and effective conservation strategies require a thorough understanding of the processes that drive variation in individual growth and survival, particularly within life stages that are subject to high mortality. For fragmented marine populations it is also important to consider how processes driving variation performance may vary through space and time. In this study I assess the interaction of two primary factors driving juvenile demography: benthic habitat composition and larval history traits, in a temperate reef fish, Forsterygion lapillum (the common triplefin). It is well understood that juveniles of many marine organisms are closely associated with structured nearshore habitats as they provide resources (refuge and food sources) that are critical for juvenile growth and/or survival. Nursery habitats are often assessed using measures of fitness of juveniles inhabiting them (e.g. rates of growth). However individual fitness measures may not only be indicative of conditions experienced in the benthic phase, but also an individual's prior history. Recent evidence suggests that variation in larval traits at settlement (e.g., size and age at settlement, larval growth rate) can impact on subsequent ecological performance (e.g., feeding ability and/or predator avoidance) and therefore influence subsequent fitness (i.e. rates of growth and/or probabilities of survival). I used otolith microstructure to assess separate and joint effects of habitat composition and larval traits on the growth of young F. lapillum. Both macroalgal composition of habitat patches and larval traits affected juvenile growth rates, and results suggested that habitat composition may have the potential to mediate fitness-related advantages that may accrue to certain individuals as a result of paternal effects and/or larval dispersal history. Quantifying spatio-temporal variability in the post-settlement fitness of Individuals with that differ in larval traits is essential for effective spatial management of marine populations. I further explore the joint effects of macroalgal composition and larval traits, within the context of additional spatial and temporal environmental variation. Results provide direct evidence that habitat can mediate the strength of carryover effects, but that the impact of habitat was variable between local populations and settlement events through time. In chapter 4 of my thesis, I focus on how small-scale variation in macroalgal composition within a nursery habitat (while controlling for individual variation) can affect the strength of density dependent growth and survival rates of F. lapillum. Density-dependent survival is evident during the first 30 days after settlement, and the strength of density dependence varied as a function of macroalgal composition. Resulting variation in estimates of nursery value (i.e., the number of late-stage juveniles produced per area unit of habitat) highlight the importance of incorporating local scale variation in juvenile demography into assessments of nursery habitat. Lastly, I assess a potential strategy of fishes to persist in a wide range of benthic environments. The ability to adjust traits (i.e., phenotypic plasticity) may allow organisms that encounter a range of unpredictable environmental conditions to maximise fitness within a single generation. In chapter 5 I explore patterns of variation in morphology of juvenile F. lapillum from two different subpopulations and from different macroalgal habitats. I evaluate possible evidence for constraints on morphological variation arising from variation in growth rate prior to and following settlement. Results suggest that for organisms with complex life cycles, variation in growth rates experienced during dispersal may constrain plasticity in later stages.</p>

Tropicalization of temperate reef fish communities depends on urchin herbivory and thermal response diversity

Global declines in structurally complex habitats are reshaping both land and seascapes in directions that affect biological communities’ responses to warming. Here, we test whether widespread loss of kelp habitats through sea urchin overgrazing systematically changes warming sensitivity of fish communities. Community thermal affinity shifts related to habitat were assessed by simulating and comparing fish communities from 2,271 surveys across 15 ecoregions. We find that fishes in kelp and urchin barrens differ in realized thermal affinities and range sizes, but only in regions where species pools have high variability in species’ thermal affinities. Barrens on warm-temperate reefs host relatively more warm-affinity fish species than neighbouring kelp beds, highlighting acceleration of tropicalization processes facilitated by urchin herbivory. By contrast, proportionally more cool-affinity fishes colonize barrens at high temperate latitudes, contributing to community lags with ocean warming in these regions. Our findings implicate urchins as drivers of ecological change, in part by affecting biological resilience to warming.

Pelagic larval growth rate impacts benthic settlement and survival of a temperate reef fish

Larvae of marine reef organisms settling into benthic habitats may vary greatly in individual quality. We evaluated potential effects of variation in larval growth rate (1 metric of quality) on larval duration, size-at-settlement, and post-settlement survival of recently settled kelp bass Paralabrax clathratus. We sampled kelp bass daily and weekly from standardized collectors located near the Wrigley Institute for Environmental Studies, Santa Catalina Island, to characterize larval traits of settlers and surviving recruits. Using growth models to fit trajectories of larval otolith growth, we estimated instantaneous larval growth rates and found that these values were good predictors of larval duration and juvenile survival. Kelp bass that grew rapidly as larvae settled ∼8.5 d sooner than the slowest growing individuals; both groups had similar sized individuals at settlement, but fast growing larvae experienced enhanced survival during the first 5 d after settlement relative to slower growing larvae. There is growing evidence suggesting that larval experience continues to exert demographic consequences on subsequent life stages. This helps to explain some of the spatial and temporal variability that characterizes recruitment in marine systems.

Spatially variable larval histories may shape recruitment rates of a temperate reef fish

Several long-standing hypotheses purport variation in recruitment to be positively correlated with pelagic environmental conditions that enhance larval growth, survival, and/or delivery to recruitment sites. However, the relationship between recruitment intensity and larval environmental conditions (or more directly, larval condition) is difficult to evaluate and poorly known for most species. We evaluate this relationship for the reef fish Forsterygion lapillum that commonly inhabits rocky reefs throughout New Zealand. We quantified variation in recruitment of F. lapillum using a nested sampling design, and found that the largest source of variation was between 2 nearby regions (a semi-enclosed harbour and an adjacent open coast system). We estimated 'settler condition' as the composite of residual body mass and 2 measurements of larval growth (reconstructed from otolith microstructure) and found that recruitment intensity was positively correlated with settler condition for sites within the harbour, but negatively correlated with settler condition for sites on the open coast. Mean pelagic larval duration of recruits to the harbour was ̃3 d shorter than recruits to the open coast. These results suggest that larval experience and relationships between recruitment and settler condition are spatially variable. We speculate that (1) larval retention within a productive embayment facilitates a positive relationship between recruitment and settler condition while (2) dispersal through a less productive environment drives a negative relationship for replenishment on the open coast. These putative differences may have important implications for patterns of recruitment, the strength of post-settlement density-dependent interactions, and dynamics of local populations.

Regional variation in larval retention and dispersal drives recruitment patterns in a temperate reef fish

Although there is wide appreciation for the influence of variable larval transport on recruitment, few studies have investigated the relationship between recruitment intensity and the origin(s) of recruits. We evaluate this relationship for the common triplefin Forsterygion lapillum in Wellington Harbour (a semi-enclosed bay) and the adjacent open coast of North Island, New Zealand. As the harbour is productive, with many F. lapillum adults, we predicted the harbour would be an important source of recruitment for both regions. We estimated larval origins and dispersal histories using embryonic and larval otolith chemistry, respectively, with differences in otolith elemental composition consistent with birth and/or development in either harbour or open ocean waters. In the harbour, recruitment was greatest when locally spawned larvae were retained. Although large recruitment pulses to the open coast were comprised mostly of larvae originating from the open coast, 72 % of all recruits to the south coast were actually spawned in the harbour. Dispersal of larvae from the harbour was frequent, but in low numbers, and was unrelated to opportunities for transport from the harbour. Given recent evidence for differences in larval growth between regions, we suggest that (1) faster larval growth combined with nearshore larval distributions result in high recruitment to the harbour and moderate, but demograph-ically important, dispersal from the harbour to the open coast. In contrast, we suggest that (2) the combination of strong tidal currents in Cook Strait and slower larval growth result in lower recruitment and greater transport of larvae among open coast populations. The mechanisms that underlie such context-dependent dispersal have important implications for patterns of connectivity. © Inter-Research 2010.

Regional variation in larval retention and dispersal drives recruitment patterns in a temperate reef fish

Although there is wide appreciation for the influence of variable larval transport on recruitment, few studies have investigated the relationship between recruitment intensity and the origin(s) of recruits. We evaluate this relationship for the common triplefin Forsterygion lapillum in Wellington Harbour (a semi-enclosed bay) and the adjacent open coast of North Island, New Zealand. As the harbour is productive, with many F. lapillum adults, we predicted the harbour would be an important source of recruitment for both regions. We estimated larval origins and dispersal histories using embryonic and larval otolith chemistry, respectively, with differences in otolith elemental composition consistent with birth and/or development in either harbour or open ocean waters. In the harbour, recruitment was greatest when locally spawned larvae were retained. Although large recruitment pulses to the open coast were comprised mostly of larvae originating from the open coast, 72 % of all recruits to the south coast were actually spawned in the harbour. Dispersal of larvae from the harbour was frequent, but in low numbers, and was unrelated to opportunities for transport from the harbour. Given recent evidence for differences in larval growth between regions, we suggest that (1) faster larval growth combined with nearshore larval distributions result in high recruitment to the harbour and moderate, but demograph-ically important, dispersal from the harbour to the open coast. In contrast, we suggest that (2) the combination of strong tidal currents in Cook Strait and slower larval growth result in lower recruitment and greater transport of larvae among open coast populations. The mechanisms that underlie such context-dependent dispersal have important implications for patterns of connectivity. © Inter-Research 2010.