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>

Growing up in a new world: trait divergence between rural, urban, and invasive populations of an amphibian urban invader

Cities are focal points of introduction for invasive species. Urban evolution might facilitate the success of invasive species in recipient urban habitats. Here we test this hypothesis by rearing tadpoles of a successful amphibian urban coloniser and invader in a common garden environment. We compared growth rate, morphological traits, swimming performance, and developmental rate of guttural toad tadpoles (Sclerophrys gutturalis) from native rural, native urban, and non-native urban habitats. By measuring these traits across ontogeny, we were also able to compare divergence across different origins as the tadpoles develop. The tadpoles of non-native urban origin showed significantly slower developmental rate (e.g., the proportion of tadpoles reaching Gosner stage 31 or higher was lower at age 40 days) than tadpoles of native urban origin. Yet, tadpoles did not differ in growth rate or any morphological or performance trait examined, and none of these traits showed divergent ontogenetic changes between tadpoles of different origin. These findings suggest that prior adaptation to urban habitats in larval traits likely does not play an important role in facilitating the invasion success of guttural toads into other urban habitats. Instead, we suggest that evolutionary changes in larval traits after colonization (e.g., developmental rate), together with decoupling of other traits and phenotypic plasticity might explain how this species succeeded in colonising extra-limital urban habitats.

Baby makes three: maternal, paternal, and zygotic genetic effects shape larval phenotypic evolution

The evolutionary potential of a population is shaped by the genetic architecture of its life-history traits. Early-life phenotypes are influenced by both maternal and offspring genotype, and efforts to understand life-history evolution therefore require consideration of the interactions between these separate but correlated genomes. We used a four-generation experimental pedigree to estimate the genetic architecture of early-life phenotypes in a species with dramatic variation in larval size and morphology. In the polychaete annelid Streblospio benedicti, females make either many small eggs that develop into complex larvae that feed in the plankton or few large eggs that develop into benthic juveniles without having to feed as larvae. By isolating the contributions of maternal, paternal, and zygotic genotype to larval traits, we determined that larval anatomical structures are governed by the offspring genotype at a small number of large-effect loci. Larval size is not shaped by the larva's own genotype but instead depends on loci that act in the mother, and at two genomic locations, by loci that act in the father. The overall phenotype of each larva thus depends on three separate genomes, and a population's response to selection on larval traits will reflect the interactions among them.

Patterns of Larval Development

In this chapter, we explore the different patterns of development following the hatching of the crustacean larvae. For many groups of crustaceans, the free-living, postembryonic, and prejuvenile phase is by far the most important part of their life cycle, providing the link between different life modes in successive phases (e.g., between a sessile adult life and the need for long-range planktonic dispersal). Among the aspects covered, we discuss the specific criteria for what a “larva” is, including the necessity for defining specific larval traits that are lacking in other phases of the life cycle. We examine the typical anamorphic and hemianamorphic developmental patterns based on larval examples from a wide selection of groups from Decapoda to Copepoda, Thecostraca to Branchiopoda. In these groups, we examine the most common larval development patterns (including intraspecific variability) of, for example, the zoea, furcilia, copepodite, nauplius, and cypris larvae. We also expand on the importance of the molting cycle as the main driver in larval ontogeny and evolution. Finally, we discuss some of the more general trends of crustacean larval development in light of the general patterns and latest knowledge on tetraconate and arthropod evolution.

Legacy of a Butterfly’s Parental Microbiome in Offspring Performance

ABSTRACT An insect’s phenotype can be influenced by the experiences of the parental generation. However, the effects of the parental symbiotic microbiome and host plant use on the offspring are unclear. We addressed this gap of knowledge by studying Pieris brassicae, a multivoltine butterfly species feeding on different brassicaceous plants across generations. We investigated how disturbance of the parental bacterial community by antibiotic treatment affects F1 larval traits. We tested the effects depending on whether F1 larvae are feeding on the same plant species as their parents or on a different one. The parental treatment alone had no impact on the biomass of F1 larvae feeding on the parental plant species. However, the parental treatment had a detrimental effect on F1 larval biomass when F1 larvae had a different host plant than their parents. This effect was linked to higher larval prophenoloxidase activity and greater downregulation of the major allergen gene (MA), a glucosinolate detoxification gene of P. brassicae. Bacterial abundance in untreated adult parents was high, while it was very low in F1 larvae from either parental type, and thus unlikely to directly influence larval traits. Our results suggest that transgenerational effects of the parental microbiome on the offspring’s phenotype become evident when the offspring is exposed to a transgenerational host plant shift. IMPORTANCE Resident bacterial communities are almost absent in larvae of butterflies and thus are unlikely to affect their host. In contrast, adult butterflies contain conspicuous amounts of bacteria. While the host plant and immune state of adult parental butterflies are known to affect offspring traits, it has been unclear whether also the parental microbiome imposes direct effects on the offspring. Here, we show that disturbance of the bacterial community in parental butterflies by an antibiotic treatment has a detrimental effect on those offspring larvae feeding on a different host plant than their parents. Hence, the study indicates that disturbance of an insect’s parental microbiome by an antibiotic treatment shapes how the offspring individuals can adjust themselves to a novel host plant.