Variability in water temperature affects trait-mediated survival of a newly settled coral reef fish

As animals with complex life cycles metamorphose from one stage to the next, carry-over effects from earlier stages can affect future mortality. To examine the relationship between early life history traits and survival, seven monthly cohorts of newly-settled bluehead wrasse Thalassoma bifasciatum were collected immediately after settlement and over sequential 3-day periods. Otolith analysis was used to quantify mean larval and juvenile growth rates, pelagic larval duration (PLD), and settlement size and condition of different age classes to identify the traits most important for survival. Overall, survivors tended to have shorter PLDs, to settle at smaller sizes and higher condition levels, and to exhibit faster early juvenile growth. Water temperature contributed to among-cohort variability in traits as warmer water led to faster larval and juvenile growth and shorter PLDs. Trait-specific fitness functions demonstrated that temperature can influence fitness by changing the nature of selection on each trait. Estimates of selection intensity revealed that settlement condition contributed the most to variation in fitness across cohorts, followed by juvenile growth. Frequent loss of low settlement condition individuals and occasional loss of the very highest condition fish suggest that particularly high settlement condition during the warmest temperatures may be detrimental. Not only does the quality of settlers vary over time, but selective loss of individuals with particular phenotypic traits is not pervasive and can vary with environmental conditions such as temperature.     

Environmental variability, early life-history traits, and survival of new coral reef fish recruits

For benthic marine organisms with complex life cycles, conditionsexperienced by pelagic larvae can influence juvenile survival.Trait-specific selective mortality has been documented in thelaboratory and field, yet our knowledge of the factors contributingto the existence, strength, and consistency of natural selectivemortality is limited. We compiled previously published and unpublisheddata on the common Caribbean coral reef fish, Thalassoma bifasciatum,recruiting to Barbados, West Indies, and the upper Florida Keysto examine how environmental variability during pelagic larvallife influences the distribution of early life-history traitsexhibited by new recruits. We explored how the scope of variabilityin otolith-derived traits such as larval growth, pelagic larvalduration (PLD), size and condition at settlement, and earlyjuvenile growth influences the degree to which mortality ofjuveniles is selective. At both locations, contrasting oceanographicconditions (periodic passage of large low-salinity North BrazilCurrent [NBC] rings near Barbados and seasonal variation inwater temperature at Florida) led to significant differencesin the early life-history traits of recruits. Mortality wasmost frequently selective for the two most variable traits,condition at settlement and early juvenile growth. Environmentalvariability (including variation in predation pressure and stress-inducingconditions) also likely influences juvenile mortality and consequentlythe degree to which selective loss of particular traits occurs.As we begin to better understand the spatial, temporal, andspecies-specific circumstances in which events occurring duringlarval life influence juvenile performance, studies must alsobe extended to examine how these processes are translated intoadult fitness.     

Survival against the odds: ontogenetic changes in selective pressure mediate growth-mortality trade-offs in a marine fish

For organisms with complex life cycles, variation among individuals in traits associated with survival in one life-history stage can strongly affect the performance in subsequent stages with important repercussions on population dynamics. To identify which individual attributes are the most influential in determining patterns of survival in a cohort of reef fish, we compared the characteristics of Pomacentrus amboinensis surviving early juvenile stages on the reef with those of the cohort from which they originated. Individuals were collected at hatching, the end of the planktonic phase, and two, three, four, six and eight weeks post-settlement. Information stored in the otoliths of individual fish revealed strong carry-over effects of larval condition at hatching on juvenile survival, weeks after settlement (i.e. smaller-is-better). Among the traits examined, planktonic growth history was, by far, the most influential and long-lasting trait associated with juvenile persistence in reef habitats. However, otolith increments suggested that larval growth rate may not be maintained during early juvenile life, when selective mortality swiftly reverses its direction. These changes in selective pressure may mediate growth-mortality trade-offs between predation and starvation risks during early juvenile life. Ontogenetic changes in the shape of selectivity may be a mechanism maintaining phenotypic variation in growth rate and size within a population.     

Variation in the effects of larval history on juvenile performance of a temperate reef fish

For organisms with complex life cycles, the transition between life stages can act as a significant demographic and selective bottleneck. Variation in developmental and growth rates among individuals present in one stage (e.g. larvae), due to initial differences in parental input and/or environmental conditions experienced, can propagate to future stages (e.g. juveniles), and such ‘carry‐over effects’ can shape fitness and phenotypic distributions within a population. However, variation in the strength of carry‐over effects between life stages and the intensity of selective mortality acting on intrinsic variation, and how these might be mediated by environmental variability in natural systems, is poorly known. Here, we evaluate variation in the strength to which larval growth histories can mediate juvenile performance (growth and survival), for a reef fish (Forsterygion lapillum) common to rocky reefs of New Zealand. We used otoliths to reconstruct demographic histories of recently settled fish that were sampled across cohorts, sites and microhabitats. We quantified sources of variation in the strength of carry‐over effects and selective mortality that operate on larval growth histories. We found overall that individuals that grew fast as larvae tended to experience proportional growth advantages as juveniles. However, the strength of growth advantages being maintained into the juvenile period varied among cohorts, sites and microhabitats. Specifically, a stronger growth advantage was found on some microhabitats (e.g. mixed stands of macroalgae) relative to others (e.g. monocultures of Carpophyllum maschalocarpum) for some cohorts and sites only. For other cohorts and sites, the degree of coupling between larval and juvenile growth rates was either indistinguishable between microhabitats or else not evident. Similarly, the intensity of growth‐based selective mortality varied among cohorts, sites and microhabitats: for the cohort and site where carry‐over effects differed between microhabitats, we also observed difference in the intensity to which fish with rapid larval growth rates were favoured. Overall, our results highlight how this spatial and temporal patchiness in extrinsic factors can interact with intrinsic variation of recruiting individuals to have a major influence on the resulting distribution of juveniles and their phenotypic traits.     

Variability in size-selective mortality obscures the importance of larval traits to recruitment success in a temperate marine fish

In fishes, the growth-mortality hypothesis has received broad acceptance as a driver of recruitment variability. Recruitment is likely to be lower in years when the risk of starvation and predation in the larval stage is greater, leading to higher mortality. Juvenile snapper, Pagrus auratus (Sparidae), experience high recruitment variation in Port Phillip Bay, Australia. Using a 5-year (2005, 2007, 2008, 2010, 2011) data set of larval and juvenile snapper abundances and their daily growth histories, based on otolith microstructure, we found selective mortality acted on larval size at 5 days post-hatch in 4 low and average recruitment years. The highest recruitment year (2005) was characterised by no size-selective mortality. Larval growth of the initial larval population was related to recruitment, but larval growth of the juveniles was not. Selective mortality may have obscured the relationship between larval traits of the juveniles and recruitment as fast-growing and large larvae preferentially survived in lower recruitment years and fast growth was ubiquitous in high recruitment years. An index of daily mortality within and among 3 years (2007, 2008, 2010), where zooplankton were concurrently sampled with ichthyoplankton, was related to per capita availability of preferred larval prey, providing support for the match–mismatch hypothesis. In 2010, periods of low daily mortality resulted in no selective mortality. Thus both intra- and inter-annual variability in the magnitude and occurrence of selective mortality in species with complex life cycles can obscure relationships between larval traits and population replenishment, leading to underestimation of their importance in recruitment studies.     

The impact of larval predators and competitors on the morphology and fitness of juvenile treefrogs

Studies of phenotypic plasticity typically focus on traits in single ontogenetic stages. However, plastic responses can be induced in multiple ontogenetic stages and traits induced early in ontogeny may have lasting effects. We examined how gray treefrog larvae altered their morphology in four different larval environments and whether different larval environments affected the survival, growth, development, and morphology of juvenile frogs at metamorphosis. We then reared these juveniles in terrestrial environments under high and low intraspecific competition to determine whether the initial differences in traits at metamorphosis affected subsequent survival and growth, whether the initial phenotypic differences converged over time, and whether competition in the terrestrial environment induced further phenotypic changes. Larval and juvenile environments both affected treefrog traits. Larval predators induced relatively deep tail fins and short bodies, but there was no impact on larval development. In contrast, larval competitors induced relatively short tails and long bodies, reduced larval growth, and slowed larval development. At metamorphosis, larval predators had no effect on juvenile growth or relative morphology while larval competitors produced juveniles that were smaller and possessed relatively shorter limbs and shorter bodies. After 1 month of terrestrial competition among the juvenile frogs, the initial differences in juvenile morphology did not converge. There were no differences in growth due to larval treatment but there were differences in survival. Individuals that experienced low competition as tadpoles experienced near perfect survival as juvenile frogs but individuals that experienced high competition as tadpoles suffered an 18% decrease in survival as juvenile frogs. There were also morphological responses to juvenile competition, but these changes appear to be due, at least in part, to allometric effects. Collectively, these results demonstrate that larval environments can have profound impacts on the traits and fitness of organisms later in ontogeny.     

Selection against late emergence and small offspring in Atlantic salmon (Salmo salar)

Timing of breeding and offspring size are maternal traits that may influence offspring competitive ability, dispersal, foraging, and vulnerability to predation and climatic conditions. To quantify the extent to which these maternal traits may ultimately affect an organism's fitness, we undertook laboratory and field experiments with Atlantic salmon (Salmo salar). To control for confounding effects caused by correlated traits, manipulations of the timing of fertilization combined with intraclutch comparisons were used. In the wild, a total of 1462 juveniles were marked at emergence from gravel nests. Recapture rates suggest that up to 83.5% mortality occurred during the first four months after emergence from the gravel nests, with the majority (67.5%) occurring during the initial period ending 17 days after median emergence. Moreover, the mortality was selective during this initial period, resulting in a significant phenotypic shift toward an earlier date of and an increased length at emergence. However, no significant selection differentials were detected thereafter, indicating that the critical episode of selection had occurred at emergence. Furthermore, standardized selection gradients indicated that selection was more intense on date of than on body size at emergence. Timing of emergence had additional consequences in terms of juvenile body size. Late-emerging juveniles were smaller than early-emerging ones at subsequent samplings, both in the wild and in parallel experiments conducted in seminatural stream channels, and this may affect success at subsequent size-selective episodes, such as winter mortality and reproduction. Finally, our findings also suggest that egg size had fitness consequences independent of the effects of emergence time that directly affected body size at emergence and, in turn, survival and size at later life stages. The causality of the maternal effects observed in the present study supports the hypothesis that selection on juvenile traits may play an important role in the evolution of maternal traits in natural populations.     

Characteristics of Settling Coral Reef Fish Are Related to Recruitment Timing and Success

Many marine populations exhibit high variability in the recruitment of young into the population. While environmental cycles and oceanography explain some patterns of replenishment, the role of other growth-related processes in influencing settlement and recruitment is less clear. Examination of a 65-mo. time series of recruitment of a common coral reef fish, Stegastes partitus, to the reefs of the upper Florida Keys revealed that during peak recruitment months, settlement stage larvae arriving during dark lunar phases grew faster as larvae and were larger at settlement compared to those settling during the light lunar phases. However, the strength and direction of early trait-mediated selective mortality also varied by settlement lunar phase such that the early life history traits of 2–4 week old recruit survivors that settled across the lunar cycle converged to more similar values. Similarly, within peak settlement periods, early life history traits of settling larvae and selective mortality of recruits varied by the magnitude of the settlement event: larvae settling in larger events had longer PLDs and consequently were larger at settlement than those settling in smaller pulses. Traits also varied by recruitment habitat: recruits surviving in live coral habitat (vs rubble) or areas with higher densities of adult conspecifics were those that were larger at settlement. Reef habitats, especially those with high densities of territorial conspecifics, are more challenging habitats for young fish to occupy and small settlers (due to lower larval growth and/or shorter PLDs) to these habitats have a lower chance of survival than they do in rubble habitats. Settling reef fish are not all equal and the time and location of settlement influences the likelihood that individuals will survive to contribute to the population.     

Genetic polymorphism and trade-offs in the early life-history strategy of the Pacific oyster, Crassostrea gigas (Thunberg, 1795): a quantitative genetic study

We investigated genetic variability and genetic correlations in early life-history traits of Crassostrea gigas. Larval survival, larval development rate, size at settlement and metamorphosis success were found to be substantially heritable, whereas larval growth rate and juvenile traits were not. We identified a strong positive genetic correlation between larval development rate and size at settlement, and argue that selection could optimize both age and size at settlement. However, trade-offs, resulting in costs of metamorphosing early and large, were suggested by negative genetic correlations or covariances between larval development rate/size at settlement and both metamorphosis success and juvenile survival. Moreover, size advantage at settlement disappeared with time during the juvenile stage. Finally, we observed no genetic correlations between larval and juvenile stages, implying genetic independence of life-history traits between life-stages. We suggest two possible scenarios for the maintenance of genetic polymorphism in the early life-history strategy of C. gigas.     

Early thermal history significantly affects the seasonal hyperplastic process occurring in the myotomal white muscle of <Emphasis Type="Italic">Dicentrarchus labrax</Emphasis> juveniles

The effect of early (embryonic and larval) thermal history on subsequent (juvenile) white muscle hyperplasia was studied in a teleost fish, the European sea bass (Dicentrarchus labrax L.). D. labrax, incubated and reared at constant temperatures of 13°C, 15°C or 20°C from the embryonic stage of half epiboly up to 18–19 mm in total length, were transferred to ambient seawater temperature and reared for the subsequent 14 months on commercial feed. The somatic growth of juveniles was linked to annual variations of ambient seawater temperature and inversely related to early rearing temperature, so that, after 14 months, the juveniles originally reared at low temperatures had compensated for the growth retardation experienced during early life. The white muscle growth process of juveniles was quantified after two periods of growth opportunity at ambient seawater temperature (100 and 400 days post-transfer) as well as, in order to follow total-length-dependent effects of early temperature and to discriminate total-length-independent effects of early temperature, on juveniles from the three batches sampled at six successive equivalent total lengths (31–33, 84–88, 141–145, 166–172, 196–206 and 211–220 mm). Our data demonstrate the existence of a seasonal recruitment of new white muscle fibres when seawater temperature increases and of a shrinkage of the largest white muscle fibres during the winter months. The seasonal recruitment of new white muscle fibres occurring in juveniles is linked to their early rearing temperature. Juveniles originating from low temperatures have a higher and longer capacity to recruit new white muscle fibres when seawater temperature increases, supporting their better somatic growth. This finding is discussed in relation to the early (embryonic and larval) myogenic processes of the three populations and is related to their sex ratio. This work was financed by the European Commission, FAIR Program no. PL96–1941 “Muscle Ontogeny in Sea bass and Trout”.     

Selective predation for low body condition at the larval-juvenile transition of a coral reef fish

Mortality is known to be high during the transition from larval to juvenile life stages in organisms that have complex life histories. We are only just beginning to understand the processes that influence which individuals survive this period of high mortality, and which traits may be beneficial. Here we document a field experiment that examines the selectivity of predation immediately following settlement to the juvenile population in a common tropical fish, Pomacentrus amboinensis (Pomacentridae). Newly metamorphosed fish were tagged and randomly placed onto replicated patches of natural habitat cleared of resident fishes. After exposure to transient predators for 3 days, fish were recollected and the attributes of survivors from patch reefs that sustained high mortality were compared to individuals from patch reefs that experienced low mortality. Seven characteristics of individuals, which were indicative of previous and present body condition, were compared between groups. Predation was found to be selective for fish that grew slowly in the latter third of their larval phase, were low in total lipids, and had a high standardized weight (Fultons K). Traits developed in the larval phase can strongly influence the survival of individuals over this critical transition period for organisms with complex life cycles.     

Temperature-induced shifts in selective pressure at a critical developmental transition

Selective mortality within a population, based on the phenotype of individuals, is the foundation of the theory of natural selection. We examined temperature-induced shifts in the relationships among early life history traits and survivorship over the embryonic and larval stages of a tropical damselfish, Pomacentrus amboinensis. Our experiments show that temperature determines the intensity of selective mortality, and that this changes with ontogeny. The size of energy stores determined survival through to hatching, after which egg size became a good indicator of fitness as predicted by theoretical models. Yet, the benefits associated with egg size were not uniform among test temperatures. Initial egg size positively influenced larval survival at control temperature (29 °C). However, this embryonic trait had no effect on post-hatching longevity of individuals reared at the higher (31 °C) and lower (25 °C) end of the temperature range. Overall, our findings indicate that the outcome of selective mortality is strongly dependent on the interaction between environment conditions and intrinsic developmental schedules.     

Juvenile growth and survival under dietary restriction: are males and females equal?

The effects of food availability on life-history traits may be direct or delayed and may vary between the sexes. We evaluated the effects of dietary restriction early in life on growth and survival of male and female juveniles in the common lizard ( Lacerta vivipara ) and surveyed the literature on sex-specific sensitivity to the environment in vertebrates. Juvenile lizards were reared in the laboratory during one month following birth under full feeding or under dietary restriction. They were then released in two outdoor enclosures, where we compared growth and survival between treatments during one year. Low food availability early in life led to lower body growth in a direct, but not delayed, manner. The absence of compensatory growth in juveniles that experienced dietary restriction might be explained by their reduced competitiveness. Dietary restriction had a strongly negative, delayed effect on survival up to the age of one year that was mediated by selection against smaller individuals. Effects of dietary restriction were not sex-specific, as expected from the similar energetic requirements of male and female juveniles. Hence, food availability has long-lasting consequences on life-history traits that might influence population dynamics in this species.     

Selective mortality of larval Japanese seabass in Ariake Bay, Japan

This article describes the growth, mortality, and selection patterns in early larval stage of Japanese seabass Lateolabrax japonicus in Ariake Bay, Kyushu, Japan. Japanese seabass larvae were collected from the spawning ground in December 2007, and juveniles were collected from the nursery ground in March 2008. Otoliths were analyzed to produce daily records of size-at-age and growth rate that were compared between larvae and juveniles to determine whether selective mortality occurred. A weight-specific growth coefficient (G) and instantaneous mortality coefficient (M) were computed, and the recruitment potential was evaluated from the ratio of M:G. Selection for fast-growing and bigger larvae was evident during the 5–14 days after hatching (DAH). Selective mortality acted to preferentially remove fish that were slow growing and/or relatively small members of the cohort at least during the period examined. Trends in the growth rate differences between larvae and juveniles suggested that the selection process continued beyond 14 DAH although the exact duration over which selective mortality occurred was unknown. Survivors of Japanese seabass exhibited traits consistent with all aspects of the ‘growth–mortality’ hypothesis: faster growth, larger size-at-age, and shorter larval stage duration (LSD), i.e., larvae with faster growth, bigger size-at-age and a shorter LSD selectively survived the larval period. Larvae had higher M (0.323) than G (0.159), resulting in the M:G ratio of >1.0 (M:G = 2.031), suggesting that the larval cohort was rapidly losing biomass. Future studies should look to determine the duration over which selective mortality occurs and the timing of transition between M and G (M = G) and accumulation of cohort biomass.     

The evolution of adult pollen feeding did not alter postembryonic growth in Heliconius butterflies

For many animals, the availability and provision of dietary resources can vary markedly between juvenile and adult stages, often leading to a temporal separation of nutrient acquisition and use. Juvenile developmental programs are likely limited by the energetic demands of many adult tissues and processes with early developmental origins. Enhanced dietary quality in the adult stage may, therefore, alter selection on life history and growth patterns in juvenile stages. Heliconius are unique among butterflies in actively collecting and digesting pollen grains, which provide an adult source of essential amino acids. The origin of pollen feeding has therefore previously been hypothesized to lift constraints on larval growth rates, allowing Heliconius to spend less time as larvae when they are most vulnerable to predation. By measuring larval and pupal life‐history traits across three pollen‐feeding and three nonpollen‐feeding Heliconiini, we provide the first test of this hypothesis. Although we detect significant interspecific variation in larval and pupal development, we do not find any consistent shift associated with pollen feeding. We discuss how this result may fit with patterns of nitrogen allocation, the benefits of nitrogenous stores, and developmental limitations on growth. Our results provide a framework for studies aiming to link innovations in adult Heliconius to altered selection regimes and developmental programs in early life stages.     

Disentangling the Evolution of Early and Late Life History Traits in Humans

Some aspects of human life history are unique among primates. Most notably, humans have a younger weaning age, a later age at first parturition, a shorter female reproductive period, and a longer lifespan than other living hominoid species. Obtaining a better understanding of when and how life history changed during human evolution is important to those studying the evolutionary developmental biology of extinct hominins, as life history traits pace developmental processes. Life history traits are thought to be linked via tradeoffs, such that changes in early life history traits directly affect those that follow later in life, and vice versa. However, it is also worth considering how changes to a single life history trait may indirectly affect other traits by way of modifying selective pressures acting on individuals and groups. For example, because they affect the size and demographic structure of a group, late life history traits (e.g., lifespan) may also affect the evolution of life history traits that occur earlier in life, but by modifying selective pressures acting on juveniles rather than by triggering a physiological tradeoff. This review marks an effort to begin to disentangle the ways in which early and late life history traits may affect each other both directly and indirectly. We concentrate on female life history characteristics. First, we review previous research on the evolution of the postmenopausal lifespan in women. Next we discuss recent findings concerning the relationship between the optimal length of the female reproductive period, mortality, and weaning age that show that selection favors a shorter female reproductive period in the presence of a younger weaning age. We discuss the implications this finding holds for understanding the evolution of life history traits that are of particular interest to developmental biologists.     

Linking life history traits in successive phases of a complex life cycle: effects of larval biomass on early juvenile development in an estuarine crab, Chasmagnathus granulata

In marine benthic invertebrates with complex life cycles, recruitment success, juvenile survival, and growth may be affected by variation in both maternal factors and environmental conditions prevailing during preceding embryonic or larval development. In an estuarine crab, Chasmagnathus granulata, previous investigations have shown that initial larval biomass is positively correlated with the biomass of recently extruded eggs, and it depends also on the salinity experienced during embryogenesis. Biomass at hatching has consequences for the subsequent larval development which, in this species, comprises two alternative developmental pathways with four or five zoeal instars (short or long pathway) and a megalopa. Larvae hatching with a lower than average biomass tend to develop through the long pathway and metamorphose to megalopae with higher biomass. In the present study, we show experimentally that the long pathway produces also significantly larger juveniles (crab size measured as carapace width, biomass as dry mass, carbon and nitrogen contents). Compared with juveniles originating from the short pathway, those from the long pathway showed in successive instars longer moulting cycles and larger carapace width, but lower size increments at ecdysis. In consequence, differences in size or biomass of long pathway vs short pathway crabs tended to disappear in later instars (after stage V). Furthermore, we tested in juveniles the tolerance of starvation at three salinities (5‰, 15‰, 32‰). Tolerance of starvation was significantly higher in juveniles originating from the long pathway, indicating higher energy reserves. While salinity played only a minor role for survival, it exerted significant effects on the time of moulting to the second juvenile instar, regardless of the preceding developmental pathway. The biomass of first juveniles obtained from the short pathway showed a significant positive correlation with the biomass of the freshly hatched zoea I, but not in those from the long pathway. In conclusion, the fitness of juvenile C. granulata is linked with previous developmental processes and environmental conditions during the embryonic and larval phase. Hence, a better understanding and prediction of the recruitment success of marine benthic invertebrates with a complex life cycle may require more comprehensive life‐history investigations.     

Early life histories of three sympatric sticklebacks in a salt-marsh

The effects of abiotic factors (water level, temperature, dissolved oxygen and salinity) and interspecific competition on the survival and growth of early life stages of three sympatric sticklebacks, Gasterosteus aculeatus , G. wheallandi and Pungitius pungitius , were studied in salt-marsh tide pools near Isle Verte, Quebec. Significant year-to-year variation was observed in the percentage of live eggs per nest, egg incubation times, and the growth and condition of juveniles for all three species. These differences were associated with variations in temperature, oxygen and salinity in the pools. Comparisons of egg survival and juvenile growth rates were also made for G. aculeatus living in two contrasting habitats of the salt-marsh, the harsh and variable tide pools, and a less harsh, less variable freshwater site. Egg incubation times were longer, and the percentage of live eggs per nest lower, in the river than in the pools. However, river juveniles had higher growth rates and condition factors than pool juveniles. Results of experiments to study interspecific competition between juvenile G. aculaetus and juvenile G. wheatlandi were inconsistent, and it is suggested that the effects of abiotic factors on the survival and growth of early life stages in sticklebacks are more important than competition at this site.     

Phenotypic variation and selective mortality as major drivers of recruitment variability in fishes

An individual's phenotype will usually influence its probability of survival. However, when evaluating the dynamics of populations, the role of selective mortality is not always clear. Not all mortality is selective, patterns of selective mortality may vary, and it is often unknown how selective mortality compares or interacts with other sources of mortality. As a result, there is seldom a clear expectation for how changes in the phenotypic composition of populations will translate into differences in average survival. We address these issues by evaluating how selective mortality affects recruitment of fish populations. First, we provide a quantitative review of selective mortality. Our results show that most of the mortality during early life is selective, and that variation in phenotypes can have large effects on survival. Next, we describe an analytical framework that accounts for variation in selection, while also describing the amount of selective mortality experienced by different cohorts recruiting to a single population. This framework is based on reconstructing fitness surfaces from phenotypic selection measurements, and can be employed for either single or multiple traits. Finally, we show how this framework can be integrated with models of density‐dependent survival to improve our understanding of recruitment variability and population dynamics.     

Larval growth history determines juvenile growth and survival in a tropical marine fish

Processes that occur around the transition between larval and juvenile life‐stages can have a major effect on the population dynamics of organisms with complex life cycles. We explore the roles of larval history and selective post‐settlement mortality in determining the growth and survival of newly settled individuals of the damselfish, Pomacentrus amboinensis (Pomacentridae). Specifically, we determine whether the direction and intensity of selection on the recruits differs among various size‐classes of predators. A mark‐recapture study showed that individuals who survived 9 or more days were significantly larger at settlement than those that died within the first day (12.3 vs 11.9 mm SL), when mortality was highest (25% d^?1). Censuses revealed that the species and size composition of piscivores differed markedly between two reef habitats where P. amboinensis was common. A cage experiment, conducted in both habitats, manipulated the sizes of predators that could access newly settled P. amboinensis to determine whether the resulting mortality of the recently settled fish was influenced by larval growth history or size at settlement. Ten days after the start of the experiment individuals that grew slowly in the second half of their larval life had been lost from most of the experimental treatments. Small fish were also selectively lost from the coarse‐mesh cage on the reef base. Significant positive relationships between pre‐ and post‐settlement growth rates were found in both habitats for the fine mesh cages, cage controls and open patch reefs. This relationship was reversed in the coarse mesh cages in both habitats. This growth compensation was facilitated through the action of a particular size range of predators, whose impact was disrupted or masked in the open treatments by the action of a diverse predator pool. The present study underscores the complexity of the processes that influence the early post‐transition growth and survival in organisms with complex life‐histories.