Monsoonal wet season influences the migration tendency of a catadromous fish (barramundi Lates calcarifer).

Many animals exhibit partial migration, which occurs when populations contain coexisting contingents of migratory and resident individuals. This individual-level variation in migration behaviour may drive differences in growth, age at maturity and survival. Therefore, partial migration is widely considered to play a key role in shaping population demography. Otolith chemistry and microstructural analysis were used to identify the environmental- and individual-specific factors that influence migratory behaviour in the facultatively catadromous barramundi (Lates calcarifer) at two distinct life history stages: firstly, as juveniles migrating upstream into fresh water; and secondly, as adults or sub-adults returning to the estuarine/marine spawning habitat. Monsoonal climate played an important role in determining the migration propensity of juveniles: individuals born in the driest year examined (weak monsoon) were more than twice as likely to undergo migration to freshwater than those born in the wettest (strong monsoon) year. In contrast, the ontogenetic timing of return migrations to the estuary by adults and sub-adults was highly variable and not strongly associated with the environmental parameters examined. We propose that scarce resources within saline natal habitats during lower rainfall years may provide an ecological incentive for juveniles to migrate upstream, whereas more abundant resources in higher rainfall years may promote resident life histories within estuaries. We conclude that inter-annual climatic variation, here evidenced by monsoonal strength, likely plays an important role in driving the persistence of diversified life histories within wild barramundi populations.

Growth portfolios buffer climate-linked environmental change in marine systems.

Large-scale, climate-induced synchrony in the productivity of fish populations is becoming more pronounced in the world's oceans. As synchrony increases, a population's "portfolio" of responses can be diminished, in turn reducing its resilience to strong perturbation. Here we argue that the costs and benefits of trait synchronization, such as the expression of growth rate, are context dependent. Contrary to prevailing views, synchrony among individuals could actually be beneficial for populations if growth synchrony increases during favorable conditions, and then declines under poor conditions when a broader portfolio of responses could be useful. Importantly, growth synchrony among individuals within populations has seldom been measured, despite well-documented evidence of synchrony across populations. Here, we used century-scale time series of annual otolith growth to test for changes in growth synchronization among individuals within multiple populations of a marine keystone species (Atlantic cod, Gadus morhua). On the basis of 74,662 annual growth increments recorded in 13,749 otoliths, we detected a rising conformity in long-term growth rates within five northeast Atlantic cod populations in response to both favorable growth conditions and a large-scale, multidecadal mode of climate variability similar to the East Atlantic Pattern. The within-population synchrony was distinct from the across-population synchrony commonly reported for large-scale environmental drivers. Climate-linked, among-individual growth synchrony was also identified in other Northeast Atlantic pelagic, deep-sea and bivalve species. We hypothesize that growth synchrony in good years and growth asynchrony in poorer years reflects adaptive trait optimization and bet hedging, respectively, that could confer an unexpected, but pervasive and stabilizing, impact on marine population productivity in response to large-scale environmental change.

Smaller adult fish size in warmer water is not explained by elevated metabolism.

Fish and other ectotherms living in warmer waters often grow faster as juveniles, mature earlier, but become smaller adults. Known as the temperature-size rule (TSR), this pattern is commonly attributed to higher metabolism in warmer waters, leaving fewer resources for growth. An alternative explanation focuses on growth and reproduction trade-offs across temperatures. We tested these hypotheses by measuring growth, maturation, metabolism and reproductive allocation from zebrafish populations kept at 26 and 30°C across six generations. Zebrafish growth and maturation followed TSR expectations but were not explained by baseline metabolic rate, which converged between temperature treatments after a few generations. Rather, we found that females at 30°C allocated more to reproduction, especially when maturing at the smallest sizes. We show that elevated temperatures do not necessarily increase baseline metabolism if sufficient acclimation is allowed and call for an urgent revision of modelling assumptions used to predict population and ecosystem responses to warming.

Environmental drivers of recruitment in a tropical fishery: Monsoonal effects and vulnerability to water abstraction.

Fisheries and natural water resources across the world are under increasing pressure from human activity, including fishing and irrigated agriculture. There is an urgent need for information on the climatic/hydrologic drivers of fishery productivity that can be readily applied to management. We use a generalized linear mixed model framework of catch curve regression to resolve the key climatic/hydrological drivers of recruitment in Barramundi Lates calcarifer using biochronological (otolith aging) data collected from four river-estuary systems in the Northern Territory, Australia. These models were then used to generate estimates of the year class strength (YCS) outcomes of different water abstraction scenarios (ranging from 10% to 40% abstraction per season/annum) for two of the rivers in low, moderate, and high discharge years. Barramundi YCS displayed strong interannual variation and was positively correlated with regional monsoon activity in all four rivers. River-specific analyses identified strong relationships between YCS and several river-specific hydrology variables, including wet and dry season discharge and flow duration. Water abstraction scenario models based on YCS-hydrology relationships predicted reductions of >30% in YCS in several cases, suggesting that increased water resource development in the future may pose risks for Barramundi recruitment and fishery productivity. Our study demonstrates the importance of the tropical monsoon as a driver of Barramundi recruitment and the potential for detrimental impacts of increased water abstraction on fishery productivity. The biochronological and statistical approaches we used have the potential to be broadly applied to inform policy and management of water resource and fisheries.

Lifetime movement history is associated with variable growth of a potamodromous freshwater fish.

Directional or stabilising selection should drive the expression of a dominant movement phenotype within a population. Widespread persistence of multiple movement phenotypes within wild populations, however, suggests that individuals that move (movers) and those that do not (residents) can have commensurate performance. The costs and benefits of mover and resident phenotypes remain poorly understood. Here, we explored how the presence and timing of movements are correlated with annual somatic growth rates, a useful proxy for performance because it is easily measured and rapidly reflects environmental changes. We used otolith growth measurements and stable isotope analyses to recreate growth and among-reach movement histories of a partially migrating, long-lived freshwater fish, golden perch Macquaria ambigua. We compared the association between movement and growth at two temporal scales: (a) short-term (annual) differences in growth, in the years preceding, during or following movement; and (b) long-term (lifetime) differences in growth. Overall, 59% of individuals performed at least one among-reach movement, with these individuals subsequently more likely to move repeatedly throughout their lives. Movers grew faster than residents, with this difference most pronounced in the juvenile and early adult stages, when most movements occurred. Annual growth did not, however, change immediately prior to or following a specific movement event. Among-individual variation in growth was initially higher for residents than for movers but decreased with age, at a faster rate for residents than for movers, such that levels conformed after 5 years of age. Our results indicate that lifetime movement is linked to faster growth in the early years of a fish's life. These faster growing movers are likely to be larger at a given age, leading to numerous potential benefits. However, the persistence of resident phenotypes suggests that there is likely a cost-benefit trade-off to moving. The presence of multiple movement phenotypes may contribute to the resilience of populations by buffering against naturally and anthropogenically exacerbated environmental variability.

Faster juvenile growth promotes earlier sex change in a protandrous hermaphrodite (barramundi Lates calcarifer).

The relationship between growth and sexual maturation is central to understanding the dynamics of animal populations which exhibit indeterminate growth. In sequential hermaphrodites, which undergo post-maturation sex change, the size and age at which sex change occurs directly affects reproductive output and hence population productivity. However, these traits are often labile, and may be strongly influenced by heterogenous growth and mortality rates. We analysed otolith microstructure of a protandrous (i.e., male-to-female) fish (barramundi Lates calcarifer) to examine growth in relation to individual variation in the timing of sex change. Growth trajectories of individuals with contrasting life histories were examined to elucidate the direction and extent to which growth rate influences the size and age individuals change sex. Then, the relationships between growth rate, maturation schedules and asymptotic maximum size were explored to identify potential trade-offs between age at female maturity and growth potential. Rapid growth was strongly associated with decreased age at sex change, but this was not accompanied by a decrease in size at sex change. Individuals that were caught as large females grew faster than those caught as males, suggesting that fast-growing individuals ultimately obtain higher fitness and therefore make a disproportionate contribution to population fecundity. These results indicate that individual-level variation in maturation schedules is not reflective of trade-offs between growth and reproduction. Rather, we suggest that conditions experienced during the juvenile phase are likely to be a key determinant of post-maturation fitness. These findings highlight the vulnerability of sex-changing species to future environmental change and harvest.

Synergistic effects of harvest and climate drive synchronous somatic growth within key New Zealand fisheries.

Fisheries harvest has pervasive impacts on wild fish populations, including the truncation of size and age structures, altered population dynamics and density, and modified habitat and assemblage composition. Understanding the degree to which harvest-induced impacts increase the sensitivity of individuals, populations and ultimately species to environmental change is essential to ensuring sustainable fisheries management in a rapidly changing world. Here we generated multiple long-term (44-62 years), annually resolved, somatic growth chronologies of four commercially important fishes from New Zealand's coastal and shelf waters. We used these novel data to investigate how regional- and basin-scale environmental variability, in concert with fishing activity, affected individual somatic growth rates and the magnitude of spatial synchrony among stocks. Changes in somatic growth can affect individual fitness and a range of population and fishery metrics such as recruitment success, maturation schedules and stock biomass. Across all species, individual growth benefited from a fishing-induced release of density controls. For nearshore snapper and tarakihi, regional-scale wind and temperature also additively affected growth, indicating that future climate change-induced warming and potentially strengthened winds will initially promote the productivity of more poleward populations. Fishing increased the sensitivity of deep-water hoki and ling growth to the Interdecadal Pacific Oscillation (IPO). A forecast shift to a positive IPO phase, in concert with current harvest strategies, will likely promote individual hoki and ling growth. At the species level, historical fishing practices and IPO synergized to strengthen spatial synchrony in average growth between stocks separated by 400-600 nm of ocean. Increased spatial synchrony can, however, increase the vulnerability of stocks to deleterious stochastic events. Together, our individual- and species-level results show how fishing and environmental factors can conflate to initially promote individual growth but then possibly heighten the sensitivity of stocks to environmental change.

Stage-dependent effects of river flow and temperature regimes on the growth dynamics of an apex predator.

In the world's rivers, alteration of flow is a major driver of biodiversity decline. Global warming is now affecting the thermal and hydrological regimes of rivers, compounding the threat and complicating conservation planning. To inform management under a non-stationary climate, we must improve our understanding of how flow and thermal regimes interact to affect the population dynamics of riverine biota. We used long-term growth biochronologies, spanning 34 years and 400,000 km2 , to model the growth dynamics of a long-lived, apex predator (Murray cod) as a function of factors extrinsic (river discharge; air temperature; sub-catchment) and intrinsic (age; individual) to the population. Annual growth of Murray cod showed significant, curvilinear, life-stage-specific responses to an interaction between annual discharge and temperature. Growth of early juveniles (age 1+ and 2+ years) exhibited a unimodal relationship with annual discharge, peaking near median annual discharge. Growth of late juveniles (3+ to 5+) and adults (>5+) increased with annual discharge, with the rate of increase being particularly high in adults, whose growth peaked during years with flooding. Years with very low annual discharge, as experienced during drought and under high abstraction, suppress growth rates of all Murray cod life-stages. Unimodal relationships between growth and annual temperature were evident across all life stages. Contrary to expectations of the Temperature Size Rule, the annual air temperature at which maximum growth occurred increased with age. The stage-specific response of Murray cod to annual discharge indicates that no single magnitude of annual discharge is optimal for cod populations, adding further weight to the case for maintaining and/or restoring flow variability in riverine ecosystems. With respect to climate change impacts, on balance our results indicate that the primary mechanism by which climate change threatens Murray cod growth is through alteration of river flows, not through warming annual mean temperatures per se.

Tracking the resource pulse: Movement responses of fish to dynamic floodplain habitat in a tropical river.

Natural river floodplains are among the Earth's most biologically diverse and productive ecosystems but face a range of critical threats due to human disturbance. Understanding the ecological processes that support biodiversity and productivity in floodplain rivers is essential for their future protection and rehabilitation. Fish assemblage structure on tropical river floodplains is widely considered to be driven by dispersal limitation during the wet season and by environmental filtering and interspecific interactions during the dry season. However, the individual-level movement behaviours (e.g. site attachment, nomadism, homing) that regulate dispersal of fish on floodplains are poorly understood. We combined radiotelemetry and remote sensing to examine the movement behaviour of two large-bodied fishes (barramundi Lates calcarifer, forktail catfish Neoarius leptaspis) over the flood cycle in a tropical river-floodplain system in northern Australia to: (a) quantify movement responses in relation to dynamic habitat resources at a landscape scale; and (b) determine the extent of spatial 'reshuffling' of individual fish following the wet season. Both species altered their behaviour rapidly in response to changes in the availability and distribution of aquatic habitat, with most individuals undertaking extensive movements (up to ~27 km from the tagging location) on the inundated floodplain during the wet season. Although there was considerable individual variation in movement patterns, overall barramundi distributions closely tracked the extent of floodplain primary productivity, whereas catfish distributions were most closely associated with the extent of flooded area. Most individuals of both species exhibited homing back to previously occupied dry season refugia during the wet-to-dry transition, even though other potential refugia were available in closer proximity to wet season activity areas. We postulate that homing behaviour modulates temporal variation in fish assemblage composition and abundance and limits the transfer of aquatic-derived energy and nutrients into terrestrial food webs by reducing fish mortality on drying floodplains. Our study demonstrates the importance of quantifying individual-level behaviour across the three stages of dispersal (emigration, inter-patch movement, immigration) for our understanding of how animal movement influences energetic subsidies and other large-scale ecosystem processes.

Migration to freshwater increases growth rates in a facultatively catadromous tropical fish.

Diadromy is a form of migration where aquatic organisms undergo regular movements between fresh and marine waters for the purposes of feeding and reproduction. Despite having arisen in independent lineages of fish, gastropod molluscs and crustaceans, the evolutionary drivers of diadromous migration remain contentious. We test a key aspect of the 'productivity hypothesis', which proposes that diadromy arises in response to primary productivity differentials between marine and freshwater habitats. Otolith chemistry and biochronology data are analysed in a facultatively catadromous tropical fish (barramundi, Lates calcarifer) to determine the effect of freshwater residence on growth rates. Individuals that accessed freshwater grew ~ 25% faster on average than estuarine residents in the year following migration, suggesting that catadromy provides a potential fitness advantage over non-catadromous (marine/estuarine) life histories, as predicted by the productivity hypothesis. Although diadromous barramundi exhibited faster growth than non-diadromous fish, we suggest that the relative reproductive success of diadromous and non-diadromous contingents is likely to be strongly influenced by local environmental variability such as temporal differences in river discharge, and that this may facilitate the persistence of diverse life history strategies within populations.

Fishing constrains phenotypic responses of marine fish to climate variability.

Fishing and climate change are profoundly impacting marine biota through unnatural selection and exposure to potentially stressful environmental conditions. Their effects, however, are often considered in isolation, and then only at the population level, despite there being great potential for synergistic selection on the individual. We explored how fishing and climate variability interact to affect an important driver of fishery productivity and population dynamics: individual growth rate. We projected that average growth rate would increase as waters warm, a harvest-induced release from density dependence would promote adult growth, and that fishing would increase the sensitivity of somatic growth to temperature. We measured growth increments from the otoliths of 400 purple wrasse (Notolabrius funicola), a site-attached temperate marine reef fish inhabiting an ocean warming hotspot. These were used to generate nearly two decades of annually resolved growth estimates from three populations spanning a period before and after the onset of commercial fishing. We used hierarchical models to partition variation in growth within and between individuals and populations, and attribute it to intrinsic (age, individual-specific) and extrinsic (local and regional climate, fishing) drivers. At the population scale, we detected predictable additive increases in average growth rate associated with warming and a release from density dependence. A fishing-warming synergy only became apparent at the individual scale where harvest resulted in the 50% reduction of thermal growth reaction norm diversity. This phenotypic change was primarily caused by the loss of larger individuals that showed a strong positive response to temperature change after the onset of size-selective harvesting. We speculate that the dramatic loss of individual-level biocomplexity is caused by either inadvertent fisheries selectivity based on behaviour, or the disruption of social hierarchies resulting from the selective harvesting of large, dominant and resource-rich individuals. Whatever the cause, the removal of individuals that display a positive growth response to temperature could substantially reduce species' capacity to adapt to climate change at temperatures well below those previously thought stressful.

Female ornamentation and the fecundity trade-off in a sex-role reversed pipefish.

Sexual ornaments found only in females are a rare occurrence in nature. One explanation for this is that female ornaments are costly to produce and maintain and, therefore, females must trade-off resources related to reproduction to promote ornament expression. Here, we investigate whether a trade-off exists between female ornamentation and fecundity in the sex-role reversed, wide-bodied pipefish, Stigmatopora nigra. We measured two components of the disk-shaped, ventral-striped female ornament, body width, and stripe thickness. After controlling for the influence of body size, we found no evidence of a cost of belly width or stripe thickness on female fecundity. Rather, females that have larger ornaments have higher fecundity and thus accurately advertise their reproductive value to males without incurring a cost to fecundity. We also investigated the relationship between female body size and egg size and found that larger females suffer a slight decrease in egg size and fecundity, although this decrease was independent of female ornamentation. More broadly, considered in light of similar findings in other taxa, lack of an apparent fecundity cost of ornamentation in female pipefish underscores the need to revisit theoretical assumptions concerning the evolution of female ornamentation.

Evolutionary traps and range shifts in a rapidly changing world.

Humans are altering the environment at an unprecedented rate. Although behavioural plasticity has allowed many species to respond by shifting their ranges to more favourable conditions, these rapid environmental changes may cause 'evolutionary traps', whereby animals mistakenly prefer resources that reduce their fitness. The role of evolutionary traps in influencing the fitness consequences of range shifts remains largely unexplored. Here, we review these interactions by considering how climate change may trigger maladaptive developmental pathways or increase the probability of animals encountering traps. We highlight how traps could selectively remove some phenotypes and compromise population persistence. We conclude by highlighting emerging areas of research that would improve our understanding of when interactions between evolutionary traps and range shifts are likely to be most detrimental to animals.

Environmental change drives long-term recruitment and growth variation in an estuarine fish.

How individuals respond to environmental change determines the strength and direction of biological processes like recruitment and growth that underpin population productivity. Ascertaining the relative importance of environmental factors can, however, be difficult given the numerous mechanisms through which they affect individuals. This is especially true in dynamic and complex estuarine environments. Here, we develop long-term otolith-based indices of recruitment and growth for estuary perch Percalates colonorum (Bemm River, Australia), to explore the importance of intrinsic (individual, demographic) and extrinsic (hydrologic, climatic, density-dependent) factors in driving estuarine fish productivity. Analyses involved a novel zero-inflated specification of catch curve regression and mixed effects modelling. The 39 years of recruitment and 46 years of growth data, spanning a period of environmental change including severe drought, displayed considerable inter-annual variation. Recruitment success was strongly related to high freshwater inflows during the spawning season, suggesting that these conditions act as spawning cues for adults and potentially provide favourable conditions for larvae. Individuals displayed age-dependent growth, with highest rates observed at younger ages in years characterized by warm temperatures, and to a lesser degree, greater magnitude base inflow conditions. We detected systematic among-year-class growth differences, but these were not attributable to year class strength, suggesting that environmental conditions experienced by individuals as juveniles can have long-lasting effects of greater importance to population productivity than density-dependent growth responses. The primacy of temperature in driving growth variation highlights that under-appreciated climatic variation can affect estuarine fish productivity through direct physiological and indirect food web mechanisms. We predict that climatic warming will promote individual growth in southerly populations of P. colonorum but concurrently limit recruitment due to forecast reductions in spawning season river discharge. Disparate trait responses are likely in other fishes as they respond to multiple and changing environmental drivers, making predictions of future population productivity challenging.

Intraspecific variation in the growth and survival of juvenile fish exposed to Eucalyptus leachate.

Whilst changes in freshwater assemblages along gradients of environmental stress have been relatively well studied, we know far less about intraspecific variation to these same stressors. A stressor common in fresh waters worldwide is leachates from terrestrial plants. Leachates alter the physiochemical environment of fresh waters by lowering pH and dissolved oxygen and also releasing toxic compounds such as polyphenols and tannins, all of which can be detrimental to aquatic organisms. We investigated how chronic exposure to Eucalyptus leaf leachate affected the growth and survival of juvenile southern pygmy perch (Nannoperca australis) collected from three populations with different litter inputs, hydrology and observed leachate concentrations. Chronic exposure to elevated leachate levels negatively impacted growth and survival, but the magnitude of these lethal and sublethal responses was conditional on body size and source population. Bigger fish had increased survival at high leachate levels but overall slower growth rates. Body size also varied among populations and fish from the population exposed to the lowest natural leachate concentrations had the highest average stress tolerance. Significant intraspecific variation in both growth and survival caused by Eucalyptus leachate exposure indicates that the magnitude (but not direction) of these stress responses varies across the landscape. This raises the potential for leachate-induced selection to operate at an among-population scale. The importance of body size demonstrates that the timing of leachate exposure during ontogeny is central in determining the magnitude of biological response, with early life stages being most vulnerable. Overall, we demonstrate that Eucalyptus leachates are prevalent and potent selective agents that can trigger important sublethal impacts, beyond those associated with more familiar fish kills, and reiterate that dissolved organic carbon is more than just an energy source in aquatic environments.

Spatial variation in egg size and egg number reflects trade-offs and bet-hedging in a freshwater fish.

1. Maternal reproductive investment is thought to reflect a trade-off between offspring size and fecundity, and models generally predict that mothers inhabiting adverse environments will produce fewer, larger offspring. More recently, the importance of environmental unpredictability in influencing maternal investment has been considered, with some models predicting that mothers should adopt a diversified bet-hedging strategy whilst others a conservative bet-hedging strategy. 2. We explore spatial egg size and fecundity patterns in the freshwater fish southern pygmy perch (Nannoperca australis) that inhabits a diversity of streams along gradients of environmental quality, variability and predictability. 3. Contrary to some predictions, N. australis populations inhabiting increasingly harsh streams produced more numerous and smaller eggs. Furthermore, within-female egg size variability increased as environments became more unpredictable. 4. We argue that in harsh environments or those prone to physical disturbance, sources of mortality are size independent with offspring size having only a minor influence on offspring fitness. Instead, maternal fitness is maximized by producing many small eggs, increasing the likelihood that some offspring will disperse to permanent water. We also provide empirical support for diversified bet-hedging as an adaptive strategy when future environmental quality is uncertain and suggest egg size may be a more appropriate fitness measure in stable environments characterized by size-dependent fitness. These results likely reflect spatial patterns of adaptive plasticity and bet-hedging in response to both predictable and unpredictable environmental variance and highlight the importance of considering both trait averages and variance. 5. Reproductive life-history traits can vary predictably along environmental gradients. Human activity, such as the hydrological modification of natural flow regimes, alters the form and magnitude of these gradients, and this can have both ecological and evolutionary implications for biota adapted to now non-existent natural environmental heterogeneity.