Fitness landscape of Atlantic cod shaped by harvest selection and natural selection

Harvesting may lead to evolutionary changes in life histories on a contemporary time scale, changes that could be maladaptive in natural contexts. However, our understanding of the strength and direction of harvest-induced selection versus natural selection is still limited, partly due to the difficulty of tracking the fate of individuals in the wild. Here, we present direct estimates of harvest mortality, natural mortality and site fidelity of coastal Atlantic cod (Gadus morhua) from the Norwegian Skagerrak coast. Furthermore, we present standardised selection differentials for fish body size. Estimates are obtained from acoustic telemetry, where we continuously monitored fish (n = 60) within a semi-sheltered area using a network of 25 listening stations. To obtain additional information about harvested cod, all fish (body size: 30–66 cm) were also tagged with traditional T-bar tags with a printed reward of 500 NOK (60 E). We estimate that 75% of the fish died within the study area during 1 year. Fishing mortality was markedly higher than natural mortality. Together, recreational fishers and commercial fishers caught at least 50% of the tagged fish during 1 year. Standardised selection differentials showed that fisheries targeted larger fish (i.e. favoured the survival of smaller fish), while natural selection favoured the survival of larger fish. Albeit on a small scale, we provide empirical evidence that harvesting can have a dominant influence on the fitness landscape experienced by a marine fish such as the Atlantic cod. We suggest that no-take marine reserves may help to counter evolutionary impacts of harvesting in the ocean.     

Consistent size‐independent harvest selection on fish body shape in two recreationally exploited marine species

Harvesting wild animals may exert size‐independent selection pressures on a range of morphological, life history, and behavioral traits. Most work so far has focused on selection pressures on life history traits and body size as morphological trait. We studied here how recreational fishing selects for morphological traits related to body shape, which may correlate with underlying swimming behavior. Using landmark‐based geometric morphometrics, we found consistent recreational fishing‐induced selection pressures on body shape in two recreationally exploited marine fish species. We show that individuals with larger‐sized mouths and more streamlined and elongated bodies were more vulnerable to passively operated hook‐and‐line fishing independent of the individual's body size or condition. While the greater vulnerability of individuals with larger mouth gapes can be explained by the direct physical interaction with hooks, selection against streamlined and elongated individuals could either involve a specific foraging mode or relate to underlying elevated swimming behavior. Harvesting using passive gear is common around the globe, and thus, size‐independent selection on body shape is expected to be widespread potentially leaving behind individuals with smaller oral gapes and more compact bodies. This might have repercussions for food webs by altering foraging and predation.     

Biology of extinction risk in marine fishes

We review interactions between extrinsic threats to marine fishes and intrinsic aspects of their biology that determine how populations and species respond to those threats. Information is available on the status of less than 5% of the world's approximately 15500 marine fish species, most of which are of commercial importance. By 2001, based on data from 98 North Atlantic and northeast Pacific populations, marine fishes had declined by a median 65% in breeding biomass from known historic levels; 28 populations had declined by more than 80%. Most of these declines would be sufficient to warrant a status of threatened with extinction under international threat criteria. However, this interpretation is highly controversial, in part because of a perception that marine fishes have a suite of life history characteristics, including high fecundity and large geographical ranges, which might confer greater resilience than that shown by terrestrial vertebrates. We review 15 comparative analyses that have tested for these and other life history correlates of vulnerability in marine fishes. The empirical evidence suggests that large body size and late maturity are the best predictors of vulnerability to fishing, regardless of whether differences among taxa in fishing mortality are controlled; there is no evidence that high fecundity confers increased resilience. The evidence reviewed here is of direct relevance to the diverse criteria used at global and national levels by various bodies to assess threat status of fishes. Simple life history traits can be incorporated directly into quantitative assessment criteria, or used to modify the conclusions of quantitative assessments, or used as preliminary screening criteria for assessment of the approximately 95% of marine fish species whose status has yet to be evaluated either by conservationists or fisheries scientists.     

The importance of terrestrial subsidies in stream food webs varies along a stream size gradient

Cross system subsidies of energy and materials can be a substantial fraction of food web fluxes in ecosystems, especially when autochthonous production is strongly limited by light or nutrients. We explored whether assimilation of terrestrial energy varied in specific consumer taxa collected from streams of different sizes and resource availabilities. Since headwater streams are often unproductive, we expected that inputs from surrounding terrestrial systems (i.e. leaf litter, terrestrial invertebrates) would be a more important food source for consumers than in mid‐size rivers that have more open canopies and higher amounts of primary production available for consumers. We collected basal resources, invertebrates, and fish along a gradient in stream size in the Adirondack Mountains (NY, USA) and in Trinidad and Tobago and analyzed all samples for hydrogen isotopes as a means of differentiating biomass derived from allochthonous versus autochthonous sources. We found significant differences in allochthonous energy use within individual consumer taxa, showing that some taxa range from being entirely allochthonous to entirely autochthonous depending on where they were collected on the stream size gradient (grazers and collector–gatherer functional feeding groups), while other taxa are relatively fixed in the source of energy they assimilate (shredder and predator functional feeding groups). Consistent with expectations, allochthonous energy use was positively correlated with canopy cover in both regions for most feeding groups, with individuals from small, shaded streams having a more pronounced allochthonous signal than individuals collected from larger streams with less canopy cover. However, consumers in the shredder/detritivore feeding group did not vary among sites in their allochthonous energy use, and had a mostly allochthonous signal regardless of canopy cover and algal biomass. Our results demonstrate that the importance of energy from terrestrial subsidies can vary markedly but are similar in both temperate and tropical streams, suggesting a widely consistent pattern.     

Predator-induced life-history changes and the coexistence of five taxa in a Daphnia species complex

Interspecific hybridization is common in water fleas of the Daphnia galeata species complex (e.g. D. galeata , D. cucullata , D. hyalina and their interspecific hybrids). We studied the effect of fish on the life histories of five taxa of this species complex originating from the Plußsee, northern Germany. Using four clones per taxon, we found that fish kairomones reduce size at birth and size at maturity significantly. For size at maturity larger taxa showed a significantly stronger reaction to fish kairomones than the smaller taxa. With respect to the intrinsic rate of increase, r , we compared two predation regimes (positive size selective and not selective). We found that under the fish predation regime most clones had a higher r when cultured with fish kairomones, leading to a higher r , and a stronger reaction for the smaller taxa. We conclude that fish predation might be an important factor influencing the co-occurrence of Daphnia parental taxa with their hybrids.     

Life history correlates of responses to fisheries exploitation

We use an approach based on phylogenetic comparisons to identify life history correlates of abundance trends in 18 intensively exploited fish stocks from the north-east Atlantic. After accounting for differences in fishing mortality, we show that those fishes that have decreased in abundance compared with their nearest relatives mature later, attain a larger maximum size, and exhibit significantly lower potential rates of population increase. Such trends were not evident in a more traditional cross-species analysis. This is the first phylogenetically independent evidence to link life histories with abundance trends, and provides a quantitative basis for assessing vulnerability of fish populations to exploitation. Our approach can be applied to the conservation and management of other exploited taxa.     

Selective and genetic constraints on the evolution of body size in a stream-dwelling salmonid fish

To examine constraints on evolution of larger body size in two stunted populations of brook charr (Salvelinus fontinalis) from a single river in Cape Race, Newfoundland, Canada, we measured viability selection acting on length-at-age traits, and estimated quantitative genetic parameters in situ (following reconstruction of pedigree information from microsatellite data). Furthermore we tested for phenotypic differentiation between the populations, and for association of high juvenile growth with early maturity that is predicted by life history theory. Within each population, selection differentials and estimates of heritabilities for length-at-age traits suggested that evolution of larger size is prevented by both selective and genetic constraints. Between the populations, phenotypic differentiation was found in length-at-age and age of maturation traits, whereas early maturation was associated with increased juvenile growth (relative to adult growth) both within and between populations. The results suggest an adaptive plastic response in age of maturation to juvenile growth rates that have a largely environmental basis of determination.     

Evidence for small scale variation in the vertebrate brain: mating strategy and sex affect brain size and structure in wild brown trout (Salmo trutta)

The basis for our knowledge of brain evolution in vertebrates rests heavily on empirical evidence from comparative studies at the species level. However, little is still known about the natural levels of variation and the evolutionary causes of differences in brain size and brain structure within‐species, even though selection at this level is an important initial generator of macroevolutionary patterns across species. Here, we examine how early life‐history decisions and sex are related to brain size and brain structure in wild populations using the existing natural variation in mating strategies among wild brown trout (Salmo trutta). By comparing the brains of precocious fish that remain in the river and sexually mature at a small size with those of migratory fish that migrate to the sea and sexually mature at a much larger size, we show, for the first time in any vertebrate, strong differences in relative brain size and brain structure across mating strategies. Precocious fish have larger brain size (when controlling for body size) but migratory fish have a larger cerebellum, the structure in charge of motor coordination. Moreover, we demonstrate sex‐specific differences in brain structure as female precocious fish have a larger brain than male precocious fish while males of both strategies have a larger telencephalon, the cognitive control centre, than females. The differences in brain size and structure across mating strategies and sexes thus suggest the possibility for fine scale adaptive evolution of the vertebrate brain in relation to different life histories.     

Exploitation of the same trophic link favors convergence of larval life-history strategies in complex life cycle helminths

Switching from one host to the next is a critical life-history transition in parasites with complex life cycles. Growth and mortality rates are thought to influence the optimal time and size at transmission, but these rates are difficult to measure in parasites. The parasite life cycle, in particular the trophic link along which transmission occurs, may be a reasonable proxy for these rates, leading to the hypothesis that life cycle should shape life-history strategy. We compiled data on the size and age at infectivity for trophically transmitted helminths (i.e., acanthocephalans, cestodes, and nematodes), and then categorized species into trophic links (e.g., planktonic crustaceans to fish, insects to terrestrial vertebrates, etc.). Comparative analyses that explicitly included stabilizing selection within trophic links fit the data significantly better than random walk models, indicating that parasites with different life cycles have different optimal times/sizes for host switching. The major helminth groups have often independently evolved similar life cycles, and we show that this has frequently led to convergent and/or parallel evolution of size and age at infectivity. This suggests that for particular life cycles there are universal optimal transmission strategies, applicable to widely divergent taxa, although the cases of parallelism might indicate that lineage-specific constraints sometimes prevent evolution to a single adaptive peak.     

Nest predators and the evolution of avian reproductive strategies: a comparison of Australian and New Zealand birds

Summary Nest predation has been considered an important factor in the evolution of avian life histories: smaller clutches and shorter incubation and nestling periods are expected where nest predation has significant effects on reproductive success. Unlike the Australian avifauna, terrestrial New Zealand birds have evolved in the absence of reptilian and mammalian predators. Here we compare the reproductive strategies of terrestrial native New Zealand birds with those of their Australian sister taxa. In 11 of 14 comparisons, New Zealand birds were larger than their Australian relatives, but we did not find any significant differences in reproductive tactics between the two regions, a result inconsistent with the nest predation hypothesis. We discuss several reasons why this may be so. One possibility is that selection imposed on avian life history tactics by mammalian predators following the arrival of humans in New Zealand has led to strategies similar to those adopted in Australia.     

Egg size and emergence timing affect morphology and behavior in juvenile Chinook Salmon,Oncorhynchus tshawytscha

Variation in early life history traits often leads to differentially expressed morphological and behavioral phenotypes. We investigated whether variation in egg size and emergence timing influence subsequent morphology associated with migration timing in juvenile spring Chinook Salmon, Oncorhynchus tshawytscha. Based on evidence for a positive relationship between growth rate and migration timing, we predicted that fish from small eggs and fish that emerged earlier would have similar morphology to fall migrants, while fish from large eggs and individuals that emerged later would be more similar to older spring yearling migrants. We sorted eyed embryos within females into two size categories: small and large. We collected early and late‐emerging juveniles from each egg size category. We used landmark‐based geometric morphometrics and found that egg size appears to drive morphological differences. Egg size shows evidence for an absolute rather than relative effect on body morphology. Fish from small eggs were morphologically more similar to fall migrants, while fish from large eggs were morphologically more similar to older spring yearling migrants. Previous research has shown that the body morphology of fish that prefer the surface or bottom location in a tank soon after emergence also correlates with the morphological variations between wild fall and spring migrants, respectively. We found that late‐emerging fish spent more time near the surface. Our study shows that subtle differences in early life history characteristics may correlate with a diversity of future phenotypes.     

Physiological performance and mating system in Clarkia (Onagraceae): Does phenotypic selection predict divergence between sister species?

? Premise of the study: The evolution of self-fertilization often occurs in association with other floral, life history, and fitness-related traits. A previous study found that field populations of Clarkia exilis (a predominantly autogamous selfer) and its sister species, Clarkia unguiculata (a facultative outcrosser) differ in mean photosynthetic rates and instantaneous water use efficiency (WUE(i)). Here, we investigate the strength and direction of selection on these traits in multiple populations of each taxon to determine whether natural selection may contribute to the phenotypic differences between them. ? Methods: In spring 2008, we measured instantaneous gas exchange rates in nine populations during vegetative growth (Early) and/or during flowering (Late). We conducted selection gradient analyses and estimated selection differentials within populations and across pooled conspecific populations to evaluate the strength, direction, and consistency of selection on each trait early and late in the season. ? Key results: The direction and relative strength of selection on photosynthetic rates in these taxa corresponds to the phenotypic difference between them; C. exilis has higher photosynthetic rates than C. unguiculata, as well as stronger, more consistent selection favoring rapid photosynthesis throughout the growing season. Patterns of selection on transpiration, WUE(i), and the timing of flowering progression are less consistent with phenotypic differences (or lack thereof) between taxa. ? Conclusions: We detected several examples where selection was consistent with the phenotypic divergence between sister taxa, but there were also numerous instances that were equivocal or in which selection did not predict the realized phenotypic difference between taxa.     

Invertebrate mesopredators are larger in streams with fish

Species with complex life cycles (e.g., aquatic larvae, terrestrial adults) are expected to shorten the time spent in the larval stage if mortality risks are high, a trade-off that lowers predation risk at the cost of reduced time for growth and thus smaller adult size. We tested these predictions by comparing the timing of and size at emergence for two relatively large and common invertebrate mesopredator species (Isoperla montana and Rhyacophila vibox) that inhabit small coastal streams, with and without predatory fish, in eastern Canada. Contrary to expectations based on predation risk–foraging trade-off theory, individuals of both invertebrate species tended to be larger rather than smaller in streams with fish than in fishless streams. The patterns were consistent, however, with the expected ecological effects of top predators on food webs, where fish lower abundances of invertebrate mesopredators, increasing resource availability and thus growth rates for the remaining individuals. We conclude that variation among streams in size at emergence is better explained by the impact of fish on resource availability than to behavioural or life history trade-offs occurring under risk of predation.     

Differences in dispersal- and colonization-related traits between taxa from the freshwater and the terrestrial realm

The aquatic and terrestrial realms differ in many physical properties that not only require specific physiological adaptations but also cause differences in dispersal options. We thus expect that life-history traits related to dispersal and colonization are under selection pressure because freshwater habitats are more isolated and thus more difficult to reach. We compared traits from European databases of three taxonomic groups along the passive–active dispersal gradient: plants (Plantes), snails (Mollusca: Gastropoda: Prosobranchia et Pulmonata) and hoverflies (Diptera: Syrphidae), all of which have both terrestrial and freshwater species (plants and snails) or early life stages (hoverflies). Aquatic taxa seem to be more successful long-distance dispersers than are terrestrial taxa. Our analysis also revealed lower numbers of seeds or eggs produced in the aquatic habitats. However, aquatic taxa often allocate resources to offspring guarding (vegetative propagules in plants, egg capsules in snails) and breeding-site selection (syrphids). Colonization of the aquatic realm is reinforced by increases in life span (plants), clonal spread (plants), shorter generation times (snails), selfing ability (marginal effect in pulmonate snails) or paedogenesis (two incidences in hoverflies, needs further studies). Probably, the variety of strategies reflects the different evolutionary backgrounds that elicit different combinations of trade-offs, but all traits also might increase invasibility of species.     

Ecological and evolutionary consequences of size‐selective harvesting: how much do we know?

Size-selective harvesting, where the large individuals of a particular species are preferentially taken, is common in both marine and terrestrial habitats. Preferential removal of larger individuals of a species has been shown to have a negative effect on its demography, life history and ecology, and empirical studies are increasingly documenting such impacts. But determining whether the observed changes represent evolutionary response or phenotypic plasticity remains a challenge. In addition, the problem is not recognized in most management plans for fish and marine invertebrates that still mandate a minimum size restriction. We use examples from both aquatic and terrestrial habitats to illustrate some of the biological consequences of size-selective harvesting and discuss possible future directions of research as well as changes in management policy needed to mitigate its negative biological impacts.     

Popularity and Propagule Pressure: Determinants of Introduction and Establishment of Aquarium Fish

Propagule pressure is frequently cited as an important determinant of invasion success for terrestrial taxa, but its importance for aquatic species is unclear. Using data on aquarium fishes in stores and historical records of fish introduced and established in Canadian and United States waters, we show clear relationships exist between frequency of occurrence in shops and likelihood of introduction and of establishment. Introduced and established taxa are also typically larger than those available from stores, consistent with the propagule pressure hypothesis in that larger fish may be released more frequently due to outgrowing their aquaria. Attempts to reduce the numbers of introductions may be the most practical mechanism to reduce the number of new successful invasions.     

Diel feeding periodicity, daily ration and prey selection of a riverine population of juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

The diel feeding periodicity, daily ration and prey selection of juvenile chinook salmon, Oncorhynchus tshawytscha , were studied in relation to the available prey. Maximum dry weight of food intake occurred about dawn, when mayflies were the major prey, but the greatest number of freshly eaten prey occurred during the afternoon, when chironomids and terrestrial dipterans predominated. Feeding activity at night was low, with smaller mayflies comprising up to 50% of the prey. During the day the young salmon fed selectively on chironomids and the larger mayflies, while trichopterans and terrestrial taxa were under-represented in the diet. Food consumption over the 24-h period averaged 8.3% of the fish dry body weight. Prey abundance in the drift explained about 50% of the composition of the diet. Although the fish selected larger mayflies, size apparently was not a main criterion for selection because chironomids, although smaller than mayflies, were also frequently eaten. Previous dietary experience of the fish and the diel pattern of prey abundance appear to best explain the selective feeding of juvenile chinook salmon.     

Selective Feeding of age-0 Arctic Grayling in Lake-Outlet Streams of the Northwest Territories, Canada

A paucity of information exists on the diet of Arctic grayling, Thymallus arcticus, particularly for young-of-the-year (YOY). We examined the diet of YOY Arctic grayling in relation to food availability, in the Barrenlands region of the Northwest Territories, Canada, where lake-outlet streams serve as nursery habitat for these fish. Given the small size of YOY grayling and the abundance of lake-derived microcrustacea in the drift of these lake-outlet streams, we anticipated that these prey would make up a major component of the YOY's diet. Food selectivity by YOY grayling, however, was strongly sized-biased; although microcrustacea dominated the drift, YOY primarily consumed larger taxa, especially Chironomidae and Simuliidae. Even among these taxa, grayling tended to select the larger individuals. As they grew, YOY grayling took larger numbers of both large and small prey, particularly the larger invertebrates, although prey size range did not change after mid-July. Selection of pupae and avoidance of Ephemeroptera suggest that prey characteristics other than size also contribute to selectivity by YOY grayling. The relatively limited consumption of terrestrial invertebrates and other large prey may reflect the small sizes of fish in this arctic study, as well as differences in prey availability. Despite the abundance of lake-derived prey, instream production of invertebrates should largely determine the productive capacity of Barrenlands streams as fish habitat.     

Evolutionary influences on body size in free-living and parasitic isopods

The influence of mode of life and habitat characteristics on the evolution of body size in isopods was investigated in a comparative analysis based on data from 746 free-living and parasitic species. The phylogeny of isopods allowed 24 independent comparisons to be made between higher taxa (families or superfamilies), each corresponding to a separate branching event. The evolution of parasitism was consistently associated with reductions in body size. On the contrary, invasion of freshwater habitats was consistently coupled with increases in body size. Lineages moving to higher latitudes were significantly more likely to evolve larger body sizes than those shifting toward the equator. In addition, colonizing deeper water resulted in a weak tendency to evolve larger body size. The analysis suggests that the large size of some isopod groups parasitic on fish (e.g. Cymothoidae) may have been inherited from a free-living ancestor and is not the product of directional selection toward large size and greater fecundity.     

Lifetime selection on adult body size and components of body size in a waterstrider: opposing selection and maintenance of sexual size dimorphism

Abstract.— Sexual size dimorphism (SSD), the difference in body size between males and females, is common in almost all taxa of animals and is generally assumed to be adaptive. Although sexual selection and fecundity selection alone have often been invoked to explain the evolution of SSD, more recent views indicate that the sexes must experience different lifetime selection pressures for SSD to evolve and be maintained. We estimated selection acting on male and female adult body size (total length) and components of body size in the waterstrider Aquarius remigis during three phases of life history. Opposing selection pressures for overall body size occurred in separate episodes of fitness for females in both years and for males in one year. Specific components of body size were often the targets of the selection on overall body size. When net adult fitness was estimated by combining each individual's fitnesses from all episodes, we found stabilizing selection in both sexes. In addition, the net optimum overall body size of males was smaller than that of females. However, even when components of body size had experienced opposing selection pressures in individual episodes, no components appeared to be under lifetime stabilizing selection. This is the first evidence that contemporary selection in a natural population acts to maintain female size larger than male size, the most common pattern of SSD in nature.