Influence of individual body size and variable thresholds on the incidence of a sneaker male reproductive tactic in Atlantic salmon

In the conditional strategy model, divergence in reproductive phenotypes depends on whether the individual's condition is above or below a genetically determined threshold. The relative contribution of the genetic and environmental components that lead to the expression of a reproductive tactic by an individual is not well understood. In the present field study, we determined when condition diverged between males that develop the mature parr phenotype and those that do not in Atlantic salmon (Salmo salar). We also investigated the uniformity of the threshold value in the population. We sampled mature parr and immature males at age one, of the same population at six different sites for four consecutive years. Our study provides an example of the interaction of genotype and environment on the expression of a reproductive tactic. Size was significantly greater for future mature parr than for future immature males as early as 20 days after hatching (emergence), suggesting that there may be a parental effect component in the tactic adopted, since no exogenous feeding takes place before this time. Size advantage at emergence was maintained through the next spring at age one to different degrees depending on the year, thus suggesting the presence of an environmental component of tactic expression. Our results support the contention that within the conditional strategy, the environment faced by a male and his condition at the moment of reproduction consistently predicts neither the environment faced by his offspring nor the fitness they will obtain by expressing the same tactic as their father. Furthermore, higher mean size at a site did not always translate into a higher proportion of mature parr, therefore supporting the hypothesis that thresholds vary across habitats within the same population.     

The evolutionary ecology of alternative migratory tactics in salmonid fishes

Extensive individual variation in spatial behaviour is a common feature among species that exhibit migratory life cycles. Nowhere is this more evident than in salmonid fishes; individual fish may complete their entire life cycle in freshwater streams, others may migrate variable distances at sea and yet others limit their migrations to larger rivers or lakes before returning to freshwater streams to spawn. This review presents evidence that individual variation in migratory behaviour and physiology in salmonid fishes is controlled by developmental thresholds and that part of the variation in proximal traits activating the development of alternative migratory tactics is genetically based. We summarize evidence that alternative migratory tactics co‐exist within populations and that all individuals may potentially adopt any of the alternative phenotypes. Even though intra‐specific genetic divergence of migratory tactics is uncommon, it may occur if female competition for oviposition sites results in spawning segregation of alternative phenotypes. Because of their polygenic nature, alternative migratory tactics are considered as threshold traits. Threshold traits have two characteristics: an underlying 'liability' trait that varies in a continuous fashion, and a threshold value which is responsible for the discreetness observed in phenotypic distribution. We review evidence demonstrating that body size is an adequate proxy for the liability trait controlling the decision to migrate, but that the same phenotypic outcome (anadromy or residency) may be reached by different developmental pathways. The evidence suggesting a significant heritable component in the development of alternative migratory tactics is subsequently reviewed, leading us to conclude that alternative migratory tactics have considerable potential to respond to selection and evolve. We review what is known about the proximal physiological mechanisms mediating the translation of the continuous value of the liability trait into a discontinuous migratory tactic. We conclude by identifying several avenues for future research, including testing the frequency‐dependent selection hypothesis, establishing the relative importance of adaptive phenotypic plasticity in explaining some geographic gradients in migratory behaviour and identifying the physiological and genetic basis of the switching mechanisms responsible for alternative migratory tactics.     

Alternative life histories in the Atlantic salmon: genetic covariances within the sneaker sexual tactic in males

Alternative reproductive tactics are ubiquitous in many species. Tactic expression often depends on whether an individual's condition surpasses thresholds that are responsible for activating particular developmental pathways. Two central goals in understanding the evolution of reproductive tactics are quantifying the extent to which thresholds are explained by additive genetic effects, and describing their covariation with condition-related traits. We monitored the development of early sexual maturation that leads to the sneaker reproductive tactic in Atlantic salmon (Salmo salar L.). We found evidence for additive genetic variance in the timing of sexual maturity (which is a measure of the surpassing of threshold values) and body-size traits. This suggests that selection can affect the patterns of sexual development by changing the timing of this event and/or body size. Significant levels of covariation between these traits also occurred, implying a potential for correlated responses to selection. Closer examination of genetic covariances suggests that the detected genetic variation is distributed along at least five directions of phenotypic variation. Our results show that the potential for evolution of the life-history traits constituting this reproductive phenotype is greatly influenced by their patterns of genetic covariance.     

Alternative mate-finding tactics in a non-territorial damselfly (Odonata: Coenagrionidae)

Males of the non-territorial damselfly Enallagma hageni have two alternative tactics for finding mates: (1) they search the banks of the pond for unmated females (searching tactic), or (2) wait at oviposition sites for females that resurface prematurely from underwater oviposition (waiting tactic). Although the searching tactic yielded more fertilizations than the waiting tactic, for time invested, the waiting tactic became increasingly successful later in the reproductive season due to changes in female oviposition behaviour. The two tactics can be maintained in the population because males can mate by the waiting tactic during the afternoon when few females are available to searchers. Among males visiting the breeding site an equal number of times, males mating by a mixture of tactics were as successful as males mating only by the main tactic. Because marked males were found to use both tactics, these behaviours are interpreted as evidence of behavioural plasticity within individuals, representing one conditional evolutionary strategy.     

Alternative tactics and individual reproductive success in natural associations of the burying beetle, Nicrophorus vespilloides

Alternative reproductive tactics can be maintained through differentevolutionary avenues. They can be genetically or stochasticallydetermined, in which case they must yield equal fitness, ortheir use can be conditional, in which case the fitness payoffof alternatives may differ. We attempted to assess the reproductivesuccess of alternative reproductive tactics employed by wildmale and female burying beetles in natural associations on carcassesplaced in the field. A beetle's reproductive tactic was definedby its potential involvement in care of larvae, and parentagewas assessed using oligolocus DNA fingerprinting of offspringand potential parents. Both in males and in females, alternativetactics yielded significantly different reproductive benefits:subordinate females (brood parasites) and males (satellite males)had considerably lower reproductive success than dominant oruncontested individuals. Joint breeding was too infrequent forstatistical inferences, generating intermediate offspring numbers.About 15% of offspring were sired by males not present on thecarcass, suggesting that mating away from reproductive resourcescan produce reproductive benefits to males. Our results, inconcert with the observation that beetles using one tactic canbe manipulated into employing the alternative, support the notionthat Nicrophorus vespilloides uses alternatives conditionally,opportunistically employing lower-benefit tactics when moreprofitable tactics are not available, or as additional "on-the-side"tactics to bolster reproductive success.     

Relative fitness of alternative male reproductive tactics in a mammal varies between years

1. In many species, males can use different behavioural tactics to achieve fertilization, so-called alternative reproductive tactics (ARTs). Few field studies have measured fitness consequences of ARTs under varying environmental conditions. 2. Here, we describe fitness consequences of three phenotypically plastic ARTs in the African striped mouse (Rhabdomys pumilio) and show that relative fitness of ARTs differs between years. Each year represents a different generation. 3. For the generation living under high population density, tactics differed in relative fitness in accordance with the theory of conditional strategies, with highly successful territorial breeding males having 10 times higher success than solitary roaming males and 102 times higher success than adult natally philopatric males. 4. For the generation living under intermediate population density, the territorial breeding and roaming tactics yielded similar fitness, which would be in agreement with the theory of mixed strategies. No philopatric males occurred. 5. For the generation living under low population density, roaming was the only tactic used and some roamers had very high fitness. 6. The main prediction of status-dependent selection for conditional strategies is a correlation between fitness and status, often measured as body mass, but we did not find this correlation within tactics when more than one tactic was expressed in the population. 7. Female distribution seems to have an effect on which reproductive tactics male chose: female defence polygyny when females are clumped (interference competition), but a searching tactic when females are dispersed (scramble competition). In contrast to predictions arising from theory on scramble competition, male body mass was important in determining fitness only in the year when females were dispersed, but not in other years. 8. Our results indicate that the differentiation between conditional and mixed strategies is not an absolute one. In many other species, environmental conditions might fluctuate temporally and spatially so that the normally suboptimal tactic yields similar fitness to the (usually) dominant tactic or that only a single tactic prevails. 9. We suggest the term single strategy, independent of current fitness consequences, for systems where tactics are not genetically determined, in contrast to genetically determined alternative strategies.     

Alternative reproductive tactics and the propensity of hybridization

One explanation for hybridization between species is the fitness benefits it occasionally confers to the hybridizing individuals. This explanation is possible in species that have evolved alternative male reproductive tactics: individuals with inferior tactics might be more prone to hybridization provided it increases their reproductive success and fitness. Here we experimentally tested whether the propensity of hybridization in the wild depends on male reproductive tactic in Calopteryx splendens damselflies. Counter to our expectation, it was males adopting the superior reproductive tactic (territoriality) that had greatest propensity to hybridize than males adopting the inferior tactics (sneakers and floaters). Moreover, among the territorial males, the most ornamented males had greatest propensity to hybridize whereas the pattern was reversed in the sneaker males. Our results suggest that there is fluctuating selection on male mate discrimination against heterospecific females depending on both ornament size and the male’s reproductive tactic.     

Alternative Mating Tactics and Evolutionarily Stable Strategies

Difficulties with applying Evolutionarily. Stable Strategy (ESS)methodology and terminology to alternative mating behaviors(in which some males in local populations adopt strikingly different,often non-competitive, behavioral patterns) are reviewed. Definitionsfor "tactic" (behavioral phenotype) and "strategy" (evolvedset of rules for expressing tactics) are given. Inconsistentand incorrect applications of "mixed," "pure," and "conditional"ESSs are discussed. Cases of condition-dependent alternative mating tactics arereviewed. Because most alternative behaviors are condition dependent,neither their population-wide nor individual fitness contributionsare expected to equal the fitness contributions of "primary"tactics. Individuals should, however, switch tactics at "equalfitness points." A particular conditional tactic will persistwhen its maintenance cost (genetic or physiological) is lessthan its fitness contribution. In only exceptional cases arethe fitness contributions of tactics expected to be equal: 1)genetic polymorphisms, 2) stochastic "mixed" ESSs, 3) frequency-dependentchoice and, 4) arbitrary assessment. Although alternative tacticsmay occur in cases of genetic polymorphism or genetic equipotence,most mating tactics probably occur when continuous heritablevariation in underlying conditional strategy exists. Selectionfor genetically influenced "roles" (genetic background) mayalso uncover apparent heritability.     

Elevated corticosterone levels elicit non-calling mating tactics in male toads independently of changes in circulating androgens

Competition among males for a limited number of females may result in the expression of condition-dependent alternative mating tactics. In such cases, decision rules mediating mating tactic expression are likely to be influenced by a male's external as well as internal environment. For example, experimental studies with anuran amphibians (frogs and toads) indicate that changes in the social-acoustic environment alter the probability that an individual adopts a calling versus non-calling "satellite" mating tactic. However, there is considerable variation in the behavioral responses of individuals in such studies, suggesting that physiological differences among individuals play an important role in tactic expression. For instance, recent models predict that natural elevation in adrenal glucocorticoids during vocal production alters androgen production and/or neural activity to mediate transitions between reproductive tactic expression in anuran amphibians. Using corticosteroid injections, we show that elevation in circulating corticosterone levels significantly increase the probability that free-ranging male Great Plains toads (Bufo cognatus) and Woodhouse's toads (B. woodhousii) adopt a non-calling satellite tactic. Corticosterone-induced behavioral transitions occurred rapidly (<1 h) and independently of fluctuations in circulating androgen levels, suggesting a direct effect of glucocorticoids on brain centers controlling vocal production. We discuss our findings in the context of behavioral studies that alter the social-acoustic environment to examine its influence on tactic expression.     

Relationship between metabolism, sex and reproductive tactics in young Atlantic salmon (Salmo salar L.)

Atlantic salmon can differ markedly in their growth and in the timing of reproductive maturation, leading to the dramatic contrast between the large anadromous adults and the diminutive mature male parr. This study examined the growth rates, anatomical and physiological characteristics of parr during the adoption of their discrete life histories to ascertain whether these properties can explain tactic choice. To minimise the impact of habitat differences upon these attributes, salmon were reared in the laboratory until 1.5years of age, when the "decisions" to undergo smoltification or to mature as parr had been taken. At 1.5years, both males and females showed bimodal size-frequency distributions. Neither the population of origin nor the paternal reproductive tactic influenced the "decision" to mature or the growth trajectories. Growth rate (% massday(-1) during their final 10months) and the % male and female offspring in the upper modal group were strongly correlated and varied markedly among families. Mean growth rate per family was negatively correlated with mean metabolic rate per family at emergence. Growth rate decreased as a function of parr size in January and the growth rates of upper modal fish were displaced upwards relative to those of lower modal fish. Most males in the smaller size mode matured, whereas all other fish began smoltification. Mature male parr did not differ from similarly sized female pre-smolt in routine metabolic rate, but these smaller fish had higher metabolic rates than larger male and female pre-smolts. However, mature parr differed markedly from similarly sized females and from larger male and female pre-smolts in possessing higher oxidative and lower glycolytic capacities in muscle. Overall, these data are consistent with the interpretation that growth rates dictate the distribution of parr between upper and lower modal groups. Individuals from faster growing families would be more likely to pass the threshold for smoltification and to accelerate growth, whereas those from slower growing families would remain in the lower mode. The use of metabolic capacities, e.g. metabolic rate, was linked with modal group, whereas muscle oxidative capacity was linked with male maturity. Mean family metabolic rate at emergence was negatively linked with mean growth during the subsequent year, suggesting that metabolic efficiency facilitates growth and eventually smoltification.     

Differential reproductive success and heritability of alternative reproductive tactics in wild Atlantic salmon (Salmo salar L.)

Abstract A critical step in understanding the evolution and maintenance of alternative reproductive tactics is to obtain accurate comparisons of their fitness and to determine factors influencing individual status. In this study, we first used individual multilocus genotypic information to compare reproductive success between two alternative reproductive tactics of anadromous Atlantic salmon (Salmo salar L.) in their natural environments. We also documented the effects of the quality of the rearing environment and of paternal reproductive tactics on heritability of juvenile growth, which is an important component of individual status. Results showed that large dominant salmon (multisea winter) had higher reproductive success than smaller satellite individuals (grilse). Also, there was a status difference associated with both habitat and male tactic. Overall, offspring produced in streams were bigger than those produced in the main river stretch. Grilse also produced bigger offspring than those fathered by multisea winter males. Heritability of juvenile growth was significant but varied according to quality of habitat: higher heritability estimates were observed in higher quality habitats (streams) than in lower quality habitats (main river stretch). Heritability estimates for juvenile growth varied as well, depending on male tactic, with progeny fathered by multisea winter males having higher values than those fathered by grilse. Together, these results indicate that a combination of additive genetic effects, parental life history and habitat quality will ultimately shape juvenile growth rate, which is the main determinant of status and of subsequent choice of life-history tactics.     

Tactic‐specific benefits of polyandry in Chinook salmon Oncorhynchus tshawytscha

This study examined whether polyandrous female Chinook salmon Oncorhynchus tshawytscha obtain benefits compared with monandrous females through an increase in hatching success. Both of the alternative reproductive tactics present in male O. tshawytscha (large hooknoses and small, precocious jacks) were used, such that eggs were either fertilized by a single male (from each tactic) or multiple males (using two males from the same or different tactics). The results show that fertilized eggs from the polyandrous treatments had a significantly higher hatching success than those from the monandrous treatments. It is also shown that sperm speed was positively related with offspring hatching success. Finally, there were tactic‐specific effects on the benefits females received. The inclusion of jacks in any cross resulted in offspring with higher hatching success, with the cross that involved a male from each tactic providing offspring with the highest hatching success than any other cross. This study has important implications for the evolution of multiple mating and why it is so prevalent across taxa, while also providing knowledge on the evolution of mating systems, specifically those with alternative reproductive tactics.     

Hormone levels of male African striped mice change as they switch between alternative reproductive tactics

Alternative reproductive tactics occur when individuals of the same species follow alternative ways to maximize reproductive success. Often younger and smaller males follow tactics that result in lower fitness than that of dominant larger males. The relative plasticity hypothesis predicts that hormone levels change as males change tactics, but direct tests of this hypothesis are missing. It has been demonstrated in a number of studies that males following different tactics also differ in hormone levels (unpaired data), but not that individual males change their hormone levels as they change tactic (paired data). We compared hormone levels in the same individuals before and after they changed their tactic, using field samples collected over a period of 6 years. We studied male striped mice (Rhabdomys pumilio) following three alternative reproductive tactics: 1. alloparental philopatric males; 2. solitary roaming males, and 3. group-living dominant breeders. Testosterone levels increased and corticosterone levels decreased when philopatric males became roamers or breeders. The increase in testosterone levels tended to be higher in philopatric males that became roamers than in philopatric males that became breeders. Testosterone levels decreased when roamers became breeders. Prolactin levels increased when males of any other tactic became breeders. Thus, males significantly changed their hormone profiles as they changed tactics. These results are in agreement with the hypothesis that changes in hormone levels are associated with the switch from one alternative reproductive tactic to another.     

Hormone levels of male African striped mice change as they switch between alternative reproductive tactics

Alternative reproductive tactics occur when individuals of the same species follow alternative ways to maximize reproductive success. Often younger and smaller males follow tactics that result in lower fitness than that of dominant larger males. The relative plasticity hypothesis predicts that hormone levels change as males change tactics, but direct tests of this hypothesis are missing. It has been demonstrated in a number of studies that males following different tactics also differ in hormone levels (unpaired data), but not that individual males change their hormone levels as they change tactic (paired data). We compared hormone levels in the same individuals before and after they changed their tactic, using field samples collected over a period of 6 years. We studied male striped mice (Rhabdomys pumilio) following three alternative reproductive tactics: 1. alloparental philopatric males; 2. solitary roaming males, and 3. group-living dominant breeders. Testosterone levels increased and corticosterone levels decreased when philopatric males became roamers or breeders. The increase in testosterone levels tended to be higher in philopatric males that became roamers than in philopatric males that became breeders. Testosterone levels decreased when roamers became breeders. Prolactin levels increased when males of any other tactic became breeders. Thus, males significantly changed their hormone profiles as they changed tactics. These results are in agreement with the hypothesis that changes in hormone levels are associated with the switch from one alternative reproductive tactic to another.     

The evolution of alternative mating strategies in variable environments

Summary We assessed the influence of phenotypic plasticity in age at maturity on the maintenance of alternative mating strategies in male Atlantic salmon,Salmo salar. We calculated the fitness,r, associated with the parr and the anadromous strategies, using age-specific survival data from the field and strategy-specific fertilization data from the laboratory. The fitness of each strategy depended largely on mate competition (numbers of parr per female, i.e. parr frequency) and on age at maturity. Fitness declined with increasing numbers of parr per female with equilibrium frequencies (at which the fitnesses of each strategy are equal) being within the range observed in the wild. Equilibrium parr frequencies declined with decreasing growth rate and increasing age at maturity. Within populations, the existence of multiple age-specific sets of fitness functions suggests that the fitnesses of alternative strategies are best represented as multidimensional surfaces. The points of intersection of these surfaces, whose boundaries encompass natural variation in age at maturity and mate competition, define an evolutionarily stable continuum (ESC) of strategy frequencies along which the fitnesses associated with each strategy are equal. We propose a simple model that incorporates polygenic thresholds of a largely environmentally-controlled trait (age at maturity) to provide a mechanism by which an ESC can be maintained within a population. An indirect test provides support for the prediction that growth-rate thresholds for parr maturation exist and are maintained by stabilizing selection. Evolutionarily stable continua, maintained by negative frequency-dependent selection on threshold traits, provide a theoretical basis for understanding how alternative life histories can evolve in variable environments.     

Female mate choice tactics in a resource-based mating system: field tests of alternative models

Abstract In this study we test theoretical models of female mate choice tactics in natural populations of pine engravers, Ips pini (Say) (Coleoptera: Scolytidae), a species with a resource-based mating system and high search costs. We first develop distinguishing predictions for each of four models of mate choice: random, comparison tactics, and fixed and adjustable thresholds. These predictions relate to commonly collected field data that include the visiting behavior of females and the quality of accepted and rejected mates. Using these types of data, we conclude that pine engravers use an adjustable threshold mate choice tactic because females often accepted the first male encountered, rarely revisited males, visited similar numbers of males in patches of different quality, accepted higher-quality males than those they rejected even on their first encounter with a male in a patch, and had higher acceptance thresholds in high-quality patches than in low-quality patches. This adjustable threshold tactic is consistent with a one-step decision rule and is predicted to occur in species such as pine engravers in which search costs are high and females have information about patch quality before beginning a search in a patch.     

Resource availability,mating opportunity and sexual selection intensity influence the expression of male alternative reproductive tactics

The expression of alternative reproductive tactics can be plastic and occur simultaneously depending on cues that vary spatially or temporally. For example, variation in resources and sexual selection intensity is expected to influence the pay‐off of each tactic and shape the decision of which tactic to employ. Males of the nuptial gift‐giving spider Pisaura mirabilis can adopt three tactics: offering a genuine prey gift, a ‘worthless’ non‐nutritious gift or no gift. We hypothesized that resources and/or male body condition, and mating opportunity and sexual selection intensity, vary over the course of the mating season to shape the co‐existence of alternative traits. We measured these variables in the field over two seasons, to investigate the predictions that as the mating season progresses, (i) males become more likely to employ a gift‐giving tactic, and (ii) the likelihood of switching from worthless to genuine gifts increases. Prey availability increased over the season and co‐varied with the propensity of males to employ the gift‐giving tactic, but we found no support for condition‐dependent gift giving. Males responded to an increase in female availability by increasing their mating effort (gift production). Furthermore, the frequency of genuine gift use increased with sexual selection intensity, consistent with the assumption that sperm competition intensity increases with time. Our results suggest that the frequency of alternative tactics is shaped by seasonal changes in ecological factors and sexual selection. This leads to relaxed selection for the gift‐giving tactic early in the season when females are less choosy and resources more scarce, and increased selection for genuine gifts later in the season driven by mating opportunity and risk of sperm competition.     

Do Feeding Resources Induce the Adoption of Resource Defence Polygyny in a Lekking Butterfly?

Among polygynous species, males often compete for the possession of mating sites to increase their reproductive success. Weaker individuals frequently adopt alternative non‐territorial mate‐locating tactics, but the adoption of alternative territorial tactics may also occur. Although alternative tactics with territory defence are less common in arthropods, factors that drive its adoption may provide information to understand the organization of different territorial mating systems in the group. Here we investigate the adoption of resource‐based territoriality as an alternative to a non‐resource‐based one by males of the butterfly Paryphthimoides phronius. Male P. phronius commonly defend sunny clearings lacking feeding resources in the forest edge (non‐resource‐based territoriality). However, after experimentally offering fermenting fruit in previously undefended sites, we showed that males also adopt a resource defence tactic. Males in territories with fermenting fruits apparently feed on this resource when they are not defending the territory. However, males in sites without resources did not migrate to territories with resources when given the opportunity. To our knowledge, this is the first experimental investigation to show a non‐resource‐ and an alternative resource‐based mate‐locating tactic in a butterfly. We suggest that this behavioural flexibility may represent an important step to understand the ecological factors responsible for the organization and evolution of different territorial mating systems in insects.     

Effects of Social Structure on the Behaviour and Performance of Alternative Reproductive Phenotypes in Male Rock Shrimp, Rhynchocinetes typus

Males that adopt alternative mating tactics within a conditional strategy often undergo costly morphological changes when switching to the next phenotype during ontogeny. Whether costs of changing to a subsequent reproductive phenotype are outweighed by a higher mating probability may depend on the frequencies of different phenotypes in a group of competitors. Benefits and costs associated with different phenotype frequencies depend on interactions within and between alternative phenotypes, but the underlying behavioural mechanisms have rarely been studied. Herein, we used the rock shrimp Rhynchocinetes typus as a model: ontogenetic male stages of this species differ in morphological and behavioural traits that indicate alternative reproductive phenotypes. The small, subordinate, male stage (typus) develops via several intermediate stages (intermedius) to the dominant male stage (robustus): in competitive interactions the typus males usually employ the sneaking tactic, while the robustus males invariably employ the monopolizing fighter tactic. In laboratory experiments, we manipulated phenotype frequencies to examine whether there are frequency‐dependent effects on searching behaviour, aggressiveness and mating probability. With increasing frequency of robustus males, the rate of aggressive interactions among them increased. Furthermore, robustus males increased walking velocity when more than one robustus male was present. In contrast, typus males did not adjust their searching or aggressive behaviour. The increase of aggressive interactions among robustus males provided more opportunities for typus males to seize a temporarily unguarded female. While typus males exploit fights among robustus males that produce mating opportunities for them, robustus males benefit from typus males, which reveal the presence of receptive females. We suggest that each phenotype benefits from the presence of the other phenotype and suffers costly interference among individuals of the same phenotype. Whether frequency‐dependent effects on the mating probability of subordinates also affect their ontogenetic switchpoint should be examined in future studies.     

The effects of paternal reproductive tactic and rearing environment on juvenile variation in growth as mediated through aggression and foraging behaviours of Chinook salmon (Oncorhynchus tshawytscha)

In species with indeterminate growth, differential growth rates can lead to animals adopting alternative reproductive tactics such as sneak–guard phenotypes, which is partially predicted by variation in growth during the juvenile life‐history stage. To investigate sources of growth variation, we examined the independent and joint effects of paternal reproductive tactic (G) and rearing environment (E) on juvenile growth in Chinook salmon (Oncorhynchus tshawytscha), hypothesizing G and E effects are partially mediated through differences in behaviour such as aggressive interactions and resulting foraging behaviours. We created maternal half‐sibling families with one‐half of the female's eggs fertilized by the milt of a sneaker “jack” and the other half by a guarder “hooknose”. At the exogenous feeding stage, each split‐clutch family was then divided again and reared in a rationed diet or growth‐promotion diet environment for approximately 6 months, during which growth parameters were measured. Before saltwater transfer at 9 months of age, social interactions were observed in groups of six fish of various competitor origins. We found ration restricts growth rate and juvenile mass, and evidence of genetic effects on growth depensation, where jack‐sired individuals grew less uniformly over time. These growth‐related differences influenced an individual's level of aggression, with individuals raised on a restricted diet and those whose families experienced greatest growth being most aggressive. These individuals were more likely to feed than not and feed most often. Jack‐sired individuals were additionally aggressive in the absence of food, and when raised on a rationed diet outcompeted others to feed most. These results show how individuals may achieve higher growth rates via intrinsic (G) or induced (E) aggressive behavioural phenotypes, and eventually attain the threshold body size necessary during the saltwater phase to precociously sexually mature and adopt alternative reproductive phenotypes.