Relative Reproductive Success Research Articles

Evolution of Litter Size: Proximate and Ultimate Mechanisms.

Relative reproductive success and failure are the ultimate determinants of Darwinian fitness. As such, reproductive traits and variation therein have an immediate and considerable impact on the evolutionary trajectory of lineages. Historically, significant attention has been paid to the ecological and evolutionary processes (ultimate factors) that shape the diversity and canalization of reproductive traits within groups to better our understanding of organismal diversity and population or species resilience. In contrast, the physiological systems that mediate variation within and among species (i.e., the proximate factors) in reproductive traits remain a significant black box. To-date, there is comparatively little information about how proximate mechanisms constrain or promote evolutionary potential in reproductive traits. In this mini-review, we focus on litter size in Eutherian mammals as a trait with relatively well-defined diversity (litter sizes are well-described both within and across species) and for which some genetic determinants have been identified. We discuss both the ultimate and potential proximate determinants of litter size with special attention to the breadth of physiological traits that may act as "toggle" switches for evolution of litter size. We close with a brief discussion of the role that physiological plasticity may play in the evolution of litter size and lay out several forward-looking areas for future research.

Carry-over effects of seasonal migration on reproductive success through breeding site retention in a partially migratory bird.

Understanding the maintenance and dynamics of phenotypic polymorphisms requires unpicking key ecological mechanisms shaping the fitness costs and benefits of expressing alternative phenotypes, generating selection. Seasonal migration versus year-round residence expressed in partially migratory populations represents one common polymorphism that can experience strong selection through differential reproductive success. Yet, key hypothesised pathways that could generate such selection remain to be empirically tested. One hypothesis is that migratory tactics affect subsequent reproductive success through carry-over effects on breeding site retention and resulting breeding dispersal. By remaining in breeding areas all year round, residents could retain their preferred breeding site between years, and consequently have higher reproductive success. Conversely, migrants that escape harsh non-breeding season conditions could return in better condition, with high resource holding potential, and outcompete residents to retain their site. Such effects could further depend on migration timing and vary between years. Yet, such pathways have not been quantified, precluding empirical parameterisation of partial migration theory. We used 4 years of breeding and non-breeding season data from partially migratory European shags (Gulosus aristotelis) to test whether the three most frequent migratory tactics in this population (full resident, early migrant departing soon after breeding, and late migrant departing in late autumn) differed in their breeding site retention; whether site retention predicted reproductive success; and hence whether effects of migratory tactic on reproductive success were explicable through site retention. Overall, residents were much more likely to retain their breeding site between years than both early and late migrants, and site retention was associated with increased reproductive success. Yet, these effects varied somewhat among years: late migrants were always least likely to retain their site but had variable relative reproductive success. Path analyses revealed that effects of migratory tactic on reproductive success were only partly attributable to breeding site retention. These results indicate that multiple mechanisms underlie reproductive selection on migratory tactics, potentially contributing to maintaining behavioural polymorphisms. Yet, the clear associations between migratory tactics and local breeding dispersal reveal that these movements can be strongly interlinked across seasons, shaping overall spatioseasonal dynamics in partially migratory systems.

Offspring sex ratio increases with paternal reproductive success in a colony of southern elephant seals

AbstractIn polygynous species, male reproductive success is often determined by their ability to dominate female harems. Harem‐holders sire a disproportionate number of offspring. Male dominance tends to correlate with age, but intense competition and early male mortality limit most males from achieving high social status. To maximize reproductive success despite low rank, offspring sex ratio adjustment may have evolved, favoring the sex with higher fitness. If traits influencing dominance are heritable and confer reproductive advantages, we expect fathers with higher reproductive success to produce more sons, as they are more likely to become dominant. In contrast, subordinate males with lower success may benefit from siring more daughters. We tested this hypothesis on a colony of southern elephant seals breeding on the Kerguelen Archipelago. We used genetic markers to link the paternity of 540 pups to 58 breeding males. We found that the probability of siring a son increases from 43% to 54% with paternal relative reproductive success, supporting the offspring sex ratio adjustment hypothesis. Given that various factors influence sex ratio in a population, future studies should tease apart these ecological processes (e.g., paternal dominance, maternal condition, local density, or adult sex ratio) and investigate how they interact with each other.

Demographic and genetic consequences of a steelhead supplementation program

AbstractObjectiveSupplementation of naturally spawning populations by the addition of hatchery‐spawned individuals is commonly conducted for recovery of threatened and endangered populations and to support harvest opportunities. Our objective was to evaluate whether the use of a juvenile captive broodstock and an integrated paradigm could increase returns of steelhead, the anadromous form of Rainbow Trout Oncorhynchus mykiss, while avoiding negative genetic impacts commonly associated with hatchery propagation.MethodsWe analyzed 291 genetic markers in adult steelhead returning to an integrated population in southwest Washington over the course of 15 years. Reproductive success (RS) of fish spawning in the natural environment was evaluated by origin (hatchery versus natural) and return year. Data were then pooled over years to maximize sample sizes for comparing RS estimates among groups (i.e., estimation of relative reproductive success [RRS]).ResultWe observed a weak relationship between RS and origin (male p = 0.347; female p = 0.066), but a significant relationship between RS and return year (male p < 0.001, η2 = 0.896; female p ⟨ 0.001, η2 = 0.867) (i.e., hatchery‐ and natural‐origin fish did well or poorly together each year). Estimates of RRS for fish spawned in the hatchery ranged from 2.4 to 6.4, indicating that fish spawned in the hatchery produced more returning adult progeny than did fish allowed to spawn in the natural environment. Estimates of RRS were ⟨1.0 for hatchery‐origin fish spawning in the natural environment, but the difference was nonsignificant for males and marginally significant for females. Hatchery‐origin fish exhibited reduced genetic diversity as well as evidence of increased temporal population structure.ConclusionWe conclude that the program was successful in achieving an increase in adult returns but not in avoiding negative genetic effects on the population and that any lasting impacts of supplementation remain to be determined.

Reproductive success of jack and full-size males in a wild coho salmon population.

Despite the wealth of research on Pacific salmon Oncorhynchus spp. life histories there is limited understanding of the lifetime reproductive success of males that spend less time at sea and mature at a smaller size (jacks) than full-size males. Over half of returning male spawners can be jacks in some populations, so it is crucial to understand their contribution to population productivity. We quantified adult-to-adult reproductive success (RS) of jacks and their relative reproductive success (RRS) compared to full-size males in a wild population of coho salmon in the Auke Creek watershed, Juneau, Alaska. We used genetic data from nearly all individuals (approx. 8000) returning to spawn over a decade (2009-2019) to conduct parentage analysis and calculate individual RS. The average adult-to-adult RS of jacks (mean = 0.7 and s.e. = 0.1) was less than that of full-size males (mean = 1.1 and s.e. = 0.1). Jack RRS was consistently below 1.0 but ranged widely (0.23 to 0.96). Despite their lower average success, jacks contributed substantially to the population by siring 23% of the total returning adult offspring (1033 of 4456) produced between 2009 and 2015. Our results imply that jacks can affect evolutionary and population dynamics, and are relevant to the conservation and management of Pacific salmon.

Effect of catch‐and‐release and temperature at release on reproductive success of Atlantic salmon (Salmo salar L.) in the Rimouski River, Québec, Canada

AbstractCatch‐and‐release fishing is a common conservation practice in recreational fisheries for Atlantic Salmon, although the effects on the reproductive success of caught‐and‐released fish are poorly understood. Herein, we compared the relative reproductive success of caught‐and‐released to non‐caught salmon and tested the effect of temperature at release on reproductive success in the Rimouski River, Québec, Canada. At least 83% of caught‐and‐released salmon that moved upstream of a dam successfully reproduced, including fish that have been released in water above 20°C. However, the reproductive success of caught‐and‐released female salmon was only 73% of the reproductive success of non‐caught salmon. Moreover, the increasing temperature did not affect the reproductive success of released fish that entered a trap, but fish caught at warmer temperatures were less likely to enter the trap. Our findings should be useful for evaluating the risks and benefits of catch‐and‐release, and for optimising conservation practices used for the preservation of Atlantic salmon populations.

Male Adaptive Plasticity Can Explain the Evolution of Sexual Perception Costs.

AbstractSensory perception of environmental cues has been shown to trigger plastic responses that can induce important fitness costs, including the dramatic modulation of aging across distant taxa. For example, male Drosophila melanogaster suffer a marked decrease in fitness, characterized by faster reproductive and actuarial aging, if they perceive female cues but fail to mate shortly after (aging via sexual perception). While this has been a breakthrough for our understanding of the mechanisms of aging, it raises the question of why such plastic responses evolved. Here, we used D. melanogaster to ask whether sexual perception costs may be a by-product of adaptive plastic responses to female cues. We found that (a) short-term perception (1 day) of female cues before mating opportunities increases male relative lifetime reproductive success in a competitive environment, (b) medium-term perception (3-7 days) is neutral, and (c) long-term perception (15 days) leads to reproductive costs. We then ran mathematical simulations under a wide range of sociosexual and demographic scenarios to show that such plastic male responses can be adaptive whenever mating rates fluctuate within the range experienced by D. melanogaster and other insects in the wild, suggesting that this may be a widespread strategy in nature. Finally, we show that, because the short-term benefits of plastic responses will be acquired mostly by high-quality males while long-term costs will be paid mostly by low-quality males, sexual perception can significantly magnify sexual selection (15%-27% average increase in the opportunity for selection).

Effects of stocking at the parr stage on the reproductive fitness and genetic diversity of a wild population of Atlantic salmon (Salmo salar L.).

Captive‐breeding programs are among the most adopted conservation practices to mitigate the loss of biodiversity, including genetic diversity. However, both genetic and nongenetic changes occurring in captivity can reduce the fitness of supplemented individuals, which complicate rehabilitation efforts. In the case of Atlantic salmon, the intensity of changes that occur in captivity and their impact on fitness will vary with the stocking practice adopted. In this study, we test whether salmon stocked at the parr stage have reduced reproductive success compared with their wild conspecifics and whether they contribute to increase genetic diversity in the targeted population. To do so, we use high‐throughput microsatellite sequencing of 38 loci to accurately assign 2381 offspring to a comprehensive set of possible parents from a supplemented Atlantic salmon population in Québec, Canada. Captive‐bred salmon stocked at the parr stage had fewer mates than their wild conspecifics, as well as a reduced relative reproductive success (RSS) compared with their wild counterparts. Nonetheless, in comparison with previous studies, stocking at the parr stage significantly improved RSS compared with salmon stocked as smolts and they displayed a reduction in reproductive success similar to salmon stocked as fry, which spend less time in captivity than parr. Moreover, supplementation of captive‐bred salmon significantly contributed to increasing genetic diversity. These results should contribute to informing resource managers in determining the best stocking practice to enhance Atlantic salmon populations.

Reduced relative fitness in hatchery-origin Pink Salmon in two streams in Prince William Sound, Alaska.

Previous studies generally report that hatchery‐origin Pacific Salmon (Oncorhynchus spp.) have lower relative reproductive success (RRS) than their natural‐origin counterparts. We estimated the RRS of Pink Salmon (O. gorbuscha) in Prince William Sound (PWS), Alaska using incomplete pedigrees. In contrast to other RRS studies, Pink Salmon have a short freshwater life history, freshwater habitats in PWS are largely unaltered by development, and sampling was conducted without the aid of dams or weirs resulting in incomplete sampling of spawning individuals. Pink Salmon released from large‐scale hatchery programs in PWS have interacted with wild populations for more than 15 generations. Hatchery populations were established from PWS populations but have subsequently been managed as separate broodstocks. Gene flow is primarily directional, from hatchery strays to wild populations. We used genetic‐based parentage analysis to estimate the RRS of a single generation of stray hatchery‐origin Pink Salmon in two streams, and across the odd‐ and even‐year lineages. Despite incomplete sampling, we assigned 1745 offspring to at least one parent. Reproductive success (RS), measured as sampled adult offspring that returned to their natal stream, was significantly lower for hatchery‐ vs. natural‐origin parents in both lineages, with RRS ranging from 0.03 to 0.47 for females and 0.05 to 0.86 for males. Generalized linear modeling for the even‐year lineage indicated that RRS was lower for hatchery‐origin fish, ranging from 0.42 to 0.60, after accounting for sample date (run timing), sample location within the stream, and fish length. Our results strongly suggest that hatchery‐origin strays have lower fitness in the wild. The consequences of reduced RRS on wild productivity depend on whether the mechanisms underlying reduced RRS are environmentally driven, and likely ephemeral, or genetically driven, and likely persistent across generations.

Pollinator-Mediated Indirect Effects on Plant Fecundity Revealed by Network Indices.

AbstractIndirect effects arise when one species influences how another species interacts with a third. Pollinator-mediated indirect effects are widespread in many plant communities and are often not restricted to plant species pairs. An analytical framework does not exist yet that allows for the evaluation of indirect effects through shared pollinators in a community context as well as their consequences for plant fitness. We used network indices describing pollinator sharing to assess the extent to which plant species affect and are affected by others in a pollination network from a species-rich dune community. For 23 plant species, we explore how these indices relate to plant fecundity (seeds/flower) over two years. We further linked plant traits and indices to uncover functional aspects of pollinator-mediated indirect interactions. Species frequently visited by shared pollinators showed higher fecundity and exhibited traits that increase pollinator attraction and generalization. Conversely, species whose shared pollinators frequently visited other plants had lower fecundity and more specialized traits. Thus, pollinator sharing benefited some species while others suffered reproductive disadvantages, consistent with competition. The framework developed here uses network tools to advance our understanding of how pollinator-mediated indirect interactions influence a species' relative reproductive success at the community level.

Mating systems and predictors of relative reproductive success in a Cutthroat Trout subspecies of conservation concern.

Mating systems and patterns of reproductive success in fishes play an important role in ecology and evolution. While information on the reproductive ecology of many anadromous salmonids (Oncorhynchus spp.) is well detailed, there is less information for nonanadromous species including the Yellowstone Cutthroat Trout (O. clarkii bouvieri), a subspecies of recreational angling importance and conservation concern. Using data from a parentage‐based tagging study, we described the genetic mating system of a migratory population of Yellowstone Cutthroat Trout, tested for evidence of sexual selection, and identified predictors of mating and reproductive success. The standardized variance in mating success (i.e., opportunity for sexual selection) was significantly greater for males relative to females, and while the relationship between mating success and reproductive success (i.e., Bateman gradient) was significantly positive for both sexes, a greater proportion of reproductive success was explained by mating success for males (r 2 = 0.80) than females (r 2 = 0.59). Overall, the population displayed a polygynandrous mating system, whereby both sexes experienced variation in mating success due to multiple mating, and sexual selection was variable across sexes. Tests for evidence of sexual selection indicated the interaction between mating success and total length best‐predicted relative reproductive success. We failed to detect a signal of inbreeding avoidance among breeding adults, but the group of parents that produced progeny were on average slightly less related than adults that did not produce progeny. Lastly, we estimated the effective number of breeders (N b) and effective population size (N e) and identified while N b was lower than N e, both are sufficiently high to suggest Yellowstone Cutthroat Trout in Burns Creek represent a genetically stable and diverse population.

The battle between harvest and natural selection creates small and shy fish

Harvest of fish and wildlife, both commercial and recreational, is a selective force that can induce evolutionary changes to life history and behavior. Naturally selective forces may create countering selection pressures. Assessing natural fitness represents a considerable challenge in broadcast spawners. Thus, our understanding about the relative strength of natural and fisheries selection is slim. In the field, we compared the strength and shape of harvest selection to natural selection on body size over four years and behavior over one year in a natural population of a freshwater top predator, the northern pike (Esox lucius). Natural selection was approximated by relative reproductive success via parent-offspring genetic assignments over four years. Harvest selection was measured by comparing individuals susceptible to recreational angling with individuals never captured by this gear type. Individual behavior was measured by high-resolution acoustic telemetry. Harvest and natural size selection operated with equal strength but opposing directions, and harvest size selection was consistently negative in all study years. Harvest selection also had a substantial behavioral component independent of body length, while natural behavioral selection was not documented, suggesting the potential for directional harvest selection favoring inactive, timid fish. Simulations of the outcomes of different fishing regulations showed that traditional minimum size-based harvest limits are unlikely to counteract harvest selection without being completely restrictive. Our study suggests harvest selection may be inevitable and recreational fisheries may thus favor small, inactive, shy, and difficult-to-capture fish. Increasing fractions of shy fish in angling-exploited stocks would have consequences for stock assessment and all fisheries operating with hook and line.

Equivocal evidence for a change in balance between selfing and pollinator‐mediated reproduction in annual Brassicaceae growing along a rainfall gradient

AbstractBecause many plants ensure their reproductive success using some level of self‐fertilization, it is predicted that the balance between pollinator‐mediated pollen transfer and autonomous selfing will be correlated with habitat harshness and pollinator activity. To study these correlations, we used three annual species that differ in their self‐incompatibility and compared their reproductive success in the form of fruit production and seed viability, with and without pollinators, along a rainfall gradient and in relation to pollinator activity. We found that pollinator activity decreased with increasing aridity. Although the reproductive success in the absence of pollinators fitted the species’ level of self‐incompatibility, no clear trend was detected between pollinator visits or site harshness and relative reproductive success. Only one of the species, Sinapis alba, showed a consistent increase in autonomous selfing with both increased aridity and decreased pollinator visits, a finding that was supported by an increase in the viability of seeds produced in the absence of pollinators. Nevertheless, unlike our prediction, the absolute number of seeds produced per S. alba flower increased with aridity. Our overall findings suggest that despite decreasing pollinator abundance and diversity with increasing aridity, the reproductive success of annual species, even of those with high self‐incompatibility, remains relatively high. Although there may be several mechanisms governing these results, the overall constant seed production despite environmental uncertainties may be part of the mechanism by which plants in arid ecosystems buffer environmental variations.

Corrigendum to: Captive ancestry upwardly biases estimates of relative reproductive success.

Corrigendum to: Captive ancestry upwardly biases estimates of relative reproductive success.

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.

Food-provisioning negatively affects calf survival and female reproductive success in bottlenose dolphins

Food-provisioning of wildlife can facilitate reliable up-close encounters desirable by tourists and, consequently, tour operators. Food-provisioning can alter the natural behavior of an animal, encouraging adverse behavior (e.g. begging for food handouts), and affect the reproductive success and the viability of a population. Studies linking food-provisioning to reproductive success are limited due to the lack of long-term datasets available, especially for long-lived species such as marine mammals. In Bunbury, Western Australia, a state-licensed food-provisioning program offers fish handouts to a limited number of free-ranging bottlenose dolphins (Tursiops aduncus). Coupled with long-term historical data, this small (<200 individuals), resident dolphin population has been extensively studied for over ten years, offering an opportunity to examine the effect of food-provisioning on the reproductive success of females (ntotal = 63; nprovisioned females = 8). Female reproductive success was estimated as the number of weaned calves produced per reproductive years and calf survival at year one and three years old was investigated. The mean reproductive success of provisioned and non-provisioned females was compared using Bayes factor. We also used generalized linear models (GLMs) to examine female reproductive success in relation to the occurrence of food-provisioning, begging behavior and location (within the study area). Furthermore, we examined the influence of these variables and birth order and climatic fluctuations (e.g. El Niño Southern Oscillation) on calf survival. Bayes factor analyses (Bayes factor = 6.12) and results from the best fitting GLMs showed that female reproductive success and calf survival were negatively influenced by food-provisioning. The negative effects of food-provisioning, although only affecting a small proportion of the adult females’ population (13.2%), are of concern, especially given previous work showing that this population is declining.

Host density predicts the probability of parasitism by avian brood parasites.

The spatial distribution of hosts can be a determining factor in the reproductive success of parasites. Highly aggregated hosts may offer more opportunities for reproduction but can have better defences than isolated hosts. Here we connect macro- and micro-evolutionary processes to understand the link between host density and parasitism, using avian brood parasites as a model system. We analyse data across more than 200 host species using phylogenetic comparative analyses and quantify parasitism rate and host reproductive success in relation to spatial distribution using field data collected on one host species over 6 years. Our comparative analysis reveals that hosts occurring at intermediate densities are more likely to be parasitized than colonial or widely dispersed hosts. Correspondingly, our intraspecific field data show that individuals living at moderate densities experience higher parasitism rates than individuals at either low or high densities. Moreover, we show for the first time that the effect of host density on host reproductive success varies according to the intensity of parasitism; hosts have greater reproductive success when living at high densities if parasitism rates are high, but fare better at low densities when parasitism rates are low. We provide the first evidence of the trade-off between host density and parasitism at both macro- and micro-evolutionary scales in brood parasites. This article is part of the theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.

Sexual selection and inbreeding: Two efficient ways to limit the accumulation of deleterious mutations.

Theory and empirical data showed that two processes can boost selection against deleterious mutations, thus facilitating the purging of the mutation load: inbreeding, by exposing recessive deleterious alleles to selection in homozygous form, and sexual selection, by enhancing the relative reproductive success of males with small mutation loads. These processes tend to be mutually exclusive because sexual selection is reduced under mating systems that promote inbreeding, such as self‐fertilization in hermaphrodites. We estimated the relative efficiency of inbreeding and sexual selection at purging the genetic load, using 50 generations of experimental evolution, in a hermaphroditic snail (Physa acuta). To this end, we generated lines that were exposed to various intensities of inbreeding, sexual selection (on the male function), and nonsexual selection (on the female function). We measured how these regimes affected the mutation load, quantified through the survival of outcrossed and selfed juveniles. We found that juvenile survival strongly decreased in outbred lines with reduced male selection, but not when female selection was relaxed, showing that male‐specific sexual selection does purge deleterious mutations. However, in lines exposed to inbreeding, where sexual selection was also relaxed, survival did not decrease, and even increased for self‐fertilized juveniles, showing that purging through inbreeding can compensate for the absence of sexual selection. Our results point to the further question of whether a mixed strategy combining the advantages of both mechanisms of genetic purging could be evolutionary stable.

Long-term evaluation of fitness and demographic effects of a Chinook Salmon supplementation program.

While the goal of supplementation programs is to provide positive, population‐level effects for species of conservation concern, these programs can also present an inherent fitness risk when captive‐born individuals are fully integrated into the natural population. In order to evaluate the long‐term effects of a supplementation program and estimate the demographic and phenotypic factors influencing the fitness of a threatened population of Chinook Salmon (Oncorhynchus tshawytscha), we genotyped tissue samples spanning a 19‐year period (1998–2016) to generate pedigrees from adult fish returning to Johnson Creek, Idaho, USA. We expanded upon previous estimates of relative reproductive success (RRS) to include grandparentage analyses and used generalized linear models to determine whether origin (hatchery or natural) or phenotypic traits (timing of arrival to spawning grounds, body length, and age) significantly predicted reproductive success (RS) across multiple years. Our results provide evidence that this supplementation program with 100% natural‐origin broodstock provided a long‐term demographic boost to the population (mean of 4.56 times in the first generation and mean of 2.52 times in the second generation). Overall, when spawning in nature, hatchery‐origin fish demonstrated a trend toward lower RS compared to natural‐origin fish (p < 0.05). However, when hatchery‐origin fish successfully spawned with natural‐origin fish, they had similar RS compared to natural by natural crosses (first‐generation mean hatchery by natural cross RRS = 1.11 females, 1.13 males; second‐generation mean hatchery by natural cross RRS = 1.03 females, 1.08 males). While origin, return year, and body length were significant predictors of fitness for both males and females (p < 0.05), return day was significant for males but not females (p > 0.05). These results indicate that supplementation programs that reduce the potential for genetic adaptation to captivity can be effective at increasing population abundance while limiting long‐term fitness effects on wild populations.

Maladaptive secondary sexual characteristics reduce the reproductive success of hybrids between native and non-native salmonids.

Human‐mediated hybridization between introduced and native species is one of the most serious threats to native taxa. Although field studies have attempted to quantify the relative fitness or reproductive success of parental species and their hybrids, only a few studies have unraveled the factors determining the fitness of hybrids. Here, we hypothesized that maladaptive secondary sexual characteristics may reduce fitness of hybrids between two fish species. To test this, we evaluated the reproductive success of introduced brook trout (BT: Salvelinus fontinalis), native white‐spotted charr (WSC: S. leucomaenis) and their hybrids in a natural stream in Hokkaido, Japan, where the two parental species show remarkably different male secondary sexual characteristics, such as elongated jaws and deeper bodies. We predicted that introgression from WSC is maladaptive for BT males because the BT male has more prominent secondary sexual characteristics. Our results suggest that both sexual selection and outbreeding depression in males and females significantly influence an individual's reproductive success. Our results also suggest that asymmetric introgression may increase the risks to persistence in the recipient species.