A small number of anadromous females drive reproduction in a brown trout (Salmo trutta) population in an English chalk stream

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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.     

Grandparentage assignments identify unexpected adfluvial life history tactic contributing offspring to a reintroduced population

Diversity in life history tactics contributes to the persistence of a population because it helps to protect against stochastic environments by varying individuals in space and time. However, some life history tactics may not be accounted for when assessing the demographic viability of a population. One important factor in demographic viability assessments is cohort replacement rate (CRR), which is defined as the number of future adults produced by an adult. We assessed if precocial resident males (<age‐3) and adfluvial Chinook salmon (Oncorhynchus tshawytscha), adults that reside in freshwater their entire lives, contributed offspring to a reintroduced population from 2008 to 2013. We found that 9 ± 5% of offspring with an unassigned parent remained unexplained after accounting for sources of human error. Using grandparentage assignments, we identified 31 precocial resident males and 48 probable adfluvial Chinook salmon produced by anadromous mate pairs from 2007 to 2012. Previously published CRR estimates for the 2007 and 2008 reintroduced adults, based on only anadromous returning adult offspring, were 0.40 and 0.31, respectively. By incorporating adfluvial females, we found CRR estimates increased by 17% (CRR: 0.46) and 13% (CRR: 0.35) for the 2007 and 2008 cohorts, respectively.     

Condition-dependent reproductive tactics by large and small anadromous male sockeye salmon Oncorhynchus nerka

Reproductive tactics, aggression and courtship by large and small (jack) male sockeye salmon Oncorhynchus nerka with ripe females in temporary enclosures depended on the size and behaviour of the competitor, not the size of the focal male. When paired with other jacks or with large but less aggressive males, jacks fought for access to ripe females rather than sneaking, but used sneaking tactics when paired with large, aggressive males. Dominant males of both size groups courted actively and females spawned readily with all combinations of males.     

Differential dispersal patterns of male and female masu salmon fry

Behavioural experiments using artificial and natural stream channels were undertaken to determine whether there were differences in dispersal between newly emerged male and female masu salmon. Eyed eggs from a cross with wild spawners were planted in the middle pool of an artificial channel. After emergence, more males than females moved into an upstream trap, while fewer males moved downstream. In a natural stream, eyed eggs were marked with alizarin complexone to distinguish them from naturally spawned eggs and these were planted into artificial redds. More newly emerged male fry remained at the planted site than female fry. In contrast, more females moved downstream than males. These results imply that differences in dispersal patterns between male and female masu salmon fry are genetically controlled.     

Maternal influence on early lipid content in an introduced partially anadromous population of Oncorhynchus mykiss

This study evaluated how the maternal migratory tactic in a partially anadromous population of Oncorhynchus mykiss may influence the early energetic status of their offspring. Total lipid content variation (% dry mass) of recently emerged fry caught in the Santa Cruz River, Argentina, was evaluated as a function of their maternal origin (anadromous v. resident) and fork length (L_F). Lipid content of fry decreased with L_F and was higher for offspring of anadromous mothers.     

Yearling male great tits, Parus major, suffer more strongly from cuckoldry than older males

In the majority of socially monogamous bird species, females have offspring sired by males other than their social mate as the result of extra-pair copulations. While it is widely recognised that there is considerable variation in the frequency of extra-pair paternity between species, between populations of a species and between individuals of a population, determinants of this variation are surprisingly difficult to establish. With respect to individual variation within a population, it is an important step to test for male and female correlates of cuckoldry to better understand the patterns as well as the adaptive significance of extra-pair mating behaviour. Here, we analysed patterns of extra-pair paternity in relation to male age, female age and their interaction in the great tit Parus major, a socially monogamous passerine with a moderate frequency of extra-pair paternity. Based on a large sample of 316 genotyped first broods from five successive years, we failed to demonstrate interaction effects of male and female age on both the proportion of extra-pair offspring and the likelihood that at least one extra-pair offspring is present within a brood. However, both the proportion of extra-pair offspring and the likelihood of paternity loss were higher for yearling as compared to older males, while this was not the case for yearling vs. older females. Furthermore, the proportion of extra-pair offspring within a brood decreased with increasing age of the attending male in within-individual analyses. We found a comparable effect also for attending females in within-individual analyses, but only when excluding two individuals with 100% extra-pair paternity. A female (extra-pair) mating preference for older males and/or a limited ability of yearling males to prevent cuckoldry in their broods could explain these age-related patterns of paternity loss. Effect sizes, however, were not particularly large and substantial residual variation within age categories suggests the importance of further yet unidentified determinants of variation in paternity loss in the study population.     

Effects of Paternal Reproductive Tactic on Juvenile Behaviour and Kin Recognition in Chinook Salmon (Oncorhynchus tshawytscha)

Salmonids are characterized by alternative reproductive tactics, which can lead to an asymmetry in relatedness among offspring within nests and consequently the benefit of discriminating among nestmates. In this study, we examined the effect of paternal reproductive tactic on juvenile behaviour and kin discrimination in Chinook salmon. We created maternal half‐sibling families by collecting eggs from mature females and fertilizing one‐half with the milt of a precocious 2‐yr‐old male and the other half with the milt of a non‐precocious 4‐yr‐old male. These families were reared in full‐sibling groups for approximately 9 mo, and social interactions were then observed in groups of six fish of mixed relatedness. We found evidence for kin discrimination, as significantly less aggression was directed towards related fish than unrelated fish, and the same trends were observed regardless of whether social interactions included full‐siblings or half‐siblings. These results show that familiarity is not required to recognize kin and thereby implicate phenotype matching as the mechanism of kin recognition. We also found that the offspring of 2‐yr‐old males were larger and more aggressive than the offspring of 4‐yr‐old males, which is consistent with other studies showing that precocious males are the fastest‐growing members of their cohort. However, kin‐directed behaviours did not differ between the offspring of 2‐ and 4‐yr‐old males.     

First evidences of sexual selection by mate choice in marine zooplankton

Sexual selection is potentially important in marine zooplankton, presumably the most abundant metazoans on earth, but it has never been documented. We examine the conditions for sexual selection through mate choice and describe mating preferences in relation to size in a marine zooplankter, the pelagic copepod Acartia tonsa. Males produce spermatophores at a rate (~1 day⁻¹) much lower than known female encounter rates for most of the year and the decision to mate a particular female thus implies lost future opportunities. Female egg production increases with female size, and males mating larger females therefore sire more offspring per mating event. Similarly, females encounter males more frequently than they need to mate. Large males produce larger spermatophores than small males and the offspring production of female increases with the size of the spermatophore she receives. Additionally, large spermatophores allow females to fertilize eggs for a longer period. Thus, mating with large males reduces the female’s need for frequent matings and she may sire sons that produce more offspring because size is heritable in copepods. Finally, we show that both males and females mate preferentially with large partners. This is the first demonstration of sexual selection by mate choice in a planktonic organism.     

Reduced reproductive success of hatchery coho salmon in the wild: insights into most likely mechanisms

Supplementation of wild salmonids with captive-bred fish is a common practice for both commercial and conservation purposes. However, evidence for lower fitness of captive-reared fish relative to wild fish has accumulated in recent years, diminishing the apparent effectiveness of supplementation as a management tool. To date, the mechanism(s) responsible for these fitness declines remain unknown. In this study, we showed with molecular parentage analysis that hatchery coho salmon (Oncorhynchus kisutch) had lower reproductive success than wild fish once they reproduced in the wild. This effect was more pronounced in males than in same-aged females. Hatchery spawned fish that were released as unfed fry (age 0), as well as hatchery fish raised for one year in the hatchery (released as smolts, age 1), both experienced lower lifetime reproductive success (RS) than wild fish. However, the subset of hatchery males that returned as 2-year olds (jacks) did not exhibit the same fitness decrease as males that returned as 3-year olds. Thus, we report three lines of evidence pointing to the absence of sexual selection in the hatchery as a contributing mechanism for fitness declines of hatchery fish in the wild: (i) hatchery fish released as unfed fry that survived to adulthood still had low RS relative to wild fish, (ii) age-3 male hatchery fish consistently showed a lower relative RS than female hatchery fish (suggesting a role for sexual selection), and (iii) age-2 jacks, which use a sneaker mating strategy, did not show the same declines as 3-year olds, which compete differently for females (again, implicating sexual selection).     

To breed or not to breed: causes and implications of non-breeding habit in the willow tit Parus montanus

Causes and consequences of non-breeding in willow tits were studied in northern Finland during 1986–1992. The breeding status was sex and age biased; males and yearling birds were in excess among the non-reproducers. Due to sex bias in the population it appeared detrimental for males to lose a mate, especially shortly before breeding. Lack of a mate was a important factor for males not reproducing (37% of non-breeding males) than for females (14%). Most of the non-breeding birds maintained a pair bond which only rarely broke up for the next breeding season (divorce rate 5.5%). This implies that parental incompatibility is not a possible explanation for pairs not reproducing. Males that did not breed tended to survive better than reproducing ones, whereas such a relationship was not found for females. It is possible that this sex-related difference in survival cost is attributable to quality differences among non-breeding individuals. It was especially low-quality yearling females, with low survival prospects, that were responsible for the discrepancy. The proportion of non-breeding females in the population correlated highly with clutch size and subsequent juvenile survival. It is therefore suggested that for most of these females non-breeding is a phenotypic response to low offspring value in the prevailing circumstances (inter-generational tradeoff). However, it is uncertain whether willow tits in a northern population can use breeding density as an indicator of changing survival prospects of their descendants, as suggested by Ekman and Askenmo (1986) for southern Sweden.     

Spawning characteristics of the anadromous brown trout in a small Swedish stream

The spawning pattern of the anadromous brown trout Salmo trutta was studied in Själsöån, a small stream in Gotland, Sweden, during eight winters between 1992–1993 and 1999–2000. The total length ( L _T) at spawning was normally distributed (185–890 mm) for females and multimodal for males (300, 400 and 550 mm most frequent length classes). Spawning males were significantly younger (2+ to 4+ years) than females (3+ to 5+ years). The sex‐ratio at the beginning and at the end of the spawning season favoured males. The mean ±  s . d . number of spawners was 70 ± 16 individuals per year. Migration into and out of the stream occurred between November and June. The highest number of spawning fish was found in the stream at the end of November or at the beginning of December. Migration mainly occurred during high water flow and at night. The majority of the females entered the stream and spawned the same (29·3% of all the females) or the next night (32·8% of all the females) while males may have stayed for 2 to 3 weeks (21·3% of all the males) in the stream before spawning. Males usually remained much longer in the stream (mean ±  s . d . 45 ± 56 days) than females (16 ± 30 days). Females lost more mass in the stream (mean ±  s . d . 17·3 ± 8·6%) than males (7·7 ± 9·6%). For both sexes, mass loss was positively correlated with the time spent in the stream. Only 7·3% of the males and 5·7% of the females occurred in the stream for >1 year. Spawning took place only during the night.     

Multi-male mating and female choice increase offspring growth in the spiderNeriene litigiosa(Linyphiidae)

The purpose of this controlled-breeding study was to investigate the viability consequences of female choice and sequential polyandry for offspring in a way that would separate the influences of these two aspects of female sexual behaviour. Female sierra dome spiders,Neriene litigiosa(=Linyphia litigiosa) typically mate two to three times before production of their first batch of eggs, although some females (ca 16%) mate only once. Strong fighters are preferred as first mates and principal sires. Large males that give a vigorous performance during copulatory courtship are preferred as sires among a female's secondary mates. In this study, the number of matings by free-living females was experimentally controlled and the size and copulatory vigour of all the females’ mates were recorded. At the end of the breeding season, the females were collected and their broods were obtained in captivity. Randomly chosen subsets of 28 female's offspring were reared under standard conditions during the following spring. The spiderlings were reared for 27 days in the company of siblings under conditions that would encourage the expression of genetic variation in viability, and growth and survivorship were monitored. After controlling for maternal effects (i.e. female size and oviposition date) and variance in feeding opportunities among rearing groups, both mate number and mate size were positively and independently related to offspring growth rates and the size of offspring attained after emergence from the natal cocoon. The results support the hypotheses that the preference for large males yields viability benefits to offspring and that polyandry can augment the benefits of selective mating.     

Potential fitness benefits from mate selection in the Atlantic cod (Gadus morhua)

Little evidence of benefits from female mate choice has been found when males provide no parental care or resources. Yet, good genes models of sexual selection suggest that elaborated male sexual characters are reliable signals of mate quality and that the offspring of males with elaborate sexual ornaments will perform better than those of males with less elaborate ornaments. We used cod (Gadus morhua L.), an externally fertilizing species where males provide nothing but sperm, to examine the potential of optimal mate selection with respect to offspring survival. By applying in vitro fertilizations, we crossed eight females with nine males in all possible combinations and reared each of the 72 sib groups. We found that offspring survival was dependent on which female was mated with which male and that optimal mate selection has the potential to increase mean offspring survival from 31.9 to 55.6% (a 74% increase). However, the size of the male sexual ornaments and sperm quality (i.e. sperm velocity and sperm density) could not predict offspring survival. Thus, even if there may be large fitness benefits of mate selection, we might not yet have identified the male characteristics generating high offspring survival.     

Intensive genetic assessment of the mating system and reproductive success in a semi-closed population of the mottled sculpin,Cottus bairdi

Most genetic surveys of parentage in nature sample only a small fraction of the breeding population. Here we apply microsatellite markers to deduce the genetic mating system and assess the reproductive success of females and males in an extensively collected, semi-closed stream population of the mottled sculpin fish, Cottus bairdi. In this species, males guard nest rocks where females deposit the eggs for fertilization. The potential exists for both males and females to mate with multiple partners and for males to provide parental care to genetically unrelated offspring. Four hundred and fifty-five adults and subadults, as well as 1,259 offspring from 23 nests, were genotyped at five polymorphic microsatellite loci. Multilocus maternal genotypes, deduced via genetic analyses of embryos, were reconstructed for more than 90% of the analysed nests, thus allowing both male and female reproductive success to be estimated accurately. There was no genetic evidence for cuckoldry, but one nest probably represents a takeover event. Successful males spawned with a mean of 2.8 partners, whereas each female apparently deposited her entire clutch of eggs in a single nest (mean fecundity = 66 eggs/female). On average, genetically deduced sires and dams were captured 1.6 and 9.3 metres from their respective nests, indicating little movement by breeders during the spawning season. Based on a 'genetic mark-recapture' estimate, the total number of potentially breeding adults (c. 570) was an order-of-magnitude larger than genetically based estimates of the effective number of breeders (c. 54). In addition, significantly fewer eggs per female were deposited in single than in multidam nests. Not only were perceived high-quality males spawning with multiple partners, but they were receiving more eggs from each female.     

Can non-native smallmouth bass, <Emphasis Type="Italic">Micropterus dolomieu</Emphasis>, be swamped by hatchery fish releases to increase juvenile Chinook salmon, <Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis>, survival?

One of the strategies that can be used to reduce predation impacts to valued fish species is by swamping predators with more prey than they can eat. We examined whether this approach was viable by calculating the maximum bioenergetic consumption potential of non-native smallmouth bass Micropterus dolomieu on fall Chinook salmon Oncorhynchus tshawytscha juveniles in the Yakima River throughout the spring between 1998 and 2002 and comparing those estimates to previously published estimates of fall Chinook salmon consumption. We found that the smallmouth bass population consumed fall Chinook salmon well below their bioenergetic potential. However, individual smallmouth bass that were piscivorous were eating other food items at a level near satiation. Furthermore, the maximum consumption potential was relatively low prior to mid-April, and then increased substantially to a peak in May. Predation mortality to hatchery fall Chinook salmon could be reduced within a year by releasing hatchery fall Chinook salmon that will emigrate quickly prior to mid-April, when predation potential is still very low. However, attempting to swamp predators with hatchery Chinook salmon to benefit naturally produced Chinook salmon poses uncertain benefits to natural origin fish and likely unacceptable costs to hatchery fish. Considerable swamping is occurring by other naturally produced fish species in the Yakima River such as dace Rhinichthys spp., mountain whitefish Prosopium williamsoni, and crayfish Pacificastus spp. Therefore, it is important to consider impacts to these non-target species because they could have indirect predation impacts on Chinook salmon.     

Reproductive investment patterns,sperm characteristics,and seminal plasma physiology in alternative reproductive tactics of Chinook salmon (Oncorhynchus tshawytscha)

Although alternative reproductive tactics (ARTs) are common across a range of taxa, little is known about whether the different tactics have adapted to sperm competition risk. Chinook salmon, Oncorhynchus tshawytscha, have two ARTs: large males that participate in dominance‐based hierarchies for access to spawning females, known as hooknoses, and small males that attempt to sneak fertilizations during spawning events from peripheral positions, known as jacks. Jacks continually face sperm competition risk because they always spawn in the presence of another male, whereas hooknoses face relatively low sperm competition risk because other males are not always present during spawning events. Based on the sneak‐guard model of sperm competition this asymmetry in sperm competition risk predicts that jacks ought to invest significantly more into sperm‐related traits important for sperm competition success relative to hooknoses. In the present study we report on reproductive investment patterns, sperm characteristics, and seminal plasma physiology of males that exhibit ARTs in Chinook salmon. We found that jacks invest significantly more of their somatic tissue into gonads compared with hooknoses. Sperm velocity also varied significantly between the ARTs, with jacks having significantly faster sperm than hooknoses. No significant differences in seminal plasma physiology metrics related to sperm quality were detected between the ARTs. We interpret these sperm investment patterns in light of the sneak‐guard model of sperm competition that is based on differences in sperm competition risk and alternative investment possibilities among ARTs. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Females produce larger eggs for large males in a paternal mouthbrooding fish.

When individuals receive different returns from their reproductive investment dependent on mate quality, they are expected to invest more when breeding with higher quality mates. A number of studies over the past decade have shown that females may alter their reproductive effort depending on the quality/attractiveness of their mate. However, to date, despite extensive work on parental investment, such a differential allocation has not been demonstrated in fish. Indeed, so far only two studies from any taxon have suggested that females alter the quality of individual offspring according to the quality/attractiveness of their mate. The banggai cardinal fish is an obligate paternal mouth brooder where females lay few large eggs. It has previously been shown that male size determines clutch weight irrespective of female size in this species. In this study, I investigated whether females perform more courtship displays towards larger males and whether females allocate their reproductive effort depending on the size of their mate by experimentally assigning females to either large or small males. I found that females displayed more towards larger males, thereby suggesting a female preference for larger males. Further, females produced heavier eggs and heavier clutches but not more eggs when paired with large males. My experiments show that females in this species adjust their offspring weight and, thus, presumably offspring quality according to the size of their mate.     

Comparison of potential increases in juvenile salmonid rearing habitat capacity among alternative restoration scenarios,Trinity River,California

River restoration plans often propose multiple rehabilitation actions to address key habitat impairments, but they rarely attempt to quantify the potential benefits of alternative sets of actions for targeted biota. We use geomorphic and biological analyses to estimate restoration potential under three alternative scenarios for a 64‐km section of the Trinity River, California, between the North Fork Trinity River and Lewiston Dam, which is the focus of habitat rehabilitation efforts under the Trinity River Restoration Program. The three scenarios are (1) increasing habitat quality by wood additions and alcove construction, (2) increasing habitat quantity by increasing sinuosity and side‐channel length, and (3) increasing both habitat quality and quantity. For each scenario, we used existing stream habitat and juvenile salmonid data from previous studies to estimate potential improvements in fry or pre‐smolt production. The potential increase in Oncorhynchus tshawytscha (Chinook salmon) and O. mykiss (steelhead) fry rearing capacity was 62 and 67%, respectively, for Scenario 1 (increasing habitat quality), and 36 and 44% for Scenario 2 (increasing habitat quantity). Only the most optimistic Scenario 3 (increasing both habitat quality and quantity) more than doubles potential juvenile salmonid production (112% increase in Chinook fry capacity and 107% increase in steelhead fry capacity). These quantitative predictions are useful in developing realistic restoration targets and evaluating whether proposed restoration actions can achieve the aims of a restoration program.     

A comparative study of the performance of family groups of Atlantic salmon reared in tanks and released in a river

A study was conducted in in the Dale River as a part of a stock enhancement programme. The aim was to compare growth and the incidence of precocious maturation between offspring from precocious and large maturing males, and to study genotype‐environment interactions. River and hatchery performance was compared for 5 × 2 maternal half‐sib family groups, which were stocked as 0+ juveniles or maintained in the hatchery throughout. To identify the offspring, the broodfish were characterized by DNA‐fingerprinting (eight microsatellite markers). Smolt size of 1+ hatchery‐reared smolt and fish caught in a smolt trap, and the size and incidence of precocious maturation among 1+ hatchery‐reared parr and 1+ and 2+ parr caught in the river are compared between the families.