General patterns of sexual dimorphism in graylings (Thymallus), with a comparison to other salmonid species

Among fishes, salmonids (family Salmonidae) have attracted a great deal of research attention focused on sexual dimorphism and associated selective forces. Most of this research has been directed toward anadromous and mostly semelparous salmon and trout (Oncorhynchus, Salmo), and comparatively little is known about intersexual variability in strictly iteroparous freshwater salmonids. We examined a comprehensive data set of 28 linear morphometric characters in 11 of 15 currently recognised species of grayling (Thymallinae, Thymallus), a genus consisting of iteroparous species only, to identify general patterns of intersexual morphological variability. Overall, we found that all grayling species show common sex-specific traits particularly relating to size dimensions of the dorsal, anal, pelvic and pectoral fins. Although the magnitude of sexual dimorphism differed among species, there was no significant phylogenetic signal associated with these differences across the genus. These results are discussed in terms of the assumed selection pressures driving sexual dimorphism in graylings and are compared to existing knowledge in Salmonidae as a whole where similarities and differences with both Salmoninae and Coregoninae exist. The present study provides the first detailed genus-wide comparison of sexually dimorphic phenotypic characters in graylings, and highlights the need for more large-scale comparative studies in multiple salmonid species to better understand general macroevolutionary trends among this important group of freshwater fishes.

Life history traits and dispersal shape neutral genetic diversity in metapopulations

Genetic diversity at population scale, depends on species life-history traits, population dynamics and local and global environmental factors. We first investigate the effect of lifehistory traits on the neutral genetic diversity of a single population using a deterministic mathematical model. When the population is stable, we show that semelparous species with precocious maturation and iteroparous species with delayed maturation exhibit higher diversity because their life history traits tend to balance the lifetimes of non reproductive individuals (juveniles) and adults which reproduce. Then, we extend our model to a metapopulation to investigate the additional effect of dispersal on diversity. We show that dispersal may truly modify the local effect of life history on diversity. As a result, the diversity at the global scale of the metapopulation differ from the local diversity which is only described through local life history traits of the populations. In particular, dispersal usually promotes diversity at the global metapopulation scale.

Social aging: Late-life fitness gains explain the absence of a selection shadow in ants

A key hypothesis for the occurrence of senescence is a decrease in the selection strength because of low late-life fitness – the so-called selection shadow. However, in social insects, aging is considered a plastic trait and senescence seems to be absent. By life-long tracking of 102 ant colonies, we find that queens increase sexual productivity in late life regardless of their absolute lifespan or worker investment. This indicates a genetically accommodated adaptive shift towards increasingly queen-biased caste ratios over the course of a queens’ life. Furthermore, mortality decreased with age, supporting the hypothesis that aging is adaptive. We argue that selection for late life reproduction diminishes the selection shadow of old age and leads to the apparent absence of senescence in ants, in contrast to most iteroparous species.

The various ways that anadromous salmonids use lake habitats to complete their life history

Despite the preponderance of exorheic lakes in rivers home to anadromous salmonids, little research has focused on how salmon, trout, and char use lakes as part of their anadromous life histories. The literature on this subject has so far revealed that some parr move into lakes to feed and grow before smoltification but that smolts moving through lakes tend to have high mortality in disproportion to what is observed in other habitats they migrate in or through. Adults have been observed using lakes for behavioural thermoregulation prior to spawning, and kelts of iteroparous species often exploit lakes to overwinter before returning to sea to recondition. We summarized knowledge on lakes as salmonid habitat and identified knowledge gaps about the use of lakes by anadromous salmonids related to whether lakes are barriers that structure genetics of populations, whether mortality in lakes is compensatory or additive, and whether systems with lakes have higher rates of repeat spawning among iteroparous salmonids. Human activities that alter lakes require further study to understand how changes in temperature, oxygen, ice, or circulation affect navigation and fate.

Climbing up the ladder: male reproductive behaviour changes with age in a long-lived fish

High reproductive performance is the key attribute of male fitness, especially due to the high reproductive skew among the males of most animal species. Males of long-lived iteroparous species have opportunities to improve upon their previous reproductive attempts with increasing age. We collected individual-specific reproductive behaviour and age data on a cyprinid fish, the asp (Leuciscus aspius), from 2015 to 2019. We tested whether males changed their performance over time using a unique dataset where individual performance was recorded yearly with passive telemetry. Individual fish behaviour was tracked from one to five reproductive seasons at least a year after the tagging. Fish were scored by measures of quality (first arrival time, number of visits and time spent in the reproductive grounds, and encountered proportion of males to all adult fish). In general, fish improved in the first three metrics with age, suggesting a shift towards behaviours likely to enhance reproductive success as individuals aged. A larger size at tagging was predictive of earlier fish arrival on the spawning ground in subsequent years. Our study therefore demonstrates the importance of age as a factor when considering the potential reproductive success of long-lived fish species. Significance statement High reproductive performance is the key attribute of male fitness. Males of long-lived species reproducing multiple times in their life have opportunities to improve upon their previous reproductive performance with increasing age. In this 5-year study, we tracked a large cyprinid fish with telemetry systems during their reproduction. We investigated the age-related behavioural changes in males and demonstrated the improvement of male reproductive timing and length of stay with potential repercussions for male’s reproductive output. We emphasize the importance of old and experienced individuals among the fish population, which are often targeted and selectively removed from the human-managed waters.

Inter-individual differences in the foraging behavior of breeding Adélie penguins are driven by individual quality and sex

Inter-individual differences in demographic traits of iteroparous species can arise through learning and maturation, as well as from permanent differences in individual ‘quality’ and sex-specific constraints. As the ability to acquire energy determines the resources an individual can allocate to reproduction and self-maintenance, foraging behavior is a key trait to study to better understand the mechanisms underlying these differences. So far, most seabird studies have focused on the effect of maturation and learning processes on foraging performance, while only a few have included measures of individual quality. Here, we investigated the effects of age, breeding experience, sex, and individual breeding quality on the foraging behavior and location of 83 known-age Adélie penguins at Cape Bird, Ross Sea, Antarctica. Over a 2 yr period, we showed that (1) high-quality birds dived deeper than lower quality ones, apparently catching a higher number of prey per dive and targeting different foraging locations; (2) females performed longer foraging trips and a higher number of dives compared to males; (3) there were no significant age-related differences in foraging behavior; and (4) breeding experience had a weak influence on foraging behavior. We suggest that high-quality individuals have higher physiological ability, enabling them to dive deeper and forage more effectively. Further inquiry should focus on determining the physiological differences among penguins of different quality.

How can we augment the few that remain? Using stable population dynamics to aid reintroduction planning of an iteroparous species

Restoration of depleted populations is an important method in biological conservation. Reintroduction strategies frequently aim to restore stable, increasing, self-sustaining populations. Knowledge of asymptotic system dynamics may provide advantage in selecting reintroduction strategies. We introduce interactive software that is designed to identify strategies for release of females that are immediately aligned with stable population dynamics from species represented by 2-, 3-, 4-, and 5-stage life history strategies. The software allows managers to input a matrix of interest, the desired number of breeding females, and the desired management timeline, and calls upon stable population theory to give release strategies that are in concert with both stable population status and the management goals. We demonstrate how the software can aid in assessing various strategies ahead of a hypothetical restoration. For the purpose of demonstration of the tool only, we use published vital rates of an ungulate species, but remark that the selection of species for demonstration is not central to the use of this tool. Adaption of this tool to real-life restorations of any 2-, 3-, 4-, or 5-stage iteroparous species may aid in understanding how to minimize undesirable recovery complications that may naturally arise from transient population dynamics. The software is freely available at: https//cwhl.vet.cornell.edu/tools/stapopd.

Maternal longevity and offspring sex in wild ungulates

In species with sexual size dimorphism, the offspring of the larger sex usually have greater energy requirements and may lead to greater fitness costs for parents. The effects of offspring sex on maternal longevity, however, have only been tested in humans. Human studies produced mixed results and considerable debate mainly owing to the difficulty of distinguishing the effects of sexual dimorphism from sociocultural factors. To advance this debate, we examined how the relative number of sons influenced maternal longevity in four species of free-living ungulates (Soay sheep Ovis aries ; bighorn sheep, Ovis canadensis ; red deer, Cervus elaphus ; mountain goat, Oreamnos americanus ), with high male-biased sexual size dimorphism but without complicating sociocultural variables. We found no evidence for a higher cumulative cost of sons than of daughters on maternal longevity. For a given number of offspring, most females with many sons in all four populations lived longer than females with few sons. The higher cost of sons over daughters on maternal lifespan reported by some human studies may be the exception rather than the rule in long-lived iteroparous species.

Prey productivity and predictability drive different axes of life-history variation in carnivorous marsupials

Variation in life-history strategies has usually been characterized as a single fast–slow continuum of life-history variation, in which mean lifespan increases with age at maturity as reproductive output at each breeding event declines. Analyses of plants and animals suggest that strategies of reproductive timing can vary on an independent axis, with iteroparous species at one extreme and semelparous species at the other. Insectivorous marsupials in the Family Dasyuridae have an unusually wide range of life-history strategies on both purported axes. We test and confirm that reproductive output and degree of iteroparity are independent in females across species. Variation in reproductive output per episode is associated with mean annual rainfall, which predicts food availability. Position on the iteroparity-semelparity axis is not associated with annual rainfall, but species in regions of unpredictable rainfall have longer maximum lifespans, more potential reproductive events per year, and longer breeding seasons. We suggest that these two axes of life-history variation arise because reproductive output is limited by overall food availability, and selection for high offspring survival favours concentrated breeding in seasonal environments. Longer lifespans are favoured when reproductive opportunities are dispersed over longer periods in environments with less predictable food schedules.