Intraspecific variation in sex allocation in hermaphroditic Plantago coronopus (L.)

Models for sex allocation assume that increased expenditure of resources on male function decreases the resources available for female function. Under some circumstances, a negative genetic correlation between investment in stamens and investment in ovules or seeds is expected. Moreover, if fitness returns for investment in male and female function are different with respect to size, sex allocation theory predicts size‐specific gender changes. We studied sex allocation and genetic variation for investment in stamens, ovules and seeds at both the flower and the plant level in a Dutch population of the wind‐pollinated and predominantly outcrossing Plantago coronopus. Data on biomass of floral structures, stamens, ovules, seedset and seedweight were used to calculate the average proportion of reproductive allocation invested in male function. Genetic variation and (genetic) correlations were estimated from the greenhouse‐grown progeny of maternal families, raised at two nutrient levels. The proportion of reproductive biomass invested in male function was high at flowering (0.86 at both nutrient levels) and much lower at fruiting (0.30 and 0.40 for the high and low nutrient treatment, respectively). Androecium and gynoecium mass exhibited moderately high levels of genetic variance, with broad‐sense heritabilities varying from 0.35 to 0.56. For seedweight no genetic variation was detected. Significant among‐family variation was also detected for the proportion of resources invested in male function at flowering, but not at fruiting. Phenotypic and broad‐sense genetic correlations between androecium and gynoecium mass were positive. Even after adjusting for plant size, as a measure of resource acquisition, maternal families that invested more biomass in the androecium also invested more in the gynoecium. This is consistent with the hypothesis that genetic variation for resource acquisition may in part be responsible for the overall lack of a negative correlation between male and female function. Larger plants had a more female‐biased allocation pattern, brought about by an increase in seedset and seedweight, whereas stamen biomass did not differ between small and large plants. These results are discussed in relation to size‐dependent sex allocation theory (SDS). Our results indicate that the studied population harboured substantial genetic variation for reproductive characters.     

Outcrossing hermaphroditic polychaete worms adjust their sex allocation to social conditions

Sex allocation theory predicts that simultaneous hermaphrodites shift sex allocation facultatively in response to variation in local group size. This study was performed to evaluate the relative investment in each sex function by the simultaneously hermaphroditic polychaete worm Ophryotrocha diadema and to test whether allocation to each sex depends on the number of reproductive competitors. Four experimental groups were set up (in a 2 x 2 factorial design) with small or large group size and with small or large enclosures to control for potential confounding effects of density. We measured the proportion of female and male investment in focal individuals. Results revealed that individuals regulated their reproductive output so that when reproductive competitors were present, the number of female gametes was strongly reduced and the male function increased. In contrast, under monogamy, individuals in small groups produced lower numbers of sperm but had a higher egg output than worms in large groups. Density did not affect sex allocation in our experiment. Our findings provide qualitative support for Local Mate Competition theory, but also show that the pattern of sex allocation specific to this species is more complex than expected by current theory.     

Contrasting energy allocation strategies of two sympatric Merluccius species in an upwelling system

This study investigated the somatic growth and energy allocation strategy of two sympatric hake species (Merluccius polli and Merluccius senegalensis), coexisting under the strong influence of the Mauritanian upwelling. The results revealed that ontogeny, bathymetry, geography and reproduction shaped the differences found between the condition dynamics of the two species. Aside from species‐specific differences, individuals were observed in better condition in the northernmost area (more influenced by the permanent upwelling) and in the deepest waters, probably the most favourable habitat for Merluccius spp. Both species also displayed contrasting trade‐offs in energy allocation probably due to the dissimilarity of their habitats, which favours the existence of divergent adaptive strategies in response to different ontogenic requirements. It was hypothesized that M. polli invests in mass and energy reserves while sacrificing growth, as larger sizes may not provide an ecological advantage in a deeper and more stable environment. Moreover, M. senegalensis capitalizes on a steady growth without major disruptions, enabling earlier spawning at the expense of a lower somatic mass, which is fitting to a less stable shallower environment. This study sheds new light on differences in the biological traits and life strategies of Merluccius spp., which permit their overlap in a complex upwelling system and may contribute to the long‐lasting scientific‐based management of these species.     

Reproduction of the Atlantic bobtail squid Sepiola atlantica (Cephalopoda: Sepiolidae) in northwest Spain

Atlantic bobtail squid (Sepiola atlantica) in northwest Spain show seasonal variation in population structure, with juvenile abundance peaking during summer and autumn. However, whether similar patterns exist for reproduction is unknown. Therefore, we describe the reproductive biology of 505 specimens of S. atlantica collected monthly during two consecutive years at two different sites off of Areamilla beach in the Ría de Vigo. Mature males displayed a type of sexual dimorphism previously unknown in members of this species, developing a muscular nodule in the base of each of the ventral arms over ontogeny. Reproductive output of both sexes was similar to that of other bobtail squids. Relative oocyte size (~10% mantle length) appeared to be similar to those of other bobtail squids. Females did not show evidence of having mated before complete maturity. Females of S. atlantica have group‐synchronous ovary maturation, with a positive correlation between female mantle length and ripe oocyte mass, suggesting a terminal investment strategy. Atlantic bobtail squids displayed the same seasonal patterns of reproductive traits at both sampling sites, with significant differences in reproductive activity between males and females. We consider reproductive traits in these small animals as adaptations to the coastal shelf lifestyle.     

Leaf lifespan is positively correlated with periods of leaf production and reproduction in 49 herb and shrub species

Leaf life span and plant phenology are central elements in strategies for plant carbon gain and nutrient conservation. Although few studies have found that leaf life span correlate with the patterns of leaf dynamics and reproductive output, but there have not been sufficient conclusive tests for relationships between leaf life span and plant phenological traits, the forms and strengths of such relationships are poorly understood. This study was conducted with 49 herb and shrub species collected from the eastern portion of the Tibetan Plateau and grown together in a common garden setting. We investigated leaf life span, the periods of leaf production and death, the time lag between leaf production and death, and the period of plant reproduction (i.e., flowering and fruiting). Interspecific relationships of leaf life span with leaf dynamics and reproduction period were determined. Leaf production period was far longer than leaf death period and largely reflected the interspecific variation of leaf life span. Moreover, leaf life span was positively correlated with the length of reproduction (i.e., flowering and fruiting) period. These relationships were generally consistent across different subgroups of species (herbs vs. shrubs) and indicate potentially widely applicable relationships between LLS and aboveground phenology. We concluded that leaf life span is associated not simply with the dynamics of the leaf itself but with reproduction period. The results demonstrate a plant trade‐off in resource allocation between production and reproduction and a coordinated arrangement of leaves, flowers, and fruits in their time investment. Our results provide insight into the relationship between leaf life span and plant phenology.     

Experimental evolution of ultraviolet radiation resistance in Escherichia coli

Ultraviolet (UV) light is a major cause of stress, mutation, and mortality in microorganisms, causing numerous forms of cellular damage. Nevertheless, there is tremendous variation within and among bacterial species in their sensitivity to UV light. We investigated direct and correlated responses to selection during exposure to UV. Replicate lines of Escherichia coli K12 were propagated for 600 generations, half with UV and half as a control without UV. All lines responded to selection, and we found strong positive and negative correlated responses to selection associated with increased UV resistance. Compared to Control populations, UV-selected populations increased in desiccation and starvation resistance approximately twofold but were 10 times more sensitive to hypersalinity. There was little evidence for a persistent large competitive fitness cost to UV resistance. These results suggest that natural variation in UV resistance may be maintained by trade-offs for resistance to other abiotic sources of mortality. We observed an average twofold increase in cell size by the UV-selected populations, consistent with a structural mode of adaptation to UV exposure having preadaptive and maladaptive consequences to other abiotic stresses.     

Size doesn't matter,sex does: a test for boldness in sister species of Brachyrhaphis fishes

The effect of divergent natural selection on the evolution of behavioral traits has long been a focus of behavioral ecologists. Predation, due to its ubiquity in nature and strength as a selective agent, has been considered an important environmental driver of behavior. Predation is often confounded with other environmental factors that could also play a role in behavioral evolution. For example, environments that contain predators are often more ecologically complex and “risky” (i.e., exposed and dangerous). Previous work shows that individuals from risky environments are often more bold, active, and explorative than those from low‐risk environments. To date, most comparative studies of environmentally driven behavioral divergence are limited to comparisons among populations within species that occur in divergent selective environments but neglect comparisons between species following speciation. This limits our understanding of how behavior evolves post‐speciation. The Central American live‐bearing fish genus Brachyrhaphis provides an ideal system for examining the relationship between selective environments and behavior, within and between species. Here, we test for differences in boldness between sister species B. roseni and B. terrabensis that occur in streams with and without piscivorous predators, respectively. We found that species do differ in boldness, with species that occur with predators being bolder than those that do not. Within each species, we found that sexes differed in boldness, with males being bolder than females. We also tested for a relationship between size (a surrogate for metabolic rate) and boldness, but found no size effects. Therefore, sex, not size, affects boldness. These results are consistent with the hypothesis that complex and risky environments favor individuals with more bold behavioral traits, but they are not consistent with the hypothesis that size (and therefore metabolic rate) drives divergence in boldness. Finally, our results provide evidence that behavioral trait divergence continues even after speciation is complete.     

Multi‐scale oviposition site selection in two mosquito species

1. In organisms characterised by complex life cycles, habitat selection often occurs at multiple spatial scales. For instance, female mosquitoes searching for an appropriate aquatic habitat to oviposit their eggs should also consider the characteristics of the terrestrial landscape in which it is embedded. 2. In this study, a field experiment was conducted to test for multi‐scale oviposition site selection in two mosquito species. Artificial pools were placed in two adjacent landscapes, olive plantations and a citrus orchard, mainly differing in their blooming periods and nectar availability. Pools were organised in three pairs: predatory caged fish were present in both pools, in one pool, or in none. 3. Early during the season, most of the egg rafts were laid by Culiseta longiareolata females in pools located within the blooming citrus orchard. When blooming shifted to the olive plantation, C. longiareolata become opportunistic. Culex pipiens females appeared later on during the season, when egg rafts of C. longiareolata were scarce, and they exhibited a higher selectivity to the olive plantation, although its blooming ended. In addition, the selectivity of C. pipiens to fish‐free pools was stronger than that of C. longiareolata. 4. Culex pipiens was more selective, possibly due to its high dispersal ability, which can lower movement cost and enhance the ability to gather environmental information. A trade‐off among gonotrophic cycles, combined with a shorter breeding season and limited recognition ability of the predatory fish may have reduced C. longiareolata selectivity. These differential oviposition patterns can strongly affect the population and community dynamics of both species.     

The importance of growth and mortality costs in the evolution of the optimal life history

A central assumption of life history theory is that the evolution of the component traits is determined in part by trade-offs between these traits. Whereas the existence of such trade-offs has been well demonstrated, the relative importance of these remains unclear. In this paper we use optimality theory to test the hypothesis that the trade-off between present and future fecundity induced by the costs of continued growth is a sufficient explanation for the optimal age at first reproduction, alpha, and the optimal allocation to reproduction, G, in 38 populations of perch and Arctic char. This hypothesis is rejected for both traits and we conclude that this trade-off, by itself, is an insufficient explanation for the observed values of alpha and G. Similarly, a fitness function that assumes a mortality cost to reproduction but no growth cost cannot account for the observed values of alpha. In contrast, under the assumption that fitness is maximized, the observed life histories can be accounted for by the joint action of trade-offs between growth and reproductive allocation and between mortality and reproductive allocation (Individual Juvenile Mortality model). Although the ability of the growth/mortality model to fit the data does not prove that this is the mechanism driving the evolution of the optimal age at first reproduction and allocation to reproduction, the fit does demonstrate that the hypothesis is consistent with the data and hence cannot at this time be rejected. We also examine two simpler versions of this model, one in which adult mortality is a constant proportion of juvenile mortality [Proportional Juvenile Mortality (PJM) model] and one in which the proportionality is constant within but not necessarily between species [Specific Juvenile Mortality (SSJM) model]. We find that the PJM model is unacceptable but that the SSJM model produces fits suggesting that, within the two species studied, juvenile mortality is proportional to adult mortality but the value differs between the two species.     

Development of sexual dimorphism in two sympatric skinks with different growth rates

Sexual size dimorphism (SSD) is widespread in animals, especially in lizards (Reptilia: Squamata), and is driven by fecundity selection, male–male competition, or other adaptive hypotheses. However, these selective pressures may vary through different life history periods; thus, it is essential to assess the relationship between growth and SSD. In this study, we tracked SSD dynamics between a “fading‐tail color skink” (blue tail skink whose tail is only blue during its juvenile stage: Plestiodon elegans) and a “nonfade color” tail skink (retains a blue tail throughout life: Plestiodon quadrilineatus) under a controlled experimental environment. We fitted growth curves of morphological traits (body mass, SVL, and TL) using three growth models (Logistic, Gompertz, and von Bertalanffy). We found that both skinks have male‐biased SSD as adults. Body mass has a higher goodness of fit (as represented by very high R² values) using the von Bertalanffy model than the other two models. In contrast, SVL and TL for both skinks had higher goodness of fit when using the Gompertz model. Two lizards displayed divergent life history tactics: P. elegans grows faster, matures earlier (at 65 weeks), and presents an allometric growth rate, whereas P. quadrilineatus grows slower, matures later (at 106 weeks), and presents an isometric growth rate. Our findings imply that species‐ and sex‐specific trade‐offs in the allocation of energy to growth and reproduction may cause the growth patterns to diverge, ultimately resulting in the dissimilar patterns of SSD.     

Trade-off acquisition and allocation in Gryllus firmus: a test of the Y model

Many models of life history evolution assume trade-offs between major life history traits; however, these trade-offs are often not found. The Y model predicts that variation in acquisition can mask underlying allocation trade-offs and is a major hypothesis explaining why negative relationships are not always found between traits that are predicted to trade-off with one another. Despite this model's influence on the field of life history evolution, it has rarely been properly tested. We use a model system, the wing dimorphic cricket, Gryllus firmus as a case study to test the assumptions and predictions of the Y model. By experimentally altering the acquisition regime and by estimating energy acquisition and energy allocation directly in this species, we are able to explicitly test this important model. Overall, we find strong support for the predictions of the Y model.     

Hormonal pleiotropy and the evolution of allocation trade‐offs

Recent empirical evidence suggests that trade‐off relationships can evolve, challenging the classical image of their high entrenchment. For energy reliant traits, this relationship should depend on the endocrine system that regulates resource allocation. Here, we model changes in this system by mutating the expression and conformation of its constitutive hormones and receptors. We show that the shape of trade‐offs can indeed evolve in this model through the combined action of genetic drift and selection, such that their evolutionarily expected curvature and length depend on context. In particular, the shape of a trade‐off should depend on the cost associated with resource storage, itself depending on the traded resource and on the ecological context. Despite this convergence at the phenotypic level, we show that a variety of physiological mechanisms may evolve in similar simulations, suggesting redundancy at the genetic level. This model should provide a useful framework to interpret and unify the overly complex observations of evolutionary endocrinology and evolutionary ecology.     

Elevation‐Related Differences in Parental Risk‐Taking Behavior are Associated with Cognitive Variation in Mountain Chickadees

Breeding animals must balance their current reproductive effort with potential costs to their own survival and consequently to future reproduction. Life‐history theory predicts that variation in reproductive investment should be based on fecundity and life expectancy with longer‐lived species favoring their own survival over parental investment. Recently, variation in parental risk taking was also linked with differences in cognition suggesting a potential trade‐off between cognitive ability and risk taking. Here, we tested whether mountain chickadees from two different elevations with known differences in cognitive ability differ in their parental risk taking by comparing the responses of nesting birds to a potential predator. Higher elevations are associated with shorter breeding season limiting renesting opportunities, but chickadees at high elevations also have better cognitive abilities, which might be potentially associated with better survival. Compared to lower elevations, high‐elevation chickadees laid larger clutches, showed longer latencies to return to the nest in the presence of a hawk, had lower fledging success, and exhibited higher rate of complete nest failures. Nestling development among successful nests, however, was similar between elevations. These data are not consistent with life‐history hypothesis because birds at high elevation invest more in the clutch, while at the same time take less risk when facing potential danger to themselves, which could jeopardize their current reproductive success. These data, however, are consistent with the hypothesis that better cognitive abilities might be associated with less parental risk taking.     

Partitioning of resources: the evolutionary genetics of sexual conflict over resource acquisition and allocation

Fitness depends on both the resources that individuals acquire and the allocation of those resources to traits that influence survival and reproduction. Optimal resource allocation differs between females and males as a consequence of their fundamentally different reproductive strategies. However, because most traits have a common genetic basis between the sexes, conflicting selection between the sexes over resource allocation can constrain the evolution of optimal allocation within each sex, and generate trade‐offs for fitness between them (i.e. ‘sexual antagonism’ or ‘intralocus sexual conflict’). The theory of resource acquisition and allocation provides an influential framework for linking genetic variation in acquisition and allocation to empirical evidence of trade‐offs between distinct life‐history traits. However, these models have not considered the emergence of trade‐offs within the context of sexual dimorphism, where they are expected to be particularly common. Here, we extend acquisition–allocation theory and develop a quantitative genetic framework for predicting genetically based trade‐offs between life‐history traits within sexes and between female and male fitness. Our models demonstrate that empirically measurable evidence of sexually antagonistic fitness variation should depend upon three interacting factors that may vary between populations: (1) the genetic variances and between‐sex covariances for resource acquisition and allocation traits, (2) condition‐dependent expression of resource allocation traits and (3) sex differences in selection on the allocation of resource to different fitness components.     

Mechanisms of a locally adaptive shift in allocation among growth,reproduction, and herbivore resistance in Mimulus guttatus*

Environmental gradients can drive adaptive evolutionary shifts in plant resource allocation among growth, reproduction, and herbivore resistance. However, few studies have attempted to connect these adaptations to underlying physiological and genetic mechanisms. Here, we evaluate potential mechanisms responsible for a coordinated locally adaptive shift between growth, reproduction, and herbivore defense in the yellow monkeyflower, Mimulus guttatus. Through manipulative laboratory experiments, we found that gibberellin (GA) growth hormones may play a role in the developmental divergence between perennial and annual ecotypes of M. guttatus. Further, we detected an interaction between a locally adaptive chromosomal inversion, DIV1, and GA addition. This finding is consistent with the inversion contributing to the evolutionary divergence between inland annual and coastal perennial ecotypes by reducing GA biosynthesis/activity in perennials. Finally, we found evidence that the DIV1 inversion is partially responsible for a coordinated shift in the divergence of growth, reproduction, and herbivore resistance traits between coastal perennial and inland annual M. guttatus. The inversion has already been established to have a substantial impact on the life‐history shift between long‐term growth and rapid reproduction. Here, we demonstrate that the DIV1 inversion also has sizable impacts on both the total abundance and composition of phytochemical compounds involved in herbivore resistance.     

Migratory costs and contemporary evolution of reproductive allocation in male chinook salmon

Energetically demanding migrations may impact the resources available for reproductive trait development and activity, and hence favour evolution of new investment strategies for remaining resources. We conducted a large-scale experiment to evaluate the proximate cost of migration on male reproductive investment in chinook salmon (Oncorhynchus tshawytscha) and contemporary evolution of reproductive allocation. Experimentally induced differences in migratory costs (17 km inland and 17 m elevation vs. 100 km and 430 m) influenced dorsal hump size and upper jaw length, two traits influencing male mating success that are developed during migration. Longer migration also reduced tissue energy reserves available for competition and length of breeding life. Corresponding shifts in the balance between natural and sexual selection appear to have been responsible for heritable population divergence in secondary sexual trait investment, in approximately 26 generations, following colonization of spawning sites with different migratory demands.     

Plasticity in reproduction and growth among 52 range‐wide populations of a Mediterranean conifer: adaptive responses to environmental stress

A plastic response towards enhanced reproduction is expected in stressful environments, but it is assumed to trade off against vegetative growth and efficiency in the use of available resources deployed in reproduction [reproductive efficiency (RE)]. Evidence supporting this expectation is scarce for plants, particularly for long‐lived species. Forest trees such as Mediterranean pines provide ideal models to study the adaptive value of allocation to reproduction vs. vegetative growth given their among‐population differentiation for adaptive traits and their remarkable capacity to cope with dry and low‐fertility environments. We studied 52 range‐wide Pinus halepensis populations planted into two environmentally contrasting sites during their initial reproductive stage. We investigated the effect of site, population and their interaction on vegetative growth, threshold size for female reproduction, reproductive–vegetative size relationships and RE. We quantified correlations among traits and environmental variables to identify allocation trade‐offs and ecotypic trends. Genetic variation for plasticity was high for vegetative growth, whereas it was nonsignificant for reproduction. Size‐corrected reproduction was enhanced in the more stressful site supporting the expectation for adverse conditions to elicit plastic responses in reproductive allometry. However, RE was unrelated with early reproductive investment. Our results followed theoretical predictions and support that phenotypic plasticity for reproduction is adaptive under stressful environments. Considering expectations of increased drought in the Mediterranean, we hypothesize that phenotypic plasticity together with natural selection on reproductive traits will play a relevant role in the future adaptation of forest tree species.     

Experimental tests of nest site competition in two Peromyscus species

The importance of interspecific competition for nest sites between the white-footed mouse (Peromyscus leucopus noveboracensis) and the cloudland deermouse (P. maniculatus nubiterrae) were investigated in the montane forests of southwestern Virginia over 3 years. Trials were conducted for both species using large, outdoor enclosures in order to examine: (i) nest site preference in isolation and (ii) nest site selection made in the presence of potential competitors. Both species demonstrated a strong preference for arboreal nest sites when tested without competitors present. After the introduction of a heterospecific intruder, P. leucopus often shifted to a non-arboreal nest while P. maniculatus continued to use arboreal nests. Intruding P. maniculatus displaced resident P. leucopus from P. leucopus' preferred nest sites in all 3 years while intruding P. leucopus never displaced P. maniculatus. Neither species was routinely displaced in conspecific trials. Resident P. maniculatus also excluded P. leucopus from access to preferred nesting sites in all three years while P. leucopus were only able to exclude P. maniculatus in the 3rd year. Both species exhibite relatively low frequencies of exclusion in conspecific trials with the exception of P. maniculatus which excluded high proportions of conspecific intruders in the second year. We suggest that the asymmetrical advantage enjoyed by P. maniculatus in nest site selection may result from both site-specific effects and a species-specific influence on P. leucopus. Nonetheless, the intensity of such influence varied between years, perhaps as a function of population density, and did not appear to drastically influence continued coexistence of these congeners.     

The importance of listening: juvenile allocation shifts in response to acoustic cues of the social environment

The social environment has a strong effect on the strength and direction of sexual selection. Juveniles, however, often have social cues that signal the current competitive environment which may provide cues of future competitive challenges. Here we demonstrate that juvenile crickets (Teleogryllus commodus) use the calls of surrounding adult males as a cue of the quality and density of rivals/mates they are likely to encounter. We reared hatchling crickets in six acoustic environments that varied in the density and quality of calls and demonstrate that individuals modified their development rate, phenotype and behaviour at maturity. Males matured more rapidly at a smaller size and called more when reared in a low competition environment. In contrast, males delayed maturity to grow larger when faced with an increased density of high-quality males. Females matured more rapidly when reared in a high density of high-quality males and allocated proportionately more resources towards egg production. A second experiment limiting nutrient availability demonstrates sex-specific allocation shifts in the last stadium when cues are most reliable. Our results demonstrate that the social environment significantly affects allocation strategies and phenotypes, highlighting the importance of juvenile experience and competitive context when examining fitness and selection.     

Experimental test of a trade‐off between moult and immune response in house sparrows Passer domesticus

A trade‐off between immune system and moulting is predicted in birds, given that both functions compete for resources. However, it is unclear whether such a trade‐off exists during post‐breeding moult. This study tests such a trade‐off in the house sparrow (Passer domesticus). Males injected with an antigen (lipopolysaccharide) significantly moulted slower than sham‐injected males. Moreover, males whose seventh primaries were plucked to simulate moult showed smaller immune response to phytohaemagglutinin than control males, in which seventh primaries were clipped. A trade‐off between moult speed and body mass was also found. The results show a clear trade‐off between moult and immune response in the house sparrow: immune response negatively affected moult and moult negatively affected immune response. These findings suggest that only individuals in good condition may have an efficient moult and simultaneously respond effectively in terms of immunity to pathogens, which could explain how plumage traits honestly indicate parasite resistance in birds.