Physiological response to etho-ecological stressors in male Alpine chamois: timescale matters!

From a life history perspective, glucocorticoids secreted by the neuroendocrine system, integrating different sources of stress through an adaptive feedback mechanism, may have important consequences on individual fitness. Although stress responses have been the object of several investigations, few studies have explored the role of proximate mechanisms responsible for the potential trade-offs between physiological stress and life history traits integrating social and environmental stressors. In 2011 and 2012, we collected data on faecal cortisol metabolites (FCM) in a marked male population of Alpine chamois, within the Gran Paradiso National Park (Italy). Using a model selection approach we analysed the effect of potential etho-ecological stressors such as age, social status (territorial vs. non-territorial males), minimum temperature, snow depth and precipitation on FCM variation. To correctly interpret environmentally and socially induced stress responses, we conducted model selections over multiple temporal scales defined a priori: year, cold months, spring, warm months, mating season. Over the year, FCM levels showed a negative relationship with minimum temperature, but altogether, climatic stressors had negligible effects on glucocorticoid secretion, possibly owing to good adaptations of chamois to severe weather conditions. Age was negatively related to FCM during the rut, possibly due to greater experience of older males in agonistic contests. Social status was an important determinant of FCM excretion: while both the ‘stress of subordination’ and the ‘stress of domination’ hypotheses received some support in spring and during the mating season, respectively, previous data suggest that only the latter may have detrimental fitness consequences on male chamois.     

An equal sex ratio followed by differential sex mortality causes overestimation of females in gall midges: no evidence for sex ratio regulation

Monogeny, the production of unisexual broods by individual females, has been recognized for nearly 80?years. The genetic nature of gall midges' sex determination predicts an equal numbers of male-producing and female-producing females in the populations such that the overall sex ratio is expected to be nearly 1:1. However, observations of some strictly monogenous populations with biased sex ratio, mainly toward females, have raised the question of whether gall midges are able to adjust their offspring sex ratio in response to changes in environmental conditions, and some authors have even considered sex ratio regulation as a strong force in the course of the evolution of monogeny. In this paper, first, by studying the sex ratio variations of the predatory gall midge, Aphidoletes aphidimyza within a generation, we showed that adult males emerge up to 1?day earlier and have shorter life span than females (less than 4?days and up to 6?days, respectively). Although, the sex ratio of A. aphidimyza at the time of emergence was nearly 1:1 (52.41?% males), a simple population simulation indicated that the differential mortality of sexes can lead to a female-biased sex ratio estimation (57.88?% females) under random sampling in the natural environments. Our results imply that the primary sex ratio of monogenous gall midges is nearly 1:1 and that the arrhenogenic/thelygenic gall midges are not able to alter the number of their male/female progenies in response to changes in environmental conditions.     

Does testis weight decline towards the Subarctic? A case study on the common frog, Rana temporaria

Interpopulation comparisons of variation in resource availability and in allocation patterns along altitudinal and latitudinal gradients allow insights into the mechanisms shaping the life history of animals. Patterns of between-population differences in female life history traits have been studied intensively across a wide range of taxa, but similar investigations in males have remained scarce. To study if testis weight—a measure of reproductive investment—varies on a geographical scale in anurans, we focussed on the variation in relative testis weight (RelTW) and asymmetry in 22 populations of the common frog Rana temporaria along a 1,600-km latitudinal transect across the Scandinavian peninsula. We found that RelTW decreased towards the north. Body mass and body length both had independent positive effects on testes mass. We found evidence for directional asymmetry (DA) in testis weight with the right testis being larger than the left. The level of DA in testis weight was not related to latitude, but both body mass and testes mass had independent positive effects on asymmetry. We discuss the northwards decrease in RelTW in terms of a decreased reproductive investment as a possible consequence of harsher environmental conditions, and perhaps also, weaker sexual selection in the north than in the south.     

Disentangling the effects of local and regional factors on the thermal tolerance of freshwater crustaceans

In the global warming context, we compared the thermal tolerance of several populations of the crustacean Gammarus pulex (Amphipoda: Gammaridae) along a latitudinal thermal gradient in the Rh?ne Valley. To disentangle the effect of regional (North vs. South) and local (site-specific) factors, the ecophysiological responses of populations were investigated at two levels of biological organisation: whole organism level considering body size [critical thermal maximum (CTmax), mean speed of locomotion (MS), time mobile (TM)] and organelle function level [mitochondrial respiratory control ratios (RCRs)]. CTmax and RCRs, but not MS and TM, revealed a significantly higher thermal tolerance in southern populations compared to northern ones. Nevertheless, temperatures ≥ 30°C were deleterious for all populations, suggesting that populations located in the warmer limit of the species distribution will be more threatened by climate change as they live closer to their upper thermal limits. The strong differences observed between populations indicate that the species-level thermal tolerance used in predictive models may not be informative enough to study the impact of global warming on species distributions. This work also reveals that an appropriate choice of indicators is essential to study the consequences of global warming since the response of organisms at the whole body level can be influenced by local conditions.     

Carotenoid coloration is related to fat digestion efficiency in a wild bird

Some of the most spectacular visual signals found in the animal kingdom are based on dietarily derived carotenoid pigments (which cannot be produced de novo), with a general assumption that carotenoids are limited resources for wild organisms, causing trade-offs in allocation of carotenoids to different physiological functions and ornamentation. This resource trade-off view has been recently questioned, since the efficiency of carotenoid processing may relax the trade-off between allocation toward condition or ornamentation. This hypothesis has so far received little exploratory support, since studies of digestive efficiency of wild animals are limited due to methodological difficulties. Recently, a method for quantifying the percentage of fat in fecal samples to measure digestive efficiency has been developed in birds. Here, we use this method to test if the intensity of the carotenoid-based coloration predicts digestive efficiency in a wild bird, the house finch (Haemorhous mexicanus). The redness of carotenoid feather coloration (hue) positively predicted digestion efficiency, with redder birds being more efficient at absorbing fats from seeds. We show for the first time in a wild species that digestive efficiency predicts ornamental coloration. Though not conclusive due to the correlative nature of our study, these results strongly suggest that fat extraction might be a crucial but overlooked process behind many ornamental traits.     

Plant invasion phenomenon enhances reproduction performance in an endangered spider

Current models in evolutionary ecology predict life history alterations in response to habitat suitability to optimize fitness. Only few empirical studies have demonstrated how life history traits that are expected to trade off against each other differ among environments. In Europe, many salt marshes have been recently invaded by the grass Elymus athericus. Previous studies however showed higher densities of the endangered spider Arctosa fulvolineata (Araneae: Lycosidae) in invaded salt marshes compared to natural habitats, which suggests a lower habitat suitability in the latter. The aim of this study was to determine if this emerging habitat (1) affects the amount of resource acquisition and (2) alters the balance between life history traits that are expected to trade off against each other in this stenotopic salt marsh species. As suggested by theoretical studies, an optimization of fitness by increasing egg size at the cost of decreasing fecundity in unsuitable (i.e., natural) habitats was expected. Females presenting cocoon were then collected in close invaded and natural salt marsh areas within the Mont Saint-Michel Bay (France). By considering female mass as covariate, cocoon mass, number of eggs, and egg volume were compared between both habitats. Clutch mass was strongly determined by female mass in both habitats. Clutch mass was however significantly smaller in the natural habitat compared to the invaded habitat, indicating a higher resource acquisition in the latter. When correcting for female size, fecundity was additionally increased in the invaded habitat through a significant decrease in egg size. This phenotypic response can be explained by differences in habitat structure between invaded and natural habitats: the former offers a more complex litter favoring nocturnal wanderers like A. fulvolineata. The existence of such an adaptive reproduction strategy depending on habitat suitability constitutes an original case of an invasion that favors an endangered species.     

Using many-objective trade-off analysis to help dams promote economic development,protect the poor and enhance ecological health

Allocating water to different uses implies trading off the benefits perceived by different sectors. This paper demonstrates how visualising the trade-offs implied by the best performing water management options helps balance water use benefits and find sustainable solutions. The approach consists of linking a water resources model that can simulate many management policies and track diverse measures of system performance, to a many-objective evolutionary optimisation algorithm. This generates the set of Pareto-optimal management alternatives for several simultaneous objectives. The relative performance of these efficient management alternatives is then visualised as trade-off curves or surfaces using visual analytic plots. Visually assessing trade-offs between benefits helps select policies that achieve a decision-maker-selected balance between different metrics of system performance. We apply this approach to a multi-reservoir water resource system in Brazil's semi-arid Jaguaribe basin where current water allocation procedures favour sectors with greater political power and technical knowledge. The case study identifies promising reservoir operating policies by exploring trade-offs between economic, ecological and livelihood benefits as well as traditional hydropower generation, irrigation and water supply. Results show optimised policies can increase allocations to downstream uses while increasing median land availability for the poorest farmers by 25%.     

Tail loss compromises immunity in the many-lined skink, Eutropis multifasciata

Tail autotomy incurs energetic costs, and thus, a trade-off in resource allocation may lead to compromised immunity in lizards. We tested the hypothesis that tailless lizards will favor constitutive innate immunity responses over an energetically costly inflammatory response. The influence of fasting and colorful ornamentation was also investigated. We experimentally induced tail autotomy in the lizard Eutropis multifasciata and found that inflammation was suppressed by tail loss, but not further affected by fasting; the suppressive effect of colorful ornamentation was manifested only in males, but not in females. Constitutive innate immunity was not affected by any of these factors. As expected, only costly inflammation was compromised, and a less expensive constitutive innate immunity might be favored as a competent first-line defense during energetically demanding periods. After considering conventional trade-offs among tail regeneration and reproduction, further extending these studies to incorporate disease risk and how this influences escape responses to predators and future reproduction would make worthwhile studies.     

Intrinsic worker mortality depends on behavioral caste and the queens’ presence in a social insect

According to the classic life history theory, selection for longevity depends on age-dependant extrinsic mortality and fecundity. In social insects, the common life history trade-off between fecundity and longevity appears to be reversed, as the most fecund individual, the queen, often exceeds workers in lifespan several fold. But does fecundity directly affect intrinsic mortality also in social insect workers? And what is the effect of task on worker mortality? Here, we studied how social environment and behavioral caste affect intrinsic mortality of ant workers. We compared worker survival between queenless and queenright Temnothorax longispinosus nests and demonstrate that workers survive longer under the queens’ absence. Temnothorax ant workers fight over reproduction when the queen is absent and dominant workers lay eggs. Worker fertility might therefore increase lifespan, possibly due to a positive physiological link between fecundity and longevity, or better care for fertile workers. In social insects, division of labor among workers is age-dependant with young workers caring for the brood and old ones going out to forage. We therefore expected nurses to survive longer than foragers, which is what we found. Surprisingly, inactive inside workers showed a lower survival than nurses but comparable to that of foragers. The reduced longevity of inactive workers could be due to them being older than the nurses, or due to a positive effect of activity on lifespan. Overall, our study points to behavioral caste-dependent intrinsic mortality rates and a positive association between fertility and longevity not only in queens but also in ant workers.     

Life history evolution: successes, limitations, and prospects

Life history theory tries to explain how evolution designs organisms to achieve reproductive success. The design is a solution to an ecological problem posed by the environment and subject to constraints intrinsic to the organism. Work on life histories has expanded the role of phenotypes in evolutionary theory, extending the range of predictions from genetic patterns to whole-organism traits directly connected to fitness. Among the questions answered are the following: Why are organisms small or large? Why do they mature early or late? Why do they have few or many offspring? Why do they have a short or a long life? Why must they grow old and die? The classical approach to life histories was optimization; it has had some convincing empirical success. Recently non-equilibrium approaches involving frequency-dependence, density-dependence, evolutionary game theory, adaptive dynamics, and explicit population dynamics have supplanted optimization as the preferred approach. They have not yet had as much empirical success, but there are logical reasons to prefer them, and they may soon extend the impact of life history theory into population dynamics and interspecific interactions in coevolving communities.     

Limited scope for reproductive senescence in wild populations of a short-lived fish

Senescence in wild populations was long considered negligible but current evidence suggests that it is widespread in natural populations of mammals and birds, affecting the survival and reproductive output of older individuals. In contrast, little is known about reproductive senescence in species with asymptotic growth that can keep increasing their reproductive output as they grow older and larger. Using a cross-sectional study, we tested age-related decline in fecundity and relative allocation to reproduction in five wild populations of an annual killifish, Nothobranchius furzeri (Cyprinodontiformes). We did not detect any decline in absolute female egg production over their short lifespan in the wild. Relative fecundity (egg production controlled for female body mass) tended to decrease with age. This effect was driven primarily by a single population that survived 17 weeks, almost twice as long as the median persistence of the other four study populations. There was no decrease in relative ovary mass while in males, relative testes mass actually increased with age. Intra-population variation in relative ovary mass increased in older females suggesting heterogeneity in individual trajectories of female reproductive allocation. Overall, we demonstrate that annual killifish do not experience significant age-related decline in reproductive functions during their very short lifespan in the wild despite the marked deterioration of gonad tissue detected in captivity.     

Climatic influences on the breeding biology of the agile frog (<Emphasis Type="Italic">Rana dalmatina</Emphasis>)

Severe population declines of amphibians have been shown to be attributed to climate change. Nevertheless, the various mechanisms through which climate can influence population dynamics of amphibians remain to be assessed, notably to disentangle the relative synergetic or antagonistic influences of temperature and precipitations on specific life history stages. We investigated the impact of rainfall and temperature on the egg-clutch abundance in a population of agile frog (Rana dalmatina) during 29 years (1987–2016) on 14 breeding sites located in Brenne Natural Park, France. Specifically, we examined the influence of environmental conditions occurring during five temporal windows of the year cycle corresponding to specific life history stages. Overall, our results suggest that the year-to-year fluctuations of egg-clutch abundances in Brenne Natural Park were partly dependent on local climatic conditions (rainfall and temperature). Climate seemed to influence breeding frogs during the autumn-winter period preceding reproduction. Spring and summer conditions did not influence reproduction. Additionally, we failed to detect effects of climatic conditions on newly metamorphosed individuals. Other factors such as density dependence and inter-specific interactions with introduced predators are likely to play a significant role in reproduction dynamics of the studied frog populations.     

Predicting fish population dynamics: compensation and the importance of site-specific considerations

The magnitude of the compensatory response inherent to fish population seems to be questioned with each new generation of scientists and regulators. The major reason for this is the demand by some that compensation be completely understood at the process level before compensation can be factored into population predictions. We first discuss several lines of evidence for the existence of compensation, and the reasons why compensation is so difficult to measure at the process level. We then offer life history theory as a general framework for improving our understanding of the likely magnitude of compensation in fish population dynamics. We use detailed, individual-based models and field data for yellow perch-walleye, striped bass, and a lake community to illustrate the complexity of compensation, and to show that predictions for a particular population require site-specific information. Analyses show that the life stage and magnitude of density-dependence can vary for different populations of the same species, and that population responses can depend on the arrangement of the food web. We must stop reinventing the density-dependence wheel; too many resources (both natural and monetary) are at stake. We suggest using three approaches for addressing the compensation issue: (1) life history theory for providing a sound basis for qualitatively understanding fish population dynamics and the magnitude of compensation; (2) age- and stage-structured matrix models for screening-level analyses to identify populations that may be at unacceptable risks of decline and therefore require additional analyses; and (3) individual-based models for better understanding the complexities of compensation at the process level and for impact assessment of at-risk, well-studied populations.     

Effects of environmental parameters on the chestnut gall wasp and its complex of indigenous parasitoids

The chestnut gall wasp (CGW), Dryocosmus kuriphilus, an invasive pest native to China, has caused severe yield and economic losses to chestnut production in Europe since its arrival in 2002. In Southern Italy, the complex of indigenous parasitoids colonizing CGW was monitored between 2013 and 2015, with the aim of estimating the composition of the indigenous parasitoid complex, its ability to control CGW populations, and the interactions of both factors with several measured environmental parameters. We compared results among three differently managed field types. Results showed an increase in the rate of parasitism both when the host population density was lower and in unmanaged chestnut stands with more natural conditions. The percentage of parasitism in galls was related to morphological traits of the galls and to higher seasonal temperatures, which reduced the parasitism intensity because CGW develops earlier under such conditions. The host–parasitoid mortality inside galls varied among sites and was associated mostly with rot fungi during wet spring and summer months. Parasitoid species richness was similar among the study sites, but the proportion of parasitoid species differed between orchards and unmanaged coppice stands. The timing of attack by parasitoids followed a species-specific successional sequence throughout the larva-to-adult life cycle of the CGW. These interactions should be considered in future research on trophic relationships and when modeling invasive scenarios for new pest species.     

Individual consistency in exploratory behaviour and mating tactics in male guppies

While behavioural plasticity is considered an adaptation to fluctuating social and environmental conditions, many animals also display a high level of individual consistency in their behaviour over time or across contexts (generally termed ‘personality’). However, studies of animal personalities that include sexual behaviour, or functionally distinct but correlated traits, are relatively scarce. In this study, we tested for individual behavioural consistency in courtship and exploratory behaviour in male guppies (Poecilia reticulata) in two light environments (high vs. low light intensity). Based on previous work on guppies, we predicted that males would modify their behaviour from sneak mating tactics to courtship displays under low light conditions, but also that the rank orders of courtship effort would remain unchanged (i.e. highly sexually active individuals would display relatively high levels of courtship under both light regimes). We also tested for correlations between courtship and exploratory behaviour, predicting that males that had high display rates would also be more likely to approach a novel object. Although males showed significant consistency in their exploratory and mating behaviour over time (1 week), we found no evidence that these traits constituted a behavioural syndrome. Furthermore, in contrast to previous work, we found no overall effect of the light environment on any of the behaviours measured, although males responded to the treatment on an individual-level basis, as reflected by a significant individual-by-environment interaction. The future challenge is to investigate how individual consistency across different environmental contexts relates to male reproductive success.     

Temperament in bullheads: do laboratory and field explorative behaviour variables correlate?

The relevance of temperament traits for life history strategy or productivity is increasingly acknowledged. Temperament traits are often either observed in captivity or in the wild, but studies combining both observations are very rare. We examine whether exploratory behaviour in the bullhead (Cottus perifretum), assayed under laboratory conditions, predicts this behaviour under field conditions. Forty-three PIT-tagged individuals were first assayed for exploration of a novel environment in the aquarium and then released into an unfamiliar stream stretch, where they were later relocated using a mobile antenna. Explorative behaviour assayed in the laboratory was significantly positively related to the exploration in the field, thus predicting distance moved in the field release. Both in the laboratory and in the field, explorative behaviour was not related to individual body length. When bullheads that did not leave the refuge in the aquarium (laboratory assay) and, therefore, did not explore the new environment were excluded from the analysis, the correlation between laboratory and field explorative behaviour variables became weaker. However, overall, our results illustrate that exploration rate of bullheads in isolated single-individual experiments can be used to predict this behaviour in the natural ecosystem.     

Carotenoid maintenance handicap and the physiology of carotenoid-based signalisation of health

Despite a reasonable scientific interest in sexual selection, the general principles of health signalisation via ornamental traits remain still unresolved in many aspects. This is also true for the mechanism preserving honesty of carotenoid-based signals. Although it is widely accepted that this type of ornamentation reflects an allocation trade-off between the physiological utilisation of carotenoids (mainly in antioxidative processes) and their deposition in ornaments, some recent evidence suggests more complex interactions. Here, we further develop the models currently proposed to explain the honesty of carotenoid-based signalisation of heath status by adding the handicap principle concept regulated by testosterone. We propose that under certain circumstances carotenoids may be dangerous for the organism because they easily transform into toxic cleavage products. When reserves of other protective antioxidants are insufficient, physiological trade-offs may exist between maintenance of carotenoids for ornament expression and their removal from the body. Furthermore, we suggest that testosterone which enhances ornamentation by increasing carotenoid bioavailability may also promote oxidative stress and hence lower antioxidant reserves. The presence of high levels of carotenoids required for high-quality ornament expression may therefore represent a handicap and only individuals in prime health could afford to produce elaborate colourful ornaments. Although further testing is needed, this ‘carotenoid maintenance handicap’ hypothesis may offer a new insight into the physiological aspects of the relationship between carotenoid function, immunity and ornamentation.     

Intra-specific variations of two Leymus chinensis divergence populations in Songnen Plain, Northeast China

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