Function of weaponry in females: the use of horns in intrasexual competition for resources in female Soay sheep

In many species, females show reduced expression of a trait that is under sexual selection in males, and this expression is thought to be maintained through genetic associations with the male phenotype. However, there is also the potential for the female trait to convey an advantage in intrasexual conflicts over resources. We tested this hypothesis in a feral population of Soay sheep, in which males and females have a polymorphism for horn development, producing either full (normal horned), reduced (scurred) or no (polled, females only) horns. During the lambing period, females who possessed horns were more likely to initiate and win aggressive interactions, independent of age, weight and birthing status. The occurrence of aggression was also context dependent, decreasing over the lambing period and associated with local density. Our results demonstrate that a trait that confers benefits to males during intrasexual competition for mates may also be used by females in intrasexual competition over resources: males use weaponry to gain mates, whereas females use weaponry to gain food

The effects of competition on fitness depend on the sex of both competitors

In intraspecific competition, the sex of competing individuals is likely to be important in determining the outcome of competitive interactions and the way exposure to conspecifics during development influences adult fitness traits. Previous studies have explored differences between males and females in their response to intraspecific competition. However, few have tested how the sex of the competitors, or any interactions between focal and competitor sex, influences the nature and intensity of competition. We set up larval seed beetles Callosobruchus maculatus to develop either alone or in the presence of a male or female competitor and measured a suite of traits: development time, emergence weight; male ejaculate mass, copulation duration, and lifespan; and female lifetime fecundity, offspring egg-adult survival, and lifespan. We found effects of competition and competitor sex on the development time and emergence weight of both males and females, and also of an interaction between focal and competitor sex: Females emerged lighter when competing with another female, while males did not. There was little effect of larval competition on male and female adult fitness traits, with the exception of the effect of a female competitor on a focal female's offspring survival rate. Our results highlight the importance of directly measuring the effects of competition on fitness traits, rather than distant proxies for fitness, and suggest that competition with the sex with the greater resource requirements (here females) might play a role in driving trait evolution. We also found that male-male competition during development resulted in shorter copulation times than male-female competition, a result that remained when controlling for the weight of competitors. Although it is difficult to definitively tease apart the effects of social environment and access to resources, this result suggests that something about the sex of competitors other than their size is driving this pattern.Funding for this project was provided by the Australian Research Council (FT160100149 to MLH and FT110100453 to LEBK

Effects of warming temperatures on germination responses and trade-offs between seed traits in an alpine plant

1. Climate warming may affect multiple aspects of plant life history, including important factors such as germination responses and the key trade-off between offspring size and number. As a case study to address these concepts, we used an alpine plant (waxy bluebell, Wahlenbergia ceracea; Campanulaceae) that shows plasticity to warming in seed traits and in which seed dormancy status regulates germination. We chose an alpine species because alpine environments are ecosystems particularly under threat by climate change. 2. We conducted germination assays under cool and warm temperatures using seeds produced by individuals that were grown under historical (cooler) and future (warmer) temperature scenarios. We assessed the presence of a seed size vs number trade-off, and then examined the effects of seed number and size on germination percentage, the fractions of dormant and viable seeds, and germination velocity. Further, we examined whether warming during parental growth and during germination affected these relationships. 3. We found evidence for a seed size vs number trade-off only under historical parental temperatures. Indeed, under future growth temperatures, parental plants produced fewer and smaller seeds and there was no evidence of a trade-off. However, the reductions in both seed traits under warming did not affect germination, despite correlations of seed size and number with germination traits. Warming increased germination, particularly of larger seeds, but overall it resulted in more than fourfold reductions in parental fitness. 4. Synthesis. Our study shows the importance of growth conditions when evaluating the seed size vs number trade-off. Stressful conditions, such as warmer temperatures, can restrain the ability of plants to reach optimal investment in reproduction, masking the trade-off. By analysing responses across the whole life cycle, we show here an overall detrimental effect of warming, highlighting the potential risk of climate change for W. ceracea, and, potentially, for alpine plant communities more widely.Files can be opened using Excel and analysed using R.Funding provided by: Australian Research CouncilCrossref Funder Registry ID: http://dx.doi.org/10.13039/501100000923Award Number: DP170101681Experiments were conducted using the plant species Wahlnebrgia ceracea (waxy bluebells). Two datasets were used in this manuscript. 1) Seed size vs number trade-off: Parental individuals from a total of 30 lines ('Line') were grown in growth chambers for 191 days under temperature conditions of a historical/cooler (1960–1970) or a projected future/warmer (2090–2100) climate ('Parental_Temperature'). The parental individuals were randomly assigned to one of three blocks, which corresponded to positions inside the chambers, and each block was equivalent in all chambers ('Block'). Day and night temperatures during the experiment were changed every 15 days to mimic seasonality, with the maximum day temperatures during the peak of summer being 24°C and 29°C for the historical and future parental temperatures, respectively. After 100 days since the imposition of the temperature treatments (during the peak of the summer), half of the plants were moved for 5 days to new chambers, where the temperature was 5°C above the respective treatments, i.e., 29°C and 34°C ('Heatwave'). After this time, the parental individuals were moved back to their respective historical or future temperature treatments. We collected the seeds throughout the 191 days of parental growth, and we stored them in desiccators for at least 11 weeks. After this time, we calculated seed size ('Seed_Size') as the average mass of three lots of 50 seeds divided by 50. We calculated seed number ('Seed_Number') as the ratio between the cumulative mass of the seeds produced by each parental individual and seed size. The 30 lines of the parental individuals were obtained by crossing plants that originated from seeds that were collected at the same elevation, either high or low elevation ('Elevation') in sites within Kosciuszko National Park, NSW, Australia. Therefore, 14 lines originated from high elevations and 14 lines from low elevations. 2) Germination responses - seed traits correlations: The seeds were harvested from the parental individuals grown under historical/cooler or projected future/warmer temperatures ('Parental_Temperature') (see above) from a subset of 14 lines ('Line'). These seeds were used in germination assays in the glasshouse under cool (25°C) or warm temperatures (30°C) ('Germination_Temperature'). We measured seed size ('Seed_Size') as the average mass of three lots of 50 seeds; then these seeds were sowed in agar dishes (25 seeds per dish, 2 dishes per temperature treatment from each parental individual). Seed number ('Seed_Number') was the same as above. Dishes were left under temperature treatments for 4 weeks to allow germination of the non-dormant fraction of the seeds ('Not_Dormant_Seeds') and germination was checked once per week. Then, all the dishes were moved to a cold room at 4–5°C in the dark for 4 weeks to allow cold stratification. After this time, dishes were moved back to the glasshouse under the same temperature treatments as before to allow germination of the dormant seeds. We considered seeds to be dormant ('Dormant_seeds') if they germinated during or after cold stratification or if they did not germinate at all but were still determined to be viable at the end of the experiment. We considered seed to be viable ('Viable_Seeds') if they germinated ('Germinated_Seeds') as well as the seeds that contained an endosperm but still did not germinate ('Not_Germinated_Seeds'), while we considered empty seeds as non-viable ('Not_Viable_Seeds'). Germinated and not germinated seeds (as above) were used to calculate the germination percentage. We calculated germination velocity ('Germination_Velocity') as the reciprocal of the mean germination time (germination velocity (germination (%) week-1) GV = (G1 + G2 +…+ Gn) / (G1 x T1 + G2 x T2 +…+ Gn x Tn), where Gn is the number of new germinating seeds at each sampling point, and Tn is the time between each sampling point (= one week). The files provided present the datasets in their first sheet and keys with the definitions of each term in the second sheet

Tolerance of warmer temperatures does not confer resilience to heatwaves in an Alpine herb

Climate change is generating both sustained trends in average temperatures and higher frequency and intensity of extreme events. This poses a serious threat to biodiversity, especially in vulnerable environments, like alpine systems. Phenotypic plasticity is considered to be an adaptive mechanism to cope with climate change in situ, yet studies of the plastic responses of alpine plants to high temperature stress are scarce. Future weather extremes will occur against a background of warmer temperatures, but we do not know whether acclimation to warmer average temperatures confers tolerance to extreme heatwaves. Nor do we know whether populations on an elevational gradient differ in their tolerance or plasticity in response to warming and heatwave events. We investigated the responses of a suite of functional traits of an endemic Australian alpine herb, Wahlenbergia ceracea, to combinations of predicted future (warmer) temperatures and (relative) heatwaves. We also tested whether responses differed between high- vs. low-elevation populations. When grown under warmer temperatures, W. ceracea plants showed signs of acclimation by means of higher thermal tolerance (Tcrit, T50, and Tmax). They also invested more in flower production, despite showing a concurrent reduction in photosynthetic efficiency (Fv/Fm) and suppression of seed production. Heatwaves reduced both photosynthetic efficiency and longevity. However, we found no evidence that acclimation to warmer temperatures conferred tolerance of the photosynthetic machinery to heatwaves. Instead, when exposed to heatwaves following warmer growth temperatures, plants had lower photosynthetic efficiency and underwent a severe reduction in seed production. High- and low-elevation populations and families exhibited limited genetic variation in trait means and plasticity in response to temperature. We conclude that W. ceracea shows some capacity to acclimate to warming conditions but there is no evidence that tolerance of warmer temperatures confers any resilience to heatwaves.This research was supported by the Australian Research Council (DP170101681), an International Ph.D. Scholarship to RN and an ARC Future Fellowship FT110100453 to LK. Research grants funded all research related costs (such as renting growth chambers or buying equipment), while the scholarship paid a stipend to RN

Effects of developmental and adult environments on ageing

Developmental and adult environments can interact in complex ways to influence the fitness of individuals. Most studies investigating effects of the environment on fitness focus on environments experienced and traits expressed at a single point in an organism's life. However, environments vary with time, so the effects of the environments that organisms experience at different ages may interact to affect how traits change throughout life. Here, we test whether thermal stress experienced during development leads individuals to cope better with thermal stress as adults. We manipulated temperature during both development and adulthood and measured a range of life‐history traits, including senescence, in male and female seed beetles (Callosobruchus maculatus). We found that thermal stress during development reduced adult reproductive performance of females. In contrast, life span and age‐dependent mortality were affected more by adult than developmental environments, with high adult temperatures decreasing longevity and increasing age‐dependent mortality. Aside from an interaction between developmental and adult environments to affect age‐dependent changes in male weight, we did not find any evidence of a beneficial acclimation response to developmental thermal stress. Overall, our results show that effects of developmental and adult environments can be both sex and trait specific, and that a full understanding of how environments interact to affect fitness and ageing requires the integrated study of conditions experienced during different stages of ontogeny

Winter mortality of a passerine bird increases following hotter summers and during winters with higher maximum temperatures

Climate change may influence animal population dynamics through reproduction and mortality. However, attributing changes in mortality to specific climate variables is challenging because the exact time of death is usually unknown in the wild. Here, we investigated climate effects on adult mortality in Australian superb fairy-wrens (Malurus cyaneus). Over a 27-year period, mortality outside the breeding season nearly doubled. This nonbreeding season mortality increased with lower minimum (night-time) and higher maximum (day-time) winter temperatures and with higher summer heat wave intensity. Fine-scale analysis showed that higher mortality in a given week was associated with higher maxima 2 weeks prior and lower minima in the current fortnight, indicating costs of temperature drops. Increases in summer heat waves and in winter maximum temperatures collectively explained 62.6% of the increase in mortality over the study period. Our results suggest that warming climate in both summer and winter can adversely affect survival, with potentially substantial population consequences

Winter mortality of a passerine bird increases following hotter summers and during winters with higher maximum temperatures

Climate change may influence animal population dynamics through reproduction and mortality. However, attributing changes in mortality to specific climate variables is challenging because the exact time of death is usually unknown in the wild. Here, we investigated climate effects on adult mortality in Australian superb fairy-wrens (Malurus cyaneus). Over a 27-year period, mortality outside the breeding season nearly doubled. This nonbreeding season mortality increased with lower minimum (night-time) and higher maximum (day-time) winter temperatures and with higher summer heat wave intensity. Fine-scale analysis showed that higher mortality in a given week was associated with higher maxima 2 weeks prior and lower minima in the current fortnight, indicating costs of temperature drops. Increases in summer heat waves and in winter maximum temperatures collectively explained 62.6% of the increase in mortality over the study period. Our results suggest that warming climate in both summer and winter can adversely affect survival, with potentially substantial population consequences