Effects of simulated heat waves on an experimental plant–herbivore–predator food chain

Greater climatic variability and extreme climatic events are currently emerging as two of the most important facets of climate change. Predicting the effects of extreme climatic events, such as heat waves, is a major challenge because they may affect both organisms and trophic interactions, leading to complex responses at the community level. In this study, we set up a simple three‐level food chain composed of a sweet pepper plant, Capsicum annuum; an aphid, Myzus persicae; and a ladybeetle, Coleomegilla maculata, to explore the consequences of simulated heat waves on organism performance, trophic interactions, and population dynamics. We found that (1) heat waves do not affect plant biomass, significantly reduce the abundance and fecundity of aphids, and slightly affect ladybeetle developmental time and biomass, (2) heat waves decrease the impact of ladybeetles on aphid populations but do not modify the effect of aphids on plant biomass, and (3) food chains including predatory ladybeetles are more resistant to heat waves than a simple plant–aphid association, with aphid abundance being less influenced by heat waves in the presence of C. maculata. Our results suggest that more biodiverse ecosystems with predators exerting a strong biotic control are likely to be less influenced by abiotic factors and then more resistant to extreme climatic events than impoverished ecosystems lacking predators. Our study emphasizes the importance of assessing the effects of climatic change on each trophic level as well as on trophic interactions to further our understanding of the stability, resilience, and resistance of ecological communities under climatic forcing.     

Trait‐specific consequences of inbreeding on adaptive phenotypic plasticity

Environmental changes may stress organisms and stimulate an adaptive phenotypic response. Effects of inbreeding often interact with the environment and can decrease fitness of inbred individuals exposed to stress more so than that of outbred individuals. Such an interaction may stem from a reduced ability of inbred individuals to respond plastically to environmental stress; however, this hypothesis has rarely been tested. In this study, we mimicked the genetic constitution of natural inbred populations by rearing replicate Drosophila melanogaster populations for 25 generations at a reduced population size (10 individuals). The replicate inbred populations, as well as control populations reared at a population size of 500, were exposed to a benign developmental temperature and two developmental temperatures at the lower and upper margins of their viable range. Flies developed at the three temperatures were assessed for traits known to vary across temperatures, namely abdominal pigmentation, wing size, and wing shape. We found no significant difference in phenotypic plasticity in pigmentation or in wing size between inbred and control populations, but a significantly higher plasticity in wing shape across temperatures in inbred compared to control populations. Given that the norms of reaction for the noninbred control populations are adaptive, we conclude that a reduced ability to induce an adaptive phenotypic response to temperature changes is not a general consequence of inbreeding and thus not a general explanation of inbreeding–environment interaction effects on fitness components.     

Life stage‐dependent effects of experimental heat waves on an insect herbivore

1. Worldwide concern about the consequences of climate change has prompted efforts to understand and predict the responses of populations to changes in temperature. 2. A heat wave can adversely affect organisms, may affect different life stages differently, and could decrease populations. In this study, green peach aphid [Myzus persicae (Sulzer)] nymphs, reproductive adults, and late‐reproductive adults were exposed to a heat wave, defined as 5 °C above the control temperature regime for five consecutive days. 3. The negative effects of experimental warming on development, reproduction, and survival negatively affected population growth. Nymphs and reproductive adults were more severely impacted than late‐reproductive adults. 4. Experiments designed to mimic temperature regimes can assess the direct effects of climate change on individuals and populations. Our study highlights the importance of assessing the life stage‐specific responses to heat stress.     

Insect responses to heat: physiological mechanisms,evolution and ecological implications in a warming world

Surviving changing climate conditions is particularly difficult for organisms such as insects that depend on environmental temperature to regulate their physiological functions. Insects are extremely threatened by global warming, since many do not have enough physiological tolerance even to survive continuous exposure to the current maximum temperatures experienced in their habitats. Here, we review literature on the physiological mechanisms that regulate responses to heat and provide heat tolerance in insects: (i) neuronal mechanisms to detect and respond to heat; (ii) metabolic responses to heat; (iii) thermoregulation; (iv) stress responses to tolerate heat; and (v) hormones that coordinate developmental and behavioural responses at warm temperatures. Our review shows that, apart from the stress response mediated by heat shock proteins, the physiological mechanisms of heat tolerance in insects remain poorly studied. Based on life‐history theory, we discuss the costs of heat tolerance and the potential evolutionary mechanisms driving insect adaptations to high temperatures. Some insects may deal with ongoing global warming by the joint action of phenotypic plasticity and genetic adaptation. Plastic responses are limited and may not be by themselves enough to withstand ongoing warming trends. Although the evidence is still scarce and deserves further research in different insect taxa, genetic adaptation to high temperatures may result from rapid evolution. Finally, we emphasize the importance of incorporating physiological information for modelling species distributions and ecological interactions under global warming scenarios. This review identifies several open questions to improve our understanding of how insects respond physiologically to heat and the evolutionary and ecological consequences of those responses. Further lines of research are suggested at the species, order and class levels, with experimental and analytical approaches such as artificial selection, quantitative genetics and comparative analyses.     

Integrating metabolic performance,thermal tolerance,and plasticity enables for more accurate predictions on species vulnerability to acute and chronic effects of global warming

Predicting species vulnerability to global warming requires a comprehensive, mechanistic understanding of sublethal and lethal thermal tolerances. To date, however, most studies investigating species physiological responses to increasing temperature have focused on the underlying physiological traits of either acute or chronic tolerance in isolation. Here we propose an integrative, synthetic approach including the investigation of multiple physiological traits (metabolic performance and thermal tolerance), and their plasticity, to provide more accurate and balanced predictions on species and assemblage vulnerability to both acute and chronic effects of global warming. We applied this approach to more accurately elucidate relative species vulnerability to warming within an assemblage of six caridean prawns occurring in the same geographic, hence macroclimatic, region, but living in different thermal habitats. Prawns were exposed to four incubation temperatures (10, 15, 20 and 25 °C) for 7 days, their metabolic rates and upper thermal limits were measured, and plasticity was calculated according to the concept of Reaction Norms, as well as Q₁₀ for metabolism. Compared to species occupying narrower/more stable thermal niches, species inhabiting broader/more variable thermal environments (including the invasive Palaemon macrodactylus) are likely to be less vulnerable to extreme acute thermal events as a result of their higher upper thermal limits. Nevertheless, they may be at greater risk from chronic exposure to warming due to the greater metabolic costs they incur. Indeed, a trade‐off between acute and chronic tolerance was apparent in the assemblage investigated. However, the invasive species P. macrodactylus represents an exception to this pattern, showing elevated thermal limits and plasticity of these limits, as well as a high metabolic control. In general, integrating multiple proxies for species physiological acute and chronic responses to increasing temperature helps providing more accurate predictions on species vulnerability to warming.     

The effects of phenotypic plasticity and local adaptation on forecasts of species range shifts under climate change

Species are the unit of analysis in many global change and conservation biology studies; however, species are not uniform entities but are composed of different, sometimes locally adapted, populations differing in plasticity. We examined how intraspecific variation in thermal niches and phenotypic plasticity will affect species distributions in a warming climate. We first developed a conceptual model linking plasticity and niche breadth, providing five alternative intraspecific scenarios that are consistent with existing literature. Secondly, we used ecological niche‐modeling techniques to quantify the impact of each intraspecific scenario on the distribution of a virtual species across a geographically realistic setting. Finally, we performed an analogous modeling exercise using real data on the climatic niches of different tree provenances. We show that when population differentiation is accounted for and dispersal is restricted, forecasts of species range shifts under climate change are even more pessimistic than those using the conventional assumption of homogeneously high plasticity across a species' range. Suitable population‐level data are not available for most species so identifying general patterns of population differentiation could fill this gap. However, the literature review revealed contrasting patterns among species, urging greater levels of integration among empirical, modeling and theoretical research on intraspecific phenotypic variation.     

Eutrophication effects on greenhouse gas fluxes from shallow‐lake mesocosms override those of climate warming

Fresh waters make a disproportionately large contribution to greenhouse gas (GHG) emissions, with shallow lakes being particular hot spots. Given their global prevalence, how GHG fluxes from shallow lakes are altered by climate change may have profound implications for the global carbon cycle. Empirical evidence for the temperature dependence of the processes controlling GHG production in natural systems is largely based on the correlation between seasonal temperature variation and seasonal change in GHG fluxes. However, ecosystem‐level GHG fluxes could be influenced by factors, which while varying seasonally with temperature are actually either indirectly related (e.g. primary producer biomass) or largely unrelated to temperature, for instance nutrient loading. Here, we present results from the longest running shallow‐lake mesocosm experiment which demonstrate that nutrient concentrations override temperature as a control of both the total and individual GHG flux. Furthermore, testing for temperature treatment effects at low and high nutrient levels separately showed only one, rather weak, positive effect of temperature (CH₄ flux at high nutrients). In contrast, at low nutrients, the CO₂ efflux was lower in the elevated temperature treatments, with no significant effect on CH₄ or N₂O fluxes. Further analysis identified possible indirect effects of temperature treatment. For example, at low nutrient levels, increased macrophyte abundance was associated with significantly reduced fluxes of both CH₄ and CO₂ for both total annual flux and monthly observation data. As macrophyte abundance was positively related to temperature treatment, this suggests the possibility of indirect temperature effects, via macrophyte abundance, on CH₄ and CO₂ flux. These findings indicate that fluxes of GHGs from shallow lakes may be controlled more by factors indirectly related to temperature, in this case nutrient concentration and the abundance of primary producers. Thus, at ecosystem scale, response to climate change may not follow predictions based on the temperature dependence of metabolic processes.     

Error management theory and the adaptive significance of transgenerational maternal‐stress effects on offspring phenotype

It is well established that circulating maternal stress hormones (glucocorticoids, GCs) can alter offspring phenotype. There is also a growing body of empirical work, within ecology and evolution, indicating that maternal GCs link the environment experienced by the mother during gestation with changes in offspring phenotype. These changes are considered to be adaptive if the maternal environment matches the offspring's environment and maladaptive if it does not. While these ideas are conceptually sound, we lack a testable framework that can be used to investigate the fitness costs and benefits of altered offspring phenotypes across relevant future environments. We present error management theory as the foundation for a framework that can be used to assess the adaptive potential of maternal stress hormones on offspring phenotype across relevant postnatal scenarios. To encourage rigorous testing of our framework, we provide field‐testable hypotheses regarding the potential adaptive role of maternal stress across a diverse array of taxa and life histories, as well as suggestions regarding how our framework might provide insight into past, present, and future research. This perspective provides an informed lens through which to design and interpret experiments on the effects of maternal stress, provides a framework for predicting and testing variation in maternal stress across and within taxa, and also highlights how rapid environmental change that induces maternal stress may lead to evolutionary traps.     

Non‐breeding Cackling,Ross's and Snow Geese on Baffin Island show no loss of body mass during wing moult

Non‐breeding Cackling Branta hutchinsii, Ross's Anser rossii and Lesser Snow Geese Anser caerulescens caerulescens captured during remigial moult on Baffin Island in 2015 showed no loss of body mass with moult stage, and individual variation in mass was largely explained by sex and measures of body size (tarsus length). Exceptional conditions in 2015 resulted in almost no reproductive effort or success in that year, so captured geese of all three species were likely to have been non‐breeding individuals that initiated moult early, whereas there were almost no failed or successful breeders, which would normally moult later. This suggests that in a non‐breeding year (i.e. in the absence of competition from large numbers of goslings), locally moulting geese can obtain sufficient exogenous energy to meet their needs during the flightless wing moult period without losing body mass. This also is consistent with the hypothesis that in other species of geese, accumulation of fat stores prior to, and depletion of such stores during, wing moult is adaptive and likely to be a feature of individual plasticity to meet particular needs, such as undertaking moult migration to remote sites where precise foraging and predation conditions cannot be anticipated, or where competition from more dominant individuals may restrict their access to a reliable food supply.     

Temperature and population density: interactional effects of environmental factors on phenotypic plasticity,immune defenses,and disease resistance in an insect pest

Temperature and crowding are key environmental factors mediating the transmission and epizooty of infectious disease in ectotherm animals. The host physiology may be altered in a temperature‐dependent manner and thus affects the pathogen development and course of diseases within an individual and host population, or the transmission rates (or infectivity) of pathogens shift linearly with the host population density. To our understanding, the knowledge of interactive and synergistic effects of temperature and population density on the host–pathogen system is limited. Here, we tested the interactional effects of these environmental factors on phenotypic plasticity, immune defenses, and disease resistance in the velvetbean caterpillar Anticarsia gemmatalis. Upon egg hatching, caterpillars were reared in thermostat‐controlled chambers in a 2 × 4 factorial design: density (1 or 8 caterpillars/pot) and temperature (20, 24, 28, or 32°C). Of the immune defenses assessed, encapsulation response was directly affected by none of the environmental factors; capsule melanization increased with temperature in both lone‐ and group‐reared caterpillars, although the lone‐reared ones presented the most evident response, and hemocyte numbers decreased with temperature regardless of the population density. Temperature, but not population density, affected considerably the time from inoculation to death of velvetbean caterpillar. Thus, velvetbean caterpillars succumbed to Anticarsia gemmatalis multiple nucleopolyhedrovirus (AgMNPV) more quickly at higher temperatures than at lower temperatures. As hypothesized, temperature likely affected caterpillars' movement rates, and thus the contact between conspecifics, which in turn affected the phenotypic expression of group‐reared caterpillars. Our results suggest that environmental factors, mainly temperature, strongly affect both the course of disease in velvetbean caterpillar population and its defenses against pathogens. As a soybean pest, velvetbean caterpillar may increase its damage on soybean fields under a scenario of global warming as caterpillars may reach the developmental resistance faster, and thus decrease their susceptibility to biological control by AgMNPV.     

Life‐history traits buffer against heat wave effects on predator–prey dynamics in zooplankton

In addition to an increase in mean temperature, extreme climatic events, such as heat waves, are predicted to increase in frequency and intensity with climate change, which are likely to affect organism interactions, seasonal succession, and resting stage recruitment patterns in terrestrial as well as in aquatic ecosystems. For example, freshwater zooplankton with different life‐history strategies, such as sexual or parthenogenetic reproduction, may respond differently to increased mean temperatures and rapid temperature fluctuations. Therefore, we conducted a long‐term (18 months) mesocosm experiment where we evaluated the effects of increased mean temperature (4°C) and an identical energy input but delivered through temperature fluctuations, i.e., as heat waves. We show that different rotifer prey species have specific temperature requirements and use limited and species‐specific temperature windows for recruiting from the sediment. On the contrary, co‐occurring predatory cyclopoid copepods recruit from adult or subadult resting stages and are therefore able to respond to short‐term temperature fluctuations. Hence, these different life‐history strategies affect the interactions between cyclopoid copepods and rotifers by reducing the risk of a temporal mismatch in predator–prey dynamics in a climate change scenario. Thus, we conclude that predatory cyclopoid copepods with long generation time are likely to benefit from heat waves since they rapidly “wake up” even at short temperature elevations and thereby suppress fast reproducing prey populations, such as rotifers. In a broader perspective, our findings suggest that differences in life‐history traits will affect predator–prey interactions, and thereby alter community dynamics, in a future climate change scenario.     

Dynamic size responses to climate change: prevailing effects of rising temperature drive long‐term body size increases in a semi‐arid passerine

Changes in animal body size have been widely reported as a correlate of contemporary climate change. Body size affects metabolism and fitness, so changing size has implications for resilience, yet the climatic factors that drive size variation remain poorly understood. We test the role of mean and extreme temperature, rainfall, and remotely sensed primary productivity (NDVI) as drivers of body size in a sedentary, semi‐arid Australian passerine, Ptilotula (Lichenostomus) penicillatus, over 23 years. To distinguish effects due to differential growth from changes in population composition, we analysed first‐year birds and adults separately and considered climatic variation at three temporal scales (current, previous, and preceding 5 years). The strongest effects related to temperature: in both age classes, larger size was associated with warmer mean temperatures in the previous year, contrary to Bergmann's Rule. Moreover, adults were larger in warmer breeding seasons, while first years was larger after heatwaves; these effects are more likely to be mediated through size‐dependent mortality, highlighting the role of body size in determining vulnerability to extinction. In addition to temperature, larger adult size was associated with lower primary productivity, which may reflect a trade‐off between vegetative growth and nectar production, on which adults rely. Finally, lower rainfall was associated with decreasing size in first year and adults, most likely related to decreased food availability. Overall, body size increased over 23 years, strongly in first‐year birds (2.7%) compared with adults (1%), with size outcomes a balance between competing drivers. As rainfall declined over time and productivity remained fairly stable, the temporal increase in body size appears largely driven by rising mean temperature and temperature extremes. Body size responses to environmental change are thus complex and dynamic, driven by effects on growth as well as mortality.     

Effects of temperature and nest heat exposure on nestling growth,dehydration and survival in a Mediterranean hole‐nesting passerine

Variable environments impose constraints on adaptation by modifying selection gradients unpredictably. Optimal bird development requires an adequate thermal range, outside which temperatures can alter nestling physiology, condition and survival. We studied the effect of temperature and nest heat exposure on the reproductive success of a population of double‐brooded Spotless Starlings Sturnus unicolor breeding in a nestbox colony in central Spain with a marked intra‐seasonal variation in temperature. We assessed whether the effect of temperature differed between first and second broods, thus constraining optimal nest‐site choice. Ambient temperature changed greatly during the chick‐rearing period and had a strong influence on nestling mass and all body size measures we recorded, although patterns of clutch size or nestling mortality were not influenced. This effect differed between first and second broods: nestlings were found to have longer wings and bills with increasing temperature in first broods, whereas the effect was the opposite in second broods. Ambient temperature was not related to nestling body mass or tarsus‐length in first broods, but in second broods, nestlings were lighter and had smaller tarsi with higher ambient temperatures. The exposure of nestboxes to heat influenced nestling morphology: heat exposure index was negatively related to nestling body mass and wing‐length in second broods, but not in first broods. Furthermore, there was a positive relationship between nest heat exposure and nestling dehydration. Our results suggest that optimal nest choice is constrained by varying environmental conditions in birds breeding over prolonged periods, and that there should be selection for parents to switch from sun‐exposed to sun‐protected nest‐sites as the season progresses. However, nest‐site availability and competition for sites are likely to impose constraints on this choice.     

Differential plasticity of size and mass to environmental change in a hibernating mammal

Morphological changes following changes in species' distribution and phenology have been suggested to be the third universal response to global environmental change. Although structural size and body mass result from different genetic, physiological, and ecological mechanisms, they are used interchangeably in studies evaluating population responses to environmental change. Using a 22‐year (1991–2013) dataset including 1768 individuals, we investigated the coupled dynamics of size and mass in a hibernating mammal, the Alpine marmot (Marmota marmota), in response to local environmental conditions. We (i) quantified temporal trends in both traits, (ii) determined the environmental drivers of trait dynamics, and (iii) identified the life‐history processes underlying the observed changes. Both phenotypic traits were followed through life: we focused on the initial trait value (juvenile size and mass) and later‐life development (annual change in size [Δsize] and mass [Δmass]). First, we demonstrated contrasting dynamics between size and mass over the study period. Juvenile size and subsequent Δsize showed significant declines, whereas juvenile mass and subsequent Δmass remained constant. As a consequence of smaller size associated with a similar mass, individuals were in better condition in recent years. Second, size and mass showed different sensitivities to environmental variables. Both traits benefited from early access to resources in spring, whereas Δmass, particularly in early life, also responded to summer and winter conditions. Third, the interannual variation in both traits was caused by changes in early life development. Our study supports the importance of considering the differences between size and mass responses to the environment when evaluating the mechanisms underlying population dynamics. The current practice of focusing on only one trait in population modeling can lead to misleading conclusions when evaluating species' resilience to contemporary climate change.     

The impacts of extreme and fluctuating temperatures on trait‐mediated indirect aphid–parasitoid interactions

1. Global climate change models predict an increase in the frequency and magnitude of extreme temperature events. These temperature events, heatwaves for example, will impact a wide range of physiological and behavioural processes, particularly in ectotherms, and may therefore influence interactions between species. 2. Anti‐predator responses may be more costly under more severe temperature regimes and therefore trait‐mediated disturbance could lead to high mortality or reduced reproduction under extreme and fluctuating temperature regimes. 3. We examined the impacts of extreme and fluctuating temperatures on trait‐mediated indirect interactions in an aphid–parasitoid community. 4. In treatments that isolated the effects of trait‐mediated disturbance from the effects of foraging parasitoids we found that an increase in both the amplitude and frequency of peak temperatures reduced aphid numbers and provided evidence that the cost of trait‐mediated disturbance could increase under frequent periods of high temperature. Aphid dispersal also increased with more frequent periods of high temperature. 5. In treatments where female wasps were allowed to freely forage (direct + trait‐mediated effects), there was no evidence that extreme and fluctuating temperatures influenced the wasp's foraging ability. Exposure to extreme fluctuating temperatures did not influence the offspring production of exposed wasps or the position of the mummies within the plots.     

Phenotypic plasticity in age and size at maturity and its effects on the integrated phenotypic expressions of life history traits of Cardamine flexuosa (Cruciferae)

We analyzed variation in phenotypic plasticity of life history traits between two Cardamine flexuosa populations based on differences in plasticity of age and size at maturity. C. flexuosa (Cruciferae) is a facultative, vernalization-sensitive, long-day annual, and its phenology and the phenotypic expressions of many life history traits are largely controlled by photoperiod and vernalization in natural populations. We used plants from two populations which differed in their responses to chilling and photoperiod treatments. The timing of developmental processes was changed by controlling temperature and photoperiod regimes in growth chambers. Plasticity in size at maturity was analyzed as changes in a growth trajectory using two parameters, age at maturity (Δt) and growth rate (k). Both traits showed plasticity, but differences between the populations were found mostly for Δt. Distinctive differences in size at maturity of individuals in the two populations were mainly due to different amounts of plasticity in Δt. Variations in plasticity of nine other life history traits and their associations to age and size at maturity were also analyzed. Variation for eight of the traits can be described, at least in part, as a function of age and size at maturity for both populations, and most of the variation in the total number of seeds was explained by age and size at maturity. Only age at maturity had any effect on changes in resource allocation. The nine life history traits were integrated through associated character expressions with age and size at maturity. Changes in the association between a trait and age and/or size at maturity were rather conservative compared to changes in the plasticity of a trait between the two populations. Associations with age and size at maturity are mostly explicable in terms of inherent relationships in the developmental processes, and they may limit the ecological range expansion and the adaptive evolution of plasticity in C. flexuosa. The negative correlation between reproductive allocation and age at maturity can be a cost of delaying maturation in C. flexuosa.     

Spatial and body‐size dependent response of marine pelagic communities to projected global climate change

Temperature, oxygen, and food availability directly affect marine life. Climate models project a global warming of the ocean's surface (~+3 °C), a de‐oxygenation of the ocean's interior (~?3%) and a decrease in total marine net primary production (~?8%) under the ‘business as usual’ climate change scenario (RCP8.5). We estimated the effects of these changes on biological communities using a coupled biogeochemical (PISCES) – ecosystems (APECOSM) model forced by the physical outputs of the last generation of the IPSL‐CM Earth System Model. The APECOSM model is a size‐structured bio‐energetic model that simulates the 3D dynamical distributions of three interactive pelagic communities (epipelagic, mesopelagic, and migratory) under the effects of multiple environmental factors. The PISCES‐APECOSM model ran from 1850 to 2100 under historical forcing followed by RCP8.5. Our RCP8.5 simulation highlights significant changes in the spatial distribution, biomass, and maximum body‐size of the simulated pelagic communities. Biomass and maximum body‐size increase at high latitude over the course of the century, reflecting the capacity of marine organisms to respond to new suitable environment. At low‐ and midlatitude, biomass and maximum body‐size strongly decrease. In those regions, large organisms cannot maintain their high metabolic needs because of limited and declining food availability. This resource reduction enhances the competition and modifies the biomass distribution among and within the three communities: the proportion of small organisms increases in the three communities and the migrant community that initially comprised a higher proportion of small organisms is favored. The greater resilience of small body‐size organisms resides in their capacity to fulfill their metabolic needs under reduced energy supply and is further favored by the release of predation pressure due to the decline of large organisms. These results suggest that small body‐size organisms might be more resilient to climate change than large ones.     

Carry‐over effects of conditions at the wintering grounds on breeding plumage signals in a migratory bird: roles of phenotypic plasticity and selection

To understand the consequences of ever‐changing environment on the dynamics of phenotypic traits, distinguishing between selection processes and individual plasticity is crucial. We examined individual consistency/plasticity in several male secondary sexual traits expressed during the breeding season (white wing and forehead patch size, UV reflectance of white wing patch and dorsal melanin coloration) in a migratory pied flycatcher (Ficedula hypoleuca) population over an 11‐year period. Furthermore, we studied carry‐over effects of three environmental variables (NAO, a climatic index; NDVI, a vegetation index; and rainfall) at the wintering grounds (during prebreeding moult) on the expression of these breeding plumage traits of pied flycatcher males at individual and population levels. Whereas NAO correlates negatively with moisture in West Africa, NDVI correlates positively with primary production. Forehead patch size and melanin coloration were highly consistent within individuals among years, whereas the consistency of the other two traits was moderate. Wing patch size decreased with higher NAO and increased with higher rainfall and NDVI at the individual level. Interestingly, small‐patched males suffered lower survival during high NAO winters than large‐patched males, and vice versa during low NAO winters. These counteracting processes meant that the individual‐level change was masked at the population level where no relationship was found. Our results provide a good example of how variation in the phenotypic composition of a natural population can be a result of both environment‐dependent individual plasticity and short‐term microevolution. Moreover, when plasticity and viability selection operate simultaneously, their impacts on population composition may not be evident.     

Variable strength of top‐down effects in Nothofagus forests: bird predation and insect herbivory during an ENSO event

Abstract Predators are thought to play a key role in controlling herbivory, thus having positive indirect effects on plants. However, evidence for terrestrial trophic cascades is still fragmentary, perhaps due to variation in top‐down forces created by environmental heterogeneity. We examined the magnitude of predation effects on foliar damage by chewing insects and mean leaf size, by excluding birds from saplings in ‘dry’ and ‘wet’Nothofagus pumilio forests in the northern Patagonian Andes, Argentina. The experiment lasted 2 years encompassing a severe drought during the La Niña phase of a strong El Niño/Southern Oscillation event, which was followed by unusually high background folivory levels. Insect damage was consistently higher in wet than in dry forest saplings. In the drought year (1999), bird exclusion increased folivory rates in both forests but did not affect tree leaf size. In the ensuing season (2000), leaf damage was generally twice as high as in the drought year. As a result, bird exclusion not only increased the extent of folivory but also significantly decreased sapling leaf size. The latter effect was stronger in the wet forest, suggesting compensation of leaf area loss by dry forest saplings. Overall, the magnitude of predator indirect effects depended on the response variable measured. Insectivorous birds were more effective at reducing folivory than at facilitating leaf area growth. Our results indicate that bird‐initiated trophic cascades protect N. pumilio saplings from insect damage even during years with above‐normal herbivory, and also support the view that large‐scale climatic events influence the strength of trophic cascades.