Environmental degradation amplifies species' responses to temperature variation in a trophic interaction.

Land-use and climate change are two of the primary drivers of the current biodiversity crisis. However, we lack understanding of how single-species and multispecies associations are affected by interactions between multiple environmental stressors. We address this gap by examining how environmental degradation interacts with daily stochastic temperature variation to affect individual life history and population dynamics in a host-parasitoid trophic interaction, using the Indian meal moth, Plodia interpunctella, and its parasitoid wasp Venturia canescens. We carried out a single-generation individual life-history experiment and a multigeneration microcosm experiment during which individuals and microcosms were maintained at a mean temperature of 26°C that was either kept constant or varied stochastically, at four levels of host resource degradation, in the presence or absence of parasitoids. At the individual level, resource degradation increased juvenile development time and decreased adult body size in both species. Parasitoids were more sensitive to temperature variation than their hosts, with a shorter juvenile stage duration than in constant temperatures and a longer adult life span in moderately degraded environments. Resource degradation also altered the host's response to temperature variation, leading to a longer juvenile development time at high resource degradation. At the population level, moderate resource degradation amplified the effects of temperature variation on host and parasitoid populations compared with no or high resource degradation and parasitoid overall abundance was lower in fluctuating temperatures. Top-down regulation by the parasitoid and bottom-up regulation driven by resource degradation contributed to more than 50% of host and parasitoid population responses to temperature variation. Our results demonstrate that environmental degradation can strongly affect how species in a trophic interaction respond to short-term temperature fluctuations through direct and indirect trait-mediated effects. These effects are driven by species differences in sensitivity to environmental conditions and modulate top-down (parasitism) and bottom-up (resource) regulation. This study highlights the need to account for differences in the sensitivity of species' traits to environmental stressors to understand how interacting species will respond to simultaneous anthropogenic changes.

How well can body size represent effects of the environment on demographic rates? Disentangling correlated explanatory variables.

Demographic rates are shaped by the interaction of past and current environments that individuals in a population experience. Past environments shape individual states via selection and plasticity, and fitness-related traits (e.g. individual size) are commonly used in demographic analyses to represent the effect of past environments on demographic rates. We quantified how well the size of individuals captures the effects of a population's past and current environments on demographic rates in a well-studied experimental system of soil mites. We decomposed these interrelated sources of variation with a novel method of multiple regression that is useful for understanding nonlinear relationships between responses and multicollinear explanatory variables. We graphically present the results using area-proportional Venn diagrams. Our novel method was developed by combining existing methods and expanding upon them. We showed that the strength of size as a proxy for the past environment varied widely among vital rates. For instance, in this organism with an income breeding life history, the environment had more effect on reproduction than individual size, but with substantial overlap indicating that size encompassed some of the effects of the past environment on fecundity. This demonstrates that the strength of size as a proxy for the past environment can vary widely among life-history processes within a species, and this variation should be taken into consideration in trait-based demographic or individual-based approaches that focus on phenotypic traits as state variables. Furthermore, the strength of a proxy will depend on what state variable(s) and what demographic rate is being examined; that is, different measures of body size (e.g. length, volume, mass, fat stores) will be better or worse proxies for various life-history processes.

Density-dependent immunity and parasitism risk in experimental populations of lizards naturally infested by ixodid ticks.

When effective immune defenses against parasites are costly and resources limited, individuals are expected to alter their investment in immunity in response to the risk of infection. As an ecological factor that can affect both food abundance and parasite exposure, host density can play an important role in host immunity and host-parasite interactions. High levels of intraspecific competition for food and social stress at high host density may diminish immune defenses and increase host susceptibility to parasites. At the same time, for contagious and environmentally transmitted parasites, parasite exposure often increases with host density, whereas in mobile parasites that actively search for hosts, parasite exposure can decrease with host density due to the "encounter-dilution effect." To unravel these multiple and potentially opposing effects of host density on immunity, we manipulated density of the common lizard Zootoca vivipara and measured local inflammation in response to PHA injection and levels of infestation by the tick Ixodes ricinus, a mobile ectoparasite for which we expected an encounter-dilution effect to occur. Local inflammation strongly decreased with lizard density in adults, but not in yearlings. Tick infestation (abundance and prevalence) was negatively correlated with lizard density in both age classes. Using path analyses, we found independent, direct negative density feedbacks on immunity and parasite exposure in adults, supporting the hypothesis of energy constraints and/or physiological stress acting on immunity at high density. In contrast, for yearlings, the best path model showed that density diluted exposure to parasites, which themselves down-regulated immune defenses in lizards. These results highlight the importance of investigating the pathways among host density, host immunity, and parasite infestation, while accounting for relevant individual traits such as age.

Density-dependent life history and the dynamics of small populations.

1. Small population dynamics depend importantly on the strength and shape of density dependence. Unfortunately, the lack of reliable life-history data often prevents to make accurate demographic predictions for populations regulated by density dependence. 2. We created a gradient from low to high densities in small experimental populations of common lizards (Zootoca vivipara) and investigated the shape and strength of the density dependence of life-history traits during a yearly cycle. We then analysed stochastic population dynamics using one-sex and two-sex age-structured matrix models. 3. Body growth and reproductive performances decreased with density, yearling and adult survival and body size at birth were density-independent, and juvenile survival increased with density. The density dependence of reproduction was partly explained by positive effects of body size on age at first reproduction and clutch size. 4. Parturition date decreased with density in sparse populations and then increased, providing one of the first empirical evidence of a component Allee effect in the phenology of reproduction. 5. Population growth rate (λ) was most affected by variations in juvenile and yearling survival. However, density at equilibrium was most affected by juvenile access to reproduction and yearling clutch size. 6. Stochastic simulations revealed that negative density dependence buffers the effects of initial density on extinction probability, has positive effects on the persistence of sparse populations and interacts with sex ratio fluctuations to shape extinction dynamics. 7. This study demonstrates that negative density dependence modifies the dynamics of small populations and should be investigated together with Allee effects to predict extinction risks.

Conspecifics as informers and competitors: an experimental study in foraging bumble-bees.

Conspecifics are usually considered competitors negatively affecting food intake rates. However, their presence can also inform about resource quality by providing inadvertent social information. Few studies have investigated whether foragers perceive conspecifics as informers or competitors. Here, we experimentally tested whether variation in the density of demonstrators ('none', 'low' and 'high'), whose location indicated flower profitability, affected decision-making of bumble-bees Bombus terrestris. Bumble-bees foraged on either 'simple' (two colours) or 'complex' (four colours) artificial floral communities. We found that conspecifics at low density may be used as sources of information in first flower choices, whereas they appeared as competitors over the whole foraging sequence. Low conspecific densities improved foragers' first-visit success rate in the simple environment, and decreased time to first landing, especially in the complex environment. High conspecific densities did not affect these behavioural parameters, but reduced flower constancy in both floral communities, which may alter the efficiency of pollinating visits. These results suggest that the balance of the costs and benefits of conspecific presence varies with foraging experience, floral community and density. Spatio-temporal scales could thus be an important determinant of social information use. This behavioural flexibility should allow bumble-bees to better exploit their environment.

Direct and socially-mediated effects of food availability late in life on life-history variation in a short-lived lizard.

Food availability is a major environmental factor that can influence life history within and across generations through direct effects on individual quality and indirect effects on the intensity of intra- and intercohort competition. Here, we investigated in yearling and adult common lizards (Zootoca vivipara) the immediate and delayed life-history effects of a prolonged food deprivation in the laboratory. We generated groups of fully fed or food-deprived yearlings and adults at the end of one breeding season. These lizards were released in 16 outdoor enclosures together with yearlings and adults from the same food treatment and with food-deprived or fully fed juveniles, creating four types of experimental populations. Experimental populations were then monitored during 2 years, which revealed complex effects of food on life-history trajectories. Food availability had immediate direct effects on morphology and delayed direct effects on immunocompetence and female body condition at winter emergence. Also, male annual survival rate and female growth rate and body size were affected by an interaction between direct effects of food availability and indirect effects on asymmetric competition with juveniles. Reproductive outputs were insensitive to past food availability, suggesting that female common lizards do not solely rely on stored energy to fuel reproduction. Finally, food conditions had socially-mediated intergenerational effects on early growth and survival of offspring through their effects on the intensity of competition. This study highlights the importance of social interactions among cohorts for life-history trajectories and population dynamics in stage-structured populations.

Ontogenetic trajectories of body coloration reveal its function as a multicomponent nonsenescent signal.

The understanding of developmental patterns of body coloration is challenging because of the multicomponent nature of color signals and the multiple selective pressures acting upon them, which further depend on the sex of the bearer and area of display. Pigmentary colors are thought to be strongly involved in sexual selection, while structural colors are thought to generally associate with conspecifics interactions and improve the discrimination of pigmentary colors. Yet, it remains unclear whether age dependency in each color component is consistent with their potential function. Here, we address lifelong ontogenetic variation in three color components (i.e. UV, pigmentary, and skin background colors) in a birth cohort of common lizards Zootoca vivipara across three ventral body regions (i.e. throat, chest, and belly). All three color components developed sexual dichromatism, with males displaying stronger pigmentary and UV colors but weaker skin background coloration than females. The development of color components led to a stronger sexual dichromatism on the concealed ventral region than on the throat. No consistent signs of late-life decay in color components were found except for a deceleration of UV reflectance increase with age on the throat of males. These results suggest that body color components in common lizards are primarily nonsenescent sexual signals, but that the balance between natural and sexual selection may be altered by the conspicuousness of the area of display. These results further support the view that skin coloration is a composite trait constituted of multiple color components conveying multiple signals depending on age, sex, and body location.