Top-down effects of intraspeciflic predator behavioral variation.

Among-individual variation in predator traits is ubiquitous in nature. However, variation among populations in this trait variation has been seldom considered in trophic dynamics. This has left unexplored (a) to what degree does among-individual variation in predator traits regulate prey populations and (b) to what degree do these effects vary spatially. We address these questions by examining how predator among-individual variation in functional traits shapes communities across habitats of varying structural complexity, in field conditions. We manipulated Chinese mantis (Tenodera sinensis) density (six or twelve individuals) and behavioral trait variability (activity level by movement on an open field) in experimental patches of old fields with varying habitat complexity (density of plant material). Then, we quantified their impacts on lower trophic levels, specifically prey (arthropods > 4 mm) and plant biomass. Predator behavioral variability only altered prey biomass in structurally complex plots, and this effect depended on mantis density. In the plots with the highest habitat complexity and mantis density, behaviorally variable groups decreased prey biomass by 40.3%. In complex plots with low mantis densities, low levels of behavioral variability decreased prey biomass by 32.2%. Behavioral variability and low habitat complexity also changed prey community composition, namely by increasing ant biomass by 881%. Our results demonstrate that among-individual trait variation can shape species-rich prey communities. Moreover, these effects depend on both predator density and habitat complexity. Incorporating this important facet of ecological diversity revealed normally unnoticed effects of functional traits on the structure and function of food webs.

Size-dependent colouration balances conspicuous aposematism and camouflage.

Colour is an important component of many different defensive strategies, but signal efficacy and detectability will also depend on the size of the coloured structures, and how pattern size interacts with the background. Consequently, size-dependent changes in colouration are common among many different species as juveniles and adults frequently use colour for different purposes in different environmental contexts. A widespread strategy in many species is switching from crypsis to conspicuous aposematic signalling as increasing body size can reduce the efficacy of camouflage, while other antipredator defences may strengthen. Curiously, despite being chemically defended, the gold-striped frog (Lithodytes lineatus, Leptodactylidae) appears to do the opposite, with bright yellow stripes found in smaller individuals, whereas larger frogs exhibit dull brown stripes. Here, we investigated whether size-dependent differences in colour support distinct defensive strategies. We first used visual modelling of potential predators to assess how colour contrast varied among frogs of different sizes. We found that contrast peaked in mid-sized individuals while the largest individuals had the least contrasting patterns. We then used two detection experiments with human participants to evaluate how colour and body size affected overall detectability. These experiments revealed that larger body sizes were easier to detect, but that the colours of smaller frogs were more detectable than those of larger frogs. Taken together our data support the hypothesis that the primary defensive strategy changes from conspicuous aposematism to camouflage with increasing size, implying size-dependent differences in the efficacy of defensive colouration. We discuss our data in relation to theories of size-dependent aposematism and evaluate the evidence for and against a possible size-dependent mimicry complex with sympatric poison frogs (Dendrobatidae).

Behavioral and physiological evidence that increasing group size ameliorates the impacts of social disturbance.

Intra-group social stability is important for the long-term productivity and health of social organisms. We evaluated the effect of group size on group stability in the face of repeated social perturbations using a cooperatively breeding fish, Neolamprologus pulcher In a laboratory study, we compared both the social and physiological responses of individuals from small versus large groups to the repeated removal and replacement of the most dominant group member (the breeder male), either with a new male (treatment condition) or with the same male (control condition). Individuals living in large groups were overall more resistant to instability but were seemingly slower to recover from perturbation. Members of small groups were more vulnerable to instability but recovered faster. Breeder females in smaller groups also showed greater physiological preparedness for instability following social perturbations. In sum, we discover both behavioral and physiological evidence that living in larger groups helps to dampen the impacts of social instability in this system.

Predictors of colony extinction vary by habitat type in social spiders.

Many animal societies are susceptible to mass mortality events and collapse. Elucidating how environmental pressures determine patterns of collapse is important for understanding how such societies function and evolve. Using the social spider Stegodyphus dumicola, we investigated the environmental drivers of colony extinction along two precipitation gradients across southern Africa, using the Namib and Kalahari deserts versus wetter savanna habitats to the north and east. We deployed experimental colonies (n = 242) along two ~ 800-km transects and returned to assess colony success in the field after 2 months. Specifically, we noted colony extinction events after the 2-month duration and collected environmental data on the correlates of those extinction events (e.g., evidence of ant attacks, no. of prey captured). We found that colony extinction events at desert sites were more frequently associated with attacks by predatory ants as compared with savanna sites, while colony extinctions in wetter savannas sites were more tightly associated with fungal outbreaks. Our findings support the hypothesis that environments vary in the selection pressures that they impose on social organisms, which may explain why different social phenotypes are often favored in each habitat.

Imperfect transparency and camouflage in glass frogs.

Camouflage patterns prevent detection and/or recognition by matching the background, disrupting edges, or mimicking particular background features. In variable habitats, however, a single pattern cannot match all available sites all of the time, and efficacy may therefore be reduced. Active color change provides an alternative where coloration can be altered to match local conditions, but again efficacy may be limited by the speed of change and range of patterns available. Transparency, on the other hand, creates high-fidelity camouflage that changes instantaneously to match any substrate but is potentially compromised in terrestrial environments where image distortion may be more obvious than in water. Glass frogs are one example of terrestrial transparency and are well known for their transparent ventral skin through which their bones, intestines, and beating hearts can be seen. However, sparse dorsal pigmentation means that these frogs are better described as translucent. To investigate whether this imperfect transparency acts as camouflage, we used in situ behavioral trials, visual modeling, and laboratory psychophysics. We found that the perceived luminance of the frogs changed depending on the immediate background, lowering detectability and increasing survival when compared to opaque frogs. Moreover, this change was greatest for the legs, which surround the body at rest and create a diffuse transition from background to frog luminance rather than a sharp, highly salient edge. This passive change in luminance, without significant modification of hue, suggests a camouflage strategy, "edge diffusion," distinct from both transparency and active color change.

Collective aggressiveness limits colony persistence in high- but not low-elevation sites at Amazonian social spiders.

Identifying the traits that foster group survival in contrasting environments is important for understanding local adaptation in social systems. Here, we evaluate the relationship between the aggressiveness of social spider colonies and their persistence along an elevation gradient using the Amazonian spider, Anelosimus eximius. We found that colonies of A. eximius exhibit repeatable differences in their collective aggressiveness (latency to attack prey stimuli) and that colony aggressiveness is linked with persistence in a site-specific manner. Less aggressive colonies are better able to persist at high-elevation sites, which lack colony-sustaining large-bodied prey, whereas colony aggression was not related to chance of persistence at low-elevation sites. This suggests that low aggressiveness promotes colony survival in high-elevation, prey-poor habitats, perhaps via increased tolerance to resource limitation. These data reveal that the collective phenotypes that relate to colony persistence vary by site, and thus, the path of social evolution in these environments is likely to be affected.

Experimental evidence of frequency-dependent selection on group behaviour.

Evolutionary ecologists often seek to identify the mechanisms maintaining intraspecific variation. In social animals, whole groups can exhibit between-group differences in their collective traits. We examined whether negative frequency-dependent selection (that is, a rare-type advantage) could help to maintain between-group variation. We engineered neighbourhoods of social spider colonies bearing bold or shy foraging phenotypes and monitored their fecundity in situ. We found that bold colonies enjoyed a rare-type advantage that is lost as the frequency of bold colonies in a neighbourhood increases. The success of shy colonies was not frequency dependent. These dynamics seem to be driven by a foraging advantage of bold colonies that is lost in bold neighbourhoods because prey become scarce, and shy colonies perform better than bold colonies under low-resource conditions. Thus, to understand selection on collective traits, it is insufficient to examine groups in isolation. The phenotypic environment in which groups reside and compete must also be considered.

Selection for Collective Aggressiveness Favors Social Susceptibility in Social Spiders.

Particularly socially influential individuals are present in many groups [1-8], but it is unclear whether their emergence is determined by their social influence versus the social susceptibility of others [9]. The social spider Stegodyphus dumicola shows regional variation in apparent leader-follower dynamics. We use this variation to evaluate the relative contributions of leader social influence versus follower social susceptibility in driving this social order. Using chimeric colonies that combine potential leaders and followers, we discover that leader-follower dynamics emerge from the site-specific social susceptibility of followers. We further show that the presence of leaders increases colony survival in environments where leader-follower dynamics occur. Thus, leadership is driven by the "social susceptibility" of the population majority, rather than the social influence of key group members.