Social networks and models for collective motion in animals

The theory of collective motion and the study of animal social networks have, each individually, received much attention. Currently, most models of collective motion do not consider social network structure. The implications for considering collective motion and social networks together are likely to be important. Social networks could determine how populations move in, split up into and form separate groups (social networks affecting collective motion). Conversely, collective movement could change the structure of social networks by creating social ties that did not exist previously and maintaining existing ties (collective motion affecting social networks). Thus, there is a need to combine the two areas of research and examine the relationship between network structure and collective motion. Here, we review different modelling approaches that combine social network structures and collective motion. Although many of these models have not been developed with ecology in mind, they present a current context in which a biologically relevant theory can be developed. We argue that future models in ecology should take inspiration from empirical observations and consider different mechanisms of how social preferences could be expressed in collectively moving animal groups.     

Feeding and non-feeding aggression can be induced in invasive shore crabs by altering food distribution

Human activity can have a large impact on surrounding ecosystems. For example, humans alter resource distributions for other species, potentially modifying these species competitive dynamics. These changes in local competitive processes are frequently associated with species invasions. Here, we investigate how differences in resource distribution affect competitive behaviour using the highly invasive European shore crab (Carcinus maenas). Using a controlled laboratory experiment in combination with behaviour assays and social network analysis, we show that individuals feeding in habitats with clumped food distributions are more aggressive than individuals feeding in habitats where food is evenly dispersed, and this aggression is present even on days where crabs are not feeding. Additionally, this persistent aggression can be induced, suggesting that individuals of this invasive species possess the flexibility to modify their competitive behaviours in response to differences in food distributions. Furthermore, we show how these individual responses can lead to changes in overall organisation of aggressive interactions within a population. We discuss these results in relation to how human impacts can have long-term effects on competitive behavioural strategies, and how behavioural flexibility can allow invasive species to colonise and persist in highly impacted sites such as urban ecosystems.     

Model selection and model averaging in behavioural ecology: the utility of the IT-AIC framework

Behavioural ecologists often study complex systems in which multiple hypotheses could be proposed to explain observed phenomena. For some systems, simple controlled experiments can be employed to reveal part of the complexity; often, however, observational studies that incorporate a multitude of causal factors may be the only (or preferred) avenue of study. We assess the value of recently advocated approaches to inference in both contexts. Specifically, we examine the use of information theoretic (IT) model selection using Akaike’s information criterion (AIC). We find that, for simple analyses, the advantages of switching to an IT-AIC approach are likely to be slight, especially given recent emphasis on biological rather than statistical significance. By contrast, the model selection approach embodied by IT approaches offers significant advantages when applied to problems of more complex causality. Model averaging is an intuitively appealing extension to model selection. However, we were unable to demonstrate consistent improvements in prediction accuracy when using model averaging with IT-AIC; our equivocal results suggest that more research is needed on its utility. We illustrate our arguments with worked examples from behavioural experiments.     

Biliverdin-based egg coloration is enhanced by carotenoid supplementation

Avian eggshell color is remarkably variable among and within species and its possible function has long been controversial. Female birds use biliverdin pigment to color their eggs blue and green. Although evidence is accumulating that blue-green eggshell coloration is costly to produce, the exact mechanisms underlying its expression are little studied. Biliverdin pigment is involved in important maintenance functions such as immune and antioxidant defenses and shows similar properties to carotenoid pigments. Carotenoids play a role as immunoenhancers and have the potential to ameliorate the impact of oxidative processes, although their antioxidant function could be minor for some bird species. Important life-history components can be limited by carotenoid availability, including sexual displays and fecundity. Here we explored if biliverdin-based eggshell pigmentation was affected by carotenoid availability in a seabird, by performing a carotenoid supplementation experiment in female blue-footed boobies (Sula nebouxii) after laying their first egg. In this species, blue eggshell color is associated with incubation patterns, and carotenoid availability affects the crucial trade-off between female foot ornamentation and fecundity. We found that under natural conditions, there was a decline in eggshell color with laying order, suggesting pigment limitation for females. However, carotenoid-supplemented females had enhanced second egg coloration compared to controls, and the color increase was independent of their previous pigment allocation to the first egg. Our results suggest that biliverdin-based eggshell coloration is costly to produce and can be alleviated by current carotenoid availability.     

Darwin’s “tug-of-war” vs. starlings’ “horse-racing”: how adaptations for sequential encounters drive simultaneous choice

Charles Darwin aided his private decision making by an explicit deliberation, famously deciding whether or not to marry by creating a list of points in a table with two columns: “Marry” and “Not Marry”. One hundred seventy-two years after Darwin’s wedding, we reconsider whether this process of choice, under which individuals assign values to their options and compare their relative merits at the time of choosing (the tug-of-war model), applies to our experimental animal, the European Starling, Sturnus vulgaris. We contrast this with the sequential choice model that postulates that decision-makers make no comparison between options at the time of choice. According to the latter, behaviour in simultaneous choices reflects adaptations to contexts with sequential encounters, in which the choice is whether to take an opportunity or let it pass. We postulate that, in sequential encounters, the decision-maker assigns (by learning) a subjective value to each option, reflecting its payoff relative to background opportunities. This value is expressed as latency and/or probability to accept each opportunity as opposed to keep searching. In simultaneous encounters, choice occurs through each option being processed independently, by a race between the mechanisms that generate option-specific latencies. We describe these alternative models and review data supporting the predictions of the sequential choice model.     

Sex-specific fitness consequences of dispersal in Siberian jays

In most birds, natal dispersal is female-biased, but the selective pressures leading to this pattern have rarely been explored with comprehensive data on lifetime reproductive success. In territorial birds, the benefit of philopatry should be higher for males than for females when males establish territories for which knowledge about the local environment is important. As females may use male characteristics for mate choice, and hence indirectly for territory choice, the benefit from the direct knowledge of the local environment may be lower for females than males. We tested this hypothesis using data from a long-term study of group living corvids, the Siberian jays (Perisoreus infaustus). In this species, the socially dominant offspring delay dispersal while the sub-dominant offspring leave the family group directly after reaching independence. Our results show that natal dispersal distance (a proxy for local knowledge) was related to sex and dispersal timing (a proxy for “quality”): Females and early dispersers traveled further on average than males and delayed dispersers. Furthermore, dispersal distance and timing were negatively related to the number of recruits produced over an individual’s lifetime in males, but not in females. Hence, the results support the hypothesis that the female-biased natal dispersal found in this and other bird species may come about through higher lifetime reproductive success of philopatric males than females.     

Movement and memory: different cognitive strategies are used to search for resources with different natural distributions

Recent attempts to integrate function and mechanism have resulted in an appreciation of the relevance of forager psychology to understanding the functional aspects of foraging behaviour. Conversely, an acknowledgement of the functional diversity of learning mechanisms has led to greater understanding of the adaptive nature of cognition. In this paper, we present data from three experiments suggesting that noisy miner birds use different cognitive strategies when searching for foods with different distributions. When searching for nectar, an immobile, readily depleted resource, birds spontaneously attend to fine-scale spatial information and use a spatial memory-based strategy that is efficient in a novel context and largely resistant to disruptions to movement. When searching for invertebrates, a mobile, clumped and cryptic resource, birds employ a strategy whose efficiency increases with increased task familiarity, is vulnerable to disruptions to their movement and may rely more on memory for movement rules than memory for location information. Previous reports of adapted cognition have reported performance differences between species (for example, better spatial cognitive performance in storing versus non-storing birds). Ours is the first study to demonstrate that differences in cognitive strategy (as opposed to just enhanced performance) occur within a single species in different foraging contexts. As well as providing an example of how specially adapted cognitive mechanisms might work, our data further emphasise the importance of jointly considering functional and mechanistic aspects to fully understand the adaptive complexities of behaviour.     

Two roads to two sexes: unifying gamete competition and gamete limitation in a single model of anisogamy evolution

Recent studies have revealed the importance of self-consistency in evolutionary models, particularly in the context of male–female interactions. This has been largely ignored in models of the ancestral divergence of the sexes, i.e., the evolution of anisogamy. Here, we model the evolution of anisogamy in a Fisher-consistent context, explicitly taking into account the number of interacting individuals in a typical reproductive group. We reveal an interaction between the number of adult individuals in the local mating group and the selection pressures responsible for the divergence of the sexes. The same underlying model can produce anisogamy in two different ways. Gamete competition can lead to anisogamy when it is relatively easy for gametes to find each other, but when this is more difficult and gamete competition is absent, gamete limitation can provide another route for anisogamy to evolve. In line with earlier models, organismal complexity favors anisogamy. We argue that the early contributions of Kalmus and Scudo, largely dismissed as group selectionist, are valid under certain conditions. Linking their work with the contributions of Parker helps to explain why precisely males keep producing more sperm than can ever lead to offspring: sperm could evolve to provision zygotes but this brings little profit for the effort required, because sperm would have to be equipped with provisioning ability before it is known which sperm will make it to the fertilization stage. This insight creates a logical link between paternal care under uncertain paternity (where again investment is selected against when some investment never brings about genetic benefits) and gamete size evolution.     

Species divergence and seasonal succession in rates of mate desertion in closely related Neotropical cichlid fishes

Across animal taxa, exclusive female offspring care has evolved repeatedly from biparental care, suggesting that the latter becomes evolutionarily unstable under certain conditions. Both the attributes of a species and the environment it experiences can help to predict shifts from one particular care mode to another. Nevertheless, factors inducing differences in care strategies among closely related species, or seasonal variation within species, have been subject to surprisingly little empirical testing. Here, we report the results of a field-based study that examined both among and within species variation in mate desertion in five species of closely related Nicaraguan cichlid fish in the genera Amphilophus and Amatitlania. The results show a link between female body size and male involvement in offspring care. Specifically, the larger the species the less often males were found to provide extended care. Furthermore, we found that solitary females became more common towards the end of the breeding season. We discuss the implications of this finding in the context of previous theoretical and empirical contributions regarding the frequency of offspring desertion by males.     

AIC model selection and multimodel inference in behavioral ecology: some background,observations, and comparisons

We briefly outline the information-theoretic (I-T) approaches to valid inference including a review of some simple methods for making formal inference from all the hypotheses in the model set (multimodel inference). The I-T approaches can replace the usual t tests and ANOVA tables that are so inferentially limited, but still commonly used. The I-T methods are easy to compute and understand and provide formal measures of the strength of evidence for both the null and alternative hypotheses, given the data. We give an example to highlight the importance of deriving alternative hypotheses and representing these as probability models. Fifteen technical issues are addressed to clarify various points that have appeared incorrectly in the recent literature. We offer several remarks regarding the future of empirical science and data analysis under an I-T framework.