Aggregation as a cost-reducing strategy for lycaenid larvae

If a mutualistic relationship entails providing services ata cost, selection will favor individuals that maximize the netbenefits of the interaction and minimize the costs. Larvae ofmany species of lycaenid butterflies secrete nutritious foodrewards to attending ants and, in return, receive protectionagainst predators and parasitoids. Because ants typically recruitmore workers to larger resources, by forming groups the larvaemay ensure more reliable access to ants and thereby gain betterprotection. A further consequence of aggregating should be achange of the cost-benefit relationship for individual larvae.The larger the group, the smaller a single larva's influencewill be on total ant density, which could lead to a smallerinvestment in secretion, thus reducing the per capita cost ofcooperation. In this study, die influence of ant attendance,group size, and companion quality on larval investment was investigated.The interaction between the obligately ant-dependent lycaenid,Jalmanus evagoras, and its attendant Iridomyrmax ants was manipulatedand the effect on larval secretion measured. As the level ofant attendance increased, the delivery of food rewards increased,bodi for solitary and for aggregated larvae. When aggregated,larvae provided less food rewards to ants dun when solitary,and secretion rate decreased with increasing group size. Furthermore,larvae had lower secretion rates when paired with a bigger,more attractive larva than when paired with a smaller one. Theconsiderable reduction in secretion rates for larvae in groupssuggests that gaining protection at a lower secretion cost couldbe one factor that promotes aggregation in myrmecophilous lycaenids.     

How ecology shapes exploitation: a framework to predict the behavioural response of human and animal foragers along exploration–exploitation trade‐offs

Understanding how humans and other animals behave in response to changes in their environments is vital for predicting population dynamics and the trajectory of coupled social‐ecological systems. Here, we present a novel framework for identifying emergent social behaviours in foragers (including humans engaged in fishing or hunting) in predator–prey contexts based on the exploration difficulty and exploitation potential of a renewable natural resource. A qualitative framework is introduced that predicts when foragers should behave territorially, search collectively, act independently or switch among these states. To validate it, we derived quantitative predictions from two models of different structure: a generic mathematical model, and a lattice‐based evolutionary model emphasising exploitation and exclusion costs. These models independently identified that the exploration difficulty and exploitation potential of the natural resource controls the social behaviour of resource exploiters. Our theoretical predictions were finally compared to a diverse set of empirical cases focusing on fisheries and aquatic organisms across a range of taxa, substantiating the framework's predictions. Understanding social behaviour for given social‐ecological characteristics has important implications, particularly for the design of governance structures and regulations to move exploited systems, such as fisheries, towards sustainability. Our framework provides concrete steps in this direction.     

Cooperative nest‐defence behaviour and territory quality in a resident and socially monogamous passerine

Nest defence is a fundamental aspect of parental care in secondary cavity‐nesting birds, and predation or competition for nesting sites can involve different defensive behaviours. Because habitat quality determines breeding success, we were interested in whether breeding pairs of the Eurasian nuthatch, Sitta europaea, established in more favourable environment also manifest higher probability of cooperative behaviour during their nest‐site defence. To explore this relationship, we quantified behavioural displays of both parents and analysed activity budget ethogram data from simulated territorial intrusions performed in the chick‐feeding phase with one conspecific and two different heterospecific stimuli (dummies of nuthatch, starling and woodpecker). We found that paired individuals shared their roles during nest‐site defence to a considerable extent. Males had a significantly higher number of attacks on intruders than females, and females performed more threat displays and controls of the brood than males. Multinomial analysis of the cooperative behaviour suggested that pairs in a high‐quality territory had higher probability of reciprocal substitution of different roles towards a balance between attacks, threat displays and nest controls. Contrary to this, pairs in a low‐quality territory had less likely pairwise combinations of simultaneous behavioural states that are associated with effective nest‐site defence. The difference in response probability according to territory quality was, however, highly variable in view of the stimulus that was used in simulated territorial intrusion. Because individual roles and the complex behavioural repertoire of pairs altered in response to territory quality and potential nest‐site competitor or brood predator, our results suggest that the cooperative nest‐defence behaviour could be linked to the breeding success of this year‐round territorial species living in a heterogeneous forest habitat.     

Associations between glucocorticoids and sociality across a continuum of vertebrate social behavior

The causes and consequences of individual differences in animal behavior and stress physiology are increasingly studied in wild animals, yet the possibility that stress physiology underlies individual variation in social behavior has received less attention. In this review, we bring together these study areas and focus on understanding how the activity of the vertebrate neuroendocrine stress axis (HPA‐axis) may underlie individual differences in social behavior in wild animals. We first describe a continuum of vertebrate social behaviors spanning from initial social tendencies (proactive behavior) to social behavior occurring in reproductive contexts (parental care, sexual pair‐bonding) and lastly to social behavior occurring in nonreproductive contexts (nonsexual bonding, group‐level cooperation). We then perform a qualitative review of existing literature to address the correlative and causal association between measures of HPA‐axis activity (glucocorticoid levels or GCs) and each of these types of social behavior. As expected, elevated HPA‐axis activity can inhibit social behavior associated with initial social tendencies (approaching conspecifics) and reproduction. However, elevated HPA‐axis activity may also enhance more elaborate social behavior outside of reproductive contexts, such as alloparental care behavior. In addition, the effect of GCs on social behavior can depend upon the sociality of the stressor (cause of increase in GCs) and the severity of stress (extent of increase in GCs). Our review shows that the while the associations between stress responses and sociality are diverse, the role of HPA‐axis activity behind social behavior may shift toward more facilitating and less inhibiting in more social species, providing insight into how stress physiology and social systems may co‐evolve.     

Simple foraging rules in competitive environments can generate socially structured populations

Social vertebrates commonly form foraging groups whose members repeatedly interact with one another and are often genetically related. Many species also exhibit within‐population specializations, which can range from preferences to forage in particular areas through to specializing on the type of prey they catch. However, within‐population structure in foraging groups, behavioral homogeneity in foraging behavior, and relatedness could be outcomes of behavioral interactions rather than underlying drivers. We present a simple process by which grouping among foragers emerges and is maintained across generations. We introduce agent‐based models to investigate (1) whether a simple rule (keep foraging with the same individuals when you were successful) leads to stable social community structure, and (2) whether this structure is robust to demographic changes and becomes kin‐structured over time. We find the rapid emergence of kin‐structured populations and the presence of foraging groups that control, or specialize on, a particular food resource. This pattern is strongest in small populations, mirroring empirical observations. Our results suggest that group stability can emerge as a product of network self‐organization and, in doing so, may provide the necessary conditions for the evolution of more sophisticated processes, such as social learning. This taxonomically general social process has implications for our understanding of the links between population, genetic, and social structures.     

Cheating on Cheaters Stabilizes Cooperation in Pseudomonas aeruginosa

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Stimulation of dopamine D1 receptor improves learning capacity in cooperating cleaner fish

Accurate contextual decision-making strategies are important in social environments. Specific areas in the brain are tasked to process these complex interactions and generate correct follow-up responses. The dorsolateral and dorsomedial parts of the telencephalon in the teleost fish brain are neural substrates modulated by the neurotransmitter dopamine (DA), and are part of an important neural circuitry that drives animal behaviour from the most basic actions such as learning to search for food, to properly choosing partners and managing decisions based on context. The Indo-Pacific cleaner wrasse Labroides dimidiatus is a highly social teleost fish species with a complex network of interactions with its ‘client’ reef fish. We asked if changes in DA signalling would affect individual learning ability by presenting cleaner fish two ecologically different tasks that simulated a natural situation requiring accurate decision-making. We demonstrate that there is an involvement of the DA system and D₁ receptor pathways on cleaners'' natural abilities to learn both tasks. Our results add significantly to the growing literature on the physiological mechanisms that underlie and facilitate the expression of cooperative abilities.     

Cooperation‐mediated plasticity in dispersal and colonization

Kin selection theory predicts that costly cooperative behaviors evolve most readily when directed toward kin. Dispersal plays a controversial role in the evolution of cooperation: dispersal decreases local population relatedness and thus opposes the evolution of cooperation, but limited dispersal increases kin competition and can negate the benefits of cooperation. Theoretical work has suggested that plasticity of dispersal, where individuals can adjust their dispersal decisions according to the social context, might help resolve this paradox and promote the evolution of cooperation. Here, we experimentally tested the hypothesis that conditional dispersal decisions are mediated by a cooperative strategy: we quantified the density‐dependent dispersal decisions and subsequent colonization efficiency from single cells or groups of cells among six genetic strains of the unicellular Tetrahymena thermophila that differ in their aggregation level (high, medium, and low), a behavior associated with cooperation strategy. We found that the plastic reaction norms of dispersal rate relative to density differed according to aggregation level: highly aggregative genotypes showed negative density‐dependent dispersal, whereas low‐aggregation genotypes showed maximum dispersal rates at intermediate density, and medium‐aggregation genotypes showed density‐independent dispersal with intermediate dispersal rate. Dispersers from highly aggregative genotypes had specialized long‐distance dispersal phenotypes, contrary to low‐aggregation genotypes; medium‐aggregation genotypes showing intermediate dispersal phenotype. Moreover, highly aggregation genotypes showed evidence for beneficial kin‐cooperation during dispersal. Our experimental results should help to resolve the evolutionary conflict between cooperation and dispersal: cooperative individuals are expected to avoid kin‐competition by dispersing long distances, but maintain the benefits of cooperation by dispersing in small groups.     

Is the hook of muroid rodent's sperm related to sperm train formation?

Competition between spermatozoa of rival males to gain fertilizations has led to a wide array of modifications in sperm structure and function. Sperm cells of most muroid rodents have hook‐shaped extensions in the apical–ventral tip of the head, but the function of this structure is largely unknown. These ‘hooks’ may facilitate aggregation of spermatozoa in so‐called ‘trains’, as an adaptation to sperm competition, because sperm in trains may swim faster than free‐swimming cells. However, there is controversy regarding the role of the hook in train formation, and in relation to whether it is selected by sperm competition. We examined spermatozoa from muroid rodents with varying levels of sperm competition to assess whether (i) sperm aggregates are common in these taxa, (ii) presence of a hook relates to the formation of sperm aggregations, and (iii) formation of sperm aggregations is explained by sperm competition. Our analyses in 25 muroid species revealed that > 92% of spermatozoa swim individually in all species, with the exception of the wood mouse, Apodemus sylvaticus, which has ~50% spermatozoa swimming freely. Species with hooked spermatozoa had higher sperm competition levels and longer sperm than species whose sperm lack a hook. Neither the presence of hook nor sperm competition levels were related to the percentage of sperm in aggregations. Thus, (i) sperm aggregates in muroid rodents are an exceptional trait found only in a few species, (ii) evolution of the sperm hook is associated to sperm competition levels, but (iii) the hook is unlikely to be related to the formation of sperm aggregates. The evolutionary significance of the sperm head hook thus remains elusive, and future studies should examine potential roles of this pervasive structure in sperm's hydrodynamic efficiency and sperm–female tract interactions.