Loss of flockmates weakens winter site fidelity in golden-crowned sparrows (Zonotrichia atricapilla).

Animal social interactions have an intrinsic spatial basis as many of these interactions occur in spatial proximity. This presents a dilemma when determining causality: Do individuals interact socially because they happen to share space, or do they share space because they are socially linked? We present a method that uses demographic turnover events as a natural experiment to investigate the links between social associations and space use in the context of interannual winter site fidelity in a migratory bird. We previously found that golden-crowned sparrows (Zonotrichia atricapilla) show consistent flocking relationships across years, and that familiarity between individuals influences the dynamics of social competition over resources. Using long-term data on winter social and spatial behavior across 10 y, we show that i) sparrows exhibit interannual fidelity to winter home ranges on the scale of tens of meters and ii) the precision of interannual site fidelity increases with the number of winters spent, but iii) this fidelity is weakened when sparrows lose close flockmates from the previous year. Furthermore, the effect of flockmate loss on site fidelity was higher for birds that had returned in more than 2 winters, suggesting that social fidelity may play an increasingly important role on spatial behavior across the lifetime of this migratory bird. Our study provides evidence that social relationships can influence site fidelity, and shows the potential of long-term studies for disentangling the relationship between social and spatial behavior.

Ecogeography of group size suggests differences in drivers of sociality among cooperatively breeding fairywrens.

Cooperatively breeding species exhibit a range of social behaviours associated with different costs and benefits to group living, often in association with different environmental conditions. For example, recent phylogenetic studies have collectively shown that the evolution and distribution of cooperative breeding behaviour is related to the environment. However, little is known about how environmental variation may drive differences in social systems across populations within species, and how the relationship between environmental conditions and sociality may differ across species. Here, we examine variation in social group size along a steep environmental gradient for two congeneric cooperatively breeding species of fairywrens (Maluridae) and show that they exhibit opposing ecogeographic patterns. Purple-backed fairywrens, a species in which helpers increase group productivity, have larger groups in hot, dry environments and smaller groups in cool, wet environments. By contrast, superb fairywrens, a species with helpers that do not increase group productivity despite the presence of alloparental care, exhibit the opposite trend. We suggest differences in the costs and benefits of sociality contribute to these opposing ecogeographical patterns and demonstrate that comparisons of intraspecific patterns of social variation across species can provide insight into how ecology shapes social systems.

Extreme offspring ornamentation in American coots is favored by selection within families, not benefits to conspecific brood parasites.

Offspring ornamentation typically occurs in taxa with parental care, suggesting that selection arising from social interactions between parents and offspring may underlie signal evolution. American coot babies are among the most ornamented offspring found in nature, sporting vividly orange-red natal plumage, a bright red beak, and other red parts around the face and pate. Previous plumage manipulation experiments showed that ornamented plumage is favored by strong parental choice for chicks with more extreme ornamentation but left unresolved the question as to why parents show the preference. Here we explore natural patterns of variation in coot chick plumage color, both within and between families, to understand the context of parental preference and to determine whose fitness interests are served by the ornamentation. Conspecific brood parasitism is common in coots and brood parasitic chicks could manipulate hosts by tapping into parental choice for ornamented chicks. However, counter to expectation, parasitic chicks were duller (less red) than nonparasitic chicks. This pattern is explained by color variation within families: Chick coloration increases with position in the egg-laying order, but parasitic eggs are usually the first eggs a female lays. Maternal effects influence chick coloration, but coot females do not use this mechanism to benefit the chicks they lay as parasites. However, within families, chick coloration predicts whether chicks become "favorites" when parents begin control over food distribution, implicating a role for the chick ornamentation in the parental life-history strategy, perhaps as a reliable signal of a chick's size or age.

The ecology of wealth inequality in animal societies.

Individuals vary in their access to resources, social connections and phenotypic traits, and a central goal of evolutionary biology is to understand how this variation arises and influences fitness. Parallel research on humans has focused on the causes and consequences of variation in material possessions, opportunity and health. Central to both fields of study is that unequal distribution of wealth is an important component of social structure that drives variation in relevant outcomes. Here, we advance a research framework and agenda for studying wealth inequality within an ecological and evolutionary context. This ecology of inequality approach presents the opportunity to reintegrate key evolutionary concepts as different dimensions of the link between wealth and fitness by (i) developing measures of wealth and inequality as taxonomically broad features of societies, (ii) considering how feedback loops link inequality to individual and societal outcomes, (iii) exploring the ecological and evolutionary underpinnings of what makes some societies more unequal than others, and (iv) studying the long-term dynamics of inequality as a central component of social evolution. We hope that this framework will facilitate a cohesive understanding of inequality as a widespread biological phenomenon and clarify the role of social systems as central to evolutionary biology.

Flock-species richness influences node importance and modularity in mixed-species flock networks.

Interdependencies in social groups of animals are a combination of multiple pairwise interactions. Heterospecific groups are often characterized by important species that contribute more to group initiation, maintenance or function than other species. However, in large heterospecific groups, many pairwise interactions are not realised, while others may not be biologically significant, confounding inferences about species importance. Hence, in this study, we examine context dependent changes in species importance and assortment in mixed-species bird flocks from a tropical field site in Southern India using social network analysis. Specifically, we ask how the structural importance of a species and the clustering patterns of species relationships depends on species richness in mixed-species flocks. We constructed both raw and filtered networks; while our results are largely correlated, we believe that filtered networks can provide insights into community-level importance of species in mixed-flocks while raw networks depict flock-level patterns. We find significant differences in flocks of different richness in that different species emerge as structurally important across flocks of varying richness. We also find that assortment is higher in two-species flocks and decreases with an increase in the number of species in the flock ('flock richness' hereafter). We argue that the link between structural importance of species in mixed-species flock networks and their functional significance in the community critically depends on the social context: namely, the species richness of the mixed-species flock. We propose that examining species structural importance at different flock-richness values provides insights into biologically meaningful functional roles of species. More generally, we suggest that it is important to consider context when interpreting species centrality and importance in network structure.

Juvenile rank acquisition is associated with fitness independent of adult rank.

Social rank is a significant determinant of fitness in a variety of species. The importance of social rank suggests that the process by which juveniles come to establish their position in the social hierarchy is a critical component of development. Here, we use the highly predictable process of rank acquisition in spotted hyenas to study the consequences of variation in rank acquisition in early life. In spotted hyenas, rank is 'inherited' through a learning process called 'maternal rank inheritance.' This pattern is very consistent: approximately 80% of juveniles acquire the exact rank expected under the rules of maternal rank inheritance. The predictable nature of rank acquisition in these societies allows the process of rank acquisition to be studied independently from the ultimate rank that each juvenile attains. In this study, we use Elo-deviance scores, a novel application of the Elo-rating method, to calculate each juvenile's deviation from the expected pattern of maternal rank inheritance during development. Despite variability in rank acquisition among juveniles, most of these juveniles come to attain the exact rank expected of them according to the rules of maternal rank inheritance. Nevertheless, we find that transient variation in rank acquisition in early life is associated with long-term fitness consequences for these individuals: juveniles 'underperforming' their expected ranks show reduced survival and lower lifetime reproductive success than better-performing peers, and this relationship is independent of both maternal rank and rank achieved in adulthood. We also find that multiple sources of early life adversity have cumulative, but not compounding, effects on fitness. Future work is needed to determine if variation in rank acquisition directly affects fitness, or if some other variable, such as maternal investment or juvenile condition, causes variation in both of these outcomes.

Manipulating badges of status only fools strangers.

Conflict is risky, but mechanisms that allow animals to assess dominance status without aggression can reduce such costs. Two different mechanisms of competitor assessment are expected to evolve in different contexts: badges of status are expected in larger, anonymous groups, whereas individual recognition is feasible in small, stable groups. However, both mechanisms may be important when social interactions occur both within and across stable social groups. We manipulated plumage in golden-crowned sparrows (Zonotrichia atricapilla) and found that two known badges of status - gold and black head plumage patch sizes - independently affect dominance among strangers but manipulations had no effect on dominance among familiar flockmates. Moreover, familiar flockmates showed less aggression and increased foraging relative to strangers. Our study provides clear experimental evidence that social recognition affects badge function, and suggests that variation in social contexts maintains coexistence and context-dependent use of these two dominance resolution mechanisms.

Coots use hatch order to learn to recognize and reject conspecific brood parasitic chicks.

Avian brood parasites and their hosts provide model systems for investigating links between recognition, learning, and their fitness consequences. One major evolutionary puzzle has continued to capture the attention of naturalists for centuries: why do hosts of brood parasites generally fail to recognize parasitic offspring after they have hatched from the egg, even when the host and parasitic chicks differ to almost comic degrees? One prominent theory to explain this pattern proposes that the costs of mistakenly learning to recognize the wrong offspring make recognition maladaptive. Here we show that American coots, Fulica americana, can recognize and reject parasitic chicks in their brood by using learned cues, despite the fact that the hosts and the brood parasites are of the same species. A series of chick cross-fostering experiments confirm that coots use first-hatched chicks in a brood as referents to learn to recognize their own chicks and then discriminate against later-hatched parasitic chicks in the same brood. When experimentally provided with the wrong reference chicks, coots can be induced to discriminate against their own offspring, confirming that the learning errors proposed by theory can exist. However, learning based on hatching order is reliable in naturally parasitized coot nests because host eggs hatch predictably ahead of parasite eggs. Conversely, a lack of reliable information may help to explain why the evolution of chick recognition is not more common in hosts of most interspecific brood parasites.

Multimodal signalling in the North American barn swallow: a phenotype network approach.

Complex signals, involving multiple components within and across modalities, are common in animal communication. However, decomposing complex signals into traits and their interactions remains a fundamental challenge for studies of phenotype evolution. We apply a novel phenotype network approach for studying complex signal evolution in the North American barn swallow (Hirundo rustica erythrogaster). We integrate model testing with correlation-based phenotype networks to infer the contributions of female mate choice and male-male competition to the evolution of barn swallow communication. Overall, the best predictors of mate choice were distinct from those for competition, while moderate functional overlap suggests males and females use some of the same traits to assess potential mates and rivals. We interpret model results in the context of a network of traits, and suggest this approach allows researchers a more nuanced view of trait clustering patterns that informs new hypotheses about the evolution of communication systems.

Social network structure in wintering golden-crowned sparrows is not correlated with kinship.

Stable social organization in a wide variety of organisms has been linked to kinship, which can minimize conflict due to the indirect fitness benefits from cooperating with relatives. In birds, kin selection has been mostly studied in the context of reproduction or in species that are social year round. Many birds however are migratory, and the role of kinship in the winter societies of these species is virtually unexplored. In a previous study, we discovered striking social complexity and stability in a wintering population of migratory golden-crowned sparrows (Zonotrichia atricapilla) - individuals repeatedly form close associations with the same social partners, including across multiple winters. Here, we test the possibility that kinship might be involved in these close and stable social affiliations. We examine the relationship between kinship and social structure for two of the consecutive wintering seasons from the previous study. We found no evidence that social structure was influenced by kinship. Relatedness between most pairs of individuals was at most that of first cousins (and mostly far lower). Genetic networks based on relatedness do not correspond to the social networks, and Mantel tests revealed no relationship between kinship and pairwise interaction frequency. Kinship also failed to predict social structure in more fine-grained analyses, including analyses of each sex separately (in the event that sex-biased migration might limit kin selection to one sex), and separate analyses for each social community. The complex winter societies of golden-crowned sparrows appear to be based on cooperative benefits unrelated to kin selection.

Across-year social stability shapes network structure in wintering migrant sparrows.

Migratory birds often form flocks on their wintering grounds, but important details of social structure such as the patterns of association between individuals are virtually unknown. We analysed networks of co-membership in short-term flocks for wintering golden-crowned sparrows (Zonotrichia atricapilla) across three years and discovered social complexity unsuspected for migratory songbirds. The population was consistently clustered into distinct social communities within a relatively small area (~ 7 ha). Birds returned to the same community across years, with mortality and recruitment leading to some degree of turnover in membership. These spatiotemporal patterns were explained by the combination of space use and social preference - birds that flocked together in one year flocked together again in the subsequent year more often than were expected based on degrees of home range overlap. Our results suggest that a surprising level of social fidelity across years leads to repeatable patterns of social network structure in migratory populations.

Winter GPS tagging reveals home ranges during the breeding season for a boreal-nesting migrant songbird, the Golden-crowned Sparrow.

Determining space use for species is fundamental to understanding their ecology, and tracking animals can reveal insights into their spatial ecology on home ranges and territories. Recent technological advances have led to GPS-tracking devices light enough for birds as small as ~30 g, creating novel opportunities to remotely monitor fine-scale movements and space use for these smaller species. We tested whether miniaturized GPS tags can allow us to understand space use of migratory birds away from their capture sites and sought to understand both pre-breeding space use as well as territory and habitat use on the breeding grounds. We used GPS tags to characterize home ranges on the breeding grounds for a migratory songbird with limited available breeding information, the Golden-crowned Sparrow (Zonotrichia atricapilla). Using GPS points from 23 individuals across 26 tags (three birds tagged twice), we found home ranges in Alaska and British Columbia were on average 44.1 ha (95% kernel density estimate). In addition, estimates of territory sizes based on field observations (mean 2.1 ha, 95% minimum convex polygon [MCP]) were three times smaller than 95% MCPs created using GPS tags (mean 6.5 ha). Home ranges included a variety of land cover classes, with shrubland particularly dominant (64-100% of home range cover for all but one bird). Three birds tracked twice returned to the same breeding area each year, supporting high breeding site fidelity for this species. We found reverse spring migration for five birds that flew up to 154 km past breeding destinations before returning. GPS-tracking technology allowed for critical ecological insights into this migratory species that breeds in very remote locations.

Family dynamics through time: brood reduction followed by parental compensation with aggression and favouritism.

Parental food allocation in birds has long been a focal point for life history and parent-offspring conflict theories. In asynchronously hatching species, parents are thought to either adjust brood size through death of marginal offspring (brood reduction), or feed the disadvantaged chicks to reduce the competitive hierarchy (parental compensation). Here, we show that parent American coots (Fulica americana) practice both strategies by switching from brood reduction to compensation across time. Late-hatching chicks suffer higher mortality only for the first few days after hatching. Later, parents begin to exhibit parental aggression towards older chicks and each parent favours a single chick, both of which are typically the youngest of the surviving offspring. The late-hatched survivors can equal or exceed their older siblings in size prior to independence. A mixed allocation strategy allows parents to compensate for the costs of competitive hierarchies while gaining the benefits of hatching asynchrony.

Acorn woodpecker movements and social networks change with wildfire smoke.

Climate change has contributed to increased wildfires1,2. Wildfire smoke exposes wildlife to hazards and mortality from particulate matter on a scale larger than the area impacted by fire3,4. Using automated radiotelemetry, we illustrate how smoky conditions are associated with changes in behavior of acorn woodpeckers (Melanerpes formicivorus), a flagship species of oak (Quercus spp.) savannas of western North America. On smoky days, birds spent more time at their home territory and reduced visitation to others, especially to distant territories. Associations between birds decreased, and individuals were less assorted by group in co-visitation networks, suggesting less inter-individual coordination on smoky days. We show that between 2016 and 2020, ∼14% of the acorn woodpecker population in the US experienced fire, potentially exposing on average 89.42% of the range to atmospheric smoke annually. These findings highlight how potential effects of smoke on animal behavior may be widespread and exacerbate negative impacts of increasingly common "megafires", even in fire-adapted ecosystems.

The dynamics of dominance: open questions, challenges and solutions.

Although social hierarchies are recognized as dynamic systems, they are typically treated as static entities for practical reasons. Here, we ask what we can learn from a dynamical view of dominance, and provide a research agenda for the next decades. We identify five broad questions at the individual, dyadic and group levels, exploring the causes and consequences of individual changes in rank, the dynamics underlying dyadic dominance relationships, and the origins and impacts of social instability. Although challenges remain, we propose avenues for overcoming them. We suggest distinguishing between different types of social mobility to provide conceptual clarity about hierarchy dynamics at the individual level, and emphasize the need to explore how these dynamic processes produce dominance trajectories over individual lifespans and impact selection on status-seeking behaviour. At the dyadic level, there is scope for deeper exploration of decision-making processes leading to observed interactions, and how stable but malleable relationships emerge from these interactions. Across scales, model systems where rank is manipulable will be extremely useful for testing hypotheses about dominance dynamics. Long-term individual-based studies will also be critical for understanding the impact of rare events, and for interrogating dynamics that unfold over lifetimes and generations. This article is part of the theme issue 'The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies'.

The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies.

A century ago, foundational work by Thorleif Schjelderup-Ebbe described a 'pecking order' in chicken societies, where individuals could be ordered according to their ability to exert their influence over their group-mates. Now known as dominance hierarchies, these structures have been shown to influence a plethora of individual characteristics and outcomes, situating dominance research as a pillar of the study of modern social ecology and evolution. Here, we first review some of the major questions that have been answered about dominance hierarchies in the last 100 years. Next, we introduce the contributions to this theme issue and summarize how they provide ongoing insight in the epistemology, physiology and neurobiology, hierarchical structure, and dynamics of dominance. These contributions employ the full range of research approaches available to modern biologists. Cross-cutting themes emerging from these contributions include a focus on cognitive underpinnings of dominance, the application of network-analytical approaches, and the utility of experimental rank manipulations for revealing causal relationships. Reflection on the last 100 years of dominance research reveals how Schjelderup-Ebbe's early ideas and the subsequent research helped drive a shift from an essentialist view of species characteristics to the modern recognition of rich inter-individual variation in social, behavioural and physiological phenotypes. This article is part of the theme issue 'The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies'.

DomArchive: a century of published dominance data.

Dominance behaviours have been collected for many groups of animals since 1922 and serve as a foundation for research on social behaviour and social structure. Despite a wealth of data from the last century of research on dominance hierarchies, these data are only rarely used for comparative insight. Here, we aim to facilitate comparative studies of the structure and function of dominance hierarchies by compiling published dominance interaction datasets from the last 100 years of work. This compiled archive includes 436 datasets from 190 studies of 367 unique groups (mean group size 13.8, s.d. = 13.4) of 135 different species, totalling over 243 000 interactions. These data are presented in an R package alongside relevant metadata and a tool for subsetting the archive based on biological or methodological criteria. In this paper, we explain how to use the archive, discuss potential limitations of the data, and reflect on best practices in publishing dominance data based on our experience in assembling this dataset. This archive will serve as an important resource for future comparative studies and will promote the development of general unifying theories of dominance in behavioural ecology that can be grounded in testing with empirical data. This article is part of the theme issue 'The centennial of the pecking order: current state and future prospects for the study of dominance hierarchies'.

To accept or reject heterospecific mates: behavioural decisions underlying premating isolation.

Premating isolation in animals involves decision-making processes that affect whether individuals accept or reject heterospecific mates. An integrative understanding of the behavioural processes underlying heterospecific acceptance can clarify the conditions under which premating isolation evolves. As an illustration, we review how Reeve's (Reeve 1989 Am. Nat.133, 407-435. (doi:10.1086/284926)) acceptance threshold model can help make sense of patterns of premating isolation in nature. This model derives a threshold trait value for acceptance for rejection of recipients of an action (e.g. mating) based on the fitness consequences of these decisions. We show that the maintenance of partial reproductive isolation can be an outcome of optimal acceptance thresholds, even in the face of reinforcement. We also use this model to clarify how the composition of multispecies communities can shape premating isolation. The acceptance threshold model can also be viewed as the behavioural underpinning of reproductive character displacement and cascading reinforcement. Finally, we highlight potential limitations of the acceptance threshold model with respect to investigating the role of sexual selection in speciation, and we propose that integration of behavioural models in speciation research will help us gain a full picture of the mechanisms underlying premating isolation. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.

How to learn to recognize conspecific brood parasitic offspring.

Recognition systems evolve to reduce the risk and costs of making recognition errors. Two main sources of recognition error include perceptual error (error arising from inability to discriminate between objects) and template error (error arising from using the wrong recognition template). We focus on how template error shapes host defence against avian brood parasites. Prior experiments in American coots (Fulica americana), a conspecific brood parasite, demonstrated how hosts learn to recognize brood parasitic chicks using predictable patterns of hatching order of host and parasite eggs. Here, we use these results to quantify the benefit of chick rejection as well as the cost of template error, and we then use mathematical models to explore fitness payoffs of chick recognition from different template acquisition mechanisms. We find that fitness differences between mechanisms do not fully explain aspects of the learning mechanism, such as why coots reacquire their recognition template each year. Other constraints arising from mating systems and genetic mechanisms likely influence which learning mechanism for parasitic chick recognition is optimal. Our approach highlights how mechanisms of template acquisition influence other recognition systems, including parasitic chick recognition in other brood parasite hosts. This article is part of the theme issue 'Signal detection theory in recognition systems: from evolving models to experimental tests'.