Daizaburo Shizuka

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.

Sparrows use multiple status signals in winter social flocks

Status signals are thought to reduce the potential costs of social conflict over resources by advertising social status to other group members and reducing unnecessary contests between individuals of differing abilities. Nearly all studies of status signals to date have focused on single signalling traits, and most studies that have investigated multiple traits did not examine whether different traits are used in different contexts, as is required for them to function as multiple signals. We examined the role of gold and black crown patches of wintering golden-crowned sparrows, Zonotrichia atricapilla, in determining social dominance during experimentally staged encounters between unfamiliar individuals. Specifically, we determined whether variation in weakly correlated or uncorrelated traits (crown patch size and colour, body size) differentially affected the outcome of interactions involving avoidance versus aggression. Overall, crown patch size and colour were better predictors of dominance than were body size or sex. Several traits, including both crown features and morphological traits, predicted which individual avoided the other in dyadic interactions that did not escalate beyond avoidance. However, when dyads had similar gold crown patch sizes, the interaction was more likely to escalate, leading to aggression. In contrast to avoidance interactions, the outcomes of aggressive contests were largely predicted by variation in the colour of black crown patches. Taken together, our results show that gold and black crown features operate as multiple status signals that accommodate an escalated gradient of interactions and suggest that social status involves more complexity than can be accommodated by a single signal.

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.

The network motif architecture of dominance hierarchies

The widespread existence of dominance hierarchies has been a central puzzle in social evolution, yet we lack a framework for synthesizing the vast empirical data on hierarchy structure in animal groups. We applied network motif analysis to compare the structures of dominance networks from data published over the past 80 years. Overall patterns of dominance relations, including some aspects of non-interactions, were strikingly similar across disparate group types. For example, nearly all groups exhibited high frequencies of transitive triads, whereas cycles were very rare. Moreover, pass-along triads were rare, and double-dominant triads were common in most groups. These patterns did not vary in any systematic way across taxa, study settings (captive or wild) or group size. Two factors significantly affected network motif structure: the proportion of dyads that were observed to interact and the interaction rates of the top-ranked individuals. Thus, study design (i.e. how many interactions were observed) and the behaviour of key individuals in the group could explain much of the variations we see in social hierarchies across animals. Our findings confirm the ubiquity of dominance hierarchies across all animal systems, and demonstrate that network analysis provides new avenues for comparative analyses of social hierarchies.

Improving the reliability of molecular sexing of birds using a W-specific marker.

Molecular techniques for identifying sex of birds utilize length differences between CHD‐Z and CHD‐W introns, but in some cases these methods can lead to sexing errors. Here we show that an additional W‐specific primer can be used in conjunction with a pre‐existing sexing primer pair to dramatically improve the reliability of molecular sexing methods. We illustrate the approach with American coots (Fulica americana), a species with CHD‐Z polymorphism that could not be accurately sexed using traditional methods. We developed a reverse primer GWR2 designed to sit within the intron of the W chromosome and amplify a distinctively small DNA fragment that serves as a W‐specific marker. Analysis of known‐sex individuals indicates that this W‐specific primer provides an efficient and reliable protocol to identify the sex of F. americana. The development of such sex‐specific primers will likely increase the reliability of molecular sexing methods in other birds as well. Comparisons between CHD‐Z alleles of coots and common moorhens (Gallinula chloropus) revealed that CHD‐Z polymorphism evolved separately in these two closely related species. We discuss the implications of repeated evolution of CHD‐Z polymorphisms among birds.

Using molt limits to age Western Bluebirds

Abstract Using data on the extent of prebasic molt in known-aged adult and first-winter Western Bluebirds (Sialia mexicana), we assessed the accuracy of the molt limit as a tool for ageing birds. Sixty-nine known-aged birds were examined over the winter 2002–2003 in upper Carmel Valley, California. We also examined 29 breeding males in spring 2004 to assess the accuracy of this method for ageing birds during the breeding season. All birds were aged correctly using molt limit as the defining characteristic. We found that all first-winter birds had replaced 2–8 greater secondary coverts, while adults had replaced all wing coverts. We observed no significant sexual variation in the extent of first prebasic molt. These results indicate that the extent of prebasic molt is a highly reliable means of ageing birds in this population.

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.

Experimental confirmation that avian plumage traits function as multiple status signals in winter contests

Status signals are thought to reduce costs of overt conflict over resources by advertising social status or an individuals ability to win contests. While most studies have focused on single badges of status, recent empirical work has shown that multiple status signals may exist. To provide robust evidence for multiple badges of status, an experimental manipulation is required to decouple signals from one another and from other traits linked to fighting ability. Such experimental evidence is lacking for most studies of multiple status signals to date. We previously found that two plumage traits in golden-crowned sparrows, Zonotrichia atricapilla, were correlated with social dominance in encounters between unfamiliar individuals. To confirm that each plumage patch functions as an independent status signal, we experimentally augmented the sizes of the gold crown patch and the black crown patch during encounters between unfamiliar individuals with similar premanipulation crown sizes. In nearly all cases, the individual with the artificially augmented gold or black crown was dominant during the trial and manipulations of each colour were equally successful in conferring dominance. The relative differences in crown sizes between manipulated and unmanipulated individuals in a dyad and mismatches in crown sizes of the manipulated bird led to escalation in gold trials, but these same factors were not significant for black trials. This study provides unequivocal evidence for multiple status signals: both black and gold crown patches influence social status per se and they do so independently of the other crown patch.

Measuring the robustness of network community structure using assortativity

The existence of discrete social clusters, or ‘communities’, is a common feature of social networks in human and nonhuman animals. The level of such community structure in networks is typically measured using an index of modularity, Q. While modularity quantifies the degree to which individuals associate within versus between social communities and provides a useful measure of structure in the social network, it assumes that the network has been well sampled. However, animal social network data is typically subject to sampling errors. In particular, the associations among individuals are often not sampled equally, and animal social network studies are often based on a relatively small set of observations. Here, we extend an existing framework for bootstrapping network metrics to provide a method for assessing the robustness of community assignment in social networks using a metric we call community assortativity (rcom). We use simulations to demonstrate that modularity can reliably detect the transition from random to structured associations in networks that differ in size and number of communities, while community assortativity accurately measures the level of confidence based on the detectability of associations. We then demonstrate the use of these metrics using three publicly available data sets of avian social networks. We suggest that by explicitly addressing the known limitations in sampling animal social network, this approach will facilitate more rigorous analyses of population-level structural patterns across social 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.