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. (Includes Supporting information.

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. (Includes Supporting information.

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. (Includes Supplemental Materials and Reviewers\u27 Comments.

The Role of Nestling Acoustic Experience in Song Discrimination in a Sparrow

Oscine songbirds are an ideal system for investigating how early experience affects vocal behavior. Young songbirds face a challenging task: how to recognize and selectively learn only their own species’ song, often during a time-limited window. Because birds are capable of hearing birdsong very early in life, early exposure to song could plausibly affect recognition of appropriate models; however, this idea conflicts with the traditional view that song learning occurs only after a bird leaves the nest. Thus, it remains unknown whether natural variation in acoustic exposure prior to song learning affects the template for recognition. In a population where sister species, golden-crowned and white-crowned sparrows, breed syntopically, we found that nestlings discriminate between heterospecific and conspecific song playbacks prior to the onset of song memorization. We then asked whether natural exposure to more frequent or louder heterospecific song explained any variation in golden-crowned nestling response to heterospecific song playbacks. We characterized the amount of each species’ song audible in golden-crowned sparrow nests and showed that even in a relatively small area, the ratio of heterospecific to conspecific song exposure varies from 0 to 20%. However, although many songbirds hear and respond to acoustic signals before fledging, golden-crowned sparrow nestlings that heard different amounts of heterospecific song did not behave differently in response to heterospecific playbacks. This study provides the first evidence that song discrimination at the onset of song learning is robust to the presence of closely related heterospecifics in nature, which may be an important adaptation in sympatry between potentially interbreeding taxa

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 behavior 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

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

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’

Range-wide patterns of geographic variation in songs of Golden-crowned Sparrows (Zonotrichia atricapilla)

Discrete geographic variation, or dialects, in songs of songbirds arise as a consequence of complex interactions between ecology and song learning. Four of the five species of Zonotrichia sparrows, including the model species White-crowned Sparrow (Z. leucophrys), have been studied with respect to the causes and consequences of geographic variation in song. Within White-crowned Sparrows, subspecies that migrate farther have larger range size of dialects. Here, we assessed geographic patterns of song variation in the fifth species of this genus, the Golden-crowned Sparrow (Z. atricapilla). We analyzed field-recorded songs from 2 sampling periods (1996–1998 and 2006–2013) covering most of its breeding range in western North America. All songs began with a descending whistle and most songs consisted of 3–4 phrases that contained combinations of whistles, buzzes, and trills. We identified 13 discrete song types based on unique sequences of phrase types and frequency changes between phrases. Over 90% of individuals sang 1 of 5 song types, and we found clear dialect structure composed of these 5 common song types. The geographic range of dialects spanned large distances (500 to 1,700 km), resembling the geographic structure of dialects in the long-distance migrant Gambel’s White-crowned Sparrow (Z. l. gambelli), though locations of dialect boundaries differ between species. Because both Golden-crowned Sparrows and Gambel’s White-crowned Sparrows migrate similarly long distances, our study provides support to the hypothesis that dialect range size correlates with migration distance. We found little evidence of change in dialect composition in 4 populations that were sampled 15 years apart, which suggests that the dialect structure is stable across multiple generations. Our study opens the door for further comparisons to investigate links between ecology and the emergence of song dialects in this well-studied genus

Range-wide patterns of geographic variation in songs of Golden-crowned Sparrows (\u3ci\u3eZonotrichia atricapilla\u3c/i\u3e)

Discrete geographic variation, or dialects, in songs of songbirds arise as a consequence of complex interactions between ecology and song learning. Four of the five species of Zonotrichia sparrows, including the model species White-crowned Sparrow (Z. leucophrys), have been studied with respect to the causes and consequences of geographic variation in song. Within White-crowned Sparrows, subspecies that migrate farther have larger range size of dialects. Here, we assessed geographic patterns of song variation in the fifth species of this genus, the Golden-crowned Sparrow (Z. atricapilla). We analyzed field-recorded songs from 2 sampling periods (1996–1998 and 2006–2013) covering most of its breeding range in western North America. All songs began with a descending whistle and most songs consisted of 3–4 phrases that contained combinations of whistles, buzzes, and trills. We identified 13 discrete song types based on unique sequences of phrase types and frequency changes between phrases. Over 90% of individuals sang 1 of 5 song types, and we found clear dialect structure composed of these 5 common song types. The geographic range of dialects spanned large distances (500 to 1,700 km), resembling the geographic structure of dialects in the long-distance migrant Gambel’s White-crowned Sparrow (Z. l. gambelli), though locations of dialect boundaries differ between species. Because both Golden-crowned Sparrows and Gambel’s White-crowned Sparrows migrate similarly long distances, our study provides support to the hypothesis that dialect range size correlates with migration distance. We found little evidence of change in dialect composition in 4 populations that were sampled 15 years apart, which suggests that the dialect structure is stable across multiple generations. Our study opens the door for further comparisons to investigate links between ecology and the emergence of song dialects in this well-studied genus

Range-wide patterns of geographic variation in songs of Golden-crowned Sparrows (\u3ci\u3eZonotrichia atricapilla\u3c/i\u3e)

Discrete geographic variation, or dialects, in songs of songbirds arise as a consequence of complex interactions between ecology and song learning. Four of the five species of Zonotrichia sparrows, including the model species White-crowned Sparrow (Z. leucophrys), have been studied with respect to the causes and consequences of geographic variation in song. Within White-crowned Sparrows, subspecies that migrate farther have larger range size of dialects. Here, we assessedgeographic patterns of song variation in the fifth species of this genus, the Golden-crowned Sparrow (Z. atricapilla). We analyzed field-recorded songs from 2 sampling periods (1996–1998 and 2006–2013) covering most of its breeding range in western North America. All songs began with a descending whistle and most songs consisted of 3–4 phrases that contained combinations of whistles, buzzes, and trills. We identified 13 discrete song types based on unique sequences of phrase types and frequency changes between phrases. Over 90% of individuals sang 1 of 5 song types, and we found clear dialect structure composed of these 5 common song types. The geographic range of dialects spanned large distances (500 to 1,700 km), resembling the geographic structure of dialects in the long-distance migrant Gambel’s White-crowned Sparrow (Z. l. gambelli), though locations of dialect boundaries differ between species. Because both Golden-crowned Sparrows and Gambel’s White-crowned Sparrows migrate similarly long distances, our study provides support to the hypothesis that dialect range size correlates with migration distance. We found little evidence of change in dialect composition in 4 populations that were sampled 15 years apart, which suggests that the dialect structure is stable across multiple generations. Our study opens the door for further comparisons to investigate links between ecology and the emergence of song dialects in this well-studied genus. La variación geográfica discreta, o los dialectos, en los cantos de las aves canoras se originan como consecuencia de interacciones complejas entre la ecología y el aprendizaje de los cantos. Cuatro de las 5 especies de Zonotrichia, incluyendo la especie modelo Z. leucophrys, han sido estudiadas con respecto a las causas y consecuencias de la variación geográfica en el canto. Al interior de la especie Z. leucophrys, las subespecies que migran más lejos tienen dialectos con rangos más amplios. En este trabajo evaluamos los patrones de variación del canto en la quinta especie de este género, Z. atricapilla. Analizamos cantos grabados en el campo de dos períodos de muestreo (1996–1998 y 2006–2013), cubriendo la mayor parte de su rango reproductivo en el oeste de América del Norte. Todos los cantos comienzan con un silbido descendente y la mayoría de los cantos consistieron en 3–4 frases con combinaciones de silbidos, zumbidos y trinos. Identificamos 13 tipos discretos de cantos basados en secuencias únicas de tipos de frases y en cambios de frecuencia entre frases. Más del 90% de los individuos cantaron uno de los 5 tipos de cantos, y encontramos una clara estructura del dialecto compuesta por estos 5 tipos de cantos comunes. El rango geogra´fico de los dialectos abarcó grandes distancias (500 a 1,700 km), asemeja´ndose a la estructura geográfica de los dialectos de la especie migratoria de larga distancia Z. l. gambelli, aunque las localidades de frontera de los dialectos difirieron entre especies. Debido a que Z. atricapilla y Z. l. gambelli migran ambas largas distancias, nuestro estudio apoya la hipótesis de que el tamaño del rango del dialecto se correlaciona con la distancia de migración. Encontramos poca evidencia de cambios en la composición del dialecto en cuatro poblaciones que fueron muestreadas con una diferencia de 15 años, lo que sugiere que la estructura del dialecto es estable a través de varias generaciones. Nuestro estudio abre la puerta a comparaciones adicionales para investigar los vínculos entre la ecología y la emergencia de dialectos del canto en este género bien estudiado