Effects of mistletoe (Phoradendron villosum) on California oaks.

Mistletoes are a widespread group of plants often considered to be hemiparasitic, having detrimental effects on growth and survival of their hosts. We studied the effects of the Pacific mistletoe, Phoradendron villosum, a member of a largely autotrophic genus, on three species of deciduous California oaks. We found no effects of mistletoe presence on radial growth or survivorship and detected a significant positive relationship between mistletoe and acorn production. This latter result is potentially explained by the tendency of P. villosum to be present on larger trees growing in nitrogen-rich soils or, alternatively, by a preference for healthy, acorn-producing trees by birds that potentially disperse mistletoe. Our results indicate that the negative consequences of Phoradendron presence on their hosts are negligible-this species resembles an epiphyte more than a parasite-and outweighed by the important ecosystem services mistletoe provides.

Inclusive fitness theory and eusociality.

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. Nowak et al. argue that inclusive fitness theory has been of little value in explaining the natural world, and that it has led to negligible progress in explaining the evolution of eusociality. However, we believe that their arguments are based upon a misunderstanding of evolutionary theory and a misrepresentation of the empirical literature. We will focus our comments on three general issues.

Age-based discrimination of rival males in western bluebirds.

Adaptive social behavior frequently involves discriminating between classes of individuals such as relatives versus non-relatives, older versus younger individuals, or individuals of different status. In the absence of spatial cues, this discrimination may be based on signals that correlate with fitness-related traits (e.g., older or high-status males may sing higher performance songs) or with identity, for example, when receivers distinguish and classify signalers based on their unique signal structure. Here, we examine vocal age-based discrimination in western bluebirds (Sialia mexicana), a North American songbird in which older males have a significant advantage in obtaining extra-pair fertilizations, and therefore pose a significantly higher threat to paternity than younger males. We asked whether western bluebird males showed a higher response to playback of songs of older males compared to younger males relative to their own age. We prepared song stimuli by removing three potential signals of age that have been identified as important in other species: (1) note consistency (which was achieved by playing a single instance of each note repeatedly), (2) note repertoire size, and (3) singing rate (the latter two were equalized across conditions). Even in the absence of these potential signals of age, young males responded more strongly to playback of older males' songs than to young males' songs, suggesting that they are able to discriminate between age classes relative to the threat they pose. Further research is required to determine whether this discrimination is based on individual recognition or signal features that are correlated with age.

Climatic constraints on wintering bird distributions are modified by urbanization and weather.

1. Ecologists have long been interested in the role of climate in shaping species' ranges, and in recent years, this relationship has taken on greater significance because of the need for accurate predictions of the effects of climate change on wildlife populations. Bioclimatic relationships, however, are potentially complicated by various environmental factors operating at multiple spatial and temporal scales. Here, we test the hypothesis that climatic constraints on bird distributions are modified by species-specific responses to weather, urbanization and use of supplemental food. 2. Our analyses focused on 18 bird species with data from over 3000 sites across the north-eastern United States and adjacent Canadian provinces. We use hierarchal occupancy modelling to quantify the effects of short-term weather variation and surrounding urbanization on food stress and probabilities of detection, and how these fine-scale changes modify the role that climate has on the distributions of wintering bird populations at regional scales. 3. Examining site occupancy and supplemental food use across the study region, we found that average minimum temperature was an important factor limiting bird distributions, supporting the hypothesis that the occupancy of wintering birds is limited by climatic constraints. We found that 15 of 18 species (83%) were more energetically stressed (had a higher likelihood of visiting a feeder station) as minimum temperature declined from the seasonal average. Because we found these patterns in populations that regularly visit supplemental food sites and were likely not food-limited, we suggest that resource availability is less important than climate in constraining wintering bird distributions. Across a winter season, local within-winter extinction probabilities were lower and colonization probabilities higher at warmer sites supporting the role of climate-mediated range shifts. Importantly, however, these relationships were modified by the degree of urbanization and species' abilities to persist in human-modified landscapes. 4. Our results suggest that urbanization and behavioural adaptation can modify the role of climate on bird ranges and should be included in future analyses of range shifts because of climate change.

Ecological determinants of American crow mortality due to West Nile virus during its North American sweep.

We examined the ecological factors influencing population declines in American crows (Corvus brachyrhynchos) as they were initially exposed to West Nile virus (WNV), a pathogen first detected in the US in 1999 that has since become one of North America's most prevalent vector-borne pathogens. The strongest effects were initial crow population density (denser populations were more likely to suffer declines), avian species diversity (populations in areas with high diversity were less likely to suffer a decline), human population density (populations were more likely to decline in more urban areas), and time since the pathogen's introduction to the US (populations exposed to the pathogen later in its North American sweep were less likely to suffer declines than those exposed earlier). Variables that played only a minor role included rainfall, mean maximum temperature, and total number of birds, used as a proxy for the overall reservoir competence of the community. These findings indicate that WNV declined in virulence during its rapid 5-year sweep and support the importance of the 'dilution effect' whereby a diverse host community dampens pathogen transmission and potentially slows its rate of spread. Results underscore the need for considering the entire community when trying to understand the factors shaping disease risk.

Effects of load-lightening and delayed extrapair benefits on the fitness consequences of helping behavior.

In most cooperative breeders, helping is directed at close kin, allowing helpers to gain indirect fitness benefits by increasing the reproductive success of close relatives, usually their parents. Extrapair paternity (EPP) occurs at high rates in some cooperative breeders, reducing the relatedness of helpers to the young they help raise. Even so, a son that helps is related to the brood by at least 0.25 through his mother and to within-pair young by 0.5, whereas a potential helper that has EPP in his own nest is related only to the offspring he sires and unrelated to any extrapair offspring. In birds, EPP often favors older males, which in the extreme case can result in sons being more closely related to young in their parents' nest than to young in their own nests. The fitness benefit of helping will thus be enhanced if helping lightens the workload and increases survival of helpers and their fathers, enabling them to become old, hyper-successful extrapair sires. Here, we develop and analyze a proof-of-concept model, grounded in the western bluebird (Sialia mexicana) system, demonstrating the conditions under which high population levels of EPP can generate inclusive fitness benefits of helping behavior that outweigh the costs. This model provides a new perspective on the relationship between EPP and helping behavior in cooperative breeders and suggests a strong need for empirical work to gather unprecedented data on paternity over the lifetime of helpers and their parents.

Which Moral Foundations Predict Willingness to Make Lifestyle Changes to Avert Climate Change in the USA?

Jonathan Haidt's Moral Foundations Theory identifies five moral axes that can influence human motivation to take action on vital problems like climate change. The theory focuses on five moral foundations, including compassion, fairness, purity, authority, and ingroup loyalty; these have been found to differ between liberals and conservatives as well as Democrats and Republicans. Here we show, based on the Cornell National Social Survey (USA), that valuations of compassion and fairness were strong, positive predictors of willingness to act on climate change, whereas purity had a non-significant tendency in the positive direction (p = 0.07). Ingroup loyalty and authority were not supported as important predictor variables using model selection ([Formula: see text]). Compassion and fairness were more highly valued by liberals, whereas purity, authority, and in-group loyalty were more highly valued by conservatives. As in previous studies, participants who were younger, more liberal, and reported greater belief in climate change, also showed increased willingness to act on climate change. Our research supports the potential importance of moral foundations as drivers of intentions with respect to climate change action, and suggests that compassion, fairness, and to a lesser extent, purity, are potential moral pathways for personal action on climate change in the USA.

Winter resource wealth drives delayed dispersal and family-group living in western bluebirds.

Delayed dispersal, where offspring remain with parents beyond the usual period of dependence, is the typical route leading to formation of kin-based cooperative societies. The prevailing explanations for why offspring stay home are variation in resource wealth, in which offspring of wealthy parents benefit disproportionately by staying home, and nepotism, where the tendency for parents to be less aggressive and share food with offspring makes home a superior place to wait to breed. These hypotheses are not strict alternatives, as only wealthy parents have sufficient resources to share. In western bluebirds, Sialia mexicana, sons usually delay dispersal until after winter, gaining feeding advantages through maternal nepotism in a familial winter group. Experimentally reducing resource wealth (mistletoe) by half on winter territories caused sons to disperse in summer, even though their parents remained on the territory during the winter. Only 8% of sons remained with their parents on mistletoe-removal territories compared to 50% of sons on control territories (t(9,10)=3.33, p<0.005). This study is the first to demonstrate that experimentally reducing wealth of a natural food resource reduces delayed dispersal, facilitating nepotism and family-group living. The results clarify the roles of year-round residency, resource limitation and relative wealth outside the breeding season in facilitating the formation of kin-based cooperative societies.

Spatial distribution of oak mistletoe as it relates to habits of oak woodland frugivores.

This study addresses the underlying spatial distribution of oak mistletoe, Phoradendron villosum, a hemi-parasitic plant that provides a continuous supply of berries for frugivorous birds overwintering the oak savanna habitat of California's outer coast range. As the winter community of birds consuming oak mistletoe varies from group-living territorial species to birds that roam in flocks, we asked if mistletoe volume was spatially autocorrelated at the scale of persistent territories or whether the patterns predicted by long-term territory use by western bluebirds are overcome by seed dispersal by more mobile bird species. The abundance of mistletoe was mapped on trees within a 700 ha study site in Carmel Valley, California. Spatial autocorrelation of mistletoe volume was analyzed using the variogram method and spatial distribution of oak mistletoe trees was analyzed using Ripley's K and O-ring statistics. On a separate set of 45 trees, mistletoe volume was highly correlated with the volume of female, fruit-bearing plants, indicating that overall mistletoe volume is a good predictor of fruit availability. Variogram analysis showed that mistletoe volume was spatially autocorrelated up to approximately 250 m, a distance consistent with persistent territoriality of western bluebirds and philopatry of sons, which often breed next door to their parents and are more likely to remain home when their parents have abundant mistletoe. Using Ripley's K and O-ring analyses, we showed that mistletoe trees were aggregated for distances up to 558 m, but for distances between 558 to 724 m the O-ring analysis deviated from Ripley's K in showing repulsion rather than aggregation. While trees with mistletoe were aggregated at larger distances, mistletoe was spatially correlated at a smaller distance, consistent with what is expected based on persistent group territoriality of western bluebirds in winter and the extreme philopatry of their sons.

Can evolutionary design of social networks make it easier to be 'green'?

The social Web is swiftly becoming a living laboratory for understanding human cooperation on massive scales. It has changed how we organize, socialize, and tackle problems that benefit from the efforts of a large crowd. A new, applied, behavioral ecology has begun to build on theoretical and empirical studies of cooperation, integrating research in the fields of evolutionary biology, social psychology, social networking, and citizen science. Here, we review the ways in which these disciplines inform the design of Internet environments to support collective pro-environmental behavior, tapping into proximate prosocial mechanisms and models of social evolution, as well as generating opportunities for 'field studies' to discover how we can support massive collective action and shift environmental social norms.

Benefits, motivations, and barriers related to environmental volunteerism for older adults: developing a research agenda.

Interest in civic engagement focused on the natural environment has grown dramatically, as has the population of older adults. Our article explores the potential for increased environmental volunteerism among older adults to enrich the lives of volunteers while benefitting the community and environmental quality. Curiously, this convergence has been relatively neglected by researchers and program developers. We review existing literatures on trends in volunteerism, motivations, benefits, and barriers to participation, with a special focus on elements most relevant to older adults. Based on this review, we identify a number of critical areas of research, and pose key research questions.

Development and cross-species testing of western bluebird (Sialia mexicana) microsatellite primers.

Western and eastern bluebirds (Sialia mexicana and S. sialis) are socially monogamous passerines that engage in extra-pair copulations. We obtained microsatellites from S. mexicana and optimized and characterized 15 microsatellite DNA loci in 60 individuals of this species. Primer pairs yielded an average of 13 alleles per locus in western bluebirds (range 3-35 alleles) with an average observed heterozygosity of 0.68 (range 0.27-0.88). All 15 loci also successfully amplified in S. sialis (n = 24), with an average of 11.5 alleles per locus (range 4-26) and an average observed heterozygosity of 0.59 (range 0.22-0.90).

Decoding dumping ducks.

Conspecific brood parasitism, where females of the same species lay eggs in each other's nests, is common in waterfowl, and is usually considered costly to host females, which are stuck looking after eggs and chicks that are not their own. However, since female waterfowl often exhibit an unusual propensity to nest near where they were born, there has been some uncertainty over whether, in ducks and geese, laying in nests of conspecifics really is parasitism. Do parasitic and host females tend to be related? And is parasitism actually a form of cooperation in disguise? In a population in Hudson Bay, Andersson & Waldeck (this issue) found that 'parasitic' eggs in nests of the common eider, Somateria mollissima sedentaria, are more closely related to host eggs than expected by chance. In fact, host and 'donor' eggs are more closely related than are females breeding at neighbouring nests. The Hudson Bay population of common eiders is unusual, because unlike in more benign climates, females do not tend to breed near their natal nest. Spatial proximity alone cannot account for the high relatedness between host eggs and 'dumped' or donor eggs. Instead, the high relatedness values are probably the result of active recognition, where females favour kin, either when dumping or accepting eggs. These new data, along with evidence indicating that the donor lays the first egg in the nest nearly half the time, suggest that what appears to be parasitism in common eiders may be a form of kin-based cooperation.

Family-based winter territoriality in western bluebirds, Sialia mexicana: the structure and dynamics of winter groups.

Winter residency is characteristic of the majority of cooperatively breeding birds, but the composition and dynamics of winter groups have been examined in relatively few. In 1996-1998, we examined winter territoriality in the western bluebird, a year-round resident that shows a limited degree of helping behaviour in central coastal California, U.S.A. In spring, most western bluebirds breed as socially monogamous pairs, but a small proportion of pairs (3-16%) have additional breeding-age males helping at the nest, usually assisting parents or brothers. We found that year-round residents commonly wintered in family groups that defended territories similar to those used in spring. Winter groups had an even sex ratio and formed early in the autumn, when hatch-year birds dispersed. More females than males left their natal groups to be replaced by an influx of immigrant hatch-year birds. Winter groups typically consisted of breeders and one or two sons from the prior breeding season along with one or more immigrant females. A second period of dispersal occurred in spring when winter groups broke up and most birds other than the breeding pair left the winter territory. When they bred, yearling males and females often bred with unrelated individuals from their winter groups. Sons were more likely to remain on the study area as yearlings when they wintered with both parents than when they wintered with just one parent. We suggest that young males stay the winter due to benefits of remaining in family groups on mistletoe-based winter territories. Subsequent localized dispersal of sons then leads to opportunistic kin-based interactions later in life. Copyright 2001 The Association for the Study of Animal Behaviour.