Brooding, Provisioning, and Compensatory Care in the Cooperatively Breeding Acorn Woodpecker

We analyzed reproductive investment in parental care (brooding and the provisioning of nestlings) in the acorn woodpecker (Melanerpes formicivorus), a cooperatively breeding species in which both polygynandry and helping-at-the-nest are common. As predicted based on the strategies pursued by birds of different sex and status, breeders generally invested more in parental care than helpers, and breeder females invested more than breeder males. Contrary to expectations, however, the degree to which individuals reduced their effort with increasing group size (i.e., patterns of load lightening or compensatory care) did not match overall investment. Instead, as group size increased, there was no significant difference in compensation in either brooding or provisioning among the different categories of birds. Compensation, at least by breeders, was significantly lower during the first week of the nestling period than later on, supporting the hypothesis that compensatory reduction in care is less likely when brood reduction is more common and was not affected by the acorn crop, which had no significant effect on the incidence of brood reduction despite being important to overall reproductive success. Our results offer support for the hypothesis that levels of compensation are influenced by the relative importance of brood reduction. More theoretical work, however, will be needed in order to understand the basis for patterns of compensation among individuals of different sex and status

Temporal Variability and cooperative Breeding: Testing the Bet-Hedging Hypothesis in the Acorn Woodpecker

Cooperative breeding is generally considered an adaptation to ecological constraints on dispersal and independent breeding, usually due to limited breeding opportunities. Although benefits of cooperative breeding are typically thought of in terms of increased mean reproductive success, it has recently been proposed that this phenomenonmay be a bet-hedging strategy that reduces variance in reproductive success (fecundity variance) inpopulations living inhighly variable environments. We tested this hypothesis using long-term data on the polygynandrous acorn woodpecker (Melanerpes formicivorus). In general, fecundity variance decreased with increasing sociality, at least when controlling for annual variation in ecological conditions. Nonetheless, decreased fecundity variance was insufficient to compensate for reduced per capita reproductive success of larger,more social groups, which typically suffered lower estimatedmean fitness.We did, however, find evidence that sociality in the form of larger group size resulted in increased fitness in years following a small acorn crop due to reduced fecundity variance. Bet-hedging, although not the factor driving sociality in general, may play a role in driving acorn woodpecker group living when acorns are scarce and ecological conditions are poor

Cooperative Breeding and Long-Distance Dispersal: A Test Using Vagrant Records

Cooperative breeding is generally associated with increased philopatry and sedentariness, presumably because short-distance dispersal facilitates the maintenance of kin groups. There are, however, few data on long-distance dispersal in cooperative breeders-the variable likely to be important for genetic diversification and speciation. We tested the hypothesis that cooperative breeders are less likely to engage in long-distance dispersal events by comparing records of vagrants outside their normal geographic range for matched pairs (cooperatively vs. non-cooperatively breeding) of North American species of birds. Results failed to support the hypothesis of reduced long-distance dispersal among cooperative breeders. Thus, our results counter the conclusion that the lower rate of speciation among cooperative breeding taxa found in recent analyses is a consequence of reduced vagility

Variable Helper Effects, Ecological Conditions, and the Evolution of Cooperative Breeding in the Acorn Woodpecker

The ecological conditions leading to delayed dispersal and helping behavior are generally thought to follow one of two contrasting scenarios: that conditions are stable and predictable, resulting in young being ecologically forced to remain as helpers (extrinsic constraints and the habitat saturation hypothesis), or that conditions are highly variable and unpredictable, leading to the need for helpers to raise young, at least when conditions are poor (intrinsic constraints and the hard life hypothesis). We investigated how variability in ecological conditions influences the degree to which helpers augment breeder fitness in the cooperatively breeding acorn woodpecker (Melanerpes formicivorus), a species in which the acorn crop, territory quality, and prior breeding experience all vary in ways that have important effects on fitness. We found that the relationship between ecological conditions and the probability that birds would remain as helpers was variable but that helpers generally yielded greater fitness benefits when ecological conditions were favorable, rather than unfavorable, for breeding. These results affirm the importance of extrinsic constraints to delayed dispersal and cooperative breeding in this species, despite its dependence on a highly variable and unpredictable acorn crop. Our findings also confirm that helpers can have very different fitness effects, depending on conditions, but that those effects are not necessarily greater when breeding conditions are unfavorable

Does Helping-at-the-Nest Help? The Case of the Acorn Woodpecker

Cooperative breeding groups often involve helpers-at-the-nest ; indeed, such behavior typically defines this intriguing breeding system. In few cases, however, has it been demonstrated that feeding nestlings by helpers, rather than some other behavior associated with helpers\u27 presence, leads to greater reproductive success. One prediction of the hypothesis that feeding behavior per se is responsible for the fitness benefits conferred by helpers is that there should be close congruence between the patterns of helping-at-the-nest and the fitness effects of helpers. Here we look for such a relationship in the cooperatively breeding acorn woodpecker (Melanerpes formicivorus) in order to begin to identify the behaviors of helpers that drive the increased fitness benefits they confer. In terms of young fledged, a helper male confers approximately the same fitness benefits to a group as does a helper female; more dramatically, the effects of helper males increases with increasing food supply, most importantly the prior year\u27s acorn crop on which this species depends, whereas that of helper females does not. These patterns do not match the nest-feeding patterns of helpers, which are greater for females than males and do not increase with a larger acorn crop the prior autumn. In contrast, the proportion of time helpers spend tending acorn-storage facilities (granaries) and are present in or near their home territory is greater for males than females and, at least for males, positively related to the size of the acorn crop. These results fail to support the hypothesis that the primary benefit conferred by helpers is feeding young in the nest; rather, they suggest that behaviors such as territorial defense and predator detection are more important. Understanding exactly what those behaviors are in this, and most other cooperatively breeding systems, remain to be determined

Effects of Gypsy Moth Outbreaks on North American Woodpeckers

We examined the effects of the introduced gypsy moth (Lymantria dispar) on seven species of North American woodpeckers by matching spatially explicit data on gypsy moth outbreaks with data on breeding and wintering populations. In general, we detected modest effects during outbreaks: during the breeding season one species, the Red-headed Woodpecker (Melanerpes erythrocephalus), increased over pre-outbreak levels, while during the winter one species, the Yellow-bellied Sapsucker (Sphyrapicus varius), increased and one, the Downy Woodpecker (Picoides pubescens), decreased from pre-outbreak levels. Responses following outbreaks were similarly variable, and in general we were unsuccessful at predicting population responses to outbreaks from a priori knowledge of woodpecker ecology and behavior. We did, however, find evidence that the response of at least half of the species changed over the 34-year period covered by the study: except for the Northern Flicker (Colaptes auratus), whose response to outbreaks during the winter decreased, populations generally responded more positively to outbreaks with time. This temporal response suggests that North American woodpeckers may be taking greater advantage of the resource pulse and/or habitat changes caused by outbreaks of this exotic pest now than previously, so in the future the effects of gypsy moth outbreaks on these species may increase

Testing Alternative Hypotheses for the Cause of Population Declines: The Case of the Red-Headed Woodpecker

The Red-headed Woodpecker (Melanerpes erythrocephalus) has experienced strong population declines during the past 3 decades. Using North American Breeding Bird Survey (BBS) and Audubon Christmas Bird Count (CBC) data, we investigated 4 hypotheses that may explain this decline, including: (1) interspecific competition with native Red-bellied Woodpeckers (Melanerpes carolinus) and nonnative European Starlings (Sturnus vulgaris); (2) predation by Cooper\u27s Hawks (Accipiter cooperii) and Sharp-shinned Hawks (Accipiter striatus); (3) climate change; and (4) changes in forested area within their range. In analyses of both the breeding and overwintering periods, our results indicated a role of increased accipiter populations in driving Red-headed Woodpecker declines through increased predation. We also found evidence for significant effects of warmer winter temperatures and increased forest cover, both directly and indirectly through their effects on enhancing accipiter populations. In contrast, our results failed to support the hypothesis that interspecific competition with either Red-bellied Woodpeckers or European Starlings has played a role in Red-headed Woodpecker declines. Despite considerable evidence for nest-site competition and aggression between Red-headed Woodpeckers and both Red-bellied Woodpeckers and European Starlings, these interactions do not appear to be limiting Red-headed Woodpecker populations

Workflow For Constructing Social Networks From Automated Telemetry Systems

1. Advances in datalogging technologies have provided a way to monitor the movement of individual animals at unprecedented spatial and temporal scales, both large and small. When used in conjunction with social network analyses, these data can provide insight into fine scale associative behaviors. The variety of technologies demand continuous progress in workflows to translate data streams from automated systems to social networks, based on biologically relevant metrics. 2. Here we present a workflow for generating flexible association matrices from automated radio-telemetry data that can be parsed into both spatial and temporal dimensions. These can then be used to generate and compare social networks across space and time. 3. We illustrate this workflow using data collected from an automated telemetry study of acorn woodpeckers (Melanerpes formicivorus), a cooperatively breeding bird. The data were collected continuously over two years at base stations placed within social group territories. We use this system to demonstrate how this flexible data structure can be used to answer a number of biological questions, specifically 1) how assortative are social associations at the population scale, 2) how do association patterns among territory visitors vary across territories, 3) and how does seasonality affect assortative affiliation within groups? 4. This flexible method allows one to generate social networks that can be used to ask a variety of biological questions pertinent to a wide range of animal systems, exploiting the investigative power that can be gained by using automated radio-telemetry in conjunction with social network analyses