Lifetime inclusive fitness effects of cooperative polygamy in the acorn woodpecker.

Although over 50 y have passed since W. D. Hamilton articulated kin selection and inclusive fitness as evolutionary explanations for altruistic behavior, quantifying inclusive fitness continues to be challenging. Here, using 30 y of data and two alternative methods, we outline an approach to measure lifetime inclusive fitness effects of cooperative polygamy (mate-sharing or cobreeding) in the cooperatively breeding acorn woodpecker Melanerpes formicivorus. For both sexes, the number of offspring (observed direct fitness) declined while the number of young parented by related cobreeders (observed indirect fitness effect) increased with cobreeding coalition size. Combining these two factors, the observed inclusive fitness effect of cobreeding was greater than breeding singly for males, while the pattern for females depended on whether fitness was age-weighted, as females breeding singly accrued greater fitness at younger ages than cobreeding females. Accounting for the fitness birds would have obtained by breeding singly, however, lifetime inclusive fitness effects declined with coalition size for males, but were greater for females breeding as duos compared to breeding singly, due largely to indirect fitness effects of kin. Our analyses provide a road map for, and demonstrate the importance of, quantifying indirect fitness as a powerful evolutionary force contributing to the costs and benefits of social behaviors.

Lifetime reproductive benefits of cooperative polygamy vary for males and females in the acorn woodpecker (Melanerpes formicivorus).

Cooperative breeding strategies lead to short-term direct fitness losses when individuals forfeit or share reproduction. The direct fitness benefits of cooperative strategies are often delayed and difficult to quantify, requiring data on lifetime reproduction. Here, we use a longitudinal dataset to examine the lifetime reproductive success of cooperative polygamy in acorn woodpeckers (Melanerpes formicivorus), which nest as lone pairs or share reproduction with same-sex cobreeders. We found that males and females produced fewer young per successful nesting attempt when sharing reproduction. However, males nesting in duos and trios had longer reproductive lifespans, more lifetime nesting attempts and higher lifetime reproductive success than those breeding alone. For females, cobreeding in duos increased reproductive lifespan so the lifetime reproductive success of females nesting in duos was comparable to those nesting alone and higher than those nesting in trios. These results suggest that for male duos and trios, reproductive success alone may provide sufficient fitness benefits to explain the presence of cooperative polygamy, and the benefits of cobreeding as a duo in females are higher than previously assumed. Lifetime individual fitness data are crucial to reveal the full costs and benefits of cooperative polygamy.

Correlates of bird collisions with buildings across three North American countries.

Collisions with buildings cause up to 1 billion bird fatalities annually in the United States and Canada. However, efforts to reduce collisions would benefit from studies conducted at large spatial scales across multiple study sites with standardized methods and consideration of species- and life-history-related variation and correlates of collisions. We addressed these research needs through coordinated collection of data on bird collisions with buildings at sites in the United States (35), Canada (3), and Mexico (2). We collected all carcasses and identified species. After removing records for unidentified carcasses, species lacking distribution-wide population estimates, and species with distributions overlapping fewer than 10 sites, we retained 269 carcasses of 64 species for analysis. We estimated collision vulnerability for 40 bird species with ≥2 fatalities based on their North American population abundance, distribution overlap in study sites, and sampling effort. Of 10 species we identified as most vulnerable to collisions, some have been identified previously (e.g., Black-throated Blue Warbler [Setophaga caerulescens]), whereas others emerged for the first time (e.g., White-breasted Nuthatch [Sitta carolinensis]), possibly because we used a more standardized sampling approach than past studies. Building size and glass area were positively associated with number of collisions for 5 of 8 species with enough observations to analyze independently. Vegetation around buildings influenced collisions for only 1 of those 8 species (Swainson's Thrush [Catharus ustulatus]). Life history predicted collisions; numbers of collisions were greatest for migratory, insectivorous, and woodland-inhabiting species. Our results provide new insight into the species most vulnerable to building collisions, making them potentially in greatest need of conservation attention to reduce collisions and into species- and life-history-related variation and correlates of building collisions, information that can help refine collision management.

Correlaciones de las Colisiones de Aves contra Edificios en Tres Países de América del Norte Resumen Las colisiones contra los edificios causan hasta mil millones de fatalidades de aves al año en los Estados Unidos y en Canadá. Sin embargo, los esfuerzos por reducir estas colisiones se beneficiarían con estudios realizados a grandes escalas espaciales en varios sitios de estudio con métodos estandarizados y considerando las variaciones relacionadas a la historia de vida y a la especie y las correlaciones de las colisiones. Abordamos estas necesidades de investigación por medio de una recolección coordinada de datos sobre las colisiones de aves contra edificios en los Estados Unidos (35), Canadá (3) y México (2). Recolectamos todos los cadáveres y los identificamos hasta especie. Después de retirar los registros de cadáveres no identificados, las especies sin estimaciones poblacionales a nivel distribución y las especies con distribuciones traslapadas en menos de diez sitios, nos quedamos con 269 cadáveres de 64 especies para el análisis. Estimamos la vulnerabilidad a colisiones para 40 especies con ≥2 fatalidades con base en la abundancia poblacional para América del Norte, el traslape de su distribución entre los sitios de estudio y el esfuerzo de muestreo. De las diez especies que identificamos como las más vulnerables a las colisiones, algunas han sido identificadas previamente (Setophaga caerulescens), y otras aparecieron por primera vez (Sitta carolinensis), posiblemente debido a que usamos una estrategia de muestreo más estandarizada que en los estudios previos. El tamaño del edificio y el área del vidrio estuvieron asociados positivamente con el número de colisiones para cinco de ocho especies con suficientes observaciones para ser analizadas independientemente. La vegetación alrededor de los edificios influyó sobre las colisiones solamente para una de esas ocho especies Catharus ustulatus). Las historias de vida pronosticaron las colisiones; el número de colisiones fue mayor para las especies migratorias, insectívoras y aquellas que habitan en las zonas boscosas. Nuestros resultados proporcionan una nueva perspectiva hacia las especies más vulnerables a las colisiones contra edificios, lo que las pone en una necesidad potencialmente mayor de atención conservacionista para reducir estas colisiones y de estudio de las variaciones relacionadas con la especie y la historia de vida y las correlaciones de las colisiones contra edificios, información que puede ayudar a refinar el manejo de colisiones.

Habitat Saturation Results in Joint-Nesting Female Coalitions in a Social Bird.

Joint nesting by females and cooperative polyandry-cooperatively breeding groups with a male-biased breeder sex ratio-are little-understood, rare breeding systems. We tested alternative hypotheses of factors potentially driving these phenomena in a population of joint-nesting acorn woodpeckers (Melanerpes formicivorus). During periods of high population density and thus low independent breeding opportunities, acorn woodpecker females formed joint-nesting coalitions with close kin. Coalitions were typically associated with groups with a male bias. We found strong evidence for both inter- and intrasexual conflict, as joint nesting conferred a fitness benefit to some males, a significant fitness cost to females, and no gain in per capita reproductive output for either sex. Such conflict, particularly the cost to females, may be an important reason why joint nesting is rare among cooperatively breeding taxa.

Artificial light at night confounds broad-scale habitat use by migrating birds.

With many of the world's migratory bird populations in alarming decline, broad-scale assessments of responses to migratory hazards may prove crucial to successful conservation efforts. Most birds migrate at night through increasingly light-polluted skies. Bright light sources can attract airborne migrants and lead to collisions with structures, but might also influence selection of migratory stopover habitat and thereby acquisition of food resources. We demonstrate, using multi-year weather radar measurements of nocturnal migrants across the northeastern U.S., that autumnal migrant stopover density increased at regional scales with proximity to the brightest areas, but decreased within a few kilometers of brightly-lit sources. This finding implies broad-scale attraction to artificial light while airborne, impeding selection for extensive forest habitat. Given that high-quality stopover habitat is critical to successful migration, and hindrances during migration can decrease fitness, artificial lights present a potentially heightened conservation concern for migratory bird populations.

Wild acorn woodpeckers recognize associations between individuals in other groups.

According to the social intelligence hypothesis, understanding the cognitive demands of the social environment is key to understanding the evolution of intelligence. Many important socio-cognitive abilities, however, have primarily been studied in a narrow subset of the social environment-within-group social interactions-despite the fact that between-group social interactions often have a substantial effect on fitness. In particular, triadic awareness (knowledge about the relationships and associations between others) is critical for navigating many types of complex social interactions, yet no existing study has investigated whether wild animals can track associations between members of other social groups. We investigated inter-group triadic awareness in wild acorn woodpeckers (Melanerpes formicivorus), a socially complex group-living bird. We presented woodpeckers with socially incongruous playbacks that simulated two outsiders from different groups calling together, and socially congruous playbacks that simulated two outsiders from the same group calling together. Subjects responded more quickly to the incongruous playbacks, suggesting that they were aware that the callers belonged to two different groups. This study provides the first demonstration that animals can recognize associations between members of other groups under natural circumstances, and highlights the importance of considering how inter-group social selection pressures may influence the evolution of cognition.

New Records of Ixodes affinis (Acari: Ixodidae) Parasitizing Avian Hosts in Southeastern Virginia.

Ixodes affinis Neumann (Acari: Ixodidae) is a hard-bodied tick species distributed throughout much of the southeastern United States. Although I. affinis does not parasitize humans, it is a competent vector of Borrelia burgdorferi sensu stricto, the causative-agent of Lyme disease, and thus contributes to the enzootic maintenance of this pathogen. This study presents evidence of I. affinis parasitizing five new host passerine species. During 2012-2014, 1,888 birds were captured and examined for ticks, and 18 immature I. affinis were collected from 12 birds-six Carolina Wrens (Thyrothorus ludovicianus); two Brown Thrashers (Toxostoma rufum); and one American Robin (Turdus migratorius), Eastern Towhee (Pipilo erythrophthalmus), Northern Cardinal (Cardinalis cardinalis), and White-throated Sparrow (Zonotrichia albicollis). Of 15 larvae and 3 nymphs collected, one nymph tested positive for B. burgdorferi DNA. I. affinis was found co-feeding on birds with immature Amblyomma americanum (L.), Ixodes brunneus Koch, Ixodes dentatus Marx, Ixodes scapularis Say, and Haemaphysalis leporispalustris Packard. The results of this research provide a better understanding of I. affinis hosts and identify avian taxa that may play a role in the maintenance and dispersal of this tick species.

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 phenomenon may be a bet-hedging strategy that reduces variance in reproductive success (fecundity variance) in populations living in highly 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 estimated mean 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.

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.

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.

Passerine birds as hosts for Ixodes ticks infected with Borrelia burgdorferi sensu stricto in southeastern Virginia.

The ecology of vector-borne diseases in a region can be attributed to vector-host interactions. In the United States, tick-borne pathogens are the cause of the highest number of reported vector-borne diseases. In the mid-Atlantic region of the eastern United States, tick-borne diseases such as Lyme disease, have increased in incidence, with tick-host-pathogen interactions considered a contributing factor to this increase. Ticks become infected with pathogens after taking a blood meal from a systemically infected host or through a localized infection while co-feeding on a host with other infected ticks. The host not only plays a role in pathogen acquisition by the tick, but can also facilitate dispersal of the tick locally within a region or over greater distances into new geographical ranges outside of their historical distributional range. In this study conducted in southeastern Virginia (USA), we examined the interaction between both resident and migratory bird species and Ixodes ticks, the primary vectors of Borrelia burgdorferi sensu stricto (s.s.) the main causative agent of Lyme disease on the East coast of the United States. Over a two-year period (2012-2014), 1879 passerine birds were surveyed, with 255 Ixodes ticks tested for the presence of Borrelia spp. Eighty passerine birds (4.3 %) representing 17 bird species were parasitized by at least one Ixodes tick, but only three bird species were parasitized by Ixodes ticks that tested positive for B. burgdorferi s.s. Twenty Ixodes ticks (7.8 %) tested positive for B. burgdorferi s.s. with nearly all collected from resident bird species including the Carolina wren (Thryothorus ludovicianus) and brown thrasher (Toxostoma rufum). Given that millions of birds pass through southeastern Virginia during migration, even with the low number of Ixodes ticks parasitizing passerine birds and the low prevalence of B. burgdorferi s.s. found within Ixodes ticks collected, the sheer volume of passerine birds suggests they may play a role in the maintenance and dispersal of B. burgdorferi s.s. in southeastern Virginia.

Quantifying purported competition with individual- and population-level metrics.

Competitive species interactions may contribute to population declines. Purportedly, Red-bellied Woodpeckers (Melanerpes carolinus), a common species, and Red-cockaded Woodpeckers (Picoides borealis), an endangered species, compete for roosting and nesting cavities in living pine trees. To determine whether behavioral interactions measured at the individual level manifest themselves at the population level, we conducted field experiments designed to test whether the presence of Red-bellied Woodpeckers resulted in a decrease in fitness to Red-cockaded Woodpeckers. As part of a 4-year study examining the nature of interspecific interactions in two populations of Red-cockaded Woodpeckers (one stable, the Apalachicola Ranger District; one declining, the Wakulla Ranger District) in the Apalachicola National Forest, Florida, we conducted a set of Red-bellied Woodpecker removal experiments. Paradoxically, following the removal of Red-bellied Woodpeckers, we observed decreases in Red-cockaded Woodpecker group size, proportion of nests that were successful, and proportion of individuals remaining on territories. Removal of Red-bellied Woodpeckers may have exaggerated the immigration rate of Red-bellied Woodpeckers to Red-cockaded Woodpecker territories. The Red-cockaded Woodpeckers in the Apalachicola Ranger District likely can withstand pressure from immigrating Red-bellied Woodpeckers given that their population has remained relatively stable despite the presence of Red-bellied Woodpeckers. A major factor of population persistence in the Wakulla Ranger District was the high turnover rate of adult female Red-cockaded Woodpeckers, a phenomenon that was exacerbated by removal of Red-bellied Woodpeckers. Relying solely on observations of apparently competitive interactions between individuals may not necessarily provide information about population-level outcomes. Paradoxically, removing species that appear to be competitors may harm species of concern.

Tracking the warriors and spectators of acorn woodpecker wars.

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First report of Candidatus Rickettsia mendelii in Ixodes brunneus from the United States.

Candidatus Rickettsia mendelii is a novel rickettsial species recently identified in Ixodes ricinus. In this study, Ixodes brunneus collected from wild birds (n = 77 ticks) or vegetation (n = 4 ticks) in southeastern Virginia were surveyed for rickettsial agents. Candidatus Rickettsia mendelii was confirmed in I. brunneus through sequencing of the gltA and 16S rRNA genes. This is the first report of this rickettsial species in Ixodes ticks in North America.

Wandering woodpeckers: foray behavior in a social bird.

In many cooperatively breeding taxa, nonbreeding subordinates, or helpers, use extra-territorial forays to discover dispersal opportunities. Such forays are considered energetically costly and foraying birds face aggression from conspecific members of the territories they visit. In contrast, breeders in cooperatively breeding taxa are expected to foray seldomly. We used novel tracking technologies to follow 62 acorn woodpeckers (Melanerpes formicivorus), a cooperatively breeding bird, to study extra-territorial foray behavior. Both helpers and breeders engaged in extra-territorial forays routinely and often several times per day. Helpers forayed earlier in the day and invested more time when foraying to high-quality territories. Unexpectedly, breeders forayed as often and as far as helpers. Breeders from high-quality territories forayed closer to their home territories than breeders from low-quality territories, reflecting a potential trade-off between foraying and territory defense. Such a routine pattern of extra-territorial forays in both helpers and breeders suggests that the motives behind forays differ by sex and social status and involve more than simply searching for dispersal opportunities.

A tale of two worlds: molecular ecology and population structure of the threatened Florida scrub-jay.

Elsewhere in this issue of Molecular Ecology, Coulon et al. provide a detailed analysis of population structure of the threatened Florida scrub-jay (Aphelocoma coerulescens) using genetic markers and compare it to that inferred from previous demographic surveys and observed dispersal behaviour in this species. In contrast to previous attempts at such comparisons, estimates from the two methods are reasonably congruent. Although challenges remain, Coulon et al.'s analyses demonstrate the potential for closing the gap between these alternative methodologies, and ultimately for future genetic surveys to be used confidently in conservation planning.

Local avian density influences risk of mortality from window strikes.

Up to a billion birds die per year in North America as a result of striking windows. Both transparent and reflective glass panes are a cause for concern, misleading birds by either acting as invisible, impenetrable barriers to desired resources, or reflecting those resources over a large surface area. A high number of window strikes occur during migration, but little is known about the factors of susceptibility, or whether particular avian taxa are more vulnerable than others. We report on a study of window strikes and mist-netting data at the Virginia Zoological Park (Norfolk, Virginia, USA), conducted in the autumn of 2013 and 2014. We focused on three factors likely to contribute to an individual's predisposition to collide with windows: (i) taxonomic classification, (ii) age, and (iii) migrant vs. resident status. Thrushes, dominated by the partial migrant American Robin (Turdus migratorius), were significantly less likely to strike glass than be sampled in mist nets (χ(2) = 9.21, p = 0.002), while wood-warblers (Parulidae) were more likely to strike than expected (χ(2) = 13.55, p < 0.001). The proportion of juveniles striking windows (45.4%) was not significantly different (χ(2) = 0.05, p = 0.827) than the population of juvenile birds naturally occurring at the zoo (48.8%). Migrants, however, were significantly more susceptible to window strikes than residents (χ(2) = 6.35, p = 0.012). Our results suggest that resident birds are able to learn to avoid and thus reduce their likelihood of striking windows; this intrinsic risk factor may help explain the apparent susceptibility of certain taxa to window strikes.

A critical review of twenty years' use of the resource-ratio theory.

A model of species interactions based on their use of shared resources was proposed in 1972 by Robert MacArthur and later expanded in an article (1980) and a book (1982) by David Tilman. This "resource-ratio theory" has been used to make a number of testable predictions about competition and community patterns. We reviewed 1,333 papers that cite Tilman's two publications to determine whether predictions of the resource-ratio theory have been adequately tested and to summarize their general conclusions. Most of the citations do not directly test the theory: only 26 studies provide well-designed tests of one or more predictions, resulting in 42 individual tests of predictions. Most of these tests were conducted in the laboratory or experimental microcosms and used primary producers in freshwater systems. Overall, the predictions of the resource-ratio theory were supported 75% of the time. One of the primary predictions of the model, that species dominance varies with the ratio of resource availabilities, was supported by 13 of 16 tests, but most other predictions have been insufficiently tested. We suggest that more experimental work in a variety of natural systems is seriously needed, especially studies designed to test predictions related to resource supply and consumption rates.

Parasitic helminths of red-bellied woodpeckers (Melanerpes carolinus) from the Apalachicola National Forest in Florida.

Seventy-four red-bellied woodpeckers (Melanerpes carolinus) from the Apalachicola National Forest (30 degrees 10'N, 84 degrees 40'W) in northwest Florida were examined for helminths. The most prevalent parasites were the nematode Aproctella stoddardi (11%) and the acanthocephalan Mediorhynchus centurorum (11%). New host records include Pseudaprocta samueli, A. stoddardi, Tridentocapillaria tridens, Diplotriaena americana, Dispharynx nasuta, Procyrnea pileata, Orthoskrjabinia rostellata, and Brachylaima fuscatum. The helminth fauna was characterized by low prevalences and intensities of infection and low numbers of species per bird (1.2). The frequency of prescribed burning and habitat understory flora composition did not influence the prevalences or intensities of helminths in red-bellied woodpeckers collected from 2 similar but differently managed sites within the forest.

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.