Elevated inbreeding in Heliconia tortuosa is determined by tropical forest stand age, isolation and loss of hummingbird functional diversity.

Forest conversion and habitat loss are major threats to biological diversity. Forest regeneration can mitigate the negative effects of old-growth forest loss on species diversity, but less is known about the extent to which forest loss reduces genetic diversity in remnant populations and whether secondary forests play a role in the maintenance of genetic diversity. We quantified genetic diversity in a tropical hummingbird-pollinated understorey herb, Heliconia tortuosa, across a landscape mosaic of primary and secondary forest regrowth. Using microsatellite genotypes from >850 adult and juvenile plants within 33 forest patches and extensive bird surveys, we examined the effect of contemporary and historical landscape features including forest age (primary vs. secondary forest), stand isolation and pollinator assemblages on genetic diversity and levels of inbreeding in H. tortuosa. We found that inbreeding was up to three times higher in secondary forest, and this effect was amplified with reductions in primary forest in the surrounding landscape through reduced observed heterozygosity in isolated fragments. Inbreeding in forest patches was negatively correlated with the local frequency of specialist long-distance foraging traplining hummingbirds. Traplining hummingbirds therefore appear to facilitate mating among unrelated plants-an inference we tested using empirically parameterized simulations. Higher levels of inbreeding in H. tortuosa are therefore associated with reduced functional diversity of hummingbirds in secondary forests and forest patches isolated from primary forests. Our findings suggest a cryptic consequence of primary forest loss and secondary forest regeneration through the disruption of mutualistic interactions resulting in the erosion of genetic diversity in a common understorey plant.

Building a better baseline to estimate 160 years of avian population change and create historically informed conservation targets.

Globally, anthropogenic land-cover change has been dramatic over the last few centuries and is frequently invoked as a major cause of wildlife population declines. Baseline data currently used to assess population trends, however, began well after major changes to the landscape. In the United States and Canada, breeding bird population trends are assessed by the North American Breeding Bird Survey, which began in the 1960s. Estimates of distribution and abundance prior to major habitat alteration would add historical perspective to contemporary trends and allow for historically based conservation targets. We used a hindcasting framework to estimate change in distribution and abundance of 7 bird species in the Willamette Valley, Oregon (United States). After reconciling classification schemes of current and 1850s reconstructed land cover, we used multiscale species distribution models and hierarchical distance sampling models to predict spatially explicit densities in the modern and historical landscapes. We estimated that since the 1850s, White-breasted Nuthatch (Sitta carolinensis) and Western Meadowlark (Sturnella neglecta) populations, 2 species sensitive to fragmentation of oak woodlands and grasslands, declined by 93% and 97%, respectively. Five other species we estimated nearly stable or increasing populations, despite steep regional declines since the 1960s. Based on these estimates, we developed historically based conservation targets for amount of habitat, population, and density for each species. Hindcasted reconstructions provide historical perspective for assessing contemporary trends and allow for historically based conservation targets that can inform current management.

Construcción de una Mejor Línea Base para Estimar 160 Años de Cambio en la Población de Aves y Crear Objetivos de Conservación Orientados Históricamente Resumen A nivel mundial, el cambio antropogénico en la cobertura del suelo ha sido dramático durante los últimos siglos y frecuentemente se le considera una de las principales causas de las declinaciones en la población de fauna. A pesar de esto, se comenzó a registrar los datos de línea base que se usan actualmente para evaluar las tendencias poblacionales mucho después de cambios importantes en el paisaje. En los Estados Unidos de América y en Canadá, las tendencias poblacionales de las aves reproductoras se evalúan por medio del Censo de Aves Reproductoras, el cual comenzó en la década de 1960. Los estimados de la distribución y abundancia previas a las alteraciones importantes en el hábitat le proporcionarían a este censo una perspectiva histórica de las tendencias contemporáneas y permitirían el establecimiento de objetivos de conservación con fundamentos históricos. Usamos un marco de trabajo de análisis retrospectivo para estimar el cambio en la distribución y abundancia de siete especies de aves en el Valle Willamette en Oregon (E.U.A). Después de reconciliar los esquemas de clasificación de la cobertura de suelo actual y la reconstruida para la década de 1850, usamos modelos multiescala de distribución de especies y modelos de muestreo de distancia jerárquica para predecir las densidades espacialmente explícitas en los paisajes históricos y modernos. Estimamos que las poblaciones de Sitta carolinensis y Sturnella neglecta, dos poblaciones sensibles a la fragmentación de los bosques de roble y de los pastizales, han declinado en un 93% y 97% desde la década de 1850, respectivamente. Otras cinco especies han tenido poblaciones casi estables o en crecimiento, a pesar de las declinaciones abruptas desde la década de 1960. Las reconstrucciones por medio de análisis retrospectivos proporcionan una perspectiva histórica para la evaluación de las tendencias contemporáneas y permiten el establecimiento de objetivos con bases históricas que pueden orientar al manejo que ocurre hoy en día.

Songbird feathers as indicators of mercury exposure: high variability and low predictive power suggest limitations.

Although feathers are commonly used to monitor mercury (Hg) in avian populations, their reliability as a sampling matrix has not been thoroughly assessed for many avian species, including most songbirds (Order Passeriformes). To better understand relationships between total Hg (THg) concentrations in feathers and other tissues for birds in the thrush and sparrow families, we (1) examined variation in THg concentrations among tissues, including feathers from six different tracts, nails, liver, and muscle; (2) tested relationships between THg concentrations in the various feather tracts and those in internal tissues from the same birds, to assess the predictive power of feather THg, and; (3) compared these relationships to those between THg concentrations in nails and internal tissues, to assess the viability of nails as a non-lethal sampling alternative. THg concentrations in all feather tracts and nails were consistently higher than those in the liver and muscle, and THg was higher in the thrushes than the sparrows. When comparing feather tracts, we observed high variation within some individuals, suggesting that estimates of Hg exposure could vary depending on which feather was sampled. Despite this variation, feather type had little effect on the predictive power of feather THg concentrations, which ranged from extremely weak in the sparrows (0.09 ≤ R2 ≤ 0.16) to moderate (0.29 ≤ R2 ≤ 0.42) in the thrushes. Alternatively, we found that nail samples better predicted internal tissue THg concentrations in both the thrushes (0.44 ≤ R2 ≤ 0.48) and sparrows (0.70 ≤ R2 ≤ 0.78). Nails have been used to monitor Hg in mammals and reptiles, but their reliability as a sampling matrix for monitoring Hg in avian populations has yet to be assessed for most taxa. While nails exhibit stronger relationships to internal tissue THg concentrations, they may not be an effective sampling option for all avian species because the collection of sizable nail samples could harm living birds, particularly small songbirds. However, this method may be reasonable for retrospective museum studies. Overall, our results suggest that, despite their current use in the literature, feathers are not a suitable sampling matrix for Hg monitoring in some songbird species.

Tropical forest fragmentation limits pollination of a keystone understory herb.

Loss of native vegetation cover is thought to be a major driver of declines in pollination success worldwide. However, it is not well known whether reducing the fragmentation of remaining vegetation can ameliorate these negative effects. We tested the independent effects of composition vs. configuration on the reproductive success of a keystone tropical forest herb (Heliconia tortuosa). To do this we designed a large-scale mensurative experiment that independently varied connected forest-patch size (configuration) and surrounding amount of forest (composition). In each patch, we tested whether pollen tubes, fruit, and seed set were associated with these landscape variables. We also captured hummingbirds as an indication of pollinator availability in a subset of patches according to the same design. We found evidence for an effect of configuration on seed set of H. tortuosa, but not on other aspects of plant reproduction; proportion of seeds produced increased 40% across the gradient in patch size we observed (0.64 to > 1300 ha), independent of the amount of forest in the surrounding landscape at both local and landscape scales. We also found that the availability of pollinators was dependent upon forest configuration; hummingbird capture rates increased three and one-half times across the patch size gradient, independent of forest amount. Finally, pollinator availability was strongly positively correlated with seed set. We hypothesize that the effects of configuration on plant fitness that we observed are due to reduced pollen quality resulting from altered hummingbird availability and/or movement behavior. Our results suggest that prioritizing larger patches of tropical forest may be particularly important for conservation of this species.

Functional connectivity experiments reflect routine movement behavior of a tropical hummingbird species.

Translocation experiments, in which researchers displace animals and then monitor their movements to return home, are commonly used as tools to assess functional connectivity of fragmented landscapes. Such experiments are purported to have important advantages of being time efficient and of standardizing "motivation" to move across individuals. Yet, we lack tests of whether movement behavior of translocated birds reflects natural behavior of unmanipulated birds. We compared the routine movement behavior of a tropical hummingbird, the Green Hermit (Phaethornis guy), to that of experimentally translocated individuals. We tested for differences in site selection patterns during movement at two spatial scales (point and path levels). We also compared movement rates between treatments. Behaviors documented during translocation experiments reflected those observed during routine movements. At the point level, both translocated and non-translocated birds showed similar levels of preference for mature tropical forest. At the path level, step selection functions showed both translocated and non-translocated hummingbirds avoiding movement across non-forested matrix and selecting streams as movement corridors. Movement rates were generally higher during translocation experiments. However, the negative influence of forest cover on movement rates was proportionately similar in translocation and routine movement treatments. We report the first evidence showing that movement behavior of birds during translocation experiments is similar to their natural movement behavior. Therefore, translocation experiments may be reliable tools to address effects of landscape structure on animal movement. We observed consistent selection of landscape elements between translocated and non-translocated birds, indicating that both routine and translocation movement studies lead to similar conclusions regarding the effect of landscape structure and forest composition on functional connectivity. Our observation that hummingbirds avoid non-forest matrix and select riparian corridors also provides a potential mechanism for pollen limitation in fragmented tropical forest.

Supplemental structured surveys and pre-existing detection models improve fine-scale density and population estimation with opportunistic community science data.

Density and population estimates aid in conservation and stakeholder communication. While free and broadly available community science data can effectively inform species distribution models, they often lack the information necessary to estimate imperfect detection and area sampled, thus limiting their use in fine-scale density modeling. We used structured distance-sampling surveys to model detection probability and calculate survey-specific detection offsets in community science models. We estimated density and population for 16 songbird species under three frameworks: (1) a fixed framework that assumes perfect detection within a specified survey radius, (2) an independent framework that calculates offsets from an independent source, and (3) a calibration framework that calculates offsets from supplemental surveys. Within the calibration framework, we examined the effects of calibration dataset size and data pooling. Estimates of density and population size were consistently biased low in the fixed framework. The independent and calibration frameworks produced reliable estimates for some species, but biased estimates for others, indicating discrepancies in detection probability between structured and community science surveys. The calibration framework produced reliable population estimates with as few as 10 calibration surveys with positive detections. Data pooling dramatically decreased bias. This study provides conservationists and managers with a cost-effective method of estimating density and population.

Integrated distance sampling models for simple point counts.

Point counts (PCs) are widely used in biodiversity surveys but, despite numerous advantages, simple PCs suffer from several problems: detectability, and therefore abundance, is unknown; systematic spatiotemporal variation in detectability yields biased inferences, and unknown survey area prevents formal density estimation and scaling-up to the landscape level. We introduce integrated distance sampling (IDS) models that combine distance sampling (DS) with simple PC or detection/nondetection (DND) data to capitalize on the strengths and mitigate the weaknesses of each data type. Key to IDS models is the view of simple PC and DND data as aggregations of latent DS surveys that observe the same underlying density process. This enables the estimation of separate detection functions, along with distinct covariate effects, for all data types. Additional information from repeat or time-removal surveys, or variable survey duration, enables the separate estimation of the availability and perceptibility components of detectability with DS and PC data. IDS models reconcile spatial and temporal mismatches among data sets and solve the above-mentioned problems of simple PC and DND data. To fit IDS models, we provide JAGS code and the new "IDS()" function in the R package unmarked. Extant citizen-science data generally lack the information necessary to adjust for detection biases, but IDS models address this shortcoming, thus greatly extending the utility and reach of these data. In addition, they enable formal density estimation in hybrid designs, which efficiently combine DS with distance-free, point-based PC or DND surveys. We believe that IDS models have considerable scope in ecology, management, and monitoring.

A re-survey in 2019-2021 of winter bird communities in the Oregon Coast Range, USA, initially surveyed in 1968-1970.

Background: Long-term changes in bird populations during winter are poorly evaluated in many parts of the world. We re-surveyed forest bird communities during winter, 2019-2021, in seven large plots originally surveyed from 1968 through to 1970, near Corvallis, Oregon, USA by Stanley Anderson, a graduate student at Oregon State University in the 1960s. Anderson counted birds and measured forest plant communities within the forests dominated by Douglas-fir (Pseudotsugamenziesii) in the Coast Range Mountain foothills. His thesis included plot locations, summaries of vegetation characteristics and point estimates of bird densities for each plot. To our knowledge, the Anderson data represent the oldest structured survey of Pacific Northwest winter forest bird communities with density estimates. Given the paucity of similar data, we re-surveyed his plots after aligning methods with his and adding modern components (distance and time interval sampling) to facilitate comparisons of changes in abundances. We preserved more extensive metadata than were preserved from Anderson's surveys, including georeferenced point count survey locations to facilitate more precisely repeatable future re-surveys. New information: Original surveys of winter bird populations in the Pacific Northwest, USA, based on georeferenced locations within seven large plots originally surveyed, 1968-1970. In addition to raw count data of all bird species detected, we include information from distance sampling and time-interval sampling methods. To our knowledge, this is one of the only structured surveys of winter forest bird populations in the Coast Range Mountains, USA.

A Framework for Investigating Rules of Life Across Disciplines.

Clearly and usefully defining the Rules of Life has long been an attractive yet elusive prospect for biologists. Life persists because requirements for existence and successful transmission of hereditary information are met. These requirements are met through mechanisms adopted by organisms, which produce solutions to environmentally imposed constraints on life. Yet, constraints and their suites of potential solutions are typically context-specific, operating at specific levels of organization, or holons, and having cascading effects across multiple levels, or the holarchy. We explore the idea that the interaction of constraints, mechanisms, and requirements within and across levels of organization may produce rules of life that can be productively defined. Although we stop short of listing specific rules, we provide a conceptual framework within which progress towards identifying rules might be made.

Influence of proximity to a geographical range limit on the physiology of a tropical bird.

1. Species' geographical ranges can be limited by a variety of biotic and abiotic factors. Physiological challenge in response to unsuitable environmental conditions can establish limits to geographical ranges. 2. We studied the physiology of Song Wrens (Cyphorhinus phaeocephalus) across their geographical range on the isthmus of Panama, an area characterized by a strong rainfall gradient. Wrens are common on the Caribbean slope of the isthmus where annual rainfall is greatest, but wren abundance declines towards the south as annual rainfall declines. Song Wrens are completely absent from the driest third of the isthmus. 3. We searched for the existence of a physiologically induced distribution limit by measuring body condition (an integrative measure of energy balance), hematocrit (% packed red blood cells in a given blood sample), and corticosterone levels (CORT, a steroid hormone that regulates the availability of energy and the endocrine stress response) in males and females. We caught birds by luring them into nets when they responded to playback of conspecific song. 4. Wrens living in drier habitat near the geographical range edge were significantly more likely to have abnormally low hematocrit scores. Baseline CORT levels were negatively associated with rainfall in one of our three best-fit path models, indicating potential energetic challenge in some individuals. Maximum CORT levels during a 60-min period of restraint correlated significantly only with sex, being higher in females. Birds with the poorest body condition lived at the dry end of the gradient. Birds on the wet end of the gradient responded fastest to conspecific song. 5. Environmental conditions vary across geographical ranges and may influence the physiological conditions of organisms, thereby enforcing limits to species' distributions. Highly specialized species, such as birds of the rain forest understory, may be especially susceptible to environmental variation associated with changing climatic conditions.

Differential reliance on aquatic prey subsidies influences mercury exposure in riparian arachnids and songbirds.

Cross-ecosystem subsidies move substantial amounts of nutrients between ecosystems. Emergent aquatic insects are a particularly important prey source for riparian songbirds but may also move aquatic contaminants, such as mercury (Hg), to riparian food webs. While many studies focus on species that eat primarily emergent aquatic insects, we instead study riparian songbirds with flexible foraging strategies, exploiting both aquatic and terrestrial prey sources. The goal in this study is to trace reliance on aquatic prey sources and correlate it to Hg concentrations in common riparian arachnids (Families Tetragnathidae, Opiliones, and Salticidae) and songbirds (Common Yellowthroat Geothlypis trichas, Spotted Towhee Pipilo maculatus, Swainson's Thrush Catharus ustulatus, Song Sparrow Melospiza melodia, and Yellow Warbler Setophaga petechia). We used stable isotopes of δ13C and δ15N and Bayesian mixing models in MixSIAR to determine the reliance of riparian predators on aquatic prey sources. Using mixed effects models, we found that arachnid families varied in their reliance on aquatic prey sources. While songbird species varied in their reliance on aquatic prey sources, songbirds sampled earlier in the season consistently relied more on aquatic prey sources than those sampled later in the season. For both arachnids and songbirds, we found a positive correlation between the amount of the aquatic prey source in their diet and their Hg concentrations. While the seasonal pulse of aquatic prey to terrestrial ecosystems is an important source of nutrients to riparian species, our results show that aquatic prey sources are linked with higher Hg exposure. For songbirds, reliance on aquatic prey sources early in the breeding season (and subsequent higher Hg exposure) coincides with timing of egg laying and development, both of which may be impacted by Hg exposure.

Erosion of tropical bird diversity over a century is influenced by abundance, diet and subtle climatic tolerances.

Human alteration of landscapes leads to attrition of biodiversity. Recommendations for maximizing retention of species richness typically focus on protection and preservation of large habitat patches. Despite a century of protection from human disturbance, 27% of the 228 bird species initially detected on Barro Colorado Island (BCI), Panama, a large hilltop forest fragment isolated by waters of Gatun Lake, are now absent. Lost species were more likely to be initially uncommon and terrestrial insectivores. Analyses of the regional avifauna, exhaustively inventoried and mapped across 24 subregions, identified strong geographical discontinuities in species distributions associated with a steep transisthmian rainfall gradient. Having lost mostly species preferring humid forests, the BCI species assemblage continues to shift from one originally typical of wetter forests toward one now resembling bird communities in drier forests. Even when habitat remnants are large and protected for 100 years, altered habitat characteristics resulting from isolation produce non-random loss of species linked with their commonness, dietary preferences and subtle climatic sensitivities.

The influence of rare birds on observer effort and subsequent rarity discovery in the American birdwatching community.

Birdwatching is a rapidly growing pastime, increasingly involving the pursuit of rare birds as birders build lists of species they encounter. We expected reports of rare bird discoveries to quickly draw birders to locations, and that the increased attention at those locations would decay over time. We hypothesized that magnitude of draw and rates of decay would vary depending on characteristics of the species and the geographic locations where rarities were discovered. Discoveries of additional rarities might affect both the draw and decay, so we also quantified empirical evidence for the Patagonia Picnic Table Effect (PPTE), a social feedback loop where rarity discoveries are presumed to lead to additional rarity discoveries because of the elevated levels of birder activity once an initial rarity is reported. Although commonly invoked, supporting evidence for the PPTE hypothesis is anecdotal. We used 10 years of eBird data (2008-2017) in the United States to (1) understand birding activity when rarities were reported and the factors associated with draw and decay, and (2) assess the frequency at which initial rarity discoveries lead to reports of additional rarities. Across 273 rarity events, birder effort, as indexed by numbers of eBird checklists, increased above the pre-event baseline level, with the magnitude of draw varying geographically. We found no indication that draw was influenced by species identity or rarity-level, but latitude and distance to small airport proved important in drawing additional eBirders to rare bird discoveries. Mean draw of rarities and mean number of checklists from the same locations prior to each rarity discovery grew through the ten years, suggesting an increased influence of eBird on birder behavior in general. Decay rates in birder effort were more gradual in rare bird events with longer durations. Effort declined below baseline-levels after rarities went undetected, suggesting, "location-fatigue" following rarity events. Results did not support the PPTE hypothesis. Controlling for site-specific circumstances, birders had no better chance of finding additional rarities during events than at times outside events. Our results emphasize that eBird checklist quantity at rarity events follows a predictable but variable pattern of draw and decay influenced by location and time since rarity discovery; that birders have statistically similar chances of finding rarities during normal "baseline" birding activities as they do when known rarities are present; and that eBird represents a largely untapped resource for studying factors that influence levels of birding activity.

Lianas maintain insectivorous bird abundance and diversity in a neotropical forest.

The spatial habitat heterogeneity hypothesis posits that habitat complexity increases the abundance and diversity of species. In tropical forests, lianas add substantial habitat heterogeneity and complexity throughout the vertical forest profile, which may maintain animal abundance and diversity. The effects of lianas on tropical animal communities, however, remain poorly understood. We propose that lianas have a positive effect on animals by enhancing habitat complexity. Lianas may have a particularly strong influence on the forest bird community, providing nesting substrate, protection from predators, and nutrition (food). Understory insectivorous birds, which forage for insects that specialize on lianas, may particularly benefit. Alternatively, it is possible that lianas have a negative effect on forest birds by increasing predator abundances and providing arboreal predators with travel routes with easy access to bird nests. We tested the spatial habitat heterogeneity hypothesis on bird abundance and diversity by removing lianas, thus reducing forest complexity, using a large-scale experimental approach in a lowland tropical forest in the Republic of Panama. We found that removing lianas decreased total bird abundance by 78.4% and diversity by 77.4% after 8 months, and by 40.0% and 51.7%, respectively, after 20 months. Insectivorous bird abundance and diversity 8 months after liana removal were 91.8% and 89.5% lower, respectively, indicating that lianas positively influence insectivorous birds. The effects of liana removal persisted longer for insectivorous birds than other birds, with 77.3% lower abundance and 76.2% lower diversity after 20 months. Liana removal also altered bird community composition, creating two distinct communities in the control and removal plots, with disproportionate effects on insectivores. Our findings demonstrate that lianas have a strong positive influence on the bird community, particularly for insectivorous birds in the forest understory. Lianas may maintain bird abundance and diversity by increasing habitat complexity, habitat heterogeneity, and resource availability.

Development syndromes in New World temperate and tropical songbirds.

We studied avian development in 49 to 153 species of temperate and tropical New World passerine birds to determine how growth rates, and incubation and nestling periods, varied in relation to other life-history traits. We collected growth data and generated unbiased mass and tarsus growth rate estimates (mass n = 92 species, tarsus n = 49 species), and measured incubation period (n = 151) and nestling period (n = 153), which we analyzed with respect to region, egg mass, adult mass, clutch size, parental care type, nest type, daily nest predation rate (DMR), and nest height. We investigated covariation of life-history and natural-history attributes with the four development traits after controlling for phylogeny. Species in our lowland tropical sample grew 20% (incubation period), 25% (mass growth rate), and 26% (tarsus growth rate) more slowly than in our temperate sample. Nestling period did not vary with respect to latitude, which suggests that tropical songbirds fledge in a less well-developed state than temperate species. Suboscine species typically exhibited slower embryonic and post-embryonic growth than oscine passerines regardless of their breeding region. This pattern of slow development in tropical species could reflect phylogenetic effects based on unknown physiological attributes. Time-dependent nest mortality was unrelated to nestling mass growth rate, tarsus growth rate, and incubation period, but was significantly associated with nestling period. This suggests that nest predation, the predominant cause of nest loss in songbirds, does not exert strong selection on physiologically constrained traits, such as embryonic and post-embryonic growth, among our samples of temperate and lowland tropical songbird species. Nestling period, which is evolutionarily more labile than growth rate, was significantly shorter in birds exposed to higher rates of nest loss and nesting at lower heights, among other traits. Differences in life-history variation across latitudes provide insight into how unique ecological characteristics of each region influence physiological processes of passerines, and thus, how they can shape the evolution of life histories. While development traits clearly vary with respect to latitude, trait distributions overlap broadly. Life-history and natural history associations differ for each development trait, which suggests that unique selective pressures or constraints influence the evolution of each trait.

Ecological and life-history factors influencing the evolution of maternal antibody allocation: a phylogenetic comparison.

Maternally derived yolk antibodies provide neonates with immune protection in early life at negligible cost to mothers. However, developmental effects on the neonate's future immunity are potentially costly and thus could limit yolk antibody deposition. The benefits to neonatal immunity must be balanced against costs, which may depend on neonate vulnerability to pathogens, developmental trajectories and the immunological strategies best suited to a species' pace of life. We measured yolk antibodies and life-history features of 23 species of small Neotropical birds and assessed the evidence for each of several hypotheses for life history and ecological effects on the evolution of yolk antibody levels. Developmental period and yolk antibodies are negatively related, which possibly reflect the importance of humoral immune priming through antigen exposure, and selection to avoid autoimmunity, in species with a slower pace of life. There is also a strong relationship between body size and yolk antibody concentration, suggesting that larger species are architecturally equipped to produce and transfer higher concentrations of antibodies. These results suggest that developmental effects of maternally derived antibodies, such as imprinting effects on B-cell diversity or autoimmune effects, are important and deserve more consideration in future research.

Predicting declines in avian species richness under nonrandom patterns of habitat loss in a neotropical landscape.

One of the key concerns in conservation is to document and predict the effects of habitat loss on species richness. To do this, the species-area relationship (SAR) is frequently used. That relationship assumes random patterns of habitat loss and species distributions. In nature, however, species distribution patterns are usually nonrandom, influenced by biotic and abiotic factors. Likewise, socioeconomic and environmental factors influence habitat loss and are not randomly distributed across landscapes. We used a recently developed SAR model that accounts for nonrandomness to predict rates of bird species loss in fragmented forests of the Panama Canal region, an area that was historically covered in forest but now has 53% forest cover. Predicted species loss was higher than that predicted by the standard SAR. Furthermore, a species loss threshold was evident when remaining forest cover declined by 25%. This level of forest cover corresponds to 40% of the historical forest cover, and our model predicts rapid species loss past that threshold. This study illustrates the importance of considering patterns of species distributions and realistic habitat loss scenarios to develop better estimates of losses in species richness. Forecasts of tropical biodiversity loss generated from simple species-area relationships may underestimate actual losses because nonrandom patterns of species distributions and habitat loss are probably not unique to the Panama Canal region.

Serum antioxidant levels in wild birds vary in relation to diet, season, life history strategy, and species.

Micronutrient antioxidants are thought to be generally important for health in many animals, but factors determining levels in individuals and species are not well understood. Diet and season are obvious environmental variables that might predict the degree to which species can accumulate such nutrients. We analyzed antioxidant levels [Trolox-equivalent antioxidant capacity (TEAC), uric acid (UA), vitamin E, and four carotenoids] in 95 bird species and compared these to species-level data on diet from the literature. Using compositional principal components analysis, we identified two main axes of diet variation: invertebrate consumption and seed-to-fruit ratio. We then examined associations between diet axes and antioxidant measures, with and without control for life-history variation and phylogeny. We also analyzed a subset of 13 species for which we had data on seasonality of antioxidant levels and diet, assessing the variance in antioxidant levels explained by seasonality, diet, and species. Unsurprisingly, there were strong associations between antioxidant levels and diet. TEAC and UA concentration were consistently positively associated with invertebrate consumption and seed-to-fruit ratio, and carotenoid concentrations (e.g. zeaxanthin and beta-carotene) were negatively associated with invertebrate consumption. However, vitamin E was not associated with diet as measured here. Importantly, there is much variation in antioxidants that is not explained by diet, and we are able to identify diet-independent effects of species, season/breeding stage, and life history on antioxidant levels. Circulating antioxidant concentrations within and across species can therefore be viewed as a function of multiple factors, including but not limited to diet, and antioxidant metabolism appears to differ across species and seasons irrespective of diet.

Idiosyncratic changes in spring arrival dates of Pacific Northwest migratory birds.

Shifts in the timing of bird migration have been associated with climatic change and species traits. However, climatic change does not affect all species or geographic locations equally. Climate in the Pacific Northwest has shifted during the last century with mean temperatures increasing by 1 °C but little change in total annual precipitation. Few long-term data on migration phenology of birds are available in the Pacific Northwest. We analyzed trends in spring arrival dates from a site in the Oregon Coast Range where nearly daily inventories of birds were conducted in 24 of 29 years. Several species showed statistically significant shifts in timing of first spring arrivals. Six of 18 species occur significantly earlier now than during the initial phase of the study. One species arrives significantly later. Eleven show no significant shifts in timing. We associated trends in spring migration phenology with regional climatic variables, weather (precipitation and temperature), traits of species such as migration strategy, foraging behavior, diet, and habitat use, and regional trends in abundance as indexed by Breeding Bird Survey data. We found no set of variables consistently correlated with avian phenological changes. Post hoc analyses of additional climate variables revealed an association of migratory arrival dates across the 18 species with rainfall totals in northern California, presumably indicating that songbird arrival dates in Oregon are slowed by spring storm systems in California. When only the six species with the most strongly advancing arrival dates were analyzed, winter maximum temperatures in the preceding three winters appeared consistently in top models, suggesting a possible role for food availability early in spring to promote the survival and successful reproduction of the earliest-arriving birds. However, additional data on food availability and avian survival and reproductive success are required to test that hypothesis. Despite the appearance of some climate variables in top models, there remains a mismatch between strongly advancing arrival dates in some songbirds and a lack of clear directional change in those climate variables. We conclude that either some previously unrecognized variable or combination of variables has affected the timing of migration in some species but not others, or the appearance of statistically significant directional changes over time can occur without being driven by consistent environmental or species-specific factors.

Why are incubation periods longer in the tropics? A common-garden experiment with house wrens reveals it is all in the egg.

Incubation periods of Neotropical birds are often longer than those of related species at temperate latitudes. We conducted a common-garden experiment to test the hypothesis that longer tropical incubation periods result from longer embryo development times rather than from different patterns of parental incubation behavior. House wrens, one of few species whose geographic range includes tropical equatorial and temperate high latitudes, have incubation periods averaging 1.2 days longer at tropical latitudes. We incubated eggs of house wrens in Illinois and Panama under identical conditions in mechanical incubators. Even after factoring out differences in egg size, tropical house wrens still required 1.33 days longer, on average, to hatch. We conclude that parental attendance patterns do not account for latitudinal differences in incubation period but that some other as yet unmeasured factor intrinsic to the egg or embryo, or both, extends development time in the tropics.