Integrating data types to estimate spatial patterns of avian migration across the Western Hemisphere.

For many avian species, spatial migration patterns remain largely undescribed, especially across hemispheric extents. Recent advancements in tracking technologies and high-resolution species distribution models (i.e., eBird Status and Trends products) provide new insights into migratory bird movements and offer a promising opportunity for integrating independent data sources to describe avian migration. Here, we present a three-stage modeling framework for estimating spatial patterns of avian migration. First, we integrate tracking and band re-encounter data to quantify migratory connectivity, defined as the relative proportions of individuals migrating between breeding and nonbreeding regions. Next, we use estimated connectivity proportions along with eBird occurrence probabilities to produce probabilistic least-cost path (LCP) indices. In a final step, we use generalized additive mixed models (GAMMs) both to evaluate the ability of LCP indices to accurately predict (i.e., as a covariate) observed locations derived from tracking and band re-encounter data sets versus pseudo-absence locations during migratory periods and to create a fully integrated (i.e., eBird occurrence, LCP, and tracking/band re-encounter data) spatial prediction index for mapping species-specific seasonal migrations. To illustrate this approach, we apply this framework to describe seasonal migrations of 12 bird species across the Western Hemisphere during pre- and postbreeding migratory periods (i.e., spring and fall, respectively). We found that including LCP indices with eBird occurrence in GAMMs generally improved the ability to accurately predict observed migratory locations compared to models with eBird occurrence alone. Using three performance metrics, the eBird + LCP model demonstrated equivalent or superior fit relative to the eBird-only model for 22 of 24 species-season GAMMs. In particular, the integrated index filled in spatial gaps for species with over-water movements and those that migrated over land where there were few eBird sightings and, thus, low predictive ability of eBird occurrence probabilities (e.g., Amazonian rainforest in South America). This methodology of combining individual-based seasonal movement data with temporally dynamic species distribution models provides a comprehensive approach to integrating multiple data types to describe broad-scale spatial patterns of animal movement. Further development and customization of this approach will continue to advance knowledge about the full annual cycle and conservation of migratory birds.

Patterns of blood mercury variation in two long-distance migratory thrushes on Mount Mansfield, Vermont.

We investigated mercury (Hg) blood concentrations in Bicknell's thrush (Catharus bicknelli) and Swainson's thrush (C. ustulatus), congeneric long-distance migratory songbirds, from 2000-2017 at a montane forest site in north-central Vermont. We analyzed variation in blood Hg of both species using mixed-effects models, incorporating atmospheric wet Hg deposition data from a nearby sampling location. Although Hg deposition varied among years and seasonally, we detected no temporal trend in either atmospheric deposition or blood Hg, nor evidence of a relationship between the two. Sampling date had the strongest effect on blood Hg concentration, which declined seasonally, followed by age and sex of the individual. The data did not support an effect of species. We believe that the absence of a clear relationship between local atmospheric deposition and thrush blood Hg concentrations suggests that Hg cycling dynamics, mechanisms of transfer, and timing of uptake by montane forest biota are complex and poorly understood. The blood Hg concentrations of ~0.07-0.1 µg/g we documented in Bicknell's and Swainson's thrush are below those found to negatively impact physiological or reproductive endpoints in other invertivorous terrestrial passerines. To better evaluate the validity of Bicknell's thrush as a bioindicator of MeHg availability in montane forest ecosystems, we recommend (1) effects-based investigations, (2) a more robust understanding of Hg and MeHg cycling, (3) more clear geospatial and temporal links between Hg deposition and biotic uptake, and (4) more thorough documentation of Hg burdens across the species' annual cycle.

Short-term effects of hurricanes Maria and Irma on forest birds of Puerto Rico.

We compared occupancy in local assemblages of birds in forested areas across Puerto Rico during a winter before (2015) and shortly after (2018) the passage of hurricanes Irma and Maria. Using dynamic community models analyzed within a Bayesian framework, we found significant changes in detectability, with some species becoming more readily detected after the storms and others becoming more difficult to detect during surveys. Changes in occupancy were equally mixed. Five species-mostly granivores and omnivores, but also Black-whiskered Vireo (Vireo altiloquus), a migratory insectivore-occupied more sites in 2018 than in 2015. Twelve species were less common after the hurricanes, including all of the obligate frugivores. Declines in site-occupancy rates were not only more common than increases, but tended to be of greater magnitude. Our results support the general conclusions that bird species respond largely independently to changes in forest structure caused by hurricanes, but that some dietary guilds, notably frugivores, are more sensitive and more likely to show changes in abundance or occupancy following strong storms.

Surveys of Forest Birds on Puerto Rico, 2015.

BACKGROUND: The island of Puerto Rico supports a diverse assemblage of breeding birds, including 16 endemic species (Raffaele et al. 1998), and provides critical wintering habitat for many North American migratory birds (Wunderle and Waide 1994). Despite being a hotspot of avian biodiversity, spatially extensive data on the distribution and abundance of birds on the island are scarce. Breeding-bird assemblages were sampled by the North American Breeding Bird Survey from 1997-2007 (Sauer et al. 2013), but comparable primary data are not available for bird assemblages present during the boreal winter. NEW INFORMATION: We provide data from one of the few spatially extensive surveys of forest birds on Puerto Rico. We sampled 211 locations in forests across the island during January-March 2015 using repeated point-count surveys. These data are suitable for use in estimating abundance, occupancy, and distribution of forest birds on Puerto Rico during the winter.

Assessing conservation status of resident and migrant birds on Hispaniola with mist-netting.

We analyzed temporal trends in mist-net capture rates of resident (n = 8) and overwintering Nearctic-Neotropical migrant (n = 3) bird species at two sites in montane broadleaf forest of the Sierra de Bahoruco, Dominican Republic, with the goal of providing quantitative information on population trends that could inform conservation assessments. We conducted sampling at least once annually during the winter months of January-March from 1997 to 2010. We found evidence of declines in capture rates for three resident species, including one species endemic to Hispaniola. Capture rate of Rufous-throated Solitaire (Myadestes genibarbis) declined by 3.9% per year (95% CL = 0%, 7.3%), Green-tailed Ground-Tanager (Microligea palustris) by 6.8% (95% CL = 3.9%, 8.8%), and Greater Antillean Bullfinch (Loxigilla violacea) by 4.9% (95% CL = 0.9%, 9.2%). Two rare and threatened endemics, Hispaniolan Highland-Tanager (Xenoligea montana) and Western Chat-Tanager (Calyptophilus tertius), showed statistically significant declines, but we have low confidence in these findings because trends were driven by exceptionally high capture rates in 1997 and varied between sites. Analyses that excluded data from 1997 revealed no trend in capture rate over the course of the study. We found no evidence of temporal trends in capture rates for any other residents or Nearctic-Neotropical migrants. We do not know the causes of the observed declines, nor can we conclude that these declines are not a purely local phenomenon. However, our findings, along with other recent reports of declines in these same species, suggest that a closer examination of their conservation status is warranted. Given the difficulty in obtaining spatially extensive, long-term estimates of population change for Hispaniolan birds, we suggest focusing on other metrics of vulnerability that are more easily quantified yet remain poorly described, such as extent of occurrence.

Transoceanic migration by a 12 g songbird.

Many fundamental aspects of migration remain a mystery, largely due to our inability to follow small animals over vast spatial areas. For more than 50 years, it has been hypothesized that, during autumn migration, blackpoll warblers (Setophaga striata) depart northeastern North America and undertake a non-stop flight over the Atlantic Ocean to either the Greater Antilles or the northeastern coast of South America. Using miniaturized light-level geolocators, we provide the first irrefutable evidence that the blackpoll warbler, a 12 g boreal forest songbird, completes an autumn transoceanic migration ranging from 2270 to 2770 km (mean ± s.d.: 2540 ± 257) and requiring up to 3 days (62 h ± 10) of non-stop flight. This is one of the longest non-stop overwater flights recorded for a songbird and confirms what has long been believed to be one of the most extraordinary migratory feats on the planet.

Avian, salamander, and forest floor mercury concentrations increase with elevation in a terrestrial ecosystem.

High-elevation ecosystems of the northeastern United States are vulnerable to deposition and environmental accumulation of atmospheric pollutants, yet little work has been done to assess mercury (Hg) concentrations in organisms occupying montane ecosystems. The authors present data on Hg concentrations in ground-foraging insectivorous songbirds, a terrestrial salamander, and forest floor horizons sampled along a forested elevational gradient from 185 m to 1273 m in the Catskill Mountains, New York, USA. Mean Hg concentrations in Catharus thrushes and the salamander Plethodon cinereus increased with elevation, as did Hg concentrations in all forest floor horizons. Mean Hg concentrations in organic soils at approximately 1200 m elevation (503.5 ± 17.7 ng/g, dry wt) were 4.4-fold greater than those at approximately 200 m. Montane ecosystems of the northeastern United States, and probably elsewhere, are exposed to higher levels of atmospheric Hg deposition as reflected in accumulation patterns in the forest floor and associated high-elevation fauna. This information can be used to parameterize and test Hg transport and bioaccumulation models of landscape-specific patterns and may serve as a monitoring tool for decision makers considering future controls on Hg emissions. Further investigation is needed into the potential effects of increased Hg concentrations on high-elevation fauna.

A winter distribution model for Bicknell's Thrush (Catharus bicknelli), a conservation tool for a threatened migratory songbird.

Conservation planning and implementation require identifying pertinent habitats and locations where protection and management may improve viability of targeted species. The winter range of Bicknell's Thrush (Catharus bicknelli), a threatened Nearctic-Neotropical migratory songbird, is restricted to the Greater Antilles. We analyzed winter records from the mid-1970s to 2009 to quantitatively evaluate winter distribution and habitat selection. Additionally, we conducted targeted surveys in Jamaica (n = 433), Cuba (n = 363), Dominican Republic (n = 1,000), Haiti (n = 131) and Puerto Rico (n = 242) yielding 179 sites with thrush presence. We modeled Bicknell's Thrush winter habitat selection and distribution in the Greater Antilles in Maxent version 3.3.1. using environmental predictors represented in 30 arc second study area rasters. These included nine landform, land cover and climatic variables that were thought a priori to have potentially high predictive power. We used the average training gain from ten model runs to select the best subset of predictors. Total winter precipitation, aspect and land cover, particularly broadleaf forests, emerged as important variables. A five-variable model that contained land cover, winter precipitation, aspect, slope, and elevation was the most parsimonious and not significantly different than the models with more variables. We used the best fitting model to depict potential winter habitat. Using the 10 percentile threshold (>0.25), we estimated winter habitat to cover 33,170 km(2), nearly 10% of the study area. The Dominican Republic contained half of all potential habitat (51%), followed by Cuba (15.1%), Jamaica (13.5%), Haiti (10.6%), and Puerto Rico (9.9%). Nearly one-third of the range was found to be in protected areas. By providing the first detailed predictive map of Bicknell's Thrush winter distribution, our study provides a useful tool to prioritize and direct conservation planning for this and other wet, broadleaf forest specialists in the Greater Antilles.

Mercury concentrations in tropical resident and migrant songbirds on Hispaniola.

Despite growing concerns over mercury (Hg) exposure to humans and wildlife on a global scale, little is known about Hg bioaccumulation in the New World tropics. From 2005 to 2011, we monitored Hg concentrations in blood of nine avian species occupying a geographic range of tropical wet broadleaf sites on the island of Hispaniola, including eight passerines (two Nearctic-Neotropical migrant and six resident species) and one top order predatory accipiter. Invertivorous songbirds were further differentiated by foraging guild, with six species of ground-foragers and two species of foliage-gleaners. Blood Hg concentrations were orders of magnitude higher in birds sampled in central and southern cloud forest sites (1,000-1,800 m elevation) than in northern and northeastern rainforest sites (50-500 m elevation), with migratory and resident species both showing 2-20× greater blood Hg concentrations in cloud forests than in rainforests. Within cloud forest sites, ground-foraging species had higher Hg concentrations than foliage-gleaning species. Top order predatory sharp-shinned hawks (Accipiter striatus) had the highest blood Hg concentrations among all species, suggesting that Hg biomagnification is occurring in terrestrial forests of Hispaniola. Two migrant songbird species overwintering on the island had higher blood Hg concentrations than have been recorded on their North American breeding grounds. Future studies should seek to elucidate sources of variation in atmospheric Hg deposition on Hispaniola and to quantify the dynamics of Hg cycling in tropical forest ecosystems, which may differ in important ways from patterns documented in temperate forest ecosystems.

Ancient islands and modern invasions: disparate phylogeographic histories among Hispaniola's endemic birds.

With its large size, complex topography and high number of avian endemics, Hispaniola appears to be a likely candidate for the in situ speciation of its avifauna, despite the worldwide rarity of avian speciation within single islands. We used multilocus comparative phylogeography techniques to examine the pattern and history of divergence in 11 endemic birds representing potential within-island speciation events. Haplotype and allele networks from mitochondrial ND2 and nuclear intron loci reveal a consistent pattern: phylogeographic divergence within or between closely related species is correlated with the likely distribution of ancient sea barriers that once divided Hispaniola into several smaller paleo-islands. Coalescent and mitochondrial clock dating of divergences indicate species-specific response to different geological events over the wide span of the island's history. We found no evidence that ecological or topographical complexity generated diversity, either by creating open niches or by restricting long-term gene flow. Thus, no true within-island speciation appears to have occurred among the species sampled on Hispaniola. Divergence events predating the merging of Hispaniola's paleo-island blocks cannot be considered in situ divergence, and postmerging divergence in response to episodic island segmentation by marine flooding probably represents in situ vicariance or interarchipelago speciation by dispersal. Our work highlights the necessity of considering island geologic history while investigating the speciation-area relationship in birds and other taxa.

Mercury bioaccumulation and trophic transfer in the terrestrial food web of a montane forest.

We investigated mercury (Hg) concentrations in a terrestrial food web in high elevation forests in Vermont. Hg concentrations increased from autotrophic organisms to herbivores < detritivores < omnivores < carnivores. Within the carnivores studied, raptors had higher blood Hg concentrations than their songbird prey. The Hg concentration in the blood of the focal study species, Bicknell's thrush (Catharus bicknelli), varied over the course of the summer in response to a diet shift related to changing availability of arthropod prey. The Bicknell's thrush food web is more detrital-based (with higher Hg concentrations) in early summer and more foliage-based (with lower Hg concentrations) during late summer. There were significant year effects in different ecosystem compartments indicating a possible connection between atmospheric Hg deposition, detrital-layer Hg concentrations, arthropod Hg concentrations, and passerine blood Hg concentrations.

Ancient differentiation in the single-island avian radiation of endemic Hispaniolan chat-tanagers (Aves: Calyptophilus).

The simple geographic structure of island systems often makes them tractable for studies of the patterns and processes of biological diversification. The Calyptophilus chat-tanagers of Hispaniola are of general evolutionary interest because their multiple lineages might have arisen on a single island, of conservation concern because several isolated populations are nearly extinct, and taxonomically ambiguous because they have been variously lumped or split into one to four species. To explore the context of diversification of the seven extant Calyptophilus populations, we conducted a multilocus coalescent analysis based on sequences of mitochondrial ND2 and three nuclear intron loci. We then compared patterns of phylogeographic genetic variation with the morphological differences that distinguish these populations. Mitochondrial haplotypes formed two reciprocally monophyletic groups separated by a large magnitude of nucleotide divergence. Intron structure largely paralleled the geographic grouping pattern of the mitochondrial DNA (mtDNA), but these groups were only reciprocally monophyletic at one of the three introns. Also, the magnitude of between-group divergence was much lower in the introns than mtDNA genealogies. Multilocus coalescent analyses inferred a nonzero divergence time between these two major geographic groups, but suggested that they have experienced a low level of gene flow. All four markers showed substantial allele sharing within each of the two groups, demonstrating that many now separated montane populations do not have long histories of isolation. Considered in concert, our multilocus phylogeographic reconstructions support the recognition of two species within the Calyptophilus complex, and raise the possibility that these taxa differentiated prior to the fusion of the two palaeo-islands that form present-day Hispaniola.

Mercury concentrations in Bicknell's thrush and other insectivorous passerines in Montane forests of northeastern North America.

Anthropogenic input of mercury (Hg) into the environment has elevated risk to fish and wildlife, particularly in northeastern North America. Investigations into the transfer and fate of Hg have focused on inhabitants of freshwater aquatic ecosystems, as these are the habitats at greatest risk for methylmercury (MeHg) biomagnification. Deviating from such an approach, we documented MeHg availability in a terrestrial montane ecosystem using a suite of insectivorous passerines. Intensive and extensive sampling of Bicknell's thrush (Catharus bicknelli) indicated significant heterogeneity in MeHg availability across 21 mountaintops in northeastern North America. Southern parts of the breeding range tended to be at greater risk than northern parts. Mean blood Hg concentrations for Bicknell's thrush at 21 distinct breeding sites ranged from 0.08 to 0.38 ug/g (ww) and at seven Greater Antillean wintering sites ranged from 0.03 to 0.42 ug/g (ww). Overall concentrations were significantly greater in wintering than in breeding areas. Mercury exposure profiles for four passerine species on Mt. Mansfield, Vermont indicated greatest MeHg uptake in Bicknell's thrush and yellow-rumped warbler (Dendroica coronata) and lowest in blackpoll warbler (Dendroica striata) and white-throated sparrow (Zonotrichia albicollis). The MeHg and total Hg ratio in blood in these four species was nearly 1:1. There was no correlation between blood and feather Hg concentrations in breeding Bicknell's thrush, in part because of apparent retention of winter Hg body burdens, within-season variation of MeHg availability, and confounding factors such as influences from age. Adult thrushes had significantly higher concentrations of feather Hg than did young-of-the-year. Although individual patterns of inter-year feather Hg concentrations were disordered, some individuals exhibited bioaccumulation of MeHg. Female blood Hg concentrations were significantly lower than males', in part because females have additional depurating mechanisms through eggs. Older male Bicknell's thrushes that breed in New England are therefore likely at greatest risk. Mechanisms for Hg methylation in montane areas without standing water are not yet fully understood. However, recent studies indicate that MeHg is present in forest tree leaves and leaf detritus; saturated soils and other moist microhabitats may also contribute to MeHg availability. Our finding of a correlation between regional litterfall Hg flux patterns and Bicknell's thrush blood Hg concentrations demonstrates on-site availability of MeHg. Further investigations into MeHg availability in montane environments are recommended to assess risk to insectivorous passerines, particularly the Bicknell's thrush.