Decomposing the spatial and temporal effects of climate on bird populations in northern European mountains.

The relationships between species abundance or occurrence versus spatial variation in climate are commonly used in species distribution models to forecast future distributions. Under "space-for-time substitution", the effects of climate variation on species are assumed to be equivalent in both space and time. Two unresolved issues of space-for-time substitution are the time period for species' responses and also the relative contributions of rapid- versus slow reactions in shaping spatial and temporal responses to climate change. To test the assumption of equivalence, we used a new approach of climate decomposition to separate variation in temperature and precipitation in Fennoscandia into spatial, temporal, and spatiotemporal components over a 23-year period (1996-2018). We compiled information on land cover, topography, and six components of climate for 1756 fixed route surveys, and we modeled annual counts of 39 bird species breeding in the mountains of Fennoscandia. Local abundance of breeding birds was associated with the spatial components of climate as expected, but the temporal and spatiotemporal climatic variation from the current and previous breeding seasons were also important. The directions of the effects of the three climate components differed within and among species, suggesting that species can respond both rapidly and slowly to climate variation and that the responses represent different ecological processes. Thus, the assumption of equivalent species' response to spatial and temporal variation in climate was seldom met in our study system. Consequently, for the majority of our species, space-for-time substitution may only be applicable once the slow species' responses to a changing climate have occurred, whereas forecasts for the near future need to accommodate the temporal components of climate variation. However, appropriate forecast horizons for space-for-time substitution are rarely considered and may be difficult to reliably identify. Accurately predicting change is challenging because multiple ecological processes affect species distributions at different temporal scales.

Regional variation in the impacts of the COVID-19 pandemic on the quantity and quality of data collected by the project eBird.

The COVID-19 pandemic has likely affected natural systems around the world; the curtailment of human activity has also affected the collection of data needed to identify the indirect effects of this pandemic on natural systems. We describe how the outbreak of COVID-19 disease, and associated stay-at-home orders in four political regions, have affected the quantity and quality of data collected by participants in one volunteer-based bird monitoring project, eBird. The four regions were selected both for their early and prolonged periods of mandated changes to human activity, and because of the high densities of observations collected. We compared the months of April 2020 with April in previous years. The most notable change was in the landscapes in which observations were made: in all but one region human-dominated landscapes were proportionally more common in the data in April 2020, and observations made near the rarer wetland habitat were less prevalent. We also found subtler changes in quantity of data collected, as well as in observer effort within observation periods. Finally, we found that these effects of COVID-19 disease varied across the political units, and thus we conclude that any analyses of eBird data will require region-specific examination of whether there have been any changes to the data collection process during the COVID-19 pandemic that would need to be taken into account.

Host population dynamics in the face of an evolving pathogen.

Interactions between hosts and pathogens are dynamic at both ecological and evolutionary levels. In the resultant 'eco-evolutionary dynamics' ecological and evolutionary processes affect each other. For example, the house finch Haemorhous mexicanus and its recently emerged pathogen, the bacterium Mycoplasma gallisepticum, form a system in which evidence suggests that changes in bacterial virulence through time enhance levels of host immunity in ways that drive the evolution of virulence in an arms race. We use data from two associated citizen science projects in order to determine whether this arms race has had any detectable effect at the population level in the north-eastern United States. We used data from two citizen science projects, based on observations of birds at bird feeders, which provide information on the long-term changes in sizes of aggregations of house finches (host population density), and the probabilities that these house finches have observable disease (disease prevalence). The initial emergence of M. gallisepticum caused a rapid halving of house finch densities; this was then followed by house finch populations remaining stable or slowly declining. Disease prevalence also decreased sharply after the initial emergence and has remained low, although with fluctuations through time. Surprisingly, while initially higher local disease prevalence was found at sites with higher local densities of finches, this relationship has reversed over time. The ability of a vertebrate host species, with a generation time of at least 1 year, to maintain stable populations in the face of evolved higher virulence of a bacterium, with generation times measurable in minutes, suggests that genetic changes in the host are insufficient to explain the observed population-level patterns. We suggest that acquired immunity plays an important role in the observed interaction between house finches and M. gallisepticum.

Rapid speciation via the evolution of pre-mating isolation in the Iberá Seedeater.

Behavioral isolation can catalyze speciation and permit the slow accumulation of additional reproductive barriers between co-occurring organisms. We illustrate how this process occurs by examining the genomic and behavioral bases of pre-mating isolation between two bird species (Sporophila hypoxantha and the recently discovered S. iberaensis) that belong to the southern capuchino seedeaters, a recent, rapid radiation characterized by variation in male plumage coloration and song. Although these two species co-occur without obvious ecological barriers to reproduction, we document behaviors indicating species recognition by song and plumage traits and strong assortative mating associated with genomic regions underlying male plumage patterning. Plumage differentiation likely originated through the reassembly of standing genetic variation, indicating how novel sexual signals may quickly arise and maintain species boundaries.

Benefits of protected areas for nonbreeding waterbirds adjusting their distributions under climate warming.

Climate warming is driving changes in species distributions and community composition. Many species have a so-called climatic debt, that is, shifts in range lag behind shifts in temperature isoclines. Inside protected areas (PAs), community changes in response to climate warming can be facilitated by greater colonization rates by warm-dwelling species, but also mitigated by lowering extirpation rates of cold-dwelling species. An evaluation of the relative importance of colonization-extirpation processes is important to inform conservation strategies that aim for both climate debt reduction and species conservation. We assessed the colonization-extirpation dynamics involved in community changes in response to climate inside and outside PAs. To do so, we used 25 years of occurrence data of nonbreeding waterbirds in the western Palearctic (97 species, 7071 sites, 39 countries, 1993-2017). We used a community temperature index (CTI) framework based on species thermal affinities to investigate species turnover induced by temperature increase. We determined whether thermal community adjustment was associated with colonization by warm-dwelling species or extirpation of cold-dwelling species by modeling change in standard deviation of the CTI (CTISD ). Using linear mixed-effects models, we investigated whether communities in PAs had lower climatic debt and different patterns of community change than communities outside PAs. For CTI and CTISD combined, communities inside PAs had more species, higher colonization, lower extirpation, and lower climatic debt (16%) than communities outside PAs. Thus, our results suggest that PAs facilitate 2 independent processes that shape community dynamics and maintain biodiversity. The community adjustment was, however, not sufficiently fast to keep pace with the large temperature increases in the central and northeastern western Palearctic. Our results underline the potential of combining CTI and CTISD metrics to improve understanding of the colonization-extirpation patterns driven by climate warming.

Beneficios de las Áreas Protegidas para las Aves Acuáticas No Reproductoras que Están Ajustando su Distribución Debido al Calentamiento Climático Resumen El calentamiento climático está generando cambios en la distribución y en la composición comunitaria de las especies. Muchas de ellas tienen una deuda climática, es decir, los cambios en la distribución se atrasan con respecto a los cambios en las isoclinas térmicas. Dentro de las áreas protegidas (APs), los cambios comunitarios como respuesta al calentamiento climático pueden facilitarse mediante tasas mayores de colonización por especies de climas cálidos, pero también pueden mitigarse al reducir las tasas de extirpación de las especies de climas fríos. Se requiere una evaluación de la importancia relativa de los procesos de colonización-extirpación para orientar las estrategias de conservación que buscan la reducción de la deuda climática y la conservación de las especies. Analizamos las dinámicas de colonización-extirpación que participan en los cambios comunitarios como respuesta al clima dentro y fuera de las APs. Para realizar lo anterior, usamos datos tomados durante 25 años de la presencia de aves acuáticas no reproductoras en el Paleártico occidental (97 especies, 7,071 sitios, 39 países, 1993-2017). Usamos un marco de trabajo del índice de temperatura comunitaria (ITC) basado en las afinidades térmicas de las especies para así investigar la rotación de especies inducida por el incremento en la temperatura. Determinamos si el ajuste térmico en la comunidad estuvo asociado con la colonización por especies de climas cálidos o con la extirpación de especies de climas fríos al modelar el cambio mediante una desviación estándar del ITC (ITCDS ). Con los modelos lineales de efectos mixtos investigamos si las comunidades dentro de las APs tenían una deuda climática más baja y patrones diferentes de cambio comunitario que las comunidades localizadas fuera de las APs. Con la combinación del ITC y deL ITCDS , las comunidades dentro de las APs tuvieron más especies, una mayor colonización, una menor extirpación y una deuda climática más baja (16%) que las comunidades fuera de las APs. Por lo tanto, nuestros resultados sugieren que las APs facilitan dos procesos independientes que moldean las dinámicas comunitarias y mantienen la biodiversidad. Sin embargo, el ajuste comunitario no fue lo suficientemente rápido para mantener el paso de los grandes incrementos en la temperatura de las regiones central y noreste del Paleártico occidental. Nuestros resultados resaltan el potencial que tiene la combinación de las medidas del ITC y del ICTDS para mejorar el entendimiento de los patrones de colonización-extirpación causados por el calentamiento climático.

Modeling avian full annual cycle distribution and population trends with citizen science data.

Information on species' distributions, abundances, and how they change over time is central to the study of the ecology and conservation of animal populations. This information is challenging to obtain at landscape scales across range-wide extents for two main reasons. First, landscape-scale processes that affect populations vary throughout the year and across species' ranges, requiring high-resolution, year-round data across broad, sometimes hemispheric, spatial extents. Second, while citizen science projects can collect data at these resolutions and extents, using these data requires appropriate analysis to address known sources of bias. Here, we present an analytical framework to address these challenges and generate year-round, range-wide distributional information using citizen science data. To illustrate this approach, we apply the framework to Wood Thrush (Hylocichla mustelina), a long-distance Neotropical migrant and species of conservation concern, using data from the citizen science project eBird. We estimate occurrence and abundance across a range of spatial scales throughout the annual cycle. Additionally, we generate intra-annual estimates of the range, intra-annual estimates of the associations between species and characteristics of the landscape, and interannual trends in abundance for breeding and non-breeding seasons. The range-wide population trajectories for Wood Thrush show a close correspondence between breeding and non-breeding seasons with steep declines between 2010 and 2013 followed by shallower rates of decline from 2013 to 2016. The breeding season range-wide population trajectory based on the independently collected and analyzed North American Breeding Bird Survey data also shows this pattern. The information provided here fills important knowledge gaps for Wood Thrush, especially during the less studied migration and non-breeding periods. More generally, the modeling framework presented here can be used to accurately capture landscape scale intra- and interannual distributional dynamics for broadly distributed, highly mobile species.

Using Semistructured Surveys to Improve Citizen Science Data for Monitoring Biodiversity.

Biodiversity is being lost at an unprecedented rate, and monitoring is crucial for understanding the causal drivers and assessing solutions. Most biodiversity monitoring data are collected by volunteers through citizen science projects, and often crucial information is lacking to account for the inevitable biases that observers introduce during data collection. We contend that citizen science projects intended to support biodiversity monitoring must gather information about the observation process as well as species occurrence. We illustrate this using eBird, a global citizen science project that collects information on bird occurrences as well as vital contextual information on the observation process while maintaining broad participation. Our fundamental argument is that regardless of what species are being monitored, when citizen science projects collect a small set of basic information about how participants make their observations, the scientific value of the data collected will be dramatically improved.

Seasonal abundance and survival of North America's migratory avifauna determined by weather radar.

Avian migration is one of Earth's largest processes of biomass transport, involving billions of birds. We estimated continental biomass flows of nocturnal avian migrants across the contiguous United States using a network of 143 weather radars. We show that, relative to biomass leaving in autumn, proportionally more biomass returned in spring across the southern United States than across the northern United States. Neotropical migrants apparently achieved higher survival during the combined migration and non-breeding period, despite an average three- to fourfold longer migration distance, compared with a more northern assemblage of mostly temperate-wintering migrants. Additional mortality expected with longer migration distances was probably offset by high survival in the (sub)tropics. Nearctic-Neotropical migrants relying on a 'higher survivorship' life-history strategy may be particularly sensitive to variations in survival on the overwintering grounds, highlighting the need to identify and conserve important non-breeding habitats.

Incomplete host immunity favors the evolution of virulence in an emergent pathogen.

Immune memory evolved to protect hosts from reinfection, but incomplete responses that allow future reinfection may inadvertently select for more-harmful pathogens. We present empirical and modeling evidence that incomplete immunity promotes the evolution of higher virulence in a natural host-pathogen system. We performed sequential infections of house finches with Mycoplasma gallisepticum strains of various levels of virulence. Virulent bacterial strains generated stronger host protection against reinfection than less virulent strains and thus excluded less virulent strains from infecting previously exposed hosts. In a two-strain model, the resulting fitness advantage selected for an almost twofold increase in pathogen virulence. Thus, the same immune systems that protect hosts from infection can concomitantly drive the evolution of more-harmful pathogens in nature.

Dynamic conservation for migratory species.

In an era of unprecedented and rapid global change, dynamic conservation strategies that tailor the delivery of habitat to when and where it is most needed can be critical for the persistence of species, especially those with diverse and dispersed habitat requirements. We demonstrate the effectiveness of such a strategy for migratory waterbirds. We analyzed citizen science and satellite data to develop predictive models of bird populations and the availability of wetlands, which we used to determine temporal and spatial gaps in habitat during a vital stage of the annual migration. We then filled those gaps using a reverse auction marketplace to incent qualifying landowners to create temporary wetlands on their properties. This approach is a cost-effective way of adaptively meeting habitat needs for migratory species, optimizes conservation outcomes relative to investment, and can be applied broadly to other conservation challenges.

Global change and the distributional dynamics of migratory bird populations wintering in Central America.

Understanding the susceptibility of highly mobile taxa such as migratory birds to global change requires information on geographic patterns of occurrence across the annual cycle. Neotropical migrants that breed in North America and winter in Central America occur in high concentrations on their non-breeding grounds where they spend the majority of the year and where habitat loss has been associated with population declines. Here, we use eBird data to model weekly patterns of abundance and occurrence for 21 forest passerine species that winter in Central America. We estimate species' distributional dynamics across the annual cycle, which we use to determine how species are currently associated with public protected areas and projected changes in climate and land-use. The effects of global change on the non-breeding grounds is characterized by decreasing precipitation, especially during the summer, and the conversion of forest to cropland, grassland, or peri-urban. The effects of global change on the breeding grounds are characterized by increasing winter precipitation, higher temperatures, and the conversion of forest to peri-urban. During spring and autumn migration, species are projected to encounter higher temperatures, forests that have been converted to peri-urban, and increased precipitation during spring migration. Based on current distributional dynamics, susceptibility to global change is characterized by the loss of forested habitats on the non-breeding grounds, warming temperatures during migration and on the breeding grounds, and declining summer rainfall on the non-breeding grounds. Public protected areas with low and medium protection status are more prevalent on the non-breeding grounds, suggesting that management opportunities currently exist to mitigate near-term non-breeding habitat losses. These efforts would affect more individuals of more species during a longer period of the annual cycle, which may create additional opportunities for species to respond to changes in habitat or phenology that are likely to develop under climate change.

Response of House Finches Recovered from Mycoplasma gallisepticum to Reinfection with a Heterologous Strain.

After recovery, house finches ( Haemorhous mexicanus) reinfected with the same Mycoplasma gallisepticum strain remain partially resistant to reinfection for at least 14 mo in that they recover from reinfection much more rapidly than do Mycoplasma gallisepticum-naïve birds. To test the response of birds to reinfection with a heterologous strain we performed two experiments. In a first experiment we exposed birds to one of three strains that differed in virulence. After they had recovered all were reinfected with the most virulent-strain available at the time of the experiment. In a second experiment we infected and later reinfected house finches with one of two Mycoplasma gallisepticum strains whereby we switched the order of the strain used. In both experiments, disease in birds reinfected with a more-virulent strain caused more-severe disease. Our data suggest that the observed increase in Mycoplasma gallisepticum virulence, once the disease has become endemic in free-ranging house finches is-in part-driven by increased resistance of recovered birds to strains of equal or lower virulence.

Patterns and biases in climate change research on amphibians and reptiles: a systematic review.

Climate change probably has severe impacts on animal populations, but demonstrating a causal link can be difficult because of potential influences by additional factors. Assessing global impacts of climate change effects may also be hampered by narrow taxonomic and geographical research foci. We review studies on the effects of climate change on populations of amphibians and reptiles to assess climate change effects and potential biases associated with the body of work that has been conducted within the last decade. We use data from 104 studies regarding the effect of climate on 313 species, from 464 species-study combinations. Climate change effects were reported in 65% of studies. Climate change was identified as causing population declines or range restrictions in half of the cases. The probability of identifying an effect of climate change varied among regions, taxa and research methods. Climatic effects were equally prevalent in studies exclusively investigating climate factors (more than 50% of studies) and in studies including additional factors, thus bolstering confidence in the results of studies exclusively examining effects of climate change. Our analyses reveal biases with respect to geography, taxonomy and research question, making global conclusions impossible. Additional research should focus on under-represented regions, taxa and questions. Conservation and climate policy should consider the documented harm climate change causes reptiles and amphibians.

A characterization of autumn nocturnal migration detected by weather surveillance radars in the northeastern USA.

Billions of birds migrate at night over North America each year. However, few studies have described the phenology of these movements, such as magnitudes, directions, and speeds, for more than one migration season and at regional scales. In this study, we characterize density, direction, and speed of nocturnally migrating birds using data from 13 weather surveillance radars in the autumns of 2010 and 2011 in the northeastern USA. After screening radar data to remove precipitation, we applied a recently developed algorithm for characterizing velocity profiles with previously developed methods to document bird migration. Many hourly radar scans contained windborne "contamination," and these scans also exhibited generally low overall reflectivities. Hourly scans dominated by birds showed nightly and seasonal patterns that differed markedly from those of low reflectivity scans. Bird migration occurred during many nights, but a smaller number of nights with large movements of birds defined regional nocturnal migration. Densities varied by date, time, and location but peaked in the second and third deciles of night during the autumn period when the most birds were migrating. Migration track (the direction to which birds moved) shifted within nights from south-southwesterly to southwesterly during the seasonal migration peaks; this shift was not consistent with a similar shift in wind direction. Migration speeds varied within nights, although not closely with wind speed. Airspeeds increased during the night; groundspeeds were highest between the second and third deciles of night, when the greatest density of birds was migrating. Airspeeds and groundspeeds increased during the fall season, although groundspeeds fluctuated considerably with prevailing winds. Significant positive correlations characterized relationships among bird densities at southern coastal radar stations and northern inland radar stations. The quantitative descriptions of broadscale nocturnal migration patterns presented here will be essential for biological and conservation applications. These descriptions help to define migration phenology in time and space, fill knowledge gaps in avian annual cycles, and are useful for monitoring long-term population trends of migrants. Furthermore, these descriptions will aid in assessing potential risks to migrants, particularly from structures with which birds collide and artificial lighting that disorients migrants.

Nocturnally migrating songbirds drift when they can and compensate when they must.

The shortest possible migratory route for birds is not always the best route to travel. Substantial research effort has established that birds in captivity are capable of orienting toward the direction of an intended goal, but efforts to examine how free-living birds use navigational information under conditions that potentially make direct flight toward that goal inefficient have been limited in spatiotemporal scales and in the number of individuals observed because of logistical and technological limitations. Using novel and recently developed techniques for analysis of Doppler polarimetric weather surveillance radar data, we examined two impediments for nocturnally migrating songbirds in eastern North America following shortest-distance routes: crosswinds and oceans. We found that migrants in flight often drifted sideways on crosswinds, but most strongly compensated for drift when near the Atlantic coast. Coastal migrants' tendency to compensate for wind drift also increased through the night, while no strong temporal differences were observed at inland sites. Such behaviors suggest that birds migrate in an adaptive way to conserve energy by assessing while airborne the degree to which they must compensate for wind drift.

Convergence of broad-scale migration strategies in terrestrial birds.

Migration is a common strategy used by birds that breed in seasonal environments. Selection for greater migration efficiency is likely to be stronger for terrestrial species whose migration strategies require non-stop transoceanic crossings. If multiple species use the same transoceanic flyway, then we expect the migration strategies of these species to converge geographically towards the most optimal solution. We test this by examining population-level migration trajectories within the Western Hemisphere for 118 migratory species using occurrence information from eBird. Geographical convergence of migration strategies was evident within specific terrestrial regions where geomorphological features such as mountains or isthmuses constrained overland migration. Convergence was also evident for transoceanic migrants that crossed the Gulf of Mexico or Atlantic Ocean. Here, annual population-level movements were characterized by clockwise looped trajectories, which resulted in faster but more circuitous journeys in the spring and more direct journeys in the autumn. These findings suggest that the unique constraints and requirements associated with transoceanic migration have promoted the spatial convergence of migration strategies. The combination of seasonal atmospheric and environmental conditions that has facilitated the use of similar broad-scale migration strategies may be especially prone to disruption under climate and land-use change.

Abundance models improve spatial and temporal prioritization of conservation resources.

Conservation prioritization requires knowledge about organism distribution and density. This information is often inferred from models that estimate the probability of species occurrence rather than from models that estimate species abundance, because abundance data are harder to obtain and model. However, occurrence and abundance may not display similar patterns and therefore development of robust, scalable, abundance models is critical to ensuring that scarce conservation resources are applied where they can have the greatest benefits. Motivated by a dynamic land conservation program, we develop and assess a general method for modeling relative abundance using citizen science monitoring data. Weekly estimates of relative abundance and occurrence were compared for prioritizing times and locations of conservation actions for migratory waterbird species in California, USA. We found that abundance estimates consistently provided better rankings of observed counts than occurrence estimates. Additionally, the relationship between abundance and occurrence was nonlinear and varied by species and season. Across species, locations prioritized by occurrence models had only 10-58% overlap with locations prioritized by abundance models, highlighting that occurrence models will not typically identify the locations of highest abundance that are vital for conservation of populations.

Taking a 'Big Data' approach to data quality in a citizen science project.

Data from well-designed experiments provide the strongest evidence of causation in biodiversity studies. However, for many species the collection of these data is not scalable to the spatial and temporal extents required to understand patterns at the population level. Only data collected from citizen science projects can gather sufficient quantities of data, but data collected from volunteers are inherently noisy and heterogeneous. Here we describe a 'Big Data' approach to improve the data quality in eBird, a global citizen science project that gathers bird observations. First, eBird's data submission design ensures that all data meet high standards of completeness and accuracy. Second, we take a 'sensor calibration' approach to measure individual variation in eBird participant's ability to detect and identify birds. Third, we use species distribution models to fill in data gaps. Finally, we provide examples of novel analyses exploring population-level patterns in bird distributions.

Can Observation Skills of Citizen Scientists Be Estimated Using Species Accumulation Curves?

Volunteers are increasingly being recruited into citizen science projects to collect observations for scientific studies. An additional goal of these projects is to engage and educate these volunteers. Thus, there are few barriers to participation resulting in volunteer observers with varying ability to complete the project's tasks. To improve the quality of a citizen science project's outcomes it would be useful to account for inter-observer variation, and to assess the rarely tested presumption that participating in a citizen science projects results in volunteers becoming better observers. Here we present a method for indexing observer variability based on the data routinely submitted by observers participating in the citizen science project eBird, a broad-scale monitoring project in which observers collect and submit lists of the bird species observed while birding. Our method for indexing observer variability uses species accumulation curves, lines that describe how the total number of species reported increase with increasing time spent in collecting observations. We find that differences in species accumulation curves among observers equates to higher rates of species accumulation, particularly for harder-to-identify species, and reveals increased species accumulation rates with continued participation. We suggest that these properties of our analysis provide a measure of observer skill, and that the potential to derive post-hoc data-derived measurements of participant ability should be more widely explored by analysts of data from citizen science projects. We see the potential for inferential results from analyses of citizen science data to be improved by accounting for observer skill.