Demographic consequences of phenological asynchrony for North American songbirds.

Changes in phenology in response to ongoing climate change have been observed in numerous taxa around the world. Differing rates of phenological shifts across trophic levels have led to concerns that ecological interactions may become increasingly decoupled in time, with potential negative consequences for populations. Despite widespread evidence of phenological change and a broad body of supporting theory, large-scale multitaxa evidence for demographic consequences of phenological asynchrony remains elusive. Using data from a continental-scale bird-banding program, we assess the impact of phenological dynamics on avian breeding productivity in 41 species of migratory and resident North American birds breeding in and around forested areas. We find strong evidence for a phenological optimum where breeding productivity decreases in years with both particularly early or late phenology and when breeding occurs early or late relative to local vegetation phenology. Moreover, we demonstrate that landbird breeding phenology did not keep pace with shifts in the timing of vegetation green-up over a recent 18-y period, even though avian breeding phenology has tracked green-up with greater sensitivity than arrival for migratory species. Species whose breeding phenology more closely tracked green-up tend to migrate shorter distances (or are resident over the entire year) and breed earlier in the season. These results showcase the broadest-scale evidence yet of the demographic impacts of phenological change. Future climate change-associated phenological shifts will likely result in a decrease in breeding productivity for most species, given that bird breeding phenology is failing to keep pace with climate change.

Comparing multiscale, presence-only habitat suitability models created with structured survey data and community science data for a rare warbler species at the southern range margin.

Golden-winged Warblers (Vermivora chrysoptera, Parulidae) are declining migrant songbirds that breed in the Great Lakes and Appalachian regions of North America. Within their breeding range, Golden-winged Warblers are found in early successional habitats adjacent to mature hardwood forest, and previous work has found that Golden-winged Warbler habitat preferences are scale-dependent. Golden-winged Warbler Working Group management recommendations were written to apply to large regions of the breeding range, but there may be localized differences in both habitat availability and preferences. Rapid declines at the southernmost extent of their breeding range in Western North Carolina necessitate investigation into landscape characteristics governing distribution in this subregion. Furthermore, with the increase in availability of community science data from platforms such as eBird, it would be valuable to know if community science data produces similar distribution models as systemic sampling data. In this study, we described patterns of Golden-winged Warbler presence in Western North Carolina by examining habitat variables at multiple spatial scales using data from standardized Audubon North Carolina (NC) playback surveys and community science data from eBird. We compared model performance and predictions between Audubon NC and eBird models and found that Golden-winged Warbler presence is associated with sites which, at a local scale (150m), have less mature forest, more young forest, more herb/shrub cover, and more road cover, and at a landscape scale (2500m), have less herb/shrub cover. Golden-winged Warbler presence is also associated with higher elevations and smaller slopes. eBird and Audubon models had similar variable importance values, response curves, and overall performance. Based on variable importance values, elevation, mature forest at the local scale, and road cover at the local scale are the primary variables driving the difference between Golden-winged Warbler breeding sites and random background sites in Western North Carolina. Additionally, our results validate the use of eBird data, since they produce species distribution modeling results that are similar to results obtained from more standardized survey methods.

Phylogenetic climatic niche conservatism in sandflies (Diptera: Phlebotominae) and their relatives.

Phylogenetic niche conservatism is a pattern in which closely related species are more similar than distant relatives in their niche-related traits. Species in the family Psychodidae show notable diversity in climatic niche, and present an opportunity to test for phylogenetic niche conservatism, which is as yet rarely studied in insects. Some species (in the subfamily Phlebotominae) transmit Leishmania parasites, responsible for the disease leishmaniasis, and their geographic range has been systematically characterized. Psychodid genus ranges can be solely tropical, confined to the temperate zones, or span both. We obtained observation site data, and associated climate data, for 234 psychodid species to understand which aspects of climate most closely predict distribution. Temperature and seasonality are strong determinants of species occurrence within the clade. Next, we built a phylogeny of Psychodidae, and found a positive relationship between pairwise genetic distance and climate niche differentiation, which indicates strong niche conservatism. This result is also supported by strong phylogenetic signals of metrics of climate differentiation. Finally, we used ancestral trait reconstruction to infer the tropicality (i.e., proportion of latitudinal range in the tropics minus the proportion of the latitudinal range in temperate areas) of ancestral species, and counted transitions to and from tropicality states. We find that tropical and temperate species produced almost entirely tropical and temperate descendant species, respectively. Taken together, our results imply that climate niches in psychodids are strongly predicted by phylogeny, and represent a formal test of a key prediction of phylogenetic niche conservatism in a clade with implications for human health.

Citizen Science in Postsecondary Education: Current Practices and Knowledge Gaps.

Citizen science involves the public in science to investigate research questions. Although citizen science facilitates learning in informal educational settings, little is known about its use or effects in postsecondary (college or university) settings. Using a literature review and a survey, we describe how and why citizen science is being used in postsecondary courses, as well as the impacts on student learning. We found that citizen science is used predominantly in biologically related fields, at diverse types of institutions, to improve student engagement and expose students to authentic research. Considerable anecdotal evidence supporting improved student learning from these experiences exists, but little empirical evidence exists to warrant any conclusion. Therefore, there is a need to rigorously assess the relationship between citizen science participation and postsecondary student learning. We highlight considerations for instructors planning to incorporate citizen science and for citizen science projects wanting to facilitate postsecondary use.

Anthropogenic drivers of avian community turnover from local to regional scales.

Anthropogenic change has altered the composition and function of ecological communities across the globe. As a result, there is a need for studies examining observed community compositional change and determining whether and how anthropogenic change drivers may be influencing that turnover. In particular, it is also important to determine to what extent community turnover is idiosyncratic or if turnover can be explained by predictable responses across species based on traits or niche characteristics. Here, we measured turnover in avian communities across North America from 1990 to 2016 in the Breeding Bird Survey using an ordination method, and modeled turnover as a function of land use and climate change drivers from local to regional scales. We also examined how turnover may be attributed to species groups, including foraging guilds, trophic groups, migratory distance, and breeding biomes. We found that at local scales, land use change explained a greater proportion of variance in turnover than climate change variables, while as scale increased, trends in temperature explained a greater proportion of variance in turnover. We also found across the study region, turnover could be attributed to one of a handful of species undergoing strong expansions or strong declines over the study time period. We did not observe consistent patterns in compositional change in any trait groups we examined except for those that included previously identified highly influential species. Our results have two important implications: First, the relative importance of different anthropogenic change drivers may vary with scale, which should be considered in studies' modeling impacts of global change on biodiversity. Second, in North American avian communities, individual species undergoing large shifts in population may drive signals in compositional change, and composite community turnover metrics should be carefully selected as a result.

The Avian Diet Database as a source of quantitative information on bird diets.

This data paper describes a compilation of 73,075 quantitative diet data records for 759 primarily North American bird species, providing standardized information not just on the diet itself, but on the context for that diet information including the year, season, location, and habitat type of each study. The methods used for collecting and cleaning these data are described, and we present tools for summarizing and visualizing diet information by bird species or prey.

Migratory strategy drives species-level variation in bird sensitivity to vegetation green-up.

Animals and plants are shifting the timing of key life events in response to climate change, yet despite recent documentation of escalating phenological change, scientists lack a full understanding of how and why phenological responses vary across space and among species. Here, we used over 7 million community-contributed bird observations to derive species-specific, spatially explicit estimates of annual spring migration phenology for 56 bird species across eastern North America. We show that changes in the spring arrival of migratory birds are coarsely synchronized with fluctuations in vegetation green-up and that the sensitivity of birds to plant phenology varied extensively. Bird arrival responded more synchronously with vegetation green-up at higher latitudes, where phenological shifts over time are also greater. Critically, species' migratory traits explained variation in sensitivity to green-up, with species that migrate more slowly, arrive earlier and overwinter further north showing greater responsiveness to earlier springs. Identifying how and why species vary in their ability to shift phenological events is fundamental to predicting species' vulnerability to climate change. Such variation in sensitivity across taxa, with long-distance neotropical migrants exhibiting reduced synchrony, may help to explain substantial declines in these species over the last several decades.

A simulation study of the use of temporal occupancy for identifying core and transient species.

Transient species, which do not maintain self-sustaining populations in a system where they are observed, are ubiquitous in nature and their presence often impacts the interpretation of ecological patterns and processes. Identifying transient species from temporal occupancy, the proportion of time a species is observed at a given site over a time series, is subject to classification errors as a result of imperfect detection and source-sink dynamics. We use a simulation-based approach to assess how often errors in detection or classification occur in order to validate the use of temporal occupancy as a metric for inferring whether a species is a core or transient member of a community. We found that low detection increases error in the classification of core species, while high habitat heterogeneity and high detection increase error in classification of transient species. These findings confirm that temporal occupancy is a valid metric for inferring whether a species can maintain a self-sustaining population, but imperfect detection, low abundance, and highly heterogeneous landscapes may yield high misclassification rates.

Genetic Diversity and Thermal Performance in Invasive and Native Populations of African Fig Flies.

During biological invasions, invasive populations can suffer losses of genetic diversity that are predicted to negatively impact their fitness/performance. Despite examples of invasive populations harboring lower diversity than conspecific populations in their native range, few studies have linked this lower diversity to a decrease in fitness. Using genome sequences, we show that invasive populations of the African fig fly, Zaprionus indianus, have less genetic diversity than conspecific populations in their native range and that diversity is proportionally lower in regions of the genome experiencing low recombination rates. This result suggests that selection may have played a role in lowering diversity in the invasive populations. We next use interspecific comparisons to show that genetic diversity remains relatively high in invasive populations of Z. indianus when compared with other closely related species. By comparing genetic diversity in orthologous gene regions, we also show that the genome-wide landscape of genetic diversity differs between invasive and native populations of Z. indianus indicating that invasion not only affects amounts of genetic diversity but also how that diversity is distributed across the genome. Finally, we use parameter estimates from thermal performance curves for 13 species of Zaprionus to show that Z. indianus has the broadest thermal niche of measured species, and that performance does not differ between invasive and native populations. These results illustrate how aspects of genetic diversity in invasive species can be decoupled from measures of fitness, and that a broad thermal niche may have helped facilitate Z. indianus's range expansion.

A Minimal Model for the Latitudinal Diversity Gradient Suggests a Dominant Role for Ecological Limits.

The latitudinal diversity gradient (LDG) is one of Earth's most iconic biodiversity patterns and still one of the most debated. Explanations for the LDG are often categorized into three broad pathways in which the diversity gradient is created by (1) differential diversification rates, (2) differential carrying capacities (ecological limits), or (3) differential time to accumulate species across latitude. Support for these pathways has, however, been mostly verbally expressed. Here, we present a minimal model to clarify the essential assumptions of the three pathways and explore the sensitivity of diversity dynamics to these pathways. We find that an LDG arises most easily from a gradient in ecological limits compared with a gradient in the time for species accumulation or diversification rate in most modeled scenarios. Differential diversification rates create a stronger LDG than ecological limits only when speciation and dispersal rates are low, but then the predicted LDG seems weaker than the observed LDG. Moreover, range dynamics may reduce an LDG created by a gradient in diversification rates or time for species accumulation, but they cannot reduce an LDG induced by differential ecological limits. We conclude that our simple model provides a null prediction for the effectiveness of the three LDG pathways and can thus aid discussions about the causal mechanisms underlying the LDG or motivate more complex models to confirm or falsify our findings.

The Latitudinal Diversity Gradient: Novel Understanding through Mechanistic Eco-evolutionary Models.

The latitudinal diversity gradient (LDG) is one of the most widely studied patterns in ecology, yet no consensus has been reached about its underlying causes. We argue that the reasons for this are the verbal nature of existing hypotheses, the failure to mechanistically link interacting ecological and evolutionary processes to the LDG, and the fact that empirical patterns are often consistent with multiple explanations. To address this issue, we synthesize current LDG hypotheses, uncovering their eco-evolutionary mechanisms, hidden assumptions, and commonalities. Furthermore, we propose mechanistic eco-evolutionary modeling and an inferential approach that makes use of geographic, phylogenetic, and trait-based patterns to assess the relative importance of different processes for generating the LDG.

The proportion of core species in a community varies with spatial scale and environmental heterogeneity.

Ecological communities are composed of a combination of core species that maintain local viable populations and transient species that occur infrequently due to dispersal from surrounding regions. Preliminary work indicates that while core and transient species are both commonly observed in community surveys of a wide range of taxonomic groups, their relative prevalence varies substantially from one community to another depending upon the spatial scale at which the community was characterized and its environmental context. We used a geographically extensive dataset of 968 bird community time series to quantitatively describe how the proportion of core species in a community varies with spatial scale and environmental heterogeneity. We found that the proportion of core species in an assemblage increased with spatial scale in a positive decelerating fashion with a concomitant decrease in the proportion of transient species. Variation in the shape of this scaling relationship between sites was related to regional environmental heterogeneity, with lower proportions of core species at a given scale associated with high environmental heterogeneity. Understanding this influence of scale and environmental heterogeneity on the proportion of core species may help resolve discrepancies between studies of biotic interactions, resource availability, and mass effects conducted at different scales, because the importance of these and other ecological processes are expected to differ substantially between core and transient species.

The prevalence and impact of transient species in ecological communities.

Transient species occur infrequently in a community over time and do not maintain viable local populations. Because transient species interact differently than non-transients with their biotic and abiotic environment, it is important to characterize the prevalence of these species and how they impact our understanding of ecological systems. We quantified the prevalence and impact of transient species in communities using data on over 19,000 community time series spanning an array of ecosystems, taxonomic groups, and spatial scales. We found that transient species are a general feature of communities regardless of taxa or ecosystem. The proportion of these species decreases with increasing spatial scale leading to a need to control for scale in comparative work. Removing transient species from analyses influences the form of a suite of commonly studied ecological patterns including species-abundance distributions, species-energy relationships, species-area relationships, and temporal turnover. Careful consideration should be given to whether transient species are included in analyses depending on the theoretical and practical relevance of these species for the question being studied.

Aligning the Measurement of Microbial Diversity with Macroecological Theory.

The number of microbial operational taxonomic units (OTUs) within a community is akin to species richness within plant/animal ("macrobial") systems. A large literature documents OTU richness patterns, drawing comparisons to macrobial theory. There is, however, an unrecognized fundamental disconnect between OTU richness and macrobial theory: OTU richness is commonly estimated on a per-individual basis, while macrobial richness is estimated per-area. Furthermore, the range or extent of sampled environmental conditions can strongly influence a study's outcomes and conclusions, but this is not commonly addressed when studying OTU richness. Here we (i) propose a new sampling approach that estimates OTU richness per-mass of soil, which results in strong support for species energy theory, (ii) use data reduction to show how support for niche conservatism emerges when sampling across a restricted range of environmental conditions, and (iii) show how additional insights into drivers of OTU richness can be generated by combining different sampling methods while simultaneously considering patterns that emerge by restricting the range of environmental conditions. We propose that a more rigorous connection between microbial ecology and macrobial theory can be facilitated by exploring how changes in OTU richness units and environmental extent influence outcomes of data analysis. While fundamental differences between microbial and macrobial systems persist (e.g., species concepts), we suggest that closer attention to units and scale provide tangible and immediate improvements to our understanding of the processes governing OTU richness and how those processes relate to drivers of macrobial species richness.

Species richness at continental scales is dominated by ecological limits.

Explaining variation in species richness among provinces and other large geographic regions remains one of the most challenging problems at the intersection of ecology and evolution. Here we argue that empirical evidence supports a model whereby ecological factors associated with resource availability regulate species richness at continental scales. Any large-scale predictive model for biological diversity must explain three robust patterns in the natural world. First, species richness for evolutionary biotas is highly correlated with resource-associated surrogate variables, including area, temperature, and productivity. Second, species richness across epochal timescales is largely stationary in time. Third, the dynamics of diversity exhibit clear and predictable responses to mass extinctions, key innovations, and other perturbations. Collectively, these patterns are readily explained by a model in which species richness is regulated by diversity-dependent feedback mechanisms. We argue that many purported tests of the ecological limits hypothesis, including branching patterns in molecular phylogenies, are inherently weak and distract from these three core patterns. We have much to learn about the complex hierarchy of processes by which local ecological interactions lead to diversity dependence at the continental scale, but the empirical evidence overwhelmingly suggests that they do.

On the processes generating latitudinal richness gradients: identifying diagnostic patterns and predictions.

We use a simulation model to examine four of the most common hypotheses for the latitudinal richness gradient and identify patterns that might be diagnostic of those four hypotheses. The hypotheses examined include (1) tropical niche conservatism, or the idea that the tropics are more diverse because a tropical clade origin has allowed more time for diversification in the tropics and has resulted in few species adapted to extra-tropical climates. (2) The ecological limits hypothesis suggests that species richness is limited by the amount of biologically available energy in a region. (3) The speciation rates hypothesis suggests that the latitudinal gradient arises from a gradient in speciation rates. (4) Finally, the tropical stability hypothesis argues that climatic fluctuations and glacial cycles in extratropical regions have led to greater extinction rates and less opportunity for specialization relative to the tropics. We found that tropical niche conservatism can be distinguished from the other three scenarios by phylogenies which are more balanced than expected, no relationship between mean root distance (MRD) and richness across regions, and a homogeneous rate of speciation across clades and through time. The energy gradient, speciation gradient, and disturbance gradient scenarios all produced phylogenies which were more imbalanced than expected, showed a negative relationship between MRD and richness, and diversity-dependence of speciation rate estimates through time. We found that the relationship between speciation rates and latitude could distinguish among these three scenarios, with no relation expected under the ecological limits hypothesis, a negative relationship expected under the speciation rates hypothesis, and a positive relationship expected under the tropical stability hypothesis. We emphasize the importance of considering multiple hypotheses and focusing on diagnostic predictions instead of predictions that are consistent with multiple hypotheses.

Range expansion and population dynamics of an invasive species: the Eurasian Collared-Dove (Streptopelia decaocto).

Invasive species offer ecologists the opportunity to study the factors governing species distributions and population growth. The Eurasian Collared-Dove (Streptopelia decaocto) serves as a model organism for invasive spread because of the wealth of abundance records and the recent development of the invasion. We tested whether a set of environmental variables were related to the carrying capacities and growth rates of individual populations by modeling the growth trajectories of individual populations of the Collared-Dove using Breeding Bird Survey (BBS) and Christmas Bird Count (CBC) data. Depending on the fit of our growth models, carrying capacity and growth rate parameters were extracted and modeled using historical, geographical, land cover and climatic predictors. Model averaging and individual variable importance weights were used to assess the strength of these predictors. The specific variables with the greatest support in our models differed between data sets, which may be the result of temporal and spatial differences between the BBS and CBC. However, our results indicate that both carrying capacity and population growth rates are related to developed land cover and temperature, while growth rates may also be influenced by dispersal patterns along the invasion front. Model averaged multivariate models explained 35-48% and 41-46% of the variation in carrying capacities and population growth rates, respectively. Our results suggest that widespread species invasions can be evaluated within a predictable population ecology framework. Land cover and climate both have important effects on population growth rates and carrying capacities of Collared-Dove populations. Efforts to model aspects of population growth of this invasive species were more successful than attempts to model static abundance patterns, pointing to a potentially fruitful avenue for the development of improved invasive distribution models.

When should species richness be energy limited, and how would we know?

Energetic constraints are fundamental to ecology and evolution, and empirical relationships between species richness and estimates of available energy (i.e. resources) have led some to suggest that richness is energetically constrained. However, the mechanism linking energy with richness is rarely specified and predictions of secondary patterns consistent with energy-constrained richness are lacking. Here, we lay out the necessary and sufficient assumptions of a causal relationship linking energy gradients to richness gradients. We then describe an eco-evolutionary simulation model that combines spatially explicit diversification with trait evolution, resource availability and assemblage-level carrying capacities. Our model identified patterns in richness and phylogenetic structure expected when a spatial gradient in energy availability determines the number of individuals supported in a given area. A comparison to patterns under alternative scenarios, in which fundamental assumptions behind energetic explanations were violated, revealed patterns that are useful for evaluating the importance of energetic constraints in empirical systems. We use a data set on rockfish (genus Sebastes) from the northeastern Pacific to show how empirical data can be coupled with model predictions to evaluate the role of energetic constraints in generating observed richness gradients.

Opposing mechanisms drive richness patterns of core and transient bird species.

Studies of biodiversity typically assume that all species are equivalent. However, some species in a community maintain viable populations in the study area, while others occur only occasionally as transient individuals. Here we show that North American bird communities can reliably be divided into core and transient species groups and that the richness of each group is driven by different processes. The richness of core species is influenced primarily by local environmental conditions, while the richness of transient species is influenced primarily by the heterogeneity of the surrounding landscape. This demonstrates that the well-known effects of the local environment and landscape heterogeneity on overall species richness are the result of two sets of processes operating differentially on core and transient species. Models of species richness should focus on explaining two distinct patterns, those of core and transient species, rather than a single pattern for the community as a whole.

Temporal turnover in the composition of tropical tree communities: functional determinism and phylogenetic stochasticity.

The degree to which turnover in biological communities is structured by deterministic or stochastic factors and the identities of influential deterministic factors are fundamental, yet unresolved, questions in ecology. Answers to these questions are particularly important for projecting the fate of forests with diverse disturbance histories worldwide. To uncover the processes governing turnover we use species-level molecular phylogenies and functional trait data sets for two long-term tropical forest plots with contrasting disturbance histories: one forest is older-growth, and one was recently disturbed. Having both phylogenetic and functional information further allows us to parse out the deterministic influences of different ecological filters. With the use of null models we find that compositional turnover was random with respect to phylogeny on average, but highly nonrandom with respect to measured functional traits. Furthermore, as predicted by a deterministic assembly process, the older-growth and disturbed forests were characterized by less than and greater than expected functional turnover, respectively. These results suggest that the abiotic environment, which changes due to succession in the disturbed forest, strongly governs the temporal dynamics of disturbed and undisturbed tropical forests. Predicting future changes in the composition of disturbed and undisturbed forests may therefore be tractable when using a functional-trait-based approach.