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Shared leadership can promote success in collaborative research networks in ecology

1. While collaborative science is becoming the norm in ecology, many ecologists participating in collaborations are less aware of the body of research that studies the processes by which collaborative teams organize and communicate. 2. Here, we discuss how we successfully used a shared leadership model in the Dry Rivers Research Coordination Network. We discuss how this model promote dour success in different stages of the project, using the Tuckman model of team development: forming, storming, norming, performing and adjourning. 3. Shared leadership in the forming phase helped us recruit a diverse membership from different scientific disciplines. In the storming and norming phases, shared leadership was especially useful in ensuring that all voices were heard in establishing group norms that promoted adhesion among and investment by RCN members. Shared leadership in the performing phase was crucial in providing opportunities for early career members to lead projects, and in the adjourning phase we reflected upon our entire collaboration to identify that shared leadership was crucial to our success, generating the thesis for this commentary. 4. It is our hope that others may find this discussion of our experience in implementing a shared leadership model useful in developing their own fruitful collaborations.

Functional Ecology

From theoretical to actual ecosystem services: mapping beneficiaries and spatial flows in ecosystem service assessments

Ecosystem services mapping and modeling has focused more on supply than demand, until recently. Whereas the potential provision of economic benefits from ecosystems to people is often quantified through ecological production functions, the use of and demand for ecosystem services has received less attention, as have the spatial flows of services from ecosystems to people. However, new modeling approaches that map and quantify service-specific sources (ecosystem capacity to provide a service), sinks (biophysical or anthropogenic features that deplete or alter service flows), users (user locations and level of demand), and spatial flows can provide a more complete understanding of ecosystem services. Through a case study in Puget Sound, Washington State, USA, we quantify and differentiate between the theoretical or in situ provision of services, i.e., ecosystems’ capacity to supply services, and their actual provision when accounting for the location of beneficiaries and the spatial connections that mediate service flows between people and ecosystems. Our analysis includes five ecosystem services: carbon sequestration and storage, riverine flood regulation, sediment regulation for reservoirs, open space proximity, and scenic viewsheds. Each ecosystem service is characterized by different beneficiary groups and means of service flow. Using the ARtificial Intelligence for Ecosystem Services (ARIES) methodology we map service supply, demand, and flow, extending on simpler approaches used by past studies to map service provision and use. With the exception of the carbon sequestration service, regions that actually provided services to people, i.e., connected to beneficiaries via flow paths, amounted to 16-66% of those theoretically capable of supplying services, i.e., all ecosystems across the landscape. These results offer a more complete understanding of the spatial dynamics of ecosystem services and their effects, and may provide a sounder basis for economic valuation and policy applications than studies that consider only theoretical service provision and/or use.

Washington

Fat or lean: adjustment of endogenous energy stores to predictable and unpredictable changes in allostatic load

1. The ability to store energy endogenously is an important ecological mechanism that allows animals to buffer predictable and unpredictable variation in allostatic load. The secretion of glucocorticoids, which reflects changes in allostatic load, is suggested to play a major role in the adjustment of endogenous stores to these varying conditions. 2. Although crucially important, the relationship between allostatic load and energy stores remains largely unexplored. Two contrasting hypotheses describe how stores may be adjusted: animals may use low allostatic loads to increase stores to a maximum possible (‘fat and fit’), or they can attain a lean physique due to fitness advantages of a low body mass (‘lean and fit’). 3. We compiled observational and experimental data available for a long-lived seabird to examine the relationship between glucocorticoids and stored energy at two life history stages (incubation and chick-rearing). Data were collected across multiple years and colonies in the North Pacific, thereby reflecting the wide range of environmental conditions birds' encounter in the marine environment. During experimental manipulations, allostatic load was minimized by supplementing food to free-living birds. 4. We found that the relationship between allostatic load and energy stores was clearly curvilinear at both life history stages. Observational data suggested that energy stores remained relatively stable under low allostatic load and decreased under high loads. Experimental data showed that birds did not maximize energy stores under favourable conditions but maintained energy stores below a physiologically attainable level. 5. Energy stores remained consistently lower during chick-rearing compared to incubation across the wide range of variations in allostatic load suggesting that stage-specific trade-offs limit the accumulation of energy during favourable environmental conditions. Secretion of glucocorticoids did not appear to mediate this shift in energy stores between the life history stages. 6. Overall, results of this study support the ‘lean and fit’ hypothesis. We conclude that increased energy stores may not necessarily reflect better environmental conditions experienced by individuals or predict their higher fitness. A major advantage of adopting a lean physique when environmental conditions allow may be the avoidance of additional energetic costs for moving a heavy body. In breeding seabirds, this advantage may be more important during chick-rearing. In the focal species, the secretion of glucocorticoids might be involved in regulation of energy stores within a life history stage but does not appear to mediate an adaptive shift in energy stores between the incubating and chick-rearing stages of reproduction.

Functional Ecology

Causal networks clarify productivity-richness interrelations, bivariate plots do not

Perhaps no other pair of variables in ecology has generated as much discussion as species richness and ecosystem productivity, as illustrated by the reactions by Pierce (2013) and others to Adler et al.'s (2011) report that empirical patterns are weak and inconsistent. Adler et al. (2011) argued we need to move beyond a focus on simplistic bivariate relationships and test mechanistic, multivariate causal hypotheses. We feel the continuing debate over productivity–richness relationships (PRRs) provides a focused context for illustrating the fundamental difficulties of using bivariate relationships to gain scientific understanding. Pierce (2013) disputes Adler et al.'s (2011) conclusion that bivariate productivity–richness relationships (PRRs) are ‘weak and variable’. He argues, instead, that relationships in the Adler et al. data are actually strong and, further, that failure to adhere to the humped-back model (HBM; sensu Grime 1979) threatens scientists' ability to advise conservationists. Here, we show that Pierce's reanalyses are invalid, that statistically significant boundary relations in the Adler et al. data are difficult to detect when proper methods are used and that his advice neither advances scientific understanding nor provides the quantitative forecasts actually needed by decision makers. We begin by examining Grimes' HBM through the lens of causal networks. We first translate the ideas contained in the HBM into a causal diagram, which shows explicitly how multiple processes are hypothesized to control biomass production and richness and their interrelationship. We then evaluate the causal diagram using structural equation modelling and example data from a published study of meadows in Finland. Formal analysis rejects the literal translation of the HBM and reveals additional processes at work. This exercise shows how the practice of abstracting systems as causal networks (i) clarifies possible hypotheses, (ii) permits explicit testing and (iii) provides more powerful and useful predictions. Building on the Finnish meadow example, we contrast the utility of bivariate plots compared with structural equation models for investigating underlying processes. Simulations illustrate the fallibility of bivariate analysis as a means of supporting one theory over another, while models based on causal networks can quantify the sensitivity of diversity patterns to both management and natural constraints. A key piece of Pierce's critique of Adler et al.'s conclusions relies on upper boundary regression which he claims to reveal strong relationships between production and richness in Adler et al.'s original data. We demonstrate that this technique shows strong associations in purely random data and is invalid for Adler et al.'s data because it depends on a uniform data distribution. We instead perform quantile regression on both the site-level summaries of the data and the plot-level data (using mixed-model quantile regression). Using a variety of nonlinear curve-fitting approaches, we were unable to detect a significant humped-shape boundary in the Adler et al. data. We reiterate that the bivariate productivity–richness relationships in Adler et al.'s data are weak and variable. We urge ecologists to consider productivity–richness relationships through the lens of causal networks to advance our understanding beyond bivariate analysis. Further, we emphasize that models based on a causal network conceptualization can also provide more meaningful guidance for conservation management than can a bivariate perspective. Measuring only two variables does not permit the evaluation of complex ideas nor resolve debates about underlying mechanisms.

Functional Ecology

Community distance sampling models allowing for imperfect detection and temporary emigration

Recent developments of community abundance models (CAMs) enable us to analyze communities subject to imperfect detection. However, existing CAMs assume spatial closure, that is, that individuals are always present in the sampling plots, which is often violated in field surveys. Violation of this assumption, such as in the presence of spatial temporary emigration, can lead to the underestimates of detection probability and overestimates of population densities and diversity metrics. Here, we propose a model that simultaneously accommodates both temporary emigration and imperfect detection by integrating CAMs and a form of hierarchical distance sampling for open populations. Expected values of species richness are obtained via the summation of occupancy (or incidence) probabilities, based on species‐level densities, across all species of the community. Simulations were used to examine the effects of spatial temporary emigration on the estimation of biological communities. We also applied the proposed model to empirical data and constructed area‐based rarefaction curves accounting for temporary emigration. Simulation experiments showed that temporary emigration can decrease the local species richness (α diversity) based on densities and increase the species turnover (β diversity). Raw species counts can overestimate or underestimate α diversity in the presence of temporary emigration, but the specific biases depend on the values of detection and emigration probabilities. Our newly proposed model yielded unbiased estimates of α, β, and γ diversity in the presence of temporary emigration. The application to empirical data suggested that accounting for temporary emigration lowered area‐based rarefaction curves because availability probabilities of individual species were estimated to be <1. Temporary emigration prevails in field surveys and has broad significance for understanding the ecology and function of biological communities and separation of imperfect detection and temporary emigration resolves long‐standing issues in the use of count data. We therefore suggest that the consideration of temporary emigration would contribute to understanding the nature and role of biological communities.

Ecosphere

Development of a multimetric index for integrated assessment of salt marsh ecosystem condition

Tools for assessing and communicating salt marsh condition are essential to guide decisions aimed at maintaining or restoring ecosystem integrity and services. Multimetric indices (MMIs) are increasingly used to provide integrated assessments of ecosystem condition. We employed a theory-based approach that considers the multivariate relationship of metrics with human disturbance to construct a salt marsh MMI for five National Parks in the northeastern USA. We quantified the degree of human disturbance for each marsh using the first principal component score from a principal components analysis of physical, chemical, and land use stressors. We then applied a metric selection algorithm to different combinations of about 45 vegetation and nekton metrics (e.g., species abundance, species richness, and ecological and functional classifications) derived from multi-year monitoring data. While MMIs derived from nekton or vegetation metrics alone were strongly correlated with human disturbance ( r values from −0.80 to −0.93), an MMI derived from both vegetation and nekton metrics yielded an exceptionally strong correlation with disturbance ( r = −0.96). Individual MMIs included from one to five metrics. The metric-assembly algorithm yielded parsimonious MMIs that exhibit the greatest possible correlations with disturbance in a way that is objective, efficient, and reproducible.

Estuaries and Coasts

Effects of an exotic prey species on a native specialist: Example of the snail kite

Despite acknowledging that exotic species can exhibit tremendous influence over native populations, few case studies have clearly demonstrated the effects of exotic prey species on native predators. We examined the effects of the recently introduced island apple snail ( Pomacea insularum ) on the foraging behavior and energetics of the endangered snail kite ( Rostrhamus sociabilis plumbeus ) in Florida. We conducted time-activity budgets: (i) on kites foraging for native Florida apple snails ( Pomacea paludosa ) in major wetland units within the kites’ range that had not been invaded by the exotic island apple snail and (ii) on kites foraging for exotic apple snails in Lake Tohopekaliga, the only major wetland utilized by the snail kite that had suffered a serious invasion of P. insularum . When foraging for P. insularum , snail kites dropped a greater proportion of snails, and they experienced increased handling times and decreased consumption rates; however, kites foraging for P. insularum also spent a smaller proportion of the day in flight. Estimates of net daily energy balances between kites feeding on P. insularum versus P. paludosa were comparable for adults, but juveniles experienced energetic deficiencies when feeding on the exotic snail. Due to this discrepancy, we hypothesize that wetlands invaded by P. insularum , such as Lake Tohopekaliga, may function as ecological traps for the snail kite in Florida by attracting breeding adults but simultaneously depressing juvenile survival. This study highlights the conservation implications and importance of elucidating the effects that exotic species have on native specialists, especially those that are endangered, because subtle influences on behavior may have significant population consequences.

Biological Conservation

Plant phenolics and absorption features in vegetation reflectance spectra near 1.66 μm

Past laboratory and field studies have quantified phenolic substances in vegetative matter from reflectance measurements for understanding plant response to herbivores and insect predation. Past remote sensing studies on phenolics have evaluated crop quality and vegetation patterns caused by bedrock geology and associated variations in soil geochemistry. We examined spectra of pure phenolic compounds, common plant biochemical constituents, dry leaves, fresh leaves, and plant canopies for direct evidence of absorption features attributable to plant phenolics. Using spectral feature analysis with continuum removal, we observed that a narrow feature at 1.66 &mu;m is persistent in spectra of manzanita, sumac, red maple, sugar maple, tea, and other species. This feature was consistent with absorption caused by aromatic C- H bonds in the chemical structure of phenolic compounds and non-hydroxylated aromatics. Because of overlapping absorption by water, the feature was weaker in fresh leaf and canopy spectra compared to dry leaf measurements. Simple linear regressions of feature depth and feature area with polyphenol concentration in tea resulted in high correlations and low errors (% phenol by dry weight) at the dry leaf ( r 2 = 0.95, RMSE = 1.0%, n = 56), fresh leaf ( r 2 = 0.79, RMSE = 2.1%, n = 56), and canopy ( r 2 = 0.78, RMSE = 1.0%, n = 13) levels of measurement. Spectra of leaves, needles, and canopies of big sagebrush and evergreens exhibited a weak absorption feature centered near 1.63 &mu;m, short ward of the phenolic compounds, possibly consistent with terpenes. This study demonstrates that subtle variation in vegetation spectra in the shortwave infrared can directly indicate biochemical constituents and be used to quantify them. Phenolics are of lesser abundance compared to the major plant constituents but, nonetheless, have important plant functions and ecological significance. Additional research is needed to advance our understanding of the spectral influences of plant phenolics and terpenes relative to dominant leaf biochemistry (water, chlorophyll, protein/nitrogen, cellulose, and lignin).

International Journal of Applied Earth Observation

Pinyon and juniper encroachment into sagebrush ecosystems impacts distribution and survival of greater sage-grouse

In sagebrush ( Artemisia spp.) ecosystems, encroachment of pinyon ( Pinus spp.) and juniper ( Juniperus spp.; hereafter, “pinyon-juniper”) trees has increased dramatically since European settlement. Understanding the impacts of this encroachment on behavioral decisions, distributions, and population dynamics of greater sage-grouse (Centrocercus urophasianus) and other sagebrush obligate species could help benefit sagebrush ecosystem management actions. We employed a novel two-stage Bayesian model that linked avoidance across different levels of pinyon-juniper cover to sage-grouse survival. Our analysis relied on extensive telemetry data collected across 6 yr and seven subpopulations within the Bi-State Distinct Population Segment (DPS), on the border of Nevada and California. The first model stage indicated avoidance behavior for all canopy cover classes on average, but individual grouse exhibited a high degree of heterogeneity in avoidance behavior of the lowest cover class (e.g., scattered isolated trees). The second stage modeled survival as a function of estimated avoidance parameters and indicated increased survival rates for individuals that exhibited avoidance of the lowest cover class. A post hoc frailty analysis revealed the greatest increase in hazard (i.e., mortality risk) occurred in areas with scattered isolated trees consisting of relatively high primary plant productivity. Collectively, these results provide clear evidence that local sage-grouse distributions and demographic rates are influenced by pinyon-juniper, especially in habitats with higher primary productivity but relatively low and seemingly benign tree cover. Such areas may function as ecological traps that convey attractive resources but adversely affect population vital rates. To increase sage-grouse survival, our model predictions support reducing actual pinyon-juniper cover as low as 1.5%, which is lower than the published target of 4.0%. These results may represent effects of pinyon-juniper cover in areas with similar ecological conditions to those of the Bi-State DPS, where populations occur at relatively high elevations and pinyon-juniper is abundant and widespread.

Rangeland Ecology and Management

Evaluation of the gap intercept method to measure rangeland connectivity

Characterizing the connectivity of materials, organisms, and energy on rangelands is critical to understanding and managing ecosystem response to disturbances. For over twenty years, scientists and rangeland managers have used the gap intercept method to monitor connectivity. However, using gap intercept measurements to infer ecosystem processes or inform management actions and conservation practices on rangelands has been limited because there are few tools and syntheses to help managers pragmatically interpret gap measurements. In this synthesis, we review the different ways the gap intercept method is conducted and highlight considerations when collecting and using gap intercept data. We discuss what is known about how gap intercept data can be used to assess wind and water erosion, biocrust abundance and composition, soil fertility, plant community dynamics, wildlife habitat characteristics, and fuel connectivity and fire behavior. Finally, we identify emerging opportunities regarding the collection and use of gap intercept data to assess rangeland connectivity, ecosystem function, and ecological processes. Through this synthesis we demonstrate the value of the gap intercept method, particularly “all-plant” (i.e., whole-community) canopy gap, in supporting assessments of rangeland condition and vulnerability, and in planning future management actions.

Rangeland Ecology & Management

The impact of grazing on plant fractal architecture and fitness of a mediterranean shrub (Anthyllis cytisoides L.)

1. We examined natural grazing by livestock (sheep and goats) on Albaida Anthyllis cytisoides L. with the aim of determining whether variation in the allometric relationships between plant parts provides a sensitive indicator of the impact of grazing. 2. The intra-individual variation in translatory symmetry with scale and increased complexity of fractal structures reflect environmental disturbance under heavy grazing pressure and lack of grazing. 3. Fitness consequences of grazing were also investigated. Grazing promotes growth and adult survival, and a drop in seed production as a consequence of consumption. In spite of that, total inclusive fitness (population rate of change) tends to increase with grazing. 4. Moderate grazing, while promoting growth, also enhances stability of vegetative structures. The favourable effect of moderate levels of herbivory on A. cytisoides is reflected in the homeostatic maintenance of its translatory symmetry and in the increased complexity of its fractal structures.

Almería

Spatiotemporal pattern of interactions between an apex predator and sympatric species

Increases in apex predator abundance can influence the behavior of sympatric species, particularly when the available habitat and/or resources are limited. We assessed the temporal and spatiotemporal interactions between Florida panthers ( Puma concolor coryi ) and six focal sympatric species in South Florida, where Florida panther abundance has increased by more than 6-fold since the 1990’s. Using camera trap data, we quantified species’ diel activity patterns, temporal overlap, and time-to-encounter (i.e., time between consecutive visits of a Florida panther and a focal species and vice versa). The Florida panther and bobcat ( Lynx rufus ) displayed a nocturnal activity pattern; the black bear ( Ursus americanus ), white-tailed deer ( Odocoileus virginianus ), wild boar ( Sus scrofa ), and wild turkey ( Meleagris gallopavo ) were mostly diurnal; and the raccoon ( Procyon lotor ) was cathemeral. Prey species and black bears minimized encounters with Florida panthers by being active during the day and displaying longer time-to-encounter, whereas Florida panthers visited a site after a prey species at higher probabilities than after competitor species, and were more likely to visit an elevated site or upland habitat. Our results suggest that interactions between Florida panthers and sympatric species in our study system are driven by species-specific behavioral responses. Gaining a better understanding of the crucial interactions driving species coexistence is important for a better understanding of the structure and function of ecological communities and help manage the potential expansion of the Florida panther into Central Florida.

Florida

Does the stress-gradient hypothesis hold water? Disentangling spatial and temporal variation in plant effects on soil moisture in dryland systems

The nature of the relationship between water limitation and facilitation has been one of the most contentious debates surrounding the stress-gradient hypothesis (SGH), which states that plant-plant interactions shift from competition to facilitation with increasing environmental stress. We take a closer look at the potential role of soil moisture in mediating plant-plant interaction outcomes by assessing effects of climate and soil texture on plant modulation of soil moisture. Using an empirically-parameterized soil moisture model, we simulated soil moisture dynamics beneath shrubs and in un-vegetated coarse and fine soils for 1000 sites in the Western United States with <700 mm mean annual precipitation. This threshold reflects the transition from dryland (<600 mm precipitation) to mesic ecosystems. Positive effects of shrubs on shallow soil moisture (i.e. the difference between shrub and interspace soil moisture) decreased along the aridity gradient when long-term average conditions were considered, contrary to expectations based on the SGH. Negative effects of shrubs on deeper soil moisture also increased with aridity. However, when extreme years were considered, positive effects of shrub on soil moisture were greatest at intermediate points along the spatial aridity gradient, consistent with a hump-backed model of plant-plant interactions. When viewed through time within a site, shrub effects on shallow soil moisture were positively related to precipitation, with more complex relationships exhibited in deeper soils Taken together, the results of this simulation study suggest that plant effects on soil moisture are predictable based on relatively general relationships between precipitation inputs and differential evaporation and transpiration rates between plant and interspace microsites that are largely driven by temperature. In particular, this study highlights the importance of differentiating between temporal and spatial variation in weather and climate, respectively, in determining plant effects on available soil moisture. Rather than focusing on the somewhat coarse-scale predictions of the SGH, it may be more beneficial to explicitly incorporate plant effects on soil moisture into predictive models of plant-plant interaction outcomes in drylands.

Functional Ecology

Enclosed nests may provide greater thermal than nest predation benefits compared with open nests across latitudes

Nest structure is thought to provide benefits that have fitness consequences for several taxa. Traditionally, reduced nest predation has been considered the primary benefit underlying evolution of nest structure, whereas thermal benefits have been considered a secondary or even non-existent factor. Yet, the relative roles of these factors on nest structures remain largely unexplored. Enclosed nests have a constructed or natural roof connected to sides that allow a restricted opening or tube entrance that provides cover in all directions except the entrance, whereas open nests are cups or platforms that are open above. We show that construction of enclosed nests is more common among songbirds (Passeriformes) in tropical and southern hemisphere regions than in north temperate regions. This geographic pattern may reflect selection from predation risk, under long-standing assumptions that nest predation rates are higher in southern regions and that enclosed nests reduce predation risk compared with open cup nests. We therefore compared nest predation rates between enclosed vs. open nests in 114 songbird species that do not nest in tree holes among five communities of coexisting birds, and for 205 non-hole-nesting species from the literature, across northern temperate, tropical, and southern hemisphere regions. Among coexisting species, enclosed nests had lower nest predation rates than open nests in two south temperate sites, but not in either of two tropical sites or a north temperate site. Nest predation did not differ between nest types at any latitude based on literature data. Among 319 species from both our field studies and the literature, enclosed nests did not show consistent benefits of reduced predation and, in fact, predation was not consistently higher in the tropics, contrary to long-standing perspectives. Thermal benefits of enclosed nests were indicated based on three indirect results. First, species that built enclosed nests were smaller than species using open nests both among coexisting species and among species from the literature. Smaller species lose heat fastest and thereby may gain important thermal benefits from reduced convective cooling. Second, eggs were warmed by parents for less time in species with enclosed nests, as can be expected if egg cooling rates are slower. Finally, species using enclosed nests exhibited enhanced growth of mass and wings compared with species using open nests, suggesting reduced thermoregulatory costs allowed increased energy for growth. Enclosed nests may therefore provide more consistent thermal than nest predation benefits, counter to long-standing perspectives.

Functional Ecology

Energetic fitness: Field metabolic rates assessed via 3D accelerometry complement conventional fitness metrics

Evaluating the fitness of organisms is an essential step towards understanding their responses to environmental change. Connections between energy expenditure and fitness have been postulated for nearly a century. However, testing this premise among wild animals is constrained by difficulties in measuring energy expenditure while simultaneously monitoring conventional fitness metrics such as survival and reproductive output. We addressed this issue by exploring the functional links between field metabolic rate (FMR), body condition, sex, age and reproductive performance in a wild population. We deployed 3D accelerometers on 115 Adélie penguins Pygoscelis adeliae during four breeding seasons at one of the largest colonies of this species, Cape Crozier, on Ross Island, Antarctica. The demography of this population has been studied for the past 18 years. From accelerometry recordings, collected for birds of known age and breeding history, we determined the vector of the dynamic body acceleration (VeDBA) and used it as a proxy for FMR. This allowed us to demonstrate relationships among FMR, a breeding quality index (BQI) and body condition. Notably, we found a significant quadratic relationship between mean VeDBA during foraging and BQI for experienced breeders, and individuals in better body condition showed lower rates of energy expenditure. We conclude that using FMR as a fitness component complementary to more conventional fitness metrics will yield greater understanding of evolutionary and conservation physiology.

Functional Ecology

State‐space modelling of the flight behaviour of a soaring bird provides new insights to migratory strategies

Characterising the spatiotemporal variation of animal behaviour can elucidate the way individuals interact with their environment and allocate energy. Increasing sophistication of tracking technologies paired with novel analytical approaches allows the characterisation of movement dynamics even when an individual is not directly observable. In this study, high‐resolution movement data collected via global positioning system (GPS) tracking in three dimensions were paired with topographical information and used in a Bayesian state‐space model to describe the flight modes of migrating golden eagles ( Aquila chrysaetos ) in eastern North America. Our model identified five functional behavioural states, two of which were previously undescribed variations on thermal soaring. The other states comprised gliding, perching and orographic soaring. States were discriminated by movement features in the horizontal (step length and turning angle) and vertical (change in altitude) planes and by the association with ridgelines promoting wind deflection. Tracked eagles spent 2%, 31%, 38%, 9% and 20% of their daytime in directed thermal soaring, gliding, convoluted thermal soaring, perching and orographic soaring, respectively. The analysis of the relative occurrence of these flight modes highlighted yearly, seasonal, age, individual and sex differences in flight strategy and performance. Particularly, less energy‐efficient orographic soaring was more frequent in autumn, when thermals were less available. Adult birds were also better at optimising energy efficiency than subadults. Our approach represents the first example of a state‐space model for bird flight mode using altitude data in conjunction with horizontal locations and is applicable to other flying organisms where similar data are available. The ability to describe animal movements in a three‐dimensional habitat is critical to advance our understanding of the functional processes driving animals’ decisions.

Functional Ecology

Migratory strategy explains differences in timing of female reproductive development in seasonally sympatric songbirds.

1. Divergent migratory strategies among populations can result in population-level differences in timing of reproduction (allochrony) and local adaptation. However, the mechanisms underlying among-population variation in timing are insufficiently understood, particularly in females. 2. We studied differences in reproductive development and its related mechanisms along the hypothalamic-pituitary-gonadal axis (HPG) in closely related migratory and sedentary (i.e., resident) female dark-eyed juncos (Junco hyemalis) living together in sympatry during early spring. We predicted that residents would be more reproductively developed than migrants. Alternatively, we predicted that females might exhibit similar reproductive development in response to the same environmental cue despite differences in migratory behavior. To compare their degree of reproductive development during seasonal sympatry and the underlying mechanisms of these differences, we collected ovarian and liver tissue in early spring prior to migration, and compared abundance of gene transcripts associated with reproduction using quantitative PCR. We also used stable hydrogen isotopes to infer relative breeding and wintering latitude of migrants. 3. We found more abundant transcripts for luteinizing hormone receptor and aromatase in addition to significantly heavier ovaries in residents than in migrants. Together these results suggest greater sensitivity and response to upstream endocrine stimulation in resident females. Transcript abundance for other receptors associated with reproduction, however, did not differ between populations. When comparing ovarian development within migrants, females with lower hydrogen isotopes (indicating higher breeding latitudes) had smaller ovaries, suggesting that longer-distance migrations may further delay reproductive development. 4. Based on differences in ovary mass and transcript abundance, we conclude that females that differ in migratory strategy also differ in timing of reproductive development. Differences in timing of reproductive development between migrant and resident females could drive reproductive isolation between populations. This study also reveals mechanisms that could drive allochrony and enable us to predict how seasonal organisms will respond to a changing world.

Functional Ecology

Nest structure affects auditory and visual detectability, but not predation risk, in a tropical songbird community

Offspring mortality varies dramatically among species with critical demographic and evolutionary ramifications, yet the causes of this variation remain unclear. Nests are widely used for breeding across taxa and thought to influence offspring mortality risk. Traditionally, more complex, enclosed nest structures are thought to reduce offspring predation by reducing the visibility of nest contents and muffling offspring sounds compared to open nests. Direct tests of the functional bases for nest structure influence on predation risk are lacking. We used experiments and 10 years of observational data to examine how nest structure influences nest predation risk in a diverse community of tropical songbirds. First, we examined how nest size was related to nest structure and nest predation rates across species. Second, we assessed how nest structure influences the detectability of nestling begging calls both in field and in laboratory settings. Finally, we examined how the acoustic properties of different nest structures influence nest predation risk. Specifically, we experimentally broadcast begging calls from open and enclosed nests to determine how auditory cues and nest structure interact to affect predation on plasticine and quail eggs. We also tested whether nest structure was associated with differences in nest predation rates between the incubation (no begging cues) and nestling (begging cues) stages. We found that enclosed nests are larger than open nests after accounting for adult size, and larger nests had increased predation rates. Moreover, enclosed nests did not consistently alter nestling begging calls in ways that reduce the likelihood of predation compared to open nests. Indeed, begging cues increased predation rates for enclosed but not open‐cup nests in our playback experiment, and nest predation rates showed greater increases after hatching in enclosed than open‐cup nests. Ultimately, enclosed nests do not necessarily provide greater predation benefits than open nests in contrast to long‐standing theory.

Functional Ecology