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USGS2023

Accounting for spatial habitat and management boundaries when estimating forest bird population distribution and density: Inferences from a soap film smoother

Birds are often obligate to specific habitats which can result in study areas with complex boundaries due to sudden changes in vegetation or other features. This can result in study areas with concave arcs or that include holes of unsuitable habitat such as lakes or agricultural fields. Spatial models used to produce species’ distribution and density estimates need to respect such boundaries to make informed decisions for species conservation and management. The soap film smoother is one model for complex study regions which controls the boundary behaviour, ensuring realistic values at the edges of the region. We apply the soap film smoother to account for boundary effects and compare it with thin plate regression spline (TPRS) smooth and design-based conventional distance sampling methods to produce abundance estimates from point-transect distance sampling collected data on Hawai‘i ‘Ākepa Loxops coccineus in the Hakalau Forest Unit of the Big Island National Wildlife Refuge Complex, Hawai‘i Island, USA. The soap film smoother predicted zero or near zero densities in the northern part of the domain and two hotspots (in the southern and central parts of the domain). Along the boundary the soap film model predicted relatively high densities where ‘Ākepa occur in the adjacent forest and near zero elsewhere. The design-based and soap film abundance estimates were nearly identical. The width of the soap film confidence interval was 16.5% and 0.8% wider than the width of the TPRS smooth and design-based confidence intervals, respectively. The peaks in predicted densities along the boundary indicates leakage by the TPRS smooth. We provide a discussion of the statistical methods, biological findings and management implications of applying soap film smoothers to estimate forest bird population status.

Hawaii↗

USGS2020

Shared functional traits explain synchronous changes in long‐term count trends of migratory raptors

Aim Assessing long‐term shifts in faunal assemblages is important to understand the consequences of ongoing global environmental change. One approach to assess drivers of assemblage changes is to identify the traits associated with synchronous shifts in count trends among species. Our research identified traits influencing trends in 73 years of count data on migrating raptors recorded in the north‐eastern USA. Location Pennsylvania, USA. Time period 1946–2018. Major taxa studied Birds of prey/raptors. Methods Migrating raptors were counted during autumn, following a standardized protocol. We used a hierarchical breakpoint model to identify when count trends shifted and to assess the role of traits in driving these trends before and after the breakpoint. Specifically, we quantified the probability of the direction (PD) of an effect of body mass, habitat or dietary specialization, migratory behaviour and susceptibility to dichlorodiphenyltrichloroethane (DDT) on count trends. Results We documented an assemblage‐wide mean shift in count trends of migrating raptors in 1974. In general, species that exhibited negative count trends before the breakpoint exhibited positive count trends afterwards. We found that traits associated with resource use (diet and habitat specialization) had high probabilities of affecting count trends, pre‐ and post‐breakpoint (> 90%). Moreover, the direction of their effects differed during both periods. Unexpectedly, other traits we evaluated, including DDT susceptibility, had relatively weaker associations with count trends. Main conclusions Trait‐based frameworks have promise for testing generalized assumptions about drivers of population trajectories. Historically, DDT was considered a key driver of changes in raptor population trends. However, our analysis suggests that other factors were also relevant. Moreover, the positive association between count trends and generalist behaviour depended on the temporal context. This result has implications for other settings where demographic trends can be linked to traits and help to identify drivers of biodiversity change.

Pennsylvania↗

USGS2023

Climate change and coastal wetland salinization: Physiological and ecological consequences for Arctic waterfowl

Coastal wetland salinization related to warming climate has the potential to impact ecological systems globally. In Alaska, the Yukon-Kuskokwim Delta (YKD) supports large concentrations of breeding water birds and is an ecologically important area for conservation of migratory bird biodiversity. On the YKD, the majority of waterfowl nest in low elevation coastal tundra where storm surges drive saline water into terrestrial wetland habitats. Because newly hatched water birds lack functional salt glands to process saline water, salinization may negatively impact their growth and survival. We investigated potential demographic impacts of wetland salinization by conducting controlled physiological experiments to determine consequences of saline water exposure to spectacled Somateria fischeri and Steller's eider Polysticta stelleri ducklings, and analysed habitat suitability based on experimentally defined tolerance thresholds and sampled wetland salinity levels. We found physiological and behavioural pathologies in <1 week old ducklings at exposure levels of 6 parts per thousand (ppt), and found some evidence of reduced growth in ducklings first exposed after 1 week of age. We conclude that the acute toxicity threshold for ducklings <1 week of age is ≤6 ppt. We found that the area of coastal tundra where wetland salinity exceeded 6 ppt varied considerably among years, suggesting that some portion of the breeding range is unsuitable for rearing ducklings in some years. However, we also found that most eiders in most years nested in areas with salinity at or exceeding the tolerance threshold for ducklings, suggesting that nesting eiders do not appear to avoid saline habitats. We suggest that localized fresh water refugia currently may allow resilience to salinization during the critical period. Understanding how species and habitats respond to climate driven changes is essential for predicting future patterns of distribution and abundance, and is necessary for making informed decisions about conservation priorities. Our study provides insights into the extent of wetland salinization in Alaska, mechanism of impact and current ecological consequences on avian communities depending on these habitats. With ongoing climate change, the probability of species crossing physiological tolerance thresholds of wetland salinity may change in the future.

Functional Ecology↗

USGS2011

Post-fledging brood and care division in the roseate tern (Sterna dougallii)

Extended post-fledging parental care is an important aspect of parental care in birds, although little studied due to logistic difficulties. Commonly, the brood is split physically (brood division) and/or preferential care is given to a subset of the brood by one parent or the other (care division). Among gulls and tern (Laridae), males and females generally share parental activities during the pre-fledging period, but the allocation of parental care after fledging is little documented. This study examined the behaviour of male and female roseate terns ( Sterna dougallii ) during the late chick-rearing and early post-fledging periods, and in particular the amount of feeds and the time spent in attendance given to individual chicks/fledglings. Pre-fledging parental care was biparental in all cases. Post-fledging parental care was dependent on the number of fledglings in the brood. Males and females continued biparental care in clutches with one surviving fledgling, while in two-fledgling clutches, males fed the A-fledgling while females fed the B-fledgling. Overall, there was no difference in attendance, only in feeds. This division of care may be influenced by the male only being certain of the paternity of the A-chick but not by chick sex.

Journal of Ethology↗

USGS1991

Observations on habitat use, breeding chronology and parental care in Bristle-thighed Curlews on the Seward Peninsula, Alaska

Results from an intensive survey in 1989 of one of the two known breeding localities of Bristle-thighed Curlews Numenius tahitiensis are reported. During the pre-nesting period birds confined most of their activity to two vegetation communities: shrub meadow tundra and low shrub/tussock tundra. During nesting more than half the Curlews seen were observed on shrub meadow tundra, whilst during brood rearing, use of low shrub/tussock tundra continued to decline in importance as birds attending young increased their use of sedge wet meadows. Despite extensive searches no nests were located; however, observations of broods indicated that nest initiation began around 25 May and that hatching occurred during the last week of June. Detailed observations are presented on the formation of four brood-groups, some of which held different combinations of unrelated adults or young. The adaptive significance of this unusual wader behaviour is discussed.

Alaska↗

USGS2016

Widespread episodic thiamine deficiency in Northern Hemisphere wildlife

Many wildlife populations are declining at rates higher than can be explained by known threats to biodiversity. Recently, thiamine (vitamin B 1 ) deficiency has emerged as a possible contributing cause. Here, thiamine status was systematically investigated in three animal classes: bivalves, ray-finned fishes, and birds. Thiamine diphosphate is required as a cofactor in at least five life-sustaining enzymes that are required for basic cellular metabolism. Analysis of different phosphorylated forms of thiamine, as well as of activities and amount of holoenzyme and apoenzyme forms of thiamine-dependent enzymes, revealed episodically occurring thiamine deficiency in all three animal classes. These biochemical effects were also linked to secondary effects on growth, condition, liver size, blood chemistry and composition, histopathology, swimming behaviour and endurance, parasite infestation, and reproduction. It is unlikely that the thiamine deficiency is caused by impaired phosphorylation within the cells. Rather, the results point towards insufficient amounts of thiamine in the food. By investigating a large geographic area, by extending the focus from lethal to sublethal thiamine deficiency, and by linking biochemical alterations to secondary effects, we demonstrate that the problem of thiamine deficiency is considerably more widespread and severe than previously reported.

Scientific Reports↗

USGS1999

Ecology and behaviour of the Alder Flycatcher Empidonax alnorum, Tyrannidae in winter

The Alder Flycatcher Empidonax alnorum (Tyrannidae) winters from Colombia and Venezuela, south to northern Argentina. We studied the species in Manu National Park, Madre de Dios, Peru. In this area the species occupies riparian zones along lowland meander rivers as well as adjacent transition zone forest in the floodplain. These habitats have a number of age-related longitudinal zones (parallel to the river margin) that are short term (I-2 year) transitory stages in the succession to floodplain forest. The fast growing plant species that make up these habitats are suitable for commercial harvest for paper pulp. We examined habitat use of the species in an attempt to identify the effects of such harvest and resulting habitat loss on these birds. The Alder Flycatcher favours two longitudinal zones where it establishes territories. The territories may be long-term or transitory, and are often occupied and defended by two individuals, generally of different age. One bird tends to be more active in territory defence than the other. Defence involves patrolling, vocal duels, and chases. Birds sally for insects but also eat large quantities of fruit.

Book chapter↗

USGS1998

Ecology and behaviour of the Alder Flycatcher Empidonax alnorum, Tyrannidae in winter

Book chapter↗

USGS2012

Invasive species management restores a plant-pollinator mutualism in Hawaii

1.The management and removal of invasive species may give rise to unanticipated changes in plant–pollinator mutualisms because they can alter the composition and functioning of plant–pollinator interactions in a variety of ways. To utilize a functional approach for invasive species management, we examined the restoration of plant–pollinator mutualisms following the large-scale removal of an invasive nectar thief and arthropod predator, Vespula pensylvanica. 2.We reduced V. pensylvanica populations in large plots managed over multiple years to examine the response of plant–pollinator mutualisms and the fruit production of a functionally important endemic Hawaiian tree species, Metrosideros polymorpha. To integrate knowledge of the invader's behaviour and the plant's mating system, we determined the efficacy of V. pensylvanica as a pollinator of M. polymorpha and quantified the dependence of M. polymorpha on animal pollination (e.g. level of self-compatibility and pollen limitation). 3.The reduction of V. pensylvanica in managed sites, when compared to unmanaged sites, resulted in a significant increase in the visitation rates of effective bee pollinators (e.g. introduced Apis mellifera and native Hylaeus spp.) and in the fruit production of M. polymorpha. 4.Apis mellifera, following the management of V. pensylvanica, appears to be acting as a substitute pollinator for M. polymorpha, replacing extinct or threatened bird and bee species in our study system. 5.Synthesis and applications. Fruit production of the native M. polymorpha was increased after management of the invasive pollinator predator V. pensylvanica; however, the main pollinators were no longer native but introduced. This research thus demonstrates the diverse impacts of introduced species on ecological function and the ambiguous role they play in restoration. We recommend incorporating ecological function and context into invasive species management as this approach may enable conservation managers to simultaneously minimize the negative and maximize the positive impacts (e.g. taxon substitution) of introduced species. Such novel restoration approaches are needed, especially in highly degraded ecosystems.

Hawai'i↗

USGS2011

Migratory movements of waterfowl in Central Asia and avian influenza emergence: Sporadic transmission of H5N1 from east to west

Waterfowl in the genera Anas and Tadorna are suspected as vectors in the long‐distance transmission of highly pathogenic avian influenza H5N1. The former Soviet Republics of Central Asia are situated at an important migratory crossroads for these and other species of birds that bridges regions where the disease is prevalent. However, waterfowl movements through Central Asia are poorly quantified. In this study, historical data derived from over 80 years of bird ringing are combined with recent satellite tracking data to delineate migration routes, movement chronology and habitat use patterns of waterfowl in relation to H5N1 outbreak locations. Results confirm migratory linkage between breeding and moulting areas in northern Kazakhstan and southern Siberia, with non‐breeding areas in the Caspian, Black and eastern Mediterranean Sea basins, as well as with South Asia. However, unlike the situation in neighbouring regions, most notably western China, H5N1 outbreaks have not been recurrent in Central Asia after they were first reported during summer 2005 and spring 2006. These findings have implications in relation to potential sampling biases, species‐specific variation in migratory behaviour and continuing regional H5N1 transmission risks.

Siberia↗

USGS2012

Relationships between yolk androgens and nest density, laying date, and laying order in Western Burrowing Owls (Athene cunicularia hypugaea)

Increases in yolk androgens within and among avian clutches have been correlated with decreased incubation time, increased aggression within a nest, increased begging behaviour, decreased immune response, and decreased life span. Although the mechanisms that lead to variability in yolk androgens within and between clutches are not completely known, yolk androgens can be a function of both social and environmental conditions. We were interested in if and how nesting density, laying date, and laying order influenced yolk androgens in Western Burrowing Owls (Athene cunicularia hypugaea (Bonaparte, 1825)) in which nest density varies considerably. In 2006 and 2007, we used radioimmunoassay to quantify the concentrations of testosterone, 5a-dihydrotestosterone, and androstenedione in the egg yolks from one early and one latelaid egg in 47 nests of Burrowing Owls located in the Morley Nelson Snake River Birds of Prey National Conservation Area in southern Idaho. Nesting density had no detectable effect on yolk androgens. Yolk androgens varied temporally and peaked in the middle of the laying season while being low before and after this time period. Within nests, late-laid eggs had higher testosterone and dihydrotestosterone than early-laid eggs; adrostendione exhibited a similar pattern in one but not both years of our study. It is possible that the seasonal pattern in yolk androgens that we observed is related to aspects of mate quality for females or declining chances of fledging success for later nesting females, whereas rises in egg androgens between early and late eggs within clutches could reflect a mechanism to assist nestlings from late-laid eggs that hatch one to several days after their siblings to better compete for resources within the nest or promote survival in the presence of larger siblings.

Canadian Journal of Zoology↗

USGS2025

Adoption of non‐related goslings and intergenerational family cohesion among Greenland White‐fronted Geese (Anser albifrons flavirostris)

Greenland White-fronted Geese Anser albifrons flavirostris exhibit prolonged parent–offspring and sibling–sibling associations, suggesting fitness advantages to such behaviour, so we used reduced representation genome sequence data to determine the degree to which marked flock members observed associating in apparent parent–offspring and sibling–sibling relationships in the field were genetically related. Among 50 bled, marked and released geese, we genetically identified members of 11 different family groups, confirming all observed male parent–offspring relationships as genetically predicted, but only 10 out of 12 (83%) possible female parent–offspring relationships (i.e. two offspring were not genetically related to the adult female in their family groups observed in the field); these two ‘adopted’ offspring were responsible for four (15%) of the cases where observed ‘siblings’ were not genetically related to other family-member first-winter birds with which they associated. One multigenerational family consisted of three genetically confirmed grandmother–mother–sibling offspring relationships, not previously reported in arctic-nesting geese, as well as one of the two ‘adopted’ first-winter geese.

Wexford Slobs↗

USGS2008

Importance of sampling design and analysis in animal population studies: a comment on Sergio et al

1. The use of predators as indicators and umbrellas in conservation has been criticized. In the Trentino region, Sergio et al. (2006; hereafter SEA) counted almost twice as many bird species in quadrats located in raptor territories than in controls. However, SEA detected astonishingly few species. We used contemporary Swiss Breeding Bird Survey data from an adjacent region and a novel statistical model that corrects for overlooked species to estimate the expected number of bird species per quadrat in that region. 2. There are two anomalies in SEA which render their results ambiguous. First, SEA detected on average only 6.8 species, whereas a value of 32 might be expected. Hence, they probably overlooked almost 80% of all species. Secondly, the precision of their mean species counts was greater in two-thirds of cases than in the unlikely case that all quadrats harboured exactly the same number of equally detectable species. This suggests that they detected consistently only a biased, unrepresentative subset of species. 3. Conceptually, expected species counts are the product of true species number and species detectability p. Plenty of factors may affect p, including date, hour, observer, previous knowledge of a site and mobbing behaviour of passerines in the presence of predators. Such differences in p between raptor and control quadrats could have easily created the observed effects. Without a method that corrects for such biases, or without quantitative evidence that species detectability was indeed similar between raptor and control quadrats, the meaning of SEA's counts is hard to evaluate. Therefore, the evidence presented by SEA in favour of raptors as indicator species for enhanced levels of biodiversity remains inconclusive. 4. Synthesis and application. Ecologists should pay greater attention to sampling design and analysis in animal population estimation. Species richness estimation means sampling a community. Samples should be representative for the community studied and the sampling fraction among communities compared should be the same on average, otherwise formal estimation approaches must be applied to avoid misleading inference.

Journal of Applied Ecology↗

USGS2024

Lead poisoning of raptors: State of the science and cross-discipline mitigation options for a global problem

Lead poisoning is an important global conservation problem for many species of wildlife, especially raptors. Despite the increasing number of individual studies and regional reviews of lead poisoning of raptors, it has been over a decade since this information has been compiled into a comprehensive global review. Here, we summarize the state of knowledge of lead poisoning of raptors, we review developments in manufacturing of non-lead ammunition, the use of which can reduce the most pervasive source of lead these birds encounter, and we compile data on voluntary and regulatory mitigation options and their associated sociological context. We support our literature review with case studies of mitigation actions, largely provided by the conservation practitioners who study or manage these efforts. Our review illustrates the growing awareness and understanding of lead exposure of raptors, and it shows that the science underpinning this understanding has expanded considerably in recent years. We also show that the political and social appetite for managing lead ammunition appears to vary substantially across administrative regions, countries, and continents. Improved understanding of the drivers of this variation could support more effective mitigation of lead exposure of wildlife. This review also shows that mitigation strategies are likely to be most effective when they are outcome driven, consider behavioural theory, local cultures, and environmental conditions, effectively monitor participation, compliance, and levels of raptor exposure, and support both environmental and human health.

Biological Reviews↗

USGS2013

Variation in body mass dynamics among sites in Black Brant Branta bernicla nigricans supports adaptivity of mass loss during moult

Birds employ varying strategies to accommodate the energetic demands of moult, one important example being changes in body mass. To understand better their physiological and ecological significance, we tested three hypotheses concerning body mass dynamics during moult. We studied Black Brant in 2006 and 2007 moulting at three sites in Alaska which varied in food availability, breeding status and whether geese undertook a moult migration. First we predicted that if mass loss during moult were simply the result of inadequate food resources then mass loss would be highest where food was least available. Secondly, we predicted that if mass loss during moult were adaptive, allowing birds to reduce activity during moult, then birds would gain mass prior to moult where feeding conditions allowed and mass loss would be positively related to mass at moult initiation. Thirdly, we predicted that if mass loss during moult were adaptive, allowing birds to regain flight sooner, then across sites and groups, mass at the end of the flightless period would converge on a theoretical optimum, i.e. the mass that permits the earliest possible return to flight. Mass loss was greatest where food was most available and thus our results did not support the prediction that mass loss resulted from inadequate food availability. Mass at moult initiation was positively related to both food availability and mass loss. In addition, among sites and years, variation in mass was high at moult initiation but greatly reduced at the end of the flightless period, appearing to converge. Thus, our results supported multiple predictions that mass loss during moult was adaptive and that the optimal moulting strategy was to gain mass prior to the flightless period, then through behavioural modifications use these body reserves to reduce activity and in so doing also reduce wing loading. Geese that undertook a moult migration initiated moult at the highest mass, indicating that they were more than able to compensate for the energetic cost of the migration. Because Brant frequently change moult sites between years in relation to breeding success, the site-specific variation in body mass dynamics we observed suggests individual plasticity in moult body mass dynamics.

Alaska↗

USGS2005

The effects of captive rearing on the behavior of newly-released whooping cranes (Grus americana)

Rearing treatments used in captivity to prepare animals for reintroduction to the wild may have a profound effect on behavior and, possibly, affect their survival after reintroduction. This study examined the behaviors of captive-reared whooping cranes (Grus americana) upon their release in Florida to determine if rearing treatments may affect the behavior of the birds and how these affect their chances of survival in the wild. Individually tagged birds were observed at the rearing facility, the U.S. Geological Survey Patuxent Wildlife Research Center in Maryland, from hatch to 20 weeks of age and at the release site in Central Florida for up to 6 weeks post release. The rearing treatments were parent reared (PR), hand reared (HR), and hand reared with exercise (HRE). Observations at the rearing facility are described in a previous paper. At the release site, each bird was observed for 5 min every morning (0700?1000 h) and late afternoon (1500?1800 h) during the 6-week study period. Our results indicated that most of the time, the n = 34 birds were foraging (46.03 ? 1.48%), followed by nonvigilant (20.89 ? 0.73%), vigilant (19.21 ? 0.72%), or performing comfort behaviors (11.61 ? 1.28%). Data were analyzed using mixed models repeated measures ANOVA. There were no significant behavioral differences between HR and HRE birds. PR birds were found in larger groups than HR birds during the first 2 weeks post release and greater than HR and HRE birds afterwards. This may be interpreted as an antipredator strategy for birds that relied on parental guidance during rearing. HR and HRE birds foraged more than PR birds during the first 2 weeks post release and PR birds were more vigilant during the first 2 weeks post release. Across rearing treatments, the percentages of time spent foraging and engaged in vigilant behaviors during rearing were positively correlated with their behavior upon release. If any of these behaviors can be demonstrated to have relevance for the survival of the whooping cranes after release then it may be possible to establish behavioral interventions to increase the frequencies of such behavior, so that they are perpetuated after release.

Applied Animal Behaviour Science↗

USGS2022

Pathways for avian influenza virus spread: GPS reveals wild waterfowl in commercial livestock facilities and connectivity with the natural wetland landscape

Zoonotic diseases are of considerable concern to the human population and viruses such as avian influenza (AIV) threaten food security, wildlife conservation and human health. Wild waterfowl and the natural wetlands they use are known AIV reservoirs, with birds capable of virus transmission to domestic poultry populations. While infection risk models have linked migration routes and AIV outbreaks, there is a limited understanding of wild waterfowl presence on commercial livestock facilities, and movement patterns linked to natural wetlands. We documented 11 wild waterfowl (three Anatidae species) in or near eight commercial livestock facilities in Washington and California with GPS telemetry data. Wild ducks used dairy and beef cattle feed lots and facility retention ponds during both day and night suggesting use for roosting and foraging. Two individuals (single locations) were observed inside poultry facility boundaries while using nearby wetlands. Ducks demonstrated high site fidelity, returning to the same areas of habitats (at livestock facilities and nearby wetlands), across months or years, showed strong connectivity with surrounding wetlands, and arrived from wetlands up to 1251 km away in the week prior. Telemetry data provides substantial advantages over observational data, allowing assessment of individual movement behaviour and wetland connectivity that has significant implications for outbreak management. Telemetry improves our understanding of risk factors for waterfowl–livestock virus transmission and helps identify factors associated with coincident space use at the wild waterfowl–domestic livestock interface. Our research suggests that even relatively small or isolated natural and artificial water or food sources in/near facilities increases the likelihood of attracting waterfowl, which has important consequences for managers attempting to minimize or prevent AIV outbreaks. Use and interpretation of telemetry data, especially in near-real-time, could provide key information for reducing virus transmission risk between waterfowl and livestock, improving protective barriers between wild and domestic species, and abating outbreaks.

Transboundary and Emerging Diseases↗

USGS2018

Animal movement models for migratory individuals and groups

Animals often exhibit changes in their behaviour during migration. Telemetry data provide a way to observe geographic position of animals over time, but not necessarily changes in the dynamics of the movement process. Continuous‐time models allow for statistical predictions of the trajectory in the presence of measurement error and during periods when the telemetry device did not record the animal's position. However, continuous‐time models capable of mimicking realistic trajectories with sufficient detail are computationally challenging to fit to large datasets. Furthermore, basic continuous‐time model specifications (e.g. Brownian motion) lack realism in their ability to capture nonstationary dynamics. We present a unified class of animal movement models that are computationally efficient and provide a suite of approaches for accommodating nonstationarity in continuous trajectories due to migration and interactions among individuals. Our approach uses process convolutions to allow for flexibility in the movement process while facilitating implementation and incorporating location uncertainty. We show how to nest convolution models to incorporate interactions among migrating individuals to account for nonstationarity and provide inference about dynamic migratory networks. We demonstrate these approaches in two case studies involving migratory birds. Specifically, we used process convolution models with temporal deformation to account for heterogeneity in individual greater white‐fronted goose migrations in Europe and Iceland, and we used nested process convolutions to model dynamic migratory networks in sandhill cranes in North America. The approach we present accounts for various forms of temporal heterogeneity in animal movement and is not limited to migratory applications. Furthermore, our models rely on well‐established principles for modelling‐dependent data and leverage modern approaches for modelling dynamic networks to help explain animal movement and social interaction.

Methods in Ecology and Evolution↗