Environmental factors influencing red knot (Calidris canutus islandica) departure times of relocation flights within the non-breeding period.

Deciding when to depart on long-distance, sometimes global, movements can be especially important for flying species. Adverse weather conditions can affect energetic flight costs and navigational ability. While departure timings and conditions have been well-studied for migratory flights to and from the breeding range, few studies have focussed on flights within the non-breeding season. Yet in some cases, overwintering ranges can be large enough that ecological barriers, and a lack of resting sites en route, may resist movement, especially in unfavorable environmental conditions. Understanding the conditions that will enable or prohibit flights within an overwintering range is particularly relevant in light of climate change, whereby increases in extreme weather events may reduce the connectivity of sites. We tracked 495 (n = 251 in 2019; n = 244 in 2020) overwintering red knots (Calidris canutus islandica) in the Dutch Wadden Sea and investigated how many departed towards the UK (on westward relocation flights), which requires flying over the North Sea. For those that departed, we used a resource selection model to determine the effect of environmental conditions on the timing of relocation flights. Specifically, we investigated the effects of wind, rain, atmospheric pressure, cloud cover, and migratory timing relative to sunset and tidal cycle, which have all been shown to be crucial to migratory departure conditions. Approximately 37% (2019) and 36% (2020) of tagged red knots departed on westward relocation flights, indicating differences between individuals' space use within the overwintering range. Red knots selected for departures between 1 and 2.5 h after sunset, approximately 4 h before high tide, with tailwinds and little cloud cover. However, rainfall and changes in atmospheric pressure appear unimportant. Our study reveals environmental conditions that are important for relocation flights across an ecological barrier, indicating potential consequences of climate change on connectivity.

Cranes soar on thermal updrafts behind cold fronts as they migrate across the sea.

Thermal soaring conditions above the sea have long been assumed absent or too weak for terrestrial migrating birds, forcing obligate soarers to take long detours and avoid sea-crossing, and facultative soarers to cross exclusively by costly flapping flight. Thus, while atmospheric convection does develop at sea and is used by some seabirds, it has been largely ignored in avian migration research. Here, we provide direct evidence for routine thermal soaring over open sea in the common crane, the heaviest facultative soarer known among terrestrial migrating birds. Using high-resolution biologging from 44 cranes tracked across their transcontinental migration over 4 years, we show that soaring performance was no different over sea than over land in mid-latitudes. Sea-soaring occurred predominantly in autumn when large water-air temperature difference followed mid-latitude cyclones. Our findings challenge a fundamental migration research paradigm and suggest that obligate soarers avoid sea-crossing not due to the absence or weakness of thermals but due to their low frequency, for which they cannot compensate with prolonged flapping. Conversely, facultative soarers other than cranes should also be able to use thermals over the sea. Marine cold air outbreaks, imperative to global energy budget and climate, may also be important for bird migration.

Robust Time-of-Arrival Location Estimation Algorithms for Wildlife Tracking.

Time-of-arrival transmitter localization systems, which use measurements from an array of sensors to estimate the location of a radio or acoustic emitter, are now widely used for tracking wildlife. Outlier measurements can severely corrupt estimated locations. This article describes a new suite of location estimation algorithms for such systems. The new algorithms detect and discard outlier time-of-arrival observations, which can be caused by non-line-of-sight propagation, radio interference, clock glitches, or an overestimation of the signal-to-noise ratio. The new algorithms also detect cases in which two locations are equally consistent with measurements and can usually select the correct one. The new algorithms can also infer approximate altitude information from a digital elevation map to improve location estimates close to one of the sensors. Finally, the new algorithms approximate the covariance matrix of location estimates in a simpler and more reliable way than the baseline algorithm. Extensive testing on real-world data involving mobile transmitters attached to wild animals demonstrates the efficacy of the new algorithms. Performance testing also shows that the new algorithms are fast and that they can easily cope with high-throughput real-time loads.

Spatial memory predicts home range size and predation risk in pheasants.

Most animals confine their activities to a discrete home range, long assumed to reflect the fitness benefits of obtaining spatial knowledge about the landscape. However, few empirical studies have linked spatial memory to home range development or determined how selection operates on spatial memory via the latter's role in mediating space use. We assayed the cognitive ability of juvenile pheasants (Phasianus colchicus) reared under identical conditions before releasing them into the wild. Then, we used high-throughput tracking to record their movements as they developed their home ranges, and determined the location, timing and cause of mortality events. Individuals with greater spatial reference memory developed larger home ranges. Mortality risk from predators was highest at the periphery of an individual's home range in areas where they had less experience and opportunity to obtain spatial information. Predation risk was lower in individuals with greater spatial memory and larger core home ranges, suggesting selection may operate on spatial memory by increasing the ability to learn about predation risk across the landscape. Our results reveal that spatial memory, determined from abstract cognitive assays, shapes home range development and variation, and suggests predation risk selects for spatial memory via experience-dependent spatial variation in mortality.

The establishment of plants following long-distance dispersal.

Long-distance dispersal (LDD) beyond the range of a species is an important driver of ecological and evolutionary patterns, but insufficient attention has been given to postdispersal establishment. In this review, we summarize current knowledge of the post-LDD establishment phase in plant colonization, identify six key determinants of establishment success, develop a general quantitative framework for post-LDD establishment, and address the major challenges and opportunities in future research. These include improving detection and understanding of LDD using novel approaches, investigating mechanisms determining post-LDD establishment success using mechanistic modeling and inference, and comparison of establishment between past and present. By addressing current knowledge gaps, we aim to further our understanding of how LDD affects plant distributions, and the long-term consequences of LDD events.

Comparing invasive and noninvasive faecal sampling in wildlife microbiome studies: A case study on wild common cranes.

In ecological and conservation studies, responsible researchers strive to obtain rich data while minimizing disturbance to wildlife and ecosystems. We assessed if samples collected noninvasively can be used for faecal microbiome research, comparing microbiota of noninvasively collected faecal samples to those collected from trapped common cranes at the same sites over the same periods. We found significant differences in faecal microbial composition (alpha and beta diversity), which likely did not result from noninvasive sample exposure to soil contaminants, as assessed by comparing bacterial oxygen use profiles. Differences might result from trapped birds' exposure to sedatives or stress. We conclude that if all samples are collected in the same manner, comparative analyses are valid, and noninvasive sampling may better represent host faecal microbiota because there are no trapping effects. Experiments with fresh and delayed sample collection can elucidate effects of environmental exposures on microbiota. Further, controlled tests of stressing or sedation may unravel how trapping affects wildlife microbiota.

Linking migration and microbiota at a major stopover site in a long-distance avian migrant.

Migration is one of the most physical and energetically demanding periods in an individual bird's life. The composition of the bird's gut or cloacal microbiota can temporarily change during migration, likely due to differences in diets, habitats and other environmental conditions experienced en route. However, how physiological condition, migratory patterns, and other drivers interact to affect microbiota composition of migratory birds is still unclear. We sampled the cloacal bacterial microbiota of a long-distance migrant, the steppe buzzard (Buteo buteo vulpinus), at an important spring stopover bottleneck in Eilat, Israel, after crossing the ca. 1800 km Sahara Desert. We examined whether diversity and composition of the cloacal microbiota varied with body condition, sex, movement patterns (i.e., arrival time and migration distance), and survival. Early arrival to Eilat was associated with better body condition, longer post-Eilat spring migration distance, higher microbial α-diversity, and differences in microbiota composition. Specifically, early arrivals had higher abundance of the phylum Synergistota and five genera, including Jonquetella and Peptococcus, whereas the phylum Proteobacteria and genus Escherichia-Shigella (as well as three other genera) were more abundant in later arrivals. While the differences in α-diversity and Escherichia-Shigella seem to be mainly driven by body condition, other compositional differences associated with arrival date could be indicators of longer migratory journeys (e.g., pre-fueling at wintering grounds or stopover habitats along the way) or migratory performance. No significant differences were found between the microbiota of surviving and non-surviving individuals. Overall, our results indicate that variation in steppe buzzard microbiota is linked to variation in migratory patterns (i.e., capture/arrival date) and body condition, highlighting the importance of sampling the microbiota of GPS-tracked individuals on multiple occasions along their migration routes to gain a more detailed understanding of the links between migration, microbiota, and health in birds.

Insectivorous bats form mobile sensory networks to optimize prey localization: The case of the common noctule bat.

Animals that depend on ephemeral, patchily distributed prey often use public information to locate resource patches. The use of public information can lead to the aggregation of foragers at prey patches, a mechanism known as local enhancement. However, when ephemeral resources are distributed over large areas, foragers may also need to increase search efficiency, and thus apply social strategies when sampling the landscape. While sensory networks of visually oriented animals have already been confirmed, we lack an understanding of how acoustic eavesdropping adds to the formation of sensory networks. Here we radio-tracked a total of 81 aerial-hawking bats at very high spatiotemporal resolution during five sessions over 3 y, recording up to 19 individuals simultaneously. Analyses of interactive flight behavior provide conclusive evidence that bats form temporary mobile sensory networks by adjusting their movements to neighboring conspecifics while probing the airspace for prey. Complementary agent-based simulations confirmed that the observed movement patterns can lead to the formation of mobile sensory networks, and that bats located prey faster when networking than when relying only on local enhancement or searching solitarily. However, the benefit of networking diminished with decreasing group size. The combination of empirical analyses and simulations elucidates how animal groups use acoustic information to efficiently locate unpredictable and ephemeral food patches. Our results highlight that declining local populations of social foragers may thus suffer from Allee effects that increase the risk of collapses under global change scenarios, like insect decline and habitat degradation.

Phylogenomics and evolutionary history of Oreocnide (Urticaceae) shed light on recent geological and climatic events in SE Asia.

Climate change and geological events have long been known to shape biodiversity, implying that these can likewise be viewed from a biological perspective. To study whether plants can shed light on this, and how they responded to climate change there, we examined Oreocnide, a genus widely distributed in SE Asia. Based on broad geographic sampling with genomic data, we employed an integrative approach of phylogenomics, molecular dating, historical biogeography, and ecological analyses. We found that Oreocnide originated in mainland East Asia and began to diversify ∼6.06 Ma, probably in response to a distinct geographic and climatic transition in East Asia at around that time, implying that the last important geological change in mainland SE Asia might be 1 Ma older than previously suggested. Around six immigration events to the islands of Malesia followed, indicating that immigration from the mainland could be an underestimated factor in the assembly of biotic communities in the region. Two detected increases of diversification rate occurred 3.13 and 1.19 Ma, which strongly implicated climatic rather than geological changes as likely drivers of diversification, with candidates being the Pliocene intensification of the East Asian monsoons, and Pleistocene climate and sea level fluctuations. Distribution modelling indicated that Pleistocene sea level and climate fluctuations were inferred to enable inter-island dispersal followed by allopatric separation, underpinning radiation in the genus. Overall, our study, based on multiple lines of evidence, linked plant diversification to the most recent climatic and geological events in SE Asia. We highlight the importance of immigration in the assembly and diversification of the SE Asian flora, and underscore the utility of plant clades, as independent lines of evidence, for reconstructing recent climatic and geological events in the SE Asian region.

First detection of avian influenza subtype H4N6 in Israel in a wild mallard (Anas platyrhynchos).

Avian influenza viruses (AIV) are a worldwide threat to animal and human health. As wild waterfowl circulate and spread these viruses around the world, investigations of AIV prevalence in wild populations are critical for understanding pathogen transmission, as well as predicting disease outbreaks in domestic animals and humans. Surveillance efforts in this study have isolated H4N6 for the first time in Israel from a faecal sample of a wild mallard (Anas platyrhynchos). Phylogenetic analyses of the HA and NA genes revealed that this strain is closely related to isolates from Europe and Asia. This Eurasian origin, together with Israel serving as an important migratory bottleneck of the mid Palearctic-African flyway, suggests a potential introduction of this strain by migratory birds. Additional phylogenetic analysis of the isolate's internal genes (PB1, PB2, PA, NP, M and NS) revealed high levels of phylogenetic relatedness with other AIV subtypes, indicating previous reassortment events. High reassortment rates are characteristic for H4N6 viruses, which, together with this subtype's ability to infect pigs and adaptability to the human receptor binding domain, raises the concern that it would potentially become zoonotic in the future. These results emphasize the importance of continuous AIV monitoring in migratory birds.

Phase Transition in a Non-Markovian Animal Exploration Model with Preferential Returns.

We study a non-Markovian and nonstationary model of animal mobility incorporating both exploration and memory in the form of preferential returns. Exact results for the probability of visiting a given number of sites are derived and a practical WKB approximation to treat the nonstationary problem is developed. A mean-field version of this model, first suggested by Song et al., [Modelling the scaling properties of human mobility, Nat. Phys. 6, 818 (2010)NPAHAX1745-247310.1038/nphys1760] was shown to well describe human movement data. We show that our generalized model adequately describes empirical movement data of Egyptian fruit bats (Rousettus aegyptiacus) when accounting for interindividual variation in the population. We also study the probability of visiting any site a given number of times and derive a mean-field equation. Our analysis yields a remarkable phase transition occurring at preferential returns which scale linearly with past visits. Following empirical evidence, we suggest that this phase transition reflects a trade-off between extensive and intensive foraging modes.

Big-data approaches lead to an increased understanding of the ecology of animal movement.

Understanding animal movement is essential to elucidate how animals interact, survive, and thrive in a changing world. Recent technological advances in data collection and management have transformed our understanding of animal "movement ecology" (the integrated study of organismal movement), creating a big-data discipline that benefits from rapid, cost-effective generation of large amounts of data on movements of animals in the wild. These high-throughput wildlife tracking systems now allow more thorough investigation of variation among individuals and species across space and time, the nature of biological interactions, and behavioral responses to the environment. Movement ecology is rapidly expanding scientific frontiers through large interdisciplinary and collaborative frameworks, providing improved opportunities for conservation and insights into the movements of wild animals, and their causes and consequences.

A guide to pre-processing high-throughput animal tracking data.

Modern, high-throughput animal tracking increasingly yields 'big data' at very fine temporal scales. At these scales, location error can exceed the animal's step size, leading to mis-estimation of behaviours inferred from movement. 'Cleaning' the data to reduce location errors is one of the main ways to deal with position uncertainty. Although data cleaning is widely recommended, inclusive, uniform guidance on this crucial step, and on how to organise the cleaning of massive datasets, is relatively scarce. A pipeline for cleaning massive high-throughput datasets must balance ease of use and computationally efficiency, in which location errors are rejected while preserving valid animal movements. Another useful feature of a pre-processing pipeline is efficiently segmenting and clustering location data for statistical methods while also being scalable to large datasets and robust to imperfect sampling. Manual methods being prohibitively time-consuming, and to boost reproducibility, pre-processing pipelines must be automated. We provide guidance on building pipelines for pre-processing high-throughput animal tracking data to prepare it for subsequent analyses. We apply our proposed pipeline to simulated movement data with location errors, and also show how large volumes of cleaned data can be transformed into biologically meaningful 'residence patches', for exploratory inference on animal space use. We use tracking data from the Wadden Sea ATLAS system (WATLAS) to show how pre-processing improves its quality, and to verify the usefulness of the residence patch method. Finally, with tracks from Egyptian fruit bats Rousettus aegyptiacus, we demonstrate the pre-processing pipeline and residence patch method in a fully worked out example. To help with fast implementation of standardised methods, we developed the R package atlastools, which we also introduce here. Our pre-processing pipeline and atlastools can be used with any high-throughput animal movement data in which the high data-volume combined with knowledge of the tracked individuals' movement capacity can be used to reduce location errors. atlastools is easy to use for beginners while providing a template for further development. The common use of simple yet robust pre-processing steps promotes standardised methods in the field of movement ecology and leads to better inferences from data.

Congruence between ocean-dispersal modelling and phylogeography explains recent evolutionary history of Cycas species with buoyant seeds.

Ocean currents play a significant role in driving the long-distance dispersal (LDD), spatial distribution and phylogeographic patterns of many organisms. Integrating phylogeographic analyses and mechanistic ocean current modelling can provide novel insights into the evolutionary history of terrestrial littoral species but has been rarely applied in this context. We focused on a group of Cycas that have buoyant seeds and occupy coastal habitats. By integrating evidence from mechanistic simulations and whole plastomic data, we examined the role of ocean circulation in shaping the phylogeography of these Cycas species. Plastomes of the studied Cycas species showed extreme conservatism, following a post-Pleistocene divergence. Phylogenies revealed three subclades, corresponding to the Pacific Ocean, Sunda Shelf and Indian Ocean. The ocean modelling results indicate that hotspots of seed stranding coincide well with the contemporary distribution of the Cycas species and that drifting trajectories from the three subclades are largely confined to separate regions. These findings suggest that ocean current systems, by driving long-distance dispersal, have shaped the distribution and phylogeography for Cycas with buoyant seeds. This study highlights how the combination of genomic data and ocean drift modelling can help explain phylogeographic patterns and diversity in terrestrial littoral ecosystems.

Drivers of change and stability in the gut microbiota of an omnivorous avian migrant exposed to artificial food supplementation.

Human activities shape resources available to wild animals, impacting diet and probably altering their microbiota and overall health. We examined drivers shaping microbiota profiles of common cranes (Grus grus) in agricultural habitats by comparing gut microbiota and crane movement patterns (GPS-tracking) over three periods of their migratory cycle, and by analysing the effect of artificially supplemented food provided as part of a crane-agriculture management programme. We sampled faecal droppings in Russia (nonsupplemented, premigration) and in Israel in late autumn (nonsupplemented, postmigration) and winter (supplemented and nonsupplemented, wintering). As supplemented food is typically homogenous, we predicted lower microbiota diversity and different composition in birds relying on supplementary feeding. We did not observe changes in microbial diversity with food supplementation, as diversity differed only in samples from nonsupplemented wintering sites. However, both food supplementation and season affected bacterial community composition and led to increased abundance of specific genera (mostly Firmicutes). Cranes from the nonsupplemented groups spent most of their time in agricultural fields, probably feeding on residual grain when available, while food-supplemented cranes spent most of their time at the feeding station. Thus, nonsupplemented and food-supplemented diets probably diverge only in winter, when crop rotation and depletion of anthropogenic resources may lead to a more variable diet in nonsupplemented sites. Our results support the role of diet in structuring bacterial communities and show that they undergo both seasonal and human-induced shifts. Movement analyses provide important clues regarding host diet and behaviour towards understanding how human-induced changes shape the gut microbiota in wild animals.

Individual environmental niches in mobile organisms.

Individual variation is increasingly recognized as a central component of ecological processes, but its role in structuring environmental niche associations remains largely unknown. Species' responses to environmental conditions are ultimately determined by the niches of single individuals, yet environmental associations are typically captured only at the level of species. Here, we develop scenarios for how individual variation may combine to define the compound environmental niche of populations, use extensive movement data to document individual environmental niche variation, test associated hypotheses of niche configuration, and examine the consistency of individual niches over time. For 45 individual white storks (Ciconia ciconia; 116 individual-year combinations), we uncover high variability in individual environmental associations, consistency of individual niches over time, and moderate to strong niche specialization. Within populations, environmental niches follow a nested pattern, with individuals arranged along a specialist-to-generalist gradient. These results reject common assumptions of individual niche equivalency among conspecifics, as well as the separation of individual niches into disparate parts of environmental space. These findings underscore the need for a more thorough consideration of individualistic environmental responses in global change research.

A role for lakes in revealing the nature of animal movement using high dimensional telemetry systems.

Movement ecology is increasingly relying on experimental approaches and hypothesis testing to reveal how, when, where, why, and which animals move. Movement of megafauna is inherently interesting but many of the fundamental questions of movement ecology can be efficiently tested in study systems with high degrees of control. Lakes can be seen as microcosms for studying ecological processes and the use of high-resolution positioning systems to triangulate exact coordinates of fish, along with sensors that relay information about depth, temperature, acceleration, predation, and more, can be used to answer some of movement ecology's most pressing questions. We describe how key questions in animal movement have been approached and how experiments can be designed to gather information about movement processes to answer questions about the physiological, genetic, and environmental drivers of movement using lakes. We submit that whole lake telemetry studies have a key role to play not only in movement ecology but more broadly in biology as key scientific arenas for knowledge advancement. New hardware for tracking aquatic animals and statistical tools for understanding the processes underlying detection data will continue to advance the potential for revealing the paradigms that govern movement and biological phenomena not just within lakes but in other realms spanning lands and oceans.