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Conserving habitat for migratory ungulates: How wide is a migration corridor?

Conserving migratory ungulates relies on the analysis of GPS collar data and associated maps of migration corridors to inform management and policy actions. Current methods for identifying migratory corridors use complex statistical models designed to account for movement uncertainty rather than estimating the amount of space required by animals to migrate. Furthermore, such methods can complicate conservation efforts by producing highly variable corridor widths and non-contiguous corridors that do not fully connect seasonal ranges. To remedy, we propose an intuitive line buffer approach for delineating individual migration corridors that is simple to implement and focuses on the functional corridor widths needed by migratory ungulates. By buffering a line that connects successive GPS locations, we can delineate individual migration corridors with consistent widths that are robust to variable parameters (GPS fix rate, travel speed, tortuosity) and provide contiguous connection between seasonal ranges. Using a combination of expert knowledge, simulation and 10-min GPS collar data collected from mule deer ( Odocoileus hemionus ) and pronghorn ( Antilocapra americana ), we suggest 400–600 m are reasonable estimates of functional migration corridor widths for individuals of those species. Synthesis and applications . Our line buffer approach is intended to simplify migration corridor delineation, improve transparency and encourage a broader discussion of functional corridor widths. These considerations help advance efforts to conserve habitat within migration corridors and prioritize conservation efforts within a single corridor or across multiple corridors.

Journal of Applied Ecology

Migration strategies supporting salmonids in Arctic Rivers: A case study of Arctic Cisco and Dolly Varden

Amphidromous fish such as Dolly Varden (Salvelinus malma) and Arctic Cisco (Coregonus autumnalis) have distinct life histories that facilitate their success in Arctic environments. Both species spawn in freshwater and make annual migrations between marine, brackish, or freshwater environments. Dolly Varden rear for one or more years in freshwater before migrating to sea whereas Arctic Cisco migrate to sea during their first summer. By contrast, Pacific salmon (Oncorhynchus spp.) spawn in freshwater, but once they smolt and go to sea they remain there until they mature and return to spawn. Salmon migrate at variable ages depending on species. Arctic marine environments offer productive food resources during summer, but during winter they are too cold for salmonids that lack antifreeze proteins. To avoid the cold sea during winter, Dolly Varden return to freshwater while Arctic Cisco overwinter in brackish estuaries. The lack of migration back to freshwater for overwintering helps explain why Pacific salmon success is limited in Arctic waters and suggests major increases in success will not be realized until Arctic seas provide suitable overwinter conditions. In this paper we contrast these migration strategies, discuss potential changes in a warming Arctic, and highlight information needs especially for juvenile fish.

Alaska

Moving across the border: Modeling migratory bat populations

The migration of animals across long distances and between multiple habitats presents a major challenge for conservation. For the migratory Mexican free-tailed bat (Tadarida brasiliensis mexicana), these challenges include identifying and protecting migratory routes and critical roosts in two countries, the United States and Mexico. Knowledge and conservation of bat migratory routes is critical in the face of increasing threats from climate change and wind turbines that might decrease migratory survival. We employ a new modeling approach for bat migration, network modeling, to simulate migratory routes between winter habitat in southern Mexico and summer breeding habitat in northern Mexico and the southwestern United States. We use the model to identify key migratory routes and the roosts of greatest conservation value to the overall population. We measure roost importance by the degree to which the overall bat population declined when the roost was removed from the model. The major migratory routes—those with the greatest number of migrants—were between winter habitat in southern Mexico and summer breeding roosts in Texas and the northern Mexican states of Sonora and Nuevo Leon. The summer breeding roosts in Texas, Sonora, and Nuevo Leon were the most important for maintaining population numbers and network structure – these are also the largest roosts. This modeling approach contributes to conservation efforts by identifying the most influential areas for bat populations, and can be used as a tool to improve our understanding of bat migration for other species. We anticipate this approach will help direct coordination of habitat protection across borders.

Ecosphere

Revealing migratory path, important stopovers and non-breeding areas of a boreal songbird in steep decline

The Olive-sided Flycatcher (Contopus cooperi) is a steeply declining aerial insectivore with one of the longest migrations of any North American passerine. We deployed light-level geolocators and archival GPS tags on breeders in boreal Alaska to determine migratory routes, important stopovers and non-breeding locations. Data from 16 individuals revealed a median 23,555 km annual journey (range: 19,387, 27,292 km) over 95 days (range: 83, 139 days) with wintering occurring in three regions of South America (NW Colombia/Ecuador, central Peru and W Brazil/S Peru). We developed a new method to identify “Important Stopovers” by quantifying intensity of use (a function of bird numbers and stop durations) along migratory routes. We identified 13 Important Stopovers that accounted for ~66% of the annual migratory period, suggestive of refueling activities. Some sites coincided with key areas previously identified for other Neotropi- cal-Nearctic migrants. Percent land “protected” at Impor- tant Stopovers, as defined by IUCN, ranged from 3.8% to 49.3% (mean [95% CI]: 17.3% [9.6, 25.0]). Total migration speed did not differ by season (median: 255 km day-1, range: 182, 295km day-1), despite greater spring travel dis- tances. Birds with longer non-breeding periods, however, migrated north faster. Climate-driven mismatches in migratory timing may be less of a concern for western than for eastern flycatcher populations, given recent con- generic analyses (C. sordidulus, C. virens). However, accel- erated high-latitude changes, may nonetheless impact boreal breeders.

Alaska

On the interface between cultural transmission, phenotypic diversity, demography and the conservation of migratory ungulates

Recent evidence indicates that green-wave surfing behaviour in ungulates and the migrations that stem from this behaviour are socially learned, culturally transmitted across generations and become more efficient via cumulative cultural evolution. But given a lack of corroborative evidence, whether ungulate migration is a cultural phenomenon remains a hypothesis deserving of further testing. In this opinion piece, we summarize the role memory and social learning play in the green-wave surfing that underlies ungulate migration, and when combined with the natural history of ungulates, we argue that the most likely mechanism for maintenance of ungulate migration is animal culture. We further our argument by providing a synopsis of processes that promote diversification of migratory behaviour and link these processes to their emergent ecological patterns, which are common in nature but have not historically been considered as potential cultural phenomena. The notion that diverse portfolios of migratory behaviour may buffer populations from environmental change emerges from this synthesis but requires empirical testing. Finally, we contend that, because the migratory behaviour of ungulates stems largely from cultural transmission as opposed to a genetic programme, the diversity of observed migratory strategies represents ‘culturally significant units’ deserving of the same conservation effort afforded to evolutionarily significant units.

Wyoming

Single receiver target localization in mobile marine acoustic telemetry

Many marine ecological studies track animal movement and migration using acoustic telemetry, in which animals are implanted with acoustic transmitters broadcasting a unique ID code. Receivers detect and decode these transmissions and provide an estimate of the animal’s location. As transmissions can be detected at distances of hundreds of meters to even several kilometers, localization performed using binary detection information is very coarse. Fine-scale localization can be performed using time difference of arrival systems, but such systems are costlier and require a greater number of receivers. We present a Bayesian method for target localization using only binary detection information collected by a single receiver mounted on a mobile platform. The approach reduces error in the location estimate by considering the distance-dependent probability of detection (detection efficiency curve), which must first be characterized. Stationary receivers are used to infer failed detections. This localization method is then used to inform waypoint selection for the mobile platform, using the second nondominated sorting genetic algorithm considering the tradeoff between the travel distance of the platform and localization error. Simulation results examine the effects of detection efficiency curve characteristics, number of transmissions, and travel distance on the localization error. The proposed method is further supported by experimental results from field trials conducted with an autonomous surface vehicle in Lake Huron.

IEEE Journal of Oceanic Engineering

Seasonally-dynamic presence-only species distribution models for a cryptic migratory bat impacted by wind energy development

Understanding seasonal distribution and movement patterns of animals that migrate long distances is an essential part of monitoring and conserving their populations. Compared to migratory birds and other more conspicuous migrants, we know very little about the movement patterns of many migratory bats. Hoary bats ( Lasiurus cinereus ), a cryptic, wide-ranging, long-distance migrant, comprise a substantial proportion of the tens to hundreds of thousands of bat fatalities estimated to occur each year at wind turbines in North America. We created seasonally-dynamic species distribution models (SDMs) from 2,753 museum occurrence records collected over five decades in North America to better understand the seasonal geographic distributions of hoary bats. We used 5 SDM approaches: logistic regression, multivariate adaptive regression splines, boosted regression trees, random forest, and maximum entropy and consolidated outputs to generate ensemble maps. These maps represent the first formal hypotheses for sex- and season-specific hoary bat distributions. Our results suggest that North American hoary bats winter in regions with relatively long growing seasons where temperatures are moderated by proximity to oceans, and then move to the continental interior for the summer. SDMs suggested that hoary bats are most broadly distributed in autumn—the season when they are most susceptible to mortality from wind turbines; this season contains the greatest overlap between potentially suitable habitat and wind energy facilities. Comparing wind-turbine fatality data to model outputs could test many predictions, such as ‘risk from turbines is highest in habitats between hoary bat summering and wintering grounds’. Although future field studies are needed to validate the SDMs, this study generated well-justified and testable hypotheses of hoary bat migration patterns and seasonal distribution.

PLoS ONE

Landscape movements by two species of migratory nectar-feeding bats (Leptonycteris) in a northern area of seasonal sympatry

Animals often migrate to exploit seasonally ephemeral food. Three species of nectar-feeding phyllostomid bats migrate north from Mexico into deserts of the United States each spring and summer to feed on blooms of columnar cactus and century plants ( Agave spp.). However, the habitat needs of these important desert pollinators are poorly understood. We followed the nighttime movements of 2 species of long-nosed bats ( Leptonycteris yerbabuenae and L. nivalis ) in an area of late-summer sympatry at the northern edges of their migratory ranges. We radio-tracked bats in extreme southwestern New Mexico during 22 nights over 2 summers and acquired location estimates for 31 individuals. Both species cohabitated 2 major day roosts that were 30 km apart and in different mountain ranges, and individual bats sometimes moved between the roosts. Sampling was opportunistic and limited, but there were no obvious qualitative differences in observed patterns of movement between species or years, or among sex, age, and reproductive groups. Both species were observed foraging most often in the mountain range that had a relatively higher observed density of presumed food plants ( Agave palmeri ); when roosting in an adjacent mountain range, bats sometimes commuted >20 km one way to forage. Contrary to evidence indicating these species partition resources farther south in Mexico, our findings suggest that L. yerbabuenae and L. nivalis seasonally share common roost and food resources during late summer in this northern area of sympatry.

Western North American Naturalist

Phenological variation in spring migration timing of adult alewife (Alosa pseudoharengus) in coastal Massachusetts

The timing of biological events in plants and animals, such as migration and reproduction, is shifting due to climate change. Anadromous fishes are particularly susceptible to these shifts as they are subject to strong seasonal cycles when transitioning between marine and freshwater habitats to spawn. We used linear models to determine the extent of phenological shifts in adult Alewife Alosa pseudoharengus as they migrated from ocean to freshwater environments during spring to spawn at 12 sites along the northeastern USA. We also evaluated broadscale oceanic and atmospheric drivers that trigger their movements from offshore to inland habitats, including sea surface temperature, North Atlantic Oscillation index, and Gulf Stream index. Run timing metrics of initiation, median (an indicator of peak run timing), end, and duration were found to vary among sites. Although most sites showed negligible shifts towards earlier timing, statistically significant changes were detected in three systems. Overall, winter sea surface temperature, spring and fall transition dates, and annual run size were the strongest predictors of run initiation and median dates, while a combination of within-season and seasonal-lag effects influenced run end and duration timing. Disparate results observed across the 12 spawning runs suggest that regional environmental processes were not consistent drivers of phenology and local environmental and ecological conditions may be more important. Additional years of data to extend time series and monitoring of Alewife timing and movements in nearshore habitats may provide important information about staging behaviors just before adults transition between ocean and freshwater habitats.

Massachusetts

An integrated path for spatial capture–recapture and animal movement modeling

Ecologists and conservation biologists increasingly rely on spatial capture–recapture (SCR) and movement modeling to study animal populations. Historically, SCR has focused on population-level processes (e.g., vital rates, abundance, density, and distribution), whereas animal movement modeling has focused on the behavior of individuals (e.g., activity budgets, resource selection, migration). Even though animal movement is clearly a driver of population-level patterns and dynamics, technical and conceptual developments to date have not forged a firm link between the two fields. Instead, movement modeling has typically focused on the individual level without providing a coherent scaling from individual- to population-level processes, whereas SCR has typically focused on the population level while greatly simplifying the movement processes that give rise to the observations underlying these models. In our view, the integration of SCR and animal movement modeling has tremendous potential for allowing ecologists to scale up from individuals to populations and advancing the types of inferences that can be made at the intersection of population, movement, and landscape ecology. Properly accounting for complex animal movement processes can also potentially reduce bias in estimators of population-level parameters, thereby improving inferences that are critical for species conservation and management. This introductory article to the Special Feature reviews recent advances in SCR and animal movement modeling, establishes a common notation, highlights potential advantages of linking individual-level (Lagrangian) movements to population-level (Eulerian) processes, and outlines a general conceptual framework for the integration of movement and SCR models. We then identify important avenues for future research, including key challenges and potential pitfalls in the developments and applications that lie ahead.

Ecology

One hundred pressing questions on the future of global fish migration science, conservation, and policy

Migration is a widespread but highly diverse component of many animal life histories. Fish migrate throughout the world's oceans, within lakes and rivers, and between the two realms, transporting matter, energy, and other species (e.g., microbes) across boundaries. Migration is therefore a process responsible for myriad ecosystem services. Many human populations depend on the presence of predictable migrations of fish for their subsistence and livelihoods. Although much research has focused on fish migration, many questions remain in our rapidly changing world. We assembled a diverse team of fundamental and applied scientists who study fish migrations in marine and freshwater environments to identify pressing unanswered questions. Our exercise revealed questions within themes related to understanding the migrating individual's internal state, navigational mechanisms, locomotor capabilities, external drivers of migration, the threats confronting migratory fish including climate change, and the role of migration. In addition, we identified key requirements for aquatic animal management, restoration, policy, and governance. Lessons revealed included the difficulties in generalizing among species and populations, and in understanding the levels of connectivity facilitated by migrating fishes. We conclude by identifying priority research needed for assuring a sustainable future for migratory fishes.

Frontiers in Ecology and Evolution

A statistical framework for modelling migration corridors

1. Management of animal populations requires spatially explicit knowledge of movement corridors, such as those used during seasonal migrations. GPS tracking data allows for mapping of corridors from directly observed movements, providing important insights, but tracking data is absent for many populations. 2. We developed a novel statistical corridor modeling approach that predicts movement corridors from cost-distance movement models fit directly to migration tracking data. Unlike existing predictive approaches, this does not require the ad-hoc transformation of habitat suitability surfaces into resistance surfaces. We tested the ability of the approach to recover parameters used to generate simulated movements. We then used GPS data from three migrating mule deer (Odocoileus hemionus) herds in Idaho and Wyoming to model corridors as a function of elevation, slope, aspect, percent shrub, date of peak green-up, snow-off date, and human footprint. We assessed the predictive ability of the fitted models using validation tracks from the same herd as well as from the other herds. 3. The approach reproduced parameters used to generate the simulated movements, predicted the corridors used by migratory populations, and described the direction, magnitude, and confidence levels of the effects of environmental variables on corridors. The effects environmental variables had on corridors differed depending on the herd. Within-herd validation indicated that fitted corridor models are more accurate at predicting migration corridors than null models, and cross-herd validation indicated that fitted models for some herds accurately predicted the observed migrations of other herds. 4. In addition to the practical benefit of mapping corridors for management, our statistical corridor modeling framework sets the stage for evaluating fundamental questions about the fitness tradeoffs, navigation, learning, fidelity, and movement constraints that influence migratory and other corridor-generating behavior. Models of predictive corridors can inform management and planning for the conservation of migrations across taxa, including the potential restoration of corridors. Our corridor modeling approach is also readily applied to non-migratory animal movements.

Idaho, Wyoming

Baylisascaris Larva Migrans

Summary Baylisascaris procyonis , the common raccoon roundworm, is the most commonly recognized cause of clinical larva migrans (LM) in animals, a condition in which an immature parasitic worm or larva migrates in a host animal’s tissues, causing obvious disease. Infection with B. procyonis is best known as a cause of fatal or severe neurologic disease that results when the larvae invade the brain, the spinal cord, or both; this condition is known as neural larva migrans (NLM). Baylisascariasis is a zoonotic disease, that is, one that is transmissible from animals to humans. In humans, B. procyonis can cause damaging visceral (VLM), ocular (OLM), and neural larva migrans. Due to the ubiquity of infected raccoons around humans, there is considerable human exposure and risk of infection with this parasite. The remarkable disease-producing capability of B. procyonis in animals and humans is one of the most significant aspects of the biology of ascarids (large roundworms) to come to light in recent years. Infection with B. procyonis has important health implications for a wide variety of free-ranging and captive wildlife, zoo animals, domestic animals, as well as human beings, on both an individual and population level. This report, eighth in the series of U.S. Geological Survey Circulars on zoonotic diseases, will help us to better understand the routes of Baylisascaris procyonis infections and how best to adequately monitor this zoonotic disease.

Circular

Miocene mammals and Central American seaways

The presence of Miocene mammals of North American affinity in the Panama Canal Zone indicates that Central America was attached to North America. That this attachment was a broad and stable land mass is shown by the close relation of the Panama Miocene herbivores to the widely distributed Miocene herbivore fauna of North America. A continuous connection existed probably throughout the Tertiary, to the west and north of the isthmian region, but the tectonically active isthmus probably was broken up into an archipelago during most of Tertiary time. Between the islands ran the Strait of Panama; from time to time parts of the isthmian area were connected to the stable land to the west, allowing eastward migration of land animals. The mammals of North American affinity in the Cucaracha Formation were found only a few kilometers from the western end of the San Blas Area, a stable land mass in eastern Panama that was separated from South America by the Bolivar Trough during most of the interval between Oligocene and Pliocene time. The Strait of Panama was a less stable barrier than the Bolivar Trough; this being so, it is likely that the San Blas Area was inhabited by land animals of North American rather than South American affinity. Thus, the disappearance of the Bolivar seaway in Pliocene time would have allowed, probably for the first time, mingling of the North and South American mammal faunas.

Panama Canal

Hydrology, phenology and the USA National Phenology Network

Phenology is the study of seasonally-recurring biological events (such as leaf-out, fruit production, and animal reproduction and migration) and how these events are influenced by environmental change. Phenological changes are some of the most sensitive biological indicators of climate change, and also affect nearly all aspects of ecosystem function. Spatially extensive patterns of phenological observations have been closely linked with climate variability. Phenology and hydrology are closely linked and affect one another across a variety of scales, from leaf intercellular spaces to the troposphere, and over periods of seconds to centuries. Ecosystem life cycles and diversity are also influenced by hydrologic processes such as floods and droughts. Therefore, understanding the relationships between hydrology and phenology is increasingly important in understanding how climate change affects biological and physical systems.

Fact Sheet

Examining speed versus selection in connectivity models using elk migration as an example

Context Landscape resistance is vital to connectivity modeling and frequently derived from resource selection functions (RSFs). RSFs estimate relative probability of use and tend to focus on understanding habitat preferences during slow, routine animal movements (e.g., foraging). Dispersal and migration, however, can produce rarer, faster movements, in which case models of movement speed rather than resource selection may be more realistic for identifying habitats that facilitate connectivity. Objective To compare two connectivity modeling approaches applied to resistance estimated from models of movement rate and resource selection. Methods Using movement data from migrating elk, we evaluated continuous time Markov chain (CTMC) and movement-based RSF models (i.e., step selection functions [SSFs]). We applied circuit theory and shortest random path (SRP) algorithms to CTMC, SSF and null (i.e., flat) resistance surfaces to predict corridors between elk seasonal ranges. We evaluated prediction accuracy by comparing model predictions to empirical elk movements. Results All connectivity models predicted elk movements well, but models applied to CTMC resistance were more accurate than models applied to SSF and null resistance. Circuit theory models were more accurate on average than SRP models. Conclusions CTMC can be more realistic than SSFs for estimating resistance for fast movements, though SSFs may demonstrate some predictive ability when animals also move slowly through corridors (e.g., stopover use during migration). High null model accuracy suggests seasonal range data may also be critical for predicting direct migration routes. For animals that migrate or disperse across large landscapes, we recommend incorporating CTMC into the connectivity modeling toolkit.

Landscape Ecology

The extra mile: Ungulate migration distance alters the use of seasonal range and exposure to anthropogenic risk

Partial migration occurs across a variety of taxa and has important ecological and evolutionary consequences. Among ungulates, studies of partially migratory populations have allowed researchers to compare and contrast performance metrics of migrants versus residents and examine how environmental factors influence the relative abundance of each. Such studies tend to characterize animals discretely as either migratory or resident, but we suggest that variable migration distances within migratory herds are an important and overlooked form of population structure, with potential consequences for animal fitness. We examined whether the variation in individual migration distances (20–264 km) within a single wintering population of mule deer ( Odocoileus hemionus ) was associated with several critical behavioral attributes of migration, including timing of migration, time allocation to seasonal ranges, and exposure to anthropogenic mortality risks. Both the timing of migration and the amount of time animals allocated to seasonal ranges varied with migration distance. Animals migrating long distances (150–250 km) initiated spring migration more than three weeks before than those migrating moderate (50–150 km) or short distances (<50 km). Across an entire year, long-distance migrants spent approximately 100 more days migrating compared to moderate- and short-distance migrants. Relatedly, winter residency of long-distance migrants was 71 d fewer than for animals migrating shorter distances. Exposure to anthropogenic mortality factors, including highways and fences, was high for long-distance migrants, whereas vulnerability to harvest was high for short- and moderate-distance migrants. By reducing the amount of time that animals spend on winter range, long-distance migration may alleviate intraspecific competition for limited forage and effectively increase carrying capacity. Clear differences in winter residency, migration duration, and risk of anthropogenic mortality among short-, moderate-, and long-distance migrants suggest fitness trade-offs may exist among migratory segments of the population. Future studies of partial migration may benefit from expanding comparisons of residents and migrants, to consider how variable migration distances of migrants may influence the costs and benefits of migration.

Wyoming

Testing four hypotheses to explain partial migration: Balancing reproductive benefits with limits to fasting endurance

Seasonal migration is ubiquitous in animals, and yet its underlying cause(s) remain poorly known. Species exhibiting short-distance altitudinal migration and intraspecific variation in migratory behavior (partial or differential migration) are ideal study systems for examining the selective pressures that affect individual migratory decisions. We used an individually marked population of yellow-eyed juncos, breeding along a 1000-m elevational gradient and migrating up and down that gradient, to examine the morphological, behavioral, and reproductive traits associated with migratory behavior. We tested the four most well-known hypotheses proposed to explain partial migration: the thermal tolerance, fasting endurance, dominance, and arrival time hypotheses. Our results indicate that: (1) limits to juncos’ fasting endurance constrain their ability to overwinter at high elevations, in support of the fasting endurance hypothesis, (2) differences in body size mediate fasting ability and are associated with variation in migratory behavior and overwinter apparent survival, (3) migratory behavior interacts with reproductive success, in partial support of the arrival time hypothesis, and (4) additional mechanisms that are not captured by the four well-known hypotheses might better explain individual variation in migratory behavior. Less migratory females achieved greater nesting success the following breeding season. Among males, nesting success influenced migratory tendency the following winter. Successful males may either migrate to a more benign winter climate without paying reproductive costs, or high levels of parental effort might physiologically constrain their ability to overwinter in harsh climates.

Behavioral Ecology and Sociobiology