Altitudinal migration by birds: a review of the literature and a comprehensive list of species

Altitudinal migration is the seasonal altitudinal movement of birds from breeding areas to non‐breeding or wintering areas at different elevations. Although this type of migration is widely reported, questions remain concerning the number of species that perform altitudinal migration, possible variation among different taxa and geographic locations in the extent of altitudinal migration, and the foraging guilds of altitudinal migrants. We conducted an extensive bibliographic survey and compiled a list of altitudinal migrant birds worldwide. We characterized species in terms of their foraging guilds because the spatial distribution of food resources along altitudinal gradients is often evoked as a driver of bird altitudinal migration. We identified 1238 species of altitudinal migrants, ~10% of the ~10,000 extant species of birds. We found a strong geographic bias in publications focusing on avian altitudinal migration toward the United States and Costa Rica, and a paucity of studies in megadiverse regions such as the Afrotropical and Indomalayan realms, and areas in the Neotropics other than Costa Rica. We also found that most species of altitudinal migrants were invertivores rather than frugivores or nectarivores. This general pattern held true for all zoogeographic realms except the Neotropics, where nectarivores and frugivores predominated among altitudinal migrants. The prevalence of invertivore birds among altitudinal migrants is not unexpected because this is the most common foraging guild among birds worldwide. Overall, we found no prevalence of any specific foraging guild among altitudinal migrants across zoogeographic regions. The results of studies to date suggest that altitudinal migration by birds may be driven by a number of factors, including access to increased food resources for breeding or molting, weather conditions, and mating and nesting opportunities. However, to better understand the mechanisms underlying altitudinal migration, broadening the geographic scope of studies is paramount, with additional study of altitudinal migration especially needed in the megadiverse tropical regions of sub‐Saharan Africa, Southeast Asia, and South America.     

Storms drive altitudinal migration in a tropical bird

Although migration is a widespread and taxonomically diverse behaviour, the ecological factors shaping migratory behaviour are poorly understood. Like other montane taxa, many birds migrate along elevational gradients in the tropics. Forty years ago, Alexander Skutch postulated that severe storms could drive birds to migrate downhill. Here, we articulate a novel mechanism that could link storms to mortality risks via reductions in foraging time and provide, to our knowledge, the first tests of this hypothesis in the White-ruffed Manakin (Corapipo altera), a small partially migratory frugivore breeding on the Atlantic slope of Costa Rica. As predicted, variation in rainfall was associated with plasma corticosterone levels, fat stores, plasma metabolites and haematocrit. By collecting data at high and low elevation sites simultaneously, we also found that high-elevation residents were more adversely affected by storms than low elevation migrants. These results, together with striking temporal capture patterns of altitudinal migrants relative to storms, provide, to our knowledge, the first evidence that weather-related risks incurred by species requiring high food intake rates can explain altitudinal migrations of tropical animals. These findings resolve conflicting evidence for and against food limitation being important in the evolution of this behaviour, and highlight how endogenous and exogenous processes influence life-history trade-offs made by individuals in the wild. Because seasonal storms are a defining characteristic of most tropical ecosystems and rainfall patterns will probably change in ensuing decades, these results have important implications for understanding the ecology, evolution and conservation of tropical animals.     

Escaping peril: perceived predation risk affects migratory propensity

Although migratory plasticity is increasingly documented, the ecological drivers of plasticity are not well understood. Predation risk can influence migratory dynamics, but whether seasonal migrants can adjust their migratory behaviour according to perceived risk is unknown. We used electronic tags to record the migration of individual roach (Rutilus rutilus), a partially migratory fish, in the wild following exposure to manipulation of direct (predator presence/absence) and indirect (high/low roach density) perceived predation risk in experimental mesocosms. Following exposure, we released fish in their lake summer habitat and monitored individual migration to connected streams over an entire season. Individuals exposed to increased perceived direct predation risk (i.e. a live predator) showed a higher migratory propensity but no change in migratory timing, while indirect risk (i.e. roach density) affected timing but not propensity showing that elevated risk carried over to alter migratory behaviour in the wild. Our key finding demonstrates predator-driven migratory plasticity, highlighting the powerful role of predation risk for migratory decision-making and dynamics.     

Does migration promote or restrict circumpolar breeding ranges of arctic birds?

Aim Migration has been suggested to promote large breeding ranges among birds because of the greater mobility of migratory compared with non‐migratory species, but migration has also been suggested to restrict breeding ranges because of evolutionary constraints imposed by the genetically based migration control programme. We aim to investigate the association between migration and the breeding ranges of both land birds and pelagic birds breeding in the Arctic region. Location The Arctic region. Methods Information on breeding and wintering ranges and migratory status of bird species breeding in the arctic tundra biome was compiled from the literature. The association between breeding range, migration distance and primary winter habitat was tested using multivariate generalized linear models and pair‐wise Mann–Whitney U‐tests. Phylogenetic effects were tested for using Mantel’s permutation tests. Results We found different relationships depending on the species’ major winter habitat. Among birds that are pelagic during winter, long‐distance migrants have the largest breeding ranges, while among terrestrial birds, residents and short‐distance migrants have the largest breeding ranges. Breeding ranges of coastal birds of all migratory distance classes are comparatively restricted. Main conclusions As a new explanation for this pattern we suggest that the possibility of colonizing large winter ranges is a key factor for the subsequent expansion of breeding ranges in arctic bird communities and possibly also in bird communities of other regions of the world. Because of the reversal in the relative extent of continents and oceans between the hemispheres, longitudinally wide winter ranges are more likely for long‐distance than short‐distance migrants among pelagic birds, while the reverse holds true for birds that use terrestrial winter habitats. For coastal birds both continents and oceans form barriers restricting colonization of extensive winter quarters and consequently also of extensive breeding ranges, regardless of the distance to the winter quarters.     

Postbreeding elevational movements of western songbirds in Northern California and Southern Oregon

Migratory species employ a variety of strategies to meet energetic demands of postbreeding molt. As such, at least a few species of western Neotropical migrants are known to undergo short‐distance upslope movements to locations where adults molt body and flight feathers (altitudinal molt migration). Given inherent difficulties in measuring subtle movements of birds occurring in western mountains, we believe that altitudinal molt migration may be a common yet poorly documented phenomenon. To examine prevalence of altitudinal molt migration, we used 29 years of bird capture data in a series of linear mixed‐effect models for nine commonly captured species that breed in northern California and southern Oregon. Candidate models were formulated a priori to examine whether elevation and distance from the coast can be used to predict abundance of breeding and molting birds. Our results suggest that long‐distance migrants such as Orange‐crowned Warbler (Oreothlypis celata) moved higher in elevation and Audubon's Warbler (Setophaga coronata) moved farther inland to molt after breeding. Conversely, for resident and short‐distance migrants, we found evidence that birds either remained on the breeding grounds until they finished molting, such as Song Sparrow (Melospiza melodia) or made small downslope movements, such as American Robin (Turdus migratorius). We conclude that altitudinal molt migration may be a common, variable, and complex behavior among western songbird communities and is related to other aspects of a species’ natural history, such as migratory strategy.     

Wing shape and migration in shorebirds: a comparative study

Migration is an energetically expensive and hazardous stage of the annual cycle of non‐resident avian species, and requires certain morphological adaptations. Wing shape is one of the morphological traits that is expected to be evolutionarily shaped by migration. Aerodynamic theory predicts that long‐distance migrants should have more pointed wings with distal primaries relatively longer than proximal primaries, an arrangement that minimizes induced drag and wing inertia, but this prediction has mostly been tested in passerine species. We applied the comparative method of phylogenetically independent contrasts to assess convergent evolution between wing shape and migration within shorebirds. We confirmed the assumption that long‐distance migrants have less rounded wings than species migrating shorter distances. Furthermore, wing roundedness negatively correlates with fat load and mean distance of migratory flights, the basic components of migration strategies. After controlling for interspecific differences in body size, we found no support for a link between wing length and migration, indicating that wing shape is a more important predictor of shorebird migratory behaviour than wing length. The results suggest that total migration distance and migratory strategy may simultaneously act on the evolution of wing shape in shorebirds, and possibly in other avian species.     

Programmed and flexible: long‐term Zugunruhe data highlight the many axes of variation in avian migratory behaviour

Studies of Zugunruhe – the ‘migratory restlessness’ behaviour of captive birds – have been integral to our understanding of animal migration, revealing an inherited propensity to migrate and an endogenous timing and navigation system. However, differences between Zugunruhe in captivity and migration in the wild call for more data, in particular on variation within and among taxa with diverse migration strategies. Here, we characterise Zugunruhe in a long‐term dataset of activity profiles from stonechats (genus Saxicola) with diverse migratory phenotypes (976 migration periods from 414 birds), using a flexible and consistent quantitative approach based on changepoint analysis. For east African, Austrian, Irish, and Siberian stonechats and hybrids, we report key inter‐population differences in the occurrence, timing, and intensity of Zugunruhe. In line with expectations, we found the highest Zugunruhe intensity in the longest‐distance migrants, more variable patterns in short‐distance migrants, and intermediate characteristics of hybrids relative to their parental groups. Inter‐population differences imply high evolutionary lability of Zugunruhe timing within a robustly structured annual cycle. However, counter to theory, Irish partial migrants showed no segregation between migrant and resident individuals, and previously reported nocturnal restlessness was confirmed for resident African stonechats. Further features of nocturnal restlessness that did not align with migratory behaviour of stonechats were juvenile nocturnal restlessness even prior to postjuvenile moult, and protandry in spring, although stonechats winter in heterosexual pairs. Importantly, Zugunruhe of all populations declined with age, and the intensity of an individual bird's Zugunruhe was correlated with activity levels during other parts of the annual cycle. Our results confirm endogenous, population‐specific migration programmes but also reveal apparent discrepancies between Zugunruhe and migration in the wild. We thus highlight both the continued potential of Zugunruhe study and the need for circumspect interpretation when using migratory restlessness to make inferences about migration in the wild.     

Partial migration in birds: tests of three hypotheses in a tropical lekking frugivore

1. Partially migratory species provide opportunities to understand which ecological factors cause some animals to migrate when others remain resident year round. Partial migration in birds has been explained by the dominance, arrival-time, and body-size hypotheses. 2. Testing these hypotheses has proven difficult due to the similarities of the predictions they make in temperate-breeding long-distance migrants. In tropical altitudinal migrants, however, these hypotheses make different predictions regarding the sex, age, and condition of migrants and residents. 3. Among white-ruffed manakins in Costa Rica, young birds were not more likely to migrate (as predicted by the dominance hypothesis), nor were females more likely to migrate (as predicted by the arrival-time hypothesis). All condition-related variables interacted with sex, together explaining much of the variation in migratory behaviour. 4. I re-articulate the body-size hypothesis in the context of tropical altitudinal bird migration, focusing explicitly on how limited foraging opportunities and differences in individual condition affect fasting ability during torrential rains. Despite ample food, the smallest birds or those stressed by parasites or moult may risk starvation at breeding elevations due to a reduction in foraging time. These results highlight how intrinsic and extrinsic factors may interact to produce observed patterns of within- and among-species variation in migratory behaviour.     

Postglacial northward expansion and genetic differentiation between migratory and sedentary populations of the broad‐tailed hummingbird (Selasphorus platycercus)

Unlike other migratory hummingbirds in North America, the broad‐tailed hummingbird (Selasphorus platycercus) exhibits both long‐distance migratory behaviour in the USA and sedentary behaviour in Mexico and Guatemala. We examined the evolution of migration linked to its northward expansion using a multiperspective approach. We analysed variation in morphology, mitochondrial and nuclear DNA, estimated migration rates between migratory and sedentary populations, compared divergence times with the occurrence of Quaternary climate events and constructed species distribution models to predict where migratory and sedentary populations resided during the Last Glacial Maximum (LGM) and Last Interglacial (LIG) events. Our results are consistent with a recent northward population expansion driven by migration from southern sedentary populations. Phylogeographical analyses and population genetics methods revealed that migratory populations in the USA and sedentary populations in Mexico of the platycercus subspecies form one admixed population, and that sedentary populations from southern Mexico and Guatemala (guatemalae) undertook independent evolutionary trajectories. Species distribution modelling revealed that the species is a niche tracker and that the climate conditions associated with modern obligate migrants in the USA were not present during the LIG, which provides indirect evidence for recent migratory behaviour in broad‐tailed hummingbirds on the temporal scale of glacial cycles. The finding that platycercus hummingbirds form one genetic population and that suitable habitat for migratory populations was observed in eastern Mexico during the LIG also suggests that the conservation of overwintering sites is crucial for obligate migratory populations currently facing climate change effects.     

A long winter for the Red Queen: rethinking the evolution of seasonal migration

This paper advances an hypothesis that the primary adaptive driver of seasonal migration is maintenance of site fidelity to familiar breeding locations. We argue that seasonal migration is therefore principally an adaptation for geographic persistence when confronted with seasonality – analogous to hibernation, freeze tolerance, or other organismal adaptations to cyclically fluctuating environments. These ideas stand in contrast to traditional views that bird migration evolved as an adaptive dispersal strategy for exploiting new breeding areas and avoiding competitors. Our synthesis is supported by a large body of research on avian breeding biology that demonstrates the reproductive benefits of breeding‐site fidelity. Conceptualizing migration as an adaptation for persistence places new emphasis on understanding the evolutionary trade‐offs between migratory behaviour and other adaptations to fluctuating environments both within and across species. Seasonality‐induced departures from breeding areas, coupled with the reproductive benefits of maintaining breeding‐site fidelity, also provide a mechanism for explaining the evolution of migration that is agnostic to the geographic origin of migratory lineages (i.e. temperate or tropical). Thus, our framework reconciles much of the conflict in previous research on the historical biogeography of migratory species. Although migratory behaviour and geographic range change fluidly and rapidly in many populations, we argue that the loss of plasticity for migration via canalization is an overlooked aspect of the evolutionary dynamics of migration and helps explain the idiosyncratic distributions and migratory routes of long‐distance migrants. Our synthesis, which revolves around the insight that migratory organisms travel long distances simply to stay in the same place, provides a necessary evolutionary context for understanding historical biogeographic patterns in migratory lineages as well as the ecological dynamics of migratory connectivity between breeding and non‐breeding locations.     

Diffuse, short and slow migration among Blue Tits

The knowledge of migration systems in long-distance regular migrants is in many cases extensive. Our understanding of the migratory characteristics of partial migrants, on the other hand, is far more rudimentary. We investigated migratory characteristics of partially migratory Blue Tits Cyanistes caeruleus using ringing recoveries of Swedish birds, to answer questions about geographic migration patterns, age-specific migrations, migration speeds and synchrony of movements. Median migration distance of Swedish Blue Tits was 82 km, with a main autumn direction in the sector between S and W (large directional scatter). Northerly and southerly populations did not differ in migration directions or distances, suggesting chain migration to be the general pattern. A larger proportion of adult Blue Tits remained near the breeding grounds during winter than was the case for juveniles. Some of the migrating birds (17%) seemed not to return in spring but stayed to breed closer to the winter area. Swedish Blue Tits show an exceptionally slow migration speed (median 13 km/day), among the slowest speeds recorded for any migrant bird. The Blue Tit represents an extreme case of diffuse, short and slow bird migration.     

Identifying critical migratory bottlenecks and high‐use areas for an endangered migratory soaring bird across three continents

Migrant birds face a number of threats throughout their annual cycle, including persecution, collision with energy infrastructure, and habitat and climate change. A key challenge for the conservation of migrants is the identification of important habitat, including migratory concentration areas, because species survival rates may be determined by events in geographically very limited areas. Remote‐tracking technology is facilitating the identification of such critical habitat, although the strategic identification of important sites and incorporation of such knowledge in conservation planning remains limited. We tracked 45 individuals of an endangered, soaring migrant (Egyptian vulture Neophron percnopterus), over 75 complete migrations that traversed three continents along the Red Sea Flyway. We summarize and contextualize migration statistics by season and age class, including migration start, midpoint, and end dates, as well as linear and cumulative migration distance, migration duration and speed, and route straightness. Then, using dynamic Brownian bridge movement models, we quantified space use to identify the most important migratory bottlenecks and high‐use areas on the flyway. These areas each accounted for < 5% of the overall movement range of the tracked birds, yet > 20% of all tracks passed through bottlenecks, and > 50% of the overall vulture time spent on migration fell within high‐use areas. The most important sites were located at the southeastern Red Sea coast and Bab‐el‐Mandeb Strait (Saudi Arabia, Yemen, Djibouti), the Suez Canal zone (Egypt), and the Gulf of Iskenderun (Turkey). Discouragingly however, none of the area within the major migratory bottlenecks was protected and < 13% of the high‐use areas were protected. This demonstrates a very concerning gap in the protected area network for migratory soaring birds along the Red Sea Flyway. Because reducing threats at migratory concentrations can be a very efficient approach to protect populations, our work provides clear guidelines where conservation investment is urgently needed to benefit as many as 35 migratory soaring‐bird species that regularly use the Red Sea Flyway.     

Insect migration: Variability and success in a capricious environment

Stochastic effects of climate and weather have a pervasive influence on the induction, performance and evolution of migration. In wing-dimorphic species, their influence on habitat quality, and on rates of development of the migrant itself, maintains variation in responses to environmental cues determining wing-form and migratory behaviour. Migrants flying above their flight boundary layer rely on winds to disperse them across landscapes in which their habitats are distributed. Patterns of distribution of habitat patches, and the influence of changing windspeeds and direction on the displacements of migrants, result in selection for variation in migratory potential at each migration. In subsequent migrations, this variation and stochastic effects of the winds on groundtracks of individual migrants ensure that their destinations ‘sample’ the landscapes they travel over. The extent and resolution of this sampling, by which migrants reach favourable habitats, depend on the components of migratory potential, their mode of inheritance, and genetic correlations between them, as well as on the characteristics of the winds on which they travel.     

Independent evolution of migration on the South American landscape in a long-distance temperate-tropical migratory bird, Swainson's flycatcher (Myiarchus swainsoni)

Aim To understand the evolution of long‐distance temperate–tropical migration in a South American bird, Swainson's flycatcher (Myiarchus swainsoni). Methods A total of 842 base pairs of the mitochondrial DNA genes ATPase 8 and 6 were sequenced from forty‐nine individuals of the M. swainsoni complex from most of its range. Analyses measured the phylogenetic signal in the data, and tools of population genetics, phylogeography and phylogeny were used to interpret the evolution of the bird and its migration on the South American landscape. Results Migratory populations in the M. swainsoni complex are not each other's closest relatives. The migratory subspecies M. s. swainsoni, which breeds in south‐eastern South America, is not closely related to the rest of the complex. The remaining migratory populations of the subspecies M. s. ferocior and two intergrade populations are extremely closely related to non‐migratory populations with which they form a well‐supported clade despite substantial morphological differentiation from each other. Within this clade of migrants and non‐migrants, net divergence across 4000 km of lowland South America is zero and most diversity is distributed among individuals not populations. Mismatch analyses and significant values of Tajima's D and Fu's Fs suggest the clade has undergone a very recent range expansion. Migration and the shifts of breeding distribution that accompanied its evolution evolved twice within what has recently been considered the polytypic species M. swainsoni. Furthermore, these shifts of range probably occurred at very different times as parts of different southward ‘pulses’ of humid, Amazonian taxa. Main conclusions Evolution of temperate‐tropical migration in the M. swainsoni complex has been spatio‐temporally layered on the South American landscape. The analysis cautions that the historical biogeography underlying a single present‐day migration system need not have been driven by a single set of environmental factors operating at one time. We suggest directions for further study of ecology and demography in zones of apparent contact between various migratory and non‐migratory populations.     

Animal migration: linking models and data beyond taxonomic limits

An international workshop on animal migration was held at the Lorentz Center in Leiden, The Netherlands, 2–6 March 2009, bringing together leading theoreticians and empiricists from the major migratory taxa, aiming at the identification of cutting-edge questions in migration research that cross taxonomic borders.     

The grand challenges of migration ecology that radar aeroecology can help answer

Many migratory species have experienced substantial declines that resulted from rapid and massive expansions of human structures and activities, habitat alterations and climate change. Migrants are also recognized as an integral component of biodiversity and provide a multitude of services and disservices that are relevant to human agriculture, economy and health. The plethora of recently published studies reflects the need for better fundamental knowledge on migrations and for better management of their ecological and human‐relevant effects. Yet, where are we in providing answers to fundamental questions and societal challenges? Engaging a broad network of researchers worldwide, we used a horizon‐scan approach to identify the most important challenges which need to be overcome in order to gain a fuller understanding of migration ecology, and which could be addressed using radar aeroecological and macroecological approaches. The top challenges include both long‐standing and novel topics, ranging from fundamental information on migration routes and phenology, orientation and navigation strategies, and the multitude of effects migrants may have on resident communities, to societal challenges, such as protecting or preventing migrant services and disservices, and the conservation of migrants in the face of environmental changes. We outline these challenges, identify the urgency of addressing them and the primary stakeholders – researchers, policy makers and practitioners, or funders of research.     

Danger management and the seasonal adjustment of migratory speed by sandpipers

The behavior of migrating birds is governed by time‐, energy‐ and danger‐minimizing strategies. The adjustment of migration speed (i.e. the rate at which distance is covered during a migration) is a behavioral tactic that might contribute to these strategic goals. Shorter stopovers and greater fuel loads increase migration speed, but both require more intensive foraging at stopovers, making migrants more vulnerable to predators. A simple numerical model shows how seasonal alterations in migration speed can lower the exposure of western sandpipers to peregrine falcons, their most important predator. The ‘caution–speed–caution’ pattern of higher migration speed in the mid‐passage period, observed in earlier work, requires that the intensive foraging necessary heightens vulnerability, and that migrants are exposed to both migrant predators as well as predators resident at migratory stopovers.     

Songbirds are resilient to hurricane disturbed habitats during spring migration

The Gulf of Mexico is a conspicuous feature of the Neotropical–Nearctic bird migration system. Traveling long distances across ecological barriers comes with considerable risks, and mortality associated with intercontinental migration may be substantial, including that caused by storms or other adverse weather events. However, little, if anything, is known about how migratory birds respond to disturbance‐induced changes in stopover habitat. Isolated, forested cheniere habitat along the northern coast of the Gulf of Mexico often concentrate migrants, during weather conditions unfavorable for northward movement or when birds are energetically stressed. We expected hurricane induced degradation of this habitat to negatively affect the abundance, propensity to stopover, and fueling trends of songbirds that stopover in coastal habitat. We used spring banding data collected in coastal Louisiana to compare migrant abundance and fueling trends before (1993–1996 and 1998–2005) and after hurricanes Rita (2006) and Ike (2009). We also characterized changes in vegetative structure before (1995) and after (2010) the hurricanes. The hurricanes caused dramatic changes to the vegetative structure, which likely decreased resources. Surprisingly, abundance, propensity to stopover, and fueling trends of most migrant species were not influenced by hurricane disturbance. Our results suggest that: 1) the function of chenieres as a refuge for migrants after completing a trans‐Gulf flight may not have changed despite significant changes to habitat and decreases in resource availability, and 2) that most migrants may be able to cope with habitat disturbance during stopover. The fact that migrants use disturbed habitat points to their conservation value along the northern coast of the Gulf of Mexico.     

The decline of Afro‐Palaearctic migrants and an assessment of potential causes

There is compelling evidence that Afro‐Palaearctic (A‐P) migrant bird populations have declined in Europe in recent decades, often to a greater degree than resident or short‐distance migrants. There appear to have been two phases of decline. The first in the 1960s–1970s, and in some cases into the early 1980s, largely affected species wintering predominantly in the arid Sahelian zone, and the second since the 1980s has mostly affected species wintering in the humid tropics and Guinea forest zone. Potential drivers of these declines are diverse and are spread across and interact within the migratory cycle. Our knowledge of declining species is generally better for the breeding than the non‐breeding parts of their life cycles, but there are significant gaps in both for many species. On the breeding grounds, degradation of breeding habitats is the factor affecting the demography of the largest number of species, particularly within agricultural systems and woodland and forests. In the non‐breeding areas, the interacting factors of anthropogenic habitat degradation and climatic conditions, particularly drought in the Sahel zone, appear to be the most important factors. Based on our synthesis of existing information, we suggest four priorities for further research: (1) use of new and emerging tracking technologies to identify migratory pathways and strategies, understand migratory connectivity and enable field research to be targeted more effectively; (2) undertake detailed field studies in sub‐Saharan Africa and at staging sites, where we understand little about distribution patterns, habitat use and foraging ecology; (3) make better use of the wealth of data from the European breeding grounds to explore spatial and temporal patterns in demographic parameters and relate these to migratory pathways and large‐scale patterns of habitat change and climatic factors; and (4) make better use of remote sensing to improve our understanding of how and where land cover is changing across these extensive areas and how this impacts A‐P migrants. This research needs to inform and underpin a flyway approach to conservation, evaluating a suite of drivers across the migratory cycle and combining this with an understanding of land management practices that integrate the needs of birds and people in these areas.