2018 - Bayly, Et Al

Bird Conservation International (2018) 28:1–26.
© BirdLife International, 2017

doi:10.1017/S0959270917000296
Major stopover regions and migratory

bottlenecks for Nearctic-Neotropical landbirds
within the Neotropics: a review
NICHOLAS J. BAYLY, KENNETH V. ROSENBERG, WENDY E. EASTON,
CAMILA GÓMEZ, JAY CARLISLE, DAVID N. EWERT, ANNA DRAKE and
LAURIE GOODRICH
Summary
Nearly 300 species of landbirds, whose populations total billions, migrate between the Neotropics
and North America. Many migratory populations are in steep decline, and migration is often
identified as the greatest source of annual mortality. Identifying birds’ needs on migration is
therefore central to designing conservation actions for Nearctic-Neotropical migratory birds; yet
migration through the Neotropics is a significant knowledge gap in our understanding of the full
annual cycle. Here, we synthesise current knowledge of Neotropical stopover regions and migra-
tory bottlenecks, focusing on long-distance, migratory landbirds that spend the boreal winter in
South America. We make the important distinction between “true” stopover—involving multi-
day refuelling stops—and rest-roost stops lasting < 24 hours, citing a growing number of studies
that show individual landbirds making long stopovers in just a few strategic areas, to accumulate
large energy reserves for long-distance flights. Based on an exhaustive literature search, we found
few published stopover studies from the Neotropics, but combined with recent tracking studies,
they describe prolonged stopovers for multiple species in the Orinoco grasslands (Llanos), the
Sierra Nevada de Santa Marta (Colombia), and the Yucatan Peninsula. Bottlenecks for diurnal
migrants are well described, with the narrowing Central American geography concentrating mil-
lions of migrating raptors at several points in SE Mexico, Costa Rica, Panama and the Darién.
However, diurnally migrating aerial insectivores remain understudied, and determining stopover/
roost sites for this steeply declining group is a priority. Despite advances in our knowledge of
migration in the Neotropics, we conclude that major knowledge gaps persist. To identify stopover
sites and habitats and the threats they face, we propose a targeted and collaborative research
agenda at an expanded network of Neotropical sites, within the context of regional conservation
planning strategies.
Introduction
Nearly 300 species of landbirds, whose combined populations represent billions of birds, migrate
between the Neotropics and North America (Martin and Finch 1995, DeGraaf and Rappole 1996,
Berlanga et al. 2010). As many as 50 of these species make an annual round trip of > 10,000 km
but the number of individuals is declining year after year, especially among those species that
migrate farthest (Sauer et al. 2014). The same is apparent in Afro-Palearctic migrants (Sanderson
et al. 2006). Global declines in migratory species have led to concerns that migration and the
capacity of our ecosystems to maintain this process are in danger of disappearing (Wilcove and
Wikelski 2008, Bauer and Hoye 2014). For many species, the majority of their long-distance
N. J. Bayly et al. 2
migrations take place south of the Tropic of Cancer, yet our understanding of migration is strongly
biased by studies in the temperate zone. Consequently the Neotropics continue to be described as
the “black box” in our knowledge of migratory landbirds (Faaborg et al. 2010a).
As early as the 1980s, results from the North American Breeding Bird Survey highlighted
widespread declines in Nearctic-Neotropical migratory landbirds (Terborgh 1980, Robbins et al.
1989), and considerable efforts have been made since to both understand and address the year-
round conservation needs of this group (Keast and Morton 1980, Hagan and Johnston 1992,
Holmes 2007, Faaborg et al. 2010a). Partners in Flight (PIF), for example, was created in 1990 to
develop strategies for the conservation of migratory landbirds throughout their annual cycle, and
today is the umbrella group for a diverse set of actors from across the Americas (Finch and Martin
1995, Rich et al. 2004, Berlanga et al. 2010, Rosenberg et al. 2016). More than 30 years of research
into Neotropical migratory birds has substantially advanced our understanding of how popula-
tions may be limited by events on the breeding grounds (Holmes et al. 1996) or the wintering
grounds (Latta and Faaborg 2002, Studds and Marra 2005, Johnson et al. 2006), and how effects
may carry over from one season to the other (Nott et al. 2002, Norris et al. 2004, Norris and
Marra 2007, Harrison et al. 2011, González-Prieto and Hobson 2013). Indeed, the formulation of
the winter limitation hypothesis (Sherry and Holmes 1995) and the subsequent collection of
empirical evidence for the limiting role of winter habitats have firmly placed Neotropical winter-
ing grounds on the research and conservation agenda (Marra 2000, Latta and Faaborg 2002,
Studds and Marra 2005). The same cannot be said, however, of the ecological needs and con-
straints of these same birds while migrating through the Neotropics (Faaborg et al. 2010a), despite
the considerable distances that many long-distance migrants must traverse there.
For some bird species, migration is by far the greatest source of mortality during their annual
cycle (Sillett and Holmes 2002, Newton 2006, Rockwell et al. 2016). For example, successive
delayed arrivals or habitat degradation at even a single major stopover site can lead to significant
declines, threatening the viability of populations across the Western Hemisphere (Baker et al.
2004). Although some en-route flexibility exists (Tottrup et al. 2008, Stanley et al. 2012), migra-
tion strategies are under strong genetic control (Gwinner 1996, Delmore and Irwin 2014), and
migrants are expected to be limited in their ability to change routes if conditions change or
stopover habitats are no longer available in the short term but may be able to adapt in the longer
term (Sutherland 1998). Recognition of the critical importance of migration stopovers and bottle-
necks has led to repeated calls for more studies and collaborations into the ecology and needs of birds
during the migration periods (Hutto 1998, Heglund and Skagen 2005, Mehlman et al. 2005, Faaborg
et al. 2010a, 2010b).
These appeals have led to more work within North America, especially along the USA Gulf
Coast (Buler et al. 2007, Moore and Buler 2011, Laughlin et al. 2013, Cohen et al. 2017), the
Pacific flyway (Carlisle et al. 2009), riparian forests in the south-west (Yong and Finch 1997, Yong
et al. 1998) and around the Great Lakes (Dunn 2001, 2002, Ewert et al. 2011). For many migra-
tory species, however, the majority of their migratory route lies south of these well-studied
areas. For example, millions of migrating landbirds converge each year on the northern Neotropics
(La Sorte et al. 2016), and species whose breeding range may stretch over 4,000 km from east to
west concentrate into areas spanning less than 100 km as a result of the funnel-shaped geography
of Central America (Bildstein 2004). Yet, despite this obvious importance, migration studies of
Nearctic-Neotropical landbirds south of the United States of America (USA) and Canada remain
rare, limiting our ability to design effective conservation strategies that address their needs
throughout the annual cycle.
Nonetheless, the last few decades have seen studies from Mexico’s Gulf Coast and the Yucatan
Peninsula (Winker 1995, Deppe and Rotenberry 2005, Johnson and Winker 2008, Bayly and
Gómez 2011, Shaw and Winker 2011), and from northern Colombia (Colorado 2010, Bayly et al.
2012a, 2013, Gómez et al. 2013, 2015), in some cases demonstrating the critical importance of
stopover sites within the Neotropics (Bayly et al. 2012a, 2013). Enormous gaps remain, however,
and in particular our knowledge of how nocturnally migrating birds use the region south of the
Stopover regions within the Neotropics 3
Yucatan Peninsula through Central America to northern South America; large Caribbean Islands;
and especially regions within South America, is poor. In addition, for over 30 species that winter
primarily in South America, much confusion persists around which regions are used during
migration because of imprecisely mapped winter distributions and the difficulty of distinguishing
migrating from wintering individuals (Remsen 2001). Some of these gaps are being partially
filled as a result of innovative lightweight tracking technologies and their application to a growing
number of Neotropical migratory landbirds (McKinnon et al. 2013a, Taylor et al. 2017). The
migration of raptors is better understood, with multiple studies detailing the main migration
routes (Martell et al. 2001, Bildstein 2004, Porras-Peñaranda et al. 2004, Colorado et al. 2006,
Ruelas Inzunza et al. 2010a, Kochert et al. 2011, Bayly et al. 2014).
While general patterns of migration speed and routes have been shown to differ between spring
and autumn migrations for many species (Gómez et al. 2013, La Sorte et al. 2014a, 2014b,
Hobson and Kardynal 2015), detailed seasonal comparisons for most species and regions are
lacking. In addition, the strategies and needs of diurnal and nocturnal migrating species are
expected to differ markedly, making generalisations across species groups difficult. Given the
critical importance of migration to annual survival and fitness (Sillett and Holmes 2002, Finch
et al. 2014) and its ecological significance for long-term population viability, our lack of knowledge
of stopover regions and migratory bottlenecks within the Neotropics poses a serious constraint
on our ability to address declines in Nearctic-Neotropical migrants (Faaborg et al. 2010a, Sheehy
et al. 2010).
In this review we synthesise current knowledge of stopover regions and migratory bottlenecks
within the Neotropics, focusing on diurnal and nocturnally migrating landbirds. Our primary
objective is to stimulate research in regions and for species that lack even baseline data. We out-
line potential threats at known and suspected stopover regions, and present an emerging picture
of the migration strategies of long-distance migratory birds in the Neotropics. We also propose
consistent terminology that, for example, can make the distinction between “true” stopover—
where birds engage in multi-day stops with significant refuelling—and rest-roost stops lasting
one day or less with no or limited refuelling. Then, based on a conceptual framework for under-
standing the needs of migratory birds, we propose a targeted and collaborative research agenda at
an expanded network of sites across the Neotropics within the context of new regional conserva-
tion planning strategies (e.g. Rosenberg et al. 2016).
The needs of migrating landbirds in the Neotropics

To successfully migrate between their breeding and over-wintering grounds, where over-wintering
is defined as the stationary period/s of the non-breeding season during the boreal winter (Table 1),
Nearctic-Neotropical migrants depend on a series of sites along the length of their migratory
route. These sites provide critical resources such as the fuel for migratory flights, safe roosting
sites, and refuges during unfavourable climatic conditions (Mehlman et al. 2005, Newton 2008).
A successful migration also depends on birds avoiding in-flight hazards such as collisions with
human infrastructure (Longcore et al. 2013) and persecution by humans (Newton 2006). Migrating
landbirds stop at multiple sites across regions, yet not all sites are of equal importance for deter-
mining the speed of migration or survival (Weber et al. 1998). Indeed, diurnal migrants typically
stop on a nightly basis between daytime flights, while nocturnal migrants generally stop on a
daily basis when migrating overland (Newton 2008). As a consequence, migrants can be encoun-
tered at numerous sites along the length of their migratory routes, making it difficult to differen-
tiate between sites used to rest between flights and those used to accumulate energy reserves
(Warnock 2010).
The lack of distinction between these two very different types of “stopover sites” has given
rise to a general perception that most migratory landbirds use numerous stopover sites during
any one migration (McKinnon et al. 2013a) and that population level movement across space
is continuous. Because of this perception, research, conservation, and management activities
N. J. Bayly et al.
Table 1. Stopover and migration terminology adopted throughout this review and its equivalence to the categories proposed by Mehlman et al. (2005), which was used as the
foundation.
Type of site or region Description Equivalent Mehlman et al.
Passage Geographical points or corridors where birds pass in continuous flight during migration N/A
Bottleneck Passage points where geography constrains migration, leading to significant concentrations of populations that N/A
could elevate threats or risks.
Rest-Roost Places associated with resting or roosting behaviours between successive migratory flights that do not involve Similar to convenience store
significant refuelling (likely applies to the majority of sites used on migration where refuelling is equivalent
to <5% of lean body mass).
Fire escape Places where migrants only make landfall when they encounter adverse conditions that prohibit onward flights; Fire escapes
sites provide vital shelter for survival but are rarely used for refuelling.
Stopover Behaviour and places associated with multi-day stops giving rise to significant refuelling (>5% of lean body mass) Full service hotel
during migration.
Staging Behaviours and places associated with prolonged stopover not entirely dedicated to refuelling, such as where large Full service hotel
numbers of birds gather, often building in numbers until favourable cues promote subsequent migration.
Moult-migration Behaviour or places where a significant portion of a species’ population pauses on migration to undergo flight N/A
feather moult before continuing on migration.
Pre-migratory Behaviour or places where significant fuelling occurs immediately following breeding or over-wintering periods Full service hotel
that may or may not involve short local movements (<100 km).
Over-wintering Behaviour or places associated with prolonged (>1 month) stationary non-breeding periods during the boreal winter N/A
season, usually between autumn and spring migration periods. Fuelling does not take place during this period.
Given the prevalence of multiple over-wintering sites for some species, may blur with pre-migratory “staging”.
4
for birds on migration have often focused on sites where large numbers of birds are present
(Moore et al. 1990, Simons et al. 2000, Dunn 2001), without considering the ecological function
of those sites. Recent studies on a range of species have challenged this paradigm, showing that in
reality, individual landbirds often make long stopovers at just three or four key areas along the
migratory route (Figure 1) (Heckscher et al. 2011, Kochert et al. 2011, Delmore et al. 2012, Fraser
et al. 2012, Stanley et al. 2012, Callo et al. 2013, McKinnon et al. 2013a, Renfrew et al. 2013), at
which they accumulate large energy reserves (Figure 2) (Bayly et al. 2012a, 2013). These findings
highlight an urgent need to identify major stopover regions (defined here as areas varying in size
from 100 km2 to 50,000 km2 with relatively uniform environmental conditions and offering a
certain set of resources) and assess the needs of birds within them. This need is magnified in the
Neotropics where the funnel-shaped geography of Mexico and Central America acts as a bottle-
neck, concentrating millions of migratory landbirds (Porras-Peñaranda et al. 2004, Batista et al.
2005, Ruelas Inzunza et al. 2010a, La Sorte et al. 2016). This also highlights a need to revise the
current stopover terminology to better define the contribution of individual sites to migration
strategies, and to this end we present an updated terminology in Table 1 that we utilize through-
out the review.
One of the major factors driving the organisation, speed and, ultimately, the success of migra-
tion is the quality of resources at stopover sites and their impact on the rate of fuel deposition
(Hedenström 2008). Indeed, with time-minimiation emerging as the main currency shaping the
evolution of migration strategies (Alerstam 2011), migratory birds are expected to seek out sites
that maximize their fuel deposition rate and, accordingly, their speed of migration (Buler et al.
2007, La Sorte et al. 2014a). It follows that one of the fundamental needs of migratory birds is
therefore the availability of high quality habitats (those supporting high fuel deposition rates), in
specific regions along their migratory route (Weber et al. 1999, Bayly et al. 2016). These habitats
Figure 1. Spring migration strategy (northward) and stopover site use of Red-eyed Vireos from
a breeding site in the north-eastern United States as revealed by geolocators recovered from nine
individuals (adapted from Callo et al. 2013). Eight of the nine individuals appeared to make a rest-
roost stop close to the tip of the Yucatan peninsula before crossing the Gulf of Mexico.
Figure 2. Individual A. Veery and B. Grey-cheeked Thrush recaptured on more than one occasion
during an autumn and spring stopover respectively, in the Sierra Nevada de Santa Marta in
northern Colombia. Birds increased significantly in body mass, providing evidence for extensive
fuelling in this region (taken from Bayly et al. 2012a, 2013).
must not only sustain high rates of fuel deposition but also offer low to moderate predation risk,
as migratory birds may avoid sites with elevated densities of predators or trade-off their foraging
rate against predator surveillance (Lindström 1990, Schmaljohann and Dierschke 2004, Pomeroy
2006, Warnock 2010). Recent work on Catharus thrushes in northern Colombia has shown that
the energy reserves acquired at key stopovers may enable birds to cover 30% or more of their
total migratory distance (Bayly et al. 2012a, Bayly et al. 2013, Gómez et al. 2017), again high-
lighting the enormous influence that individual stopover sites can have on a migratory journey
(Figure 2).
Additional sites used by migrating birds between major stopover regions (which we will refer
to as rest-roost stops hereafter) are believed to have far less impact on the speed and organistion
of migration (Alerstam 2011); nevertheless they have the potential to affect survival (Newton
2006). Energetic requirements at rest-roost stops are generally low, as individuals may only be
resting between successive nocturnal or diurnal flights, although the ability to top up reserves
may constitute a key element of the migratory strategy in certain species (Robson et al. 2001,
Delingat et al. 2006). The primary function of rest-roost stops is therefore to provide safe roosting
conditions and access to resources such as water. Sites used as rest-roost stops may assume a
greater importance when situated close to ecological barriers, such as the Gulf of Mexico or the
Caribbean Sea (Moore et al. 1990). Under optimal conditions, birds typically overfly coastal habi-
tats either side of these barriers (Lowery 1945, Moore et al. 1990) but when they encounter
unfavourable conditions such as rain or strong headwinds, these habitats may temporarily sup-
port high concentrations of birds (Simons et al. 2000). Habitats used in such emergencies, often
referred to as “fire escapes”, can also be important away from ecological barriers (Mehlman et al.
2005). For example, passing tropical storms in Central America occasionally ground migrating
raptors for days at sites where normally they would not stop (Bildstein 2004) and possibly delay
subsequent returns to the breeding grounds.
Just as stopover sites used for migratory fuelling affect the outcome of migration, the quality
of sites occupied prior to the onset of migration can also have a fundamental impact on migration
strategies (Bearhop et al. 2004, Norris and Marra 2007). For example, many species undergo pre-
migratory fuelling on or near their Neotropical overwintering grounds, and the energy reserves
gained there may enable birds to cover a significant proportion of their northbound migration
(Stutchbury et al. 2009, Heckscher et al. 2011, DeLuca et al. 2015). Further, the quality of over-
wintering habitats can influence the timing of migration, both in terms of initiation and arrival
on the breeding grounds, with carry-over effects on territory acquisition and breeding perfor-
mance (Norris et al. 2004, Studds and Marra 2005, Norris and Marra 2007, Guillemain et al.
2008). Overwintering habitats therefore likely constitute an essential element in many birds’
migration strategies but, unlike stopover habitats, they tend not to support high concentrations of
birds. As a consequence, the loss of stopover habitat along migration routes may have more severe
population consequences than the loss of overwinter habitat used for pre-migratory fuelling
(Baker et al. 2004, Newton 2004).
While moult rarely overlaps with migration, this is not true in all species and stopover or
pre-migratory fuelling sites may also be used to meet the energetic requirements of feather
replacement. Moult-migration, whereby birds undergo a complete moult along their migration
route, apparently “pausing” their migration to do so, occurs in the monsoon region of south-
western USA and north-western Mexico in several species that breed in western North America
(Leu and Thompson 2002, Rohwer et al. 2005). Energetic demands at moult-migration sites are
presumably higher than at typical stopover sites, yet the extent to which this phenomenon
occurs elsewhere in the Neotropics is unknown. Similarly, pre-alternate moults that give rise to
breeding plumages also likely occur at sites used for pre-migratory fuelling, but specific strate-
gies or needs of birds during this critical phase of the life cycle have rarely been studied. In at
least one long-distance migrant, the Bobolink Dolichonyx oryzivorus, areas used in South
America for pre-alternate moult are part of a sequence of stationary non-breeding sites used by
this species, blurring the concepts of over-wintering, stopover, or staging sites (Renfrew et al.
2013).
Current knowledge of stopover regions and migratory bottlenecks for

landbirds in the Neotropics
In this section we discuss our current knowledge of diurnal migrants and nocturnal migrants
separately, given the differing needs of these two groups; e.g. diurnal migrants both migrate and
feed by day and sleep at night, whereas nocturnal migrants migrate at night and feed and/or rest
by day. This may not hold true, however, when birds make long over-water flights that take more
than 12 hours to complete. Secondly, we discuss findings from geolocators and other tracking
technologies independently of on-the-ground field studies, in light of the qualitative differences
in the data they provide. Finally, although we discuss migrants from both eastern and western
North America, we concentrate on the preponderance of eastern breeding species whose migra-
tion routes take them through Central and northern South America and across barriers such as
the Gulf of Mexico and Caribbean Sea. In contrast, many species from western North America
overwinter in the northern extent of the Neotropics (Pacific slope of Mexico and northern Central
America; Kelly and Hutto 2005) and key stopover regions linked with survival and productivity
occur outside of our region of interest (LaManna et al. 2012, Drake et al. 2014). Nonetheless,
a handful of western species migrate to South America and likely also rely on Neotropical
stopovers.
The studies presented in this section were compiled following an ongoing and exhaustive lit-
erature search, employing the Google Scholar and Web of Science search engines. We entered the
following key words singly or in combination: fuelling, fuel load, fuel deposition rate, geolocator,
migration, migratory birds, migration strategy, stopover; in combination with geographical loca-
tions such as Neotropics, Central America, Costa Rica, Colombia, etc. In addition, the literature
cited in relevant publications was surveyed for appropriate references. In compiling and summa-
rising information, we specifically looked for papers in which evidence for stopovers such as
stopover duration or changes in fuel loads were presented. Finally, we used the general Google
search engine to discover technical reports published on the web.
Raptors and other diurnal migrants

Among the diurnally migrating species that use terrestrial habitats, perhaps the best-known
group is the raptors (in which we include falcons and New World Vultures). In addition to long-
term monitoring programmes at a variety of sites between Veracruz, Mexico (Ruelas Inzunza
et al. 2010a), and the Canal Zone in Panama (Porras-Peñaranda et al. 2004, Batista et al. 2005),
the use of satellite transmitters has revealed considerable detail with respect to routes and
stopovers (Fuller et al. 1998, Martell et al. 2001, Kochert et al. 2011).
More than 30 raptor species have been recorded migrating through Mesoamerica but the
majority of the 4 to 6 million raptors that migrate to or through the Neotropics on an annual basis
belong to just four species: Turkey Vulture Cathartes aura (2 million), Broad-winged Hawk Buteo
platypterus (2.1 million), Swainson’s Hawk Buteo swainsonii (1.2 million), and Mississippi Kite
Ictinia mississipiensis (0.2 million) (Bildstein 2004, Ruelas Inzunza et al. 2010b). Other less abun-
dant but visible species along the flyway include Osprey Pandion haliaetus, Peregrine Falcon Falco
peregrinus, and Swallow-tailed Kite Elanoides forficatus. Acting like a natural bottleneck, the nar-
rowing geography of Central America concentrates migrating raptors at several points. The most
well studied bottlenecks include the narrow gap between the Sierra Madre Oriental and the Gulf
of Mexico in Veracruz, Mexico (Ruelas Inzunza et al. 2009), the corridor between the Cordillera
de Talamanca and the Caribbean Sea in Costa Rica and western Panama (Porras-Peñaranda et al.
2004) and the Isthmus of Panama (Batista et al. 2005). Nevertheless, large numbers of migrating
raptors can be seen at several other points along the route including the Darién of Panama and
Colombia (Bayly et al. 2014), and further study will likely reveal a concentrated passage along the
Pacific coasts of Guatemala, El Salvador and Nicaragua (Bildstein 2004).
The routes taken by raptors between Mexico and South America are well-defined, with the
main flyway dividing in two in southern Mexico, after birds have passed through coastal Veracruz
(Bildstein 2004). The majority of birds cross to the Pacific coast at this point, following it down
through Guatemala, El Salvador and Nicaragua until crossing to the Caribbean lowlands in south-
ern Nicaragua and northern Costa Rica (Porras-Peñaranda et al. 2004, Kochert et al. 2011); how-
ever, Broad-winged Hawk, for example, may follow the Caribbean coast throughout. On entering
Panama, many individuals cross again to the Pacific slope, although many others continue down
the Caribbean coast, entering South America via the Darién. In South America routes are not well
studied (Colorado et al. 2006), but satellite tracks suggest that while Swainson’s Hawks continue
on an almost direct southerly route (Kochert et al. 2011), Turkey Vultures head east-south-east
towards the Llanos of Venezuela and Colombia. Many Osprey, Peregrine Falcon and Merlin Falco
columbarius do not join the main flight through Mesoamerica until its lower portion, having left
North America via the Florida peninsula and migrating over-sea via Cuba (Fuller et al. 1998,
Martell et al. 2001, Rodríguez-Santana et al. 2014). Western Cuba has also been shown to be a
major migratory route for USA-breeding Swallow-tailed Kites (Zimmerman and Meyer 2004).
Migrating raptors generally pass along the routes described above rapidly, rarely pausing to top
up fuel reserves (Smith et al. 1986, Fuller et al. 1998, Martell et al. 2001, Kochert et al. 2011);
however, a number of stopover regions have been identified. For example, while autumn migrat-
ing Swainson’s Hawks appear to accumulate much of their energy requirements at stopovers in
the southern USA, birds do make occasional stops in south-eastern Mexico, western Guatemala
and Nicaragua, north-western Colombia and in the Colombian Llanos (Kochert et al. 2011).
In contrast, the spring migration of Swainson’s Hawks does not appear to involve Neotropical
stopovers (Smith et al. 1986, Kochert et al. 2011). Swallow-tailed Kites make a prolonged stopover
on the Yucatan peninsula during autumn migration (Zimmerman and Meyer 2004), seemingly to
recover after an over-sea crossing from Florida or Cuba. The lack of information on other species
and the general notion that most raptors can complete their migration through the Mesoamerican
corridor without stopping to fuel (Smith et al. 1986), imply that their principal need may be safe
roosting sites. However, anecdotal evidence aside (Hicks et al. 1966, Bildstein and Saborio 2000,
Ruelas Inzunza et al. 2009), there is very little information regarding the use of roosting sites by
migrating raptors. Nonetheless, the highly concentrated flyway used by most raptors suggests
that they likely have as yet unidentified needs, which in part may be dictated by their diet which
impedes the en masse use of stopover sites.
Besides raptors, very little has been published on other diurnally migrating landbirds, although
these species are routinely encountered during raptor migration monitoring. Hirundines are a
highly abundant group of diurnal migrants and seasonal counts of Cliff Petrochelidon pyr-
rhonota, Barn Hirundo rustica and Bank Riparia riparia Swallows at bottlenecks in Veracruz and
the Colombian Darién involve hundreds of thousands of birds (Winkler 2006, Bayly et al. 2014).
Little is known, however, about stopover behaviour in these species. Given the declining status of
many aerial insectivores (Table 2) (Nebel et al. 2010, Rosenberg et al. 2016), determining stopover
and major roost sites for this group is a priority, especially if they adopt staging behaviours simi-
lar to those seen in Purple Martin Progne subis and Tree Swallow Tachycineta bicolor (Fraser
et al. 2013, Laughlin et al. 2013), where birds remain at sites longer than would appear necessary
for fuelling purposes. Many aerial insectivores migrate as far as the southern cone of South
America and likely have complex migration strategies and differential strategies between popula-
tions of a single species (Table 2). Other abundant yet rarely reported diurnal migrants are even
less well known, and aside from counts at concentration points in the Darién (Bayly et al. 2014),
little is known of migration or even overwintering grounds of species such as Chimney Swift
Chaetura pelagica, Black Swift Cypseloides niger, Common Nighthawk Chordeiles minor, Eastern
Kingbird Tyrannus tyrannus and Dickcissel Spiza americana.
Nocturnal migrants
The vast majority of Nearctic-Neotropical migratory landbirds (e.g. warblers, thrushes, tanagers,
cuckoos) migrate solely or primarily at night (DeGraaf and Rappole 1996, Alerstam 2009), and many
traverse vast stretches of the Caribbean basin or Central and northern South America to reach
wintering grounds in the Andes, the Amazon Basin, or farther south still. Despite the importance
of this extensive geography to annual survival, many Neotropical regions are still completely
unexplored in terms of where, when and why migratory landbirds stop (Faaborg et al. 2010b).
Table 2. Over-wintering destination and population status of diurnally migrating landbirds that winter in
South America. Season counts are based on the highest single autumn counts from a single site in the Darién
of Colombia (see Figure 4, region 3) reported in Bayly et al. (2014) or on counts from the same study site in
2014 and 2015. Three species (marked N/A) are rarely recorded at this watchsite presumably because their
migration route takes them over the Caribbean Sea or the Pacific Ocean.
Species Over-wintering Population Season high
Destination Status1 counts (Year)
2Common Nighthawk Chordeiles minor S. South America3 -58% 21,922 (2015)
Chimney Swift Chaetura pelagica W. Amazon Basin -67% 6,922 (2014)
Black Swift Cypseloides niger W. Amazon Basin -94% N/A
Eastern Kingbird Tyrannus tyrannus W. Amazon Basin -38% 243,669 (2014)
Purple Martin Progne subis S. American Lowlands4 -23% N/A
Bank Swallow Riparia riparia S. American Lowlands4 -89% 22,898 (2011)
Barn Swallow Hirundo rustica S. American Lowlands4 -38% 87,655 (2014)
Cliff Swallow Petrochelidon pyrrhonota S. South America3 +37% 158,413 (2014)
Bobolink Dolichonyx oryzivorus S. South America3 -60% N/A
Dickcissel Spiza americana N. South America -14% 30,268 (2015)
1Population loss from 1970-2014 based on Rosenberg et al. 2016.
2Nighthawks regularly initiate and terminate flights during daylight hours but may migrate primarily at
night.
3Includes regions primarily south and east of the Amazon Basin.
4Includes various lowland areas including Amazon Basin and regions to south and east.
In this section, we describe current knowledge of Neotropical stopover sites for nocturnal migrants
within six broad geographic regions (Fig. 3): (1) south-eastern Mexico and the Yucatan peninsula;
(2) northern Central America including Honduras, Nicaragua, and Guatemala; (3) Costa Rica and
Panama; (4) northern Colombia; (5) the Caribbean islands; and (6) western Mexico. Other regions in
the Neotropics are no doubt used for stopover but they remain largely unstudied.
South-eastern Mexico and the Yucatan Peninsula — Perhaps the best studied region, with several
studies from Veracruz (Winker 1995, Martínez Leyva et al. 2005, Shaw and Winker 2011), the
northern tip of the Yucatan (Deppe and Rotenberry 2005), north-eastern Belize (Bayly and
Gómez 2011, Gómez and Bayly 2011), and southern Belize (Johnson and Winker 2008). During
autumn migration, species richness (≈70 species) and abundance of migratory landbirds is high
and it is evident that the coastal geography of the region concentrates individuals of many species
that cross the Gulf of Mexico or migrate around its western edge (Winker 1995, Bayly and Gómez
2011). However, mist-netting has revealed that individuals of most species carry moderate fuel
reserves and therefore have little need to refuel at stopover sites (Winker 1995, Johnson and
Winker 2008, Bayly and Gómez 2011). Low recapture probabilities and short between-capture
durations also imply that most birds were making rest-roost stops between nocturnal flights
(Bayly and Gómez 2011).
Winker (1995) and Johnson and Winker (2008) provide evidence that birds may be able to top
up their reserves during these daytime stops but provide no evidence that birds stopover for
longer. Together these findings suggest that most migrants (19 out of 20 transient species in
north-east Belize, for example) passing through the region may accumulate sufficient energy
reserves north of the Gulf of Mexico to cross both the Gulf and the Yucatan Peninsula (Bayly and
Gómez 2011). Not all species or individuals follow this pattern and Willow Flycatchers Empidonax
traillii, for example, probably make a short refuelling stop before continuing their migration
(Gómez and Bayly 2011).
Spring migration sees a reduction in species richness in the region, and many transient species
on route from South America, appear to be carrying sufficient reserves on arrival to reach North
America without refuelling (Bayly and Gómez 2011). For a smaller number of Central American
wintering species, fuel loads, recapture rates and fuel deposition rates point to a stopover both on
Figure 3. The Neotropical region lying between the Tropic of Cancer and the Tropic of Capricorn.
Major regions discussed in the section on nocturnal migrants and referred to throughout the text
are outlined in shades of grey and named following the text.
the Yucatan Peninsula and in south-eastern Mexico, including species of concern such as Wood
Thrush Hylocichla mustelina and Kentucky Warbler Geothlypis formosa (Rogers and Odum
1966, Bayly and Gómez 2011, Shaw and Winker 2011).
Northern Central America — There are no published studies of autumn stopover site use in
northern Central America, despite the potential importance of this region for a large number of
species that cross the Gulf of Mexico (Cohen et al. 2017). A spring study focused on Cerulean
Warblers Setophaga cerulea in the mountains of northern Honduras, eastern Guatemala and
southern Belize, highlighted the importance of this area for this species of conservation concern
(Welton et al. 2012). Cerulean Warblers are rarely recorded anywhere along their migration
route between North American breeding grounds and Andean wintering grounds, but Welton
et al. (2012) documented the presence of birds during a two-week period in early April, largely on
Caribbean-facing mountains in Honduras and Guatemala. Several other warbler species, includ-
ing the ‘Near Threatened’ Golden-winged Warbler Vermivora chrysoptera, were also recorded in
this study. Given its geographic position and intriguing unpublished records submitted to the
online public database eBird (e.g. 400 Bay-breasted Warblers, Setophaga castanea, counted at a
single location in Nicaragua in April) (Sullivan et al. 2014), northern Central America will likely
prove to be a critical stopover region for many species.
Costa Rica and Panama — Stopover studies from Costa Rica and Panama are also surprisingly
rare, and aside from those dating back 50 years (Rogers and Odum 1966) and a recent study on
Swainson’s Thrush Catharus ustulatus (Wilson et al. 2008), we could find no reference to the
stopover behaviour of Nearctic-Neotropical migrants in this region. There is, however, a long-
term migration monitoring station at Tortuguero, Costa Rica (Elizondo-Camacho and Ralph
2012), and the data collected there would no doubt shed light on the use of the Caribbean lowlands
for stopovers, beyond the habitat selection inferences thus far reported (Wolfe et al. 2014).
Further, unpublished records in eBird and elsewhere suggest that a variety of species (e.g. Cerulean
Warbler) may depend on montane forest habitats in Costa Rica and Panama, especially in early
autumn. Studies of autumn migrants on the Caribbean coast of Panama found a mixture of
exhausted birds and individuals with sufficient reserves to continue their migration well into
South America (Rogers 1965, Rogers and Odum 1966). There was some evidence that fuel-
depleted birds stopped over in Panama to rebuild their energy reserves (Galindo et al. 1963). In a
spring study of Swainson’s Thrushes on the Pacific coast of Costa Rica, there was no evidence of
birds making prolonged stopovers (Wilson et al. 2008).
Northern Colombia — In contrast to the above regions, a growing body of evidence on stopover
site use in northern Colombia is emerging, with studies on Empidonax flycatchers in the Darién
(Colorado 2010) and on Catharus thrushes, vireos, and warblers in the Sierra Nevada de Santa
Marta (SNSM) and the Darién revealing new insights on the importance of this region to
migrants (Gómez et al. 2013, 2014). During autumn, Empidonax flycatchers stopping in the
Darién did not appear to build energy reserves (Colorado 2010), whereas three species of Catharus
thrushes exhibited a mixed strategy, with some individuals passing through rapidly and others
making multi-day stopovers to refuel (Gomez et al. 2014). In contrast, migrants passing through
the SNSM frequently made prolonged stopovers in both autumn and spring. For example, Veery
Catharus fuscescens on stopover in the SNSM stayed for an average of nine days in autumn, dur-
ing which time they increased their energy reserves by around 30% (Figure 2) and were subse-
quently capable of flights of up to 2,000 km towards their South American overwintering grounds
(Bayly et al. 2012a). A similar situation was found for Grey-cheeked Thrush Catharus minimus
in spring, with birds stopping in the SNSM for up to two weeks and storing sufficient energy to
reach North America (>2,500 km) without refuelling (Bayly et al. 2013). In a third species,
the Tennessee Warbler (Oreothlypis peregrina), long spring stopovers in the SNSM resulted in
smaller fuel loads compared to Grey-checked Thrushes, but they were still sufficient for flights to
northern Central America and, in some individuals, to the southern USA (Bayly et al. 2016).
These recent results from studies in northern Colombia suggest that migratory landbirds are
capable of adopting a migratory strategy similar to that of several shorebirds (Gómez et al. 2017);
i.e. using a small number of sites to store large fuel reserves for long-distance flights (Piersma
1987, Gill et al. 2009, Klaassen et al. 2011). The strategy of species like the Grey-cheeked Thrush
also has similarities with those adopted by migrating raptors, which pass over Mesoamerica with-
out the need for refuelling stops (Smith et al. 1986, Kochert et al. 2011). These strategies do not
appear to be restricted to the species studied in Colombia to date, as the results from geolocators
described below demonstrate. Another key lesson from studies in Colombia is that migratory
landbirds are far more abundant in certain stopover habitats (e.g. pre-montane forest) compared
to others and that different habitats support different fuelling opportunities and rates, which can
make the difference between individuals successfully crossing a water barrier or needing to take
a longer route or make additional stopovers (Gómez et al. 2013, 2015, Bayly et al. 2016).
Caribbean islands — Despite many long-term studies of overwintering migrants in the West
Indies, studies of stopover behaviour in the Caribbean are rare. General texts on Caribbean birds
indicate that most species migrating to South America are rarely encountered on Caribbean
islands and that most records are thought to be weather-related (Garrido and Kirkconnell 2000,
Raffaele et al. 2003, Latta et al. 2006); thus the optimal strategy seems to be to overfly the
Caribbean, rather than stopping to refuel on islands. This hypothesis is supported by a study of
Blackpoll Warbler Setophaga striata in the Dominican Republic, where birds primarily undertook
daytime rest-roost stops, often in association with unfavourable weather (Latta and Brown 1999).
Geolocators suggest a similar role for Caribbean islands (see below), but further study may reveal
stopover areas, particularly on large islands such as Cuba where information is scarce.
Western Mexico — Few studies of landbird migration stopover have occurred in western Mexico,
yet the diversity of habitats, ranging from coastal lowlands to high mountains, combined with the
large number of species migrating through or overwintering in Mexico, strongly suggests its
importance as a stopover region, especially for species breeding in the western USA and Canada
(Howell and Webb 1995). Recent studies of moult migration in north-western Mexico, in which
individuals of many species undergo an extended stopover to complete feather moult during and
after the summer monsoon season and then continue to overwintering areas (Leu and Thompson
2002, Rohwer et al. 2005, Pyle et al. 2009), is one key example showing the importance of western
Mexico to a critical life history stage.
Findings from geolocators and other tracking technologies

Geolocators have revolutionised our ability to track small landbirds during migration, revealing a
wealth of information that can be used to identify migratory routes and stopover regions.
Nonetheless, due to the relatively low precision of light-level geolocators (typically ± 100 km but
up to 400 km in the tropics) (McKinnon et al. 2013a, 2013b), it is only by combining findings
from geolocators with detailed on-the-ground field studies that habitat use and quality within
broad stopover regions can be assessed. Indeed, the array of habitats present within polygons or
confidence ellipses identified by geolocators can be extremely large. For example, in northern
Colombia, where geolocators have identified a general region used by Veery for autumn stopover
(Heckscher et al. 2011, Hobson and Kardynal 2015), it is possible to travel from snow-capped
peaks, through humid montane forest, to xeric scrub in less than 100 km. Only through detailed
mark-recapture studies within that region has the high value of foothill and pre-montane forest
been demonstrated for Veery and other Nearctic-Neotropical migrants (Bayly et al. 2012a, 2016).
Geolocators have revealed a number of unexpected patterns in the migratory strategies of small
landbirds, including faster than expected travel speeds and a general pattern of birds making
prolonged stopovers in a small number of regions (McKinnon et al. 2013a). For example, eastern-
breeding Veery were found to make multi-day stopovers in three main regions, the south-
eastern USA, northern Colombia, and the northern Amazon, when on route from a Delaware
breeding site to overwintering sites in the southern Amazon basin (Heckscher et al. 2011).
Remarkably, western-breeding Veery also converge on the same northern Colombia stopover
region in autumn (Hobson and Kardynal 2015), as results from an analysis of stable isotopes in
feathers had predicted (González-Prieto et al. 2011). Bobolink use fewer regions still, with many
birds making one long stopover in the Llanos (grasslands) of Venezuela and Colombia, while on
route between North America and their southern South American overwintering grounds
(Renfrew et al. 2013).
The pattern of long stopovers at a relatively small number of sites is emerging from landbird
studies in both the Americas and Europe (Stanley et al. 2012, Kristensen et al. 2013, Lemke et al.
2013, DeLuca et al. 2015). This strategy may be used primarily by nocturnally migrating species,
as aerial insectivores (Beason et al. 2012) and other diurnal migrant birds appear to behave dif-
ferently (Jahn et al. 2013): although Fraser et al. (2013) documented prolonged stopovers in
Purple Martin. Presumably, the ability of diurnal migrants to feed while actively migrating explains
part of this difference. Aside from revealing the widespread occurrence of long stopovers, geolo-
cators also show why landbirds were previously considered to use numerous sites/regions during
migration. Indeed, although birds move rapidly between these major stopover regions, often at
rates of 500 km/day (Callo et al. 2013, Fraser et al. 2013), flights are interrupted at a series of sites
along the route, which most likely act as diurnal or nocturnal rest-roost stops in order to avoid
migrating at unfavourable times of day (Stutchbury et al. 2009, Delmore et al. 2012).
There are still too few published geolocator studies to draw conclusions about the Neotropical
stopover regions used by the wider community of migratory landbirds, or by species with shared
wintering grounds, or even those from different breeding regions (e.g. eastern vs. western North
America). Nevertheless, several regions are now known to be important for two or more species
(Table 3). For example, Purple Martin (Fraser et al. 2013), Swainson’s Thrush (Delmore et al.
2012), and to a lesser extent Wood Thrush (Stutchbury et al. 2009) make an extended autumn
stopover on the Yucatan Peninsula, joining the list of species found to make short stops there by
banding studies (see above). Northern Colombia hosts extended stopovers by at least three spe-
cies, Veery (autumn), Red-eyed Vireo Vireo olivaceus (spring), and Swainson’s Thrush (autumn),
adding further evidence for this region’s importance to South American wintering species
(Heckscher et al. 2011, Delmore et al. 2012, Callo et al. 2013). Geolocators further indicate
that both Red-eyed Vireo (spring) and Swainson’s Thrush (autumn and spring) also make
long stopovers in northern Central America, confirming our prediction that this region
should figure prominently in migration strategies. Finally, it is worth emphasizing the criti-
cal importance of the Llanos of Venezuela and Colombia as an autumn and spring stopover
site for Bobolink (Renfrew et al. 2013).
GPS tags and coded nano-tags (radio transmitters) also hold promise for determining the pre-
cise location of new stopover sites. Nano-tags, in particular, are an excellent tool for studying
stopover behaviour and the outcome of subsequent migratory flights (Taylor et al. 2011, 2017),
such as the confirmation of non-stop flights >3,000 km by Grey-cheeked Thrush on leaving a
stopover site in Northern Colombia (Gómez et al. 2017).
Emerging picture of stopover regions in the Neotropics

The general pattern emerging from both on-the-ground field studies and the deployment of
geolocators is that long-distance migration strategies in many small landbirds are actually similar
to those of certain shorebirds (Atkinson et al. 2007, Gill et al. 2009, Lindström et al. 2016),
involving long flights and prolonged multi-day stopovers at a relatively small number of sites.
We are just beginning to understand the generality of this pattern, as well as which regions are of
greatest importance to the largest number of migratory birds.
N. J. Bayly et al.
Table 3. Evidence for major stopover regions and bottlenecks for migratory landbirds in the Neotropics. Numbers following region names relate to Fig. 3 & 4.
Region (see Fig. 3) Species Season Method Evidence for Reference
Yucatan/SE Mexico (8 & 9)
Veracruz Raptors, diurnal landbirds Autumn Observations Major bottleneck for several raptor species Ruelas I. et al. 2010a
Yucatan Peninsula Purple Martin Autumn Geolocators Stopover: mean stopover duration 16 days Fraser et al. 2013
SE Mexico Kentucky Warbler & others Spring Mist-netting Mass gains and recapture rate suggest Shaw & Winkler 2011
stopover in Kentucky & Hooded Warbler
NE Belize Magnolia & Yellow Warbler Spring Mist-netting Increased abundance, mass gains in recaptures Bayly & Gomez 2011
Northern Central America (6 & 7)
Nicaragua Red-eyed Vireo Spring Geolocators Stopover averaging 8 days Callo et al. 2013
Honduras/Guatemala Cerulean Warbler Spring Observations Stopover suspected in early-mid April Welton et al. 2012
Guatemala Swainson’s Thrush Spring Geolocators Stopover averaging 11 days Delmore et al. 2012
Panama & Costa Rica (5)
Costa Rica Raptors Both Observations Bottleneck on Caribbean coast Porras-Peñaranda et al.
2004
Panama Raptors Both Observations Bottleneck mainly on Pacific coast Bastista et al. 2005
Panama Swainson’s Thrush Spring Geolocators Stopover averaging 11 days Delmore et al. 2012
Northern Colombia (3 & 4)
Sierra Nevada de Santa Marta Veery Autumn Mist-netting & Stopover: body mass gains up to 40% of lean Bayly et al. 2012a;
(SNSM) Geolocators body mass Heckscher et al. 2012
Darién Catharus thrushes Autumn Mist-netting Short stopover in some individuals Gomez et al. 2014
Darién Diurnal migrants Autumn Observations Bottleneck for raptors, swallows, Chimney Bayly et al. 2014
Swift, Dickcissel, Common Nighthawk
SNSM Grey-cheeked Thrush & Spring Mist-netting Stopover: body mass gains of 40% and 30% Bayly et al. 2013;
Tennessee Warbler of lean body mass respectively. Bayly et al. 2016
NW Colombia Red-eyed Vireo Spring Geolocators Stopover averaging 6 days Callo et al. 2013
Amazon & Llanos (1 & 2)
Northern edge of Amazon Red-eyed Vireo & Veery Spring Geolocators Prolonged stopover at north edge of Amazon Callo et al. 2013;
Heckscher et al. 2012
Llanos Colombia / Venezuela Bobolink Both Geolocators Staging during autumn and spring Renfrew et al. 2013
14
Along the major flyways used by species breeding in eastern North America, a number of
major stopover regions are emerging (Table 3; Figure. 4), including areas inland of the Gulf Coast
and northern Florida in North America and northern Colombia in South America, both of which
are used to store sufficient energy to make unbroken flights across the Gulf of Mexico and/or the
Caribbean Sea. In particular, the Sierra Nevada de Santa Marta, an isolated massif on Colombia’s
Caribbean coast, is of considerable strategic importance for a number of species (Bayly et al.
2012a, 2016, Gómez et al. 2015). Not all birds are capable of such long-distance flights and various
points in Central America, such as the Yucatan/south-east Mexico and northern Central America
host stopover regions for certain species but we lack sufficient information to highlight regions
used by multiple species. No doubt the deployment of geolocators and GPS tags on additional
species and the establishment of migration banding stations in little-studied regions such as east-
ern Nicaragua and Honduras will reveal other critical fuelling regions.
Neotropical stopover regions for western-breeding migrants are almost completely unknown,
but sites within south-western Mexico and the Pacific slope of Central America are likely to prove
important for species on passage to South America, as they have for western populations of
Swainson’s Thrush (Ruegg and Smith 2002, Delmore et al. 2012), and may mirror findings in
north-west Mexico where moult-migration is integral to migration strategies (Rohwer et al. 2005).
Figure 4. Major Neotropical stopover regions (dashed circles) and bottlenecks (closed circles) for
migratory landbirds identified to date. 1. Northern Amazon (Veery, Red-eyed Vireo); 2. Llanos of
Venezuela and Colombia (Bobolink); 3. NW Colombia (bottleneck; also stopovers by Red-eyed
Vireo & Catharus thrushes); 4. Sierra Nevada de Santa Marta (Tennessee Warbler, Grey-checked
Thrush, Veery); 5. Costa Rica Caribbean slope & Panama Canal Zone (bottleneck for raptors);
6. Western Nicaragua (Red-eyed Vireo); 7. Highlands Honduras, Guatemala and Belize (Cerulean
Warbler); 8. Yucatan Peninsula (Purple Martin, Red-eyed Vireo, Magnolia Warbler); 9. Veracruz
(bottleneck for raptors, Kentucky and Hooded Warblers, Scissor-tailed Flycatcher); 10. SW Mexico
(Swainson’s Thrush). See Table 3 for references.
Indeed, while this region is known to hold a diverse assemblage of over-wintering migrants
(Howell and Webb 1995, Hutto 1997, Fagan and Komar 2016), it may also host stopover populations
of long-distance migrant species such as Yellow-billed Cuckoo Coccyzus americanus, Western
Wood-Pewee Contopus sordidulus and Olive-sided Flycatcher Contopus cooperi.
Threats facing landbirds on migration in the Neotropics

Just as we learn about the importance of key regions to long-distance migratory birds within the
Neotropics, new concerns are being raised about threats to specific sites and vital habitats within
those regions, as well as the predicted effects of climate change. Habitat loss continues to be the
leading threat to migratory bird populations throughout their annual cycle (Faaborg et al. 2010a,
Rosenberg et al. 2016), and in areas where birds are highly concentrated during migration stopover,
loss or degradation of critical habitats may contribute disproportionately to overall population
declines (Baker et al. 2004, Newton 2006). In addition, direct mortality of migrating birds due to
collisions with buildings, communication towers and other structures could be magnified in areas
where large proportions of species’ populations concentrate. Although astounding numbers of
birds are thought to perish through collisions in North America alone (Longcore et al. 2013),
including significant percentages of global populations of species of concern such as Golden-
winged Warbler (Arnold and Zink 2011), equivalent data from south of the USA are few and far
between (Agudelo-Álvarez et al. 2010). Nonetheless, urban and resort development along coast-
lines, and proliferation of wind-energy and communications infrastructure along coasts, narrow
landmasses, and mountain passes could pose major challenges for migratory bird conservation
(Cohen et al. 2017).
Research to date indicates that the majority of landbirds stopping over in Central and north-
ern South America rely on native forests, especially pre-montane forests on Caribbean-facing
slopes of coastal mountains, such as in the Sierra Nevada de Santa Marta in Colombia, as well
as lowland tropical wet and dry forests along the coasts (Johnson and Winker 2008, Welton et al.
2012, Gómez et al. 2015). As elsewhere in the Neotropics, these tropical forests are under severe
threat from expanding agriculture and development, and several of the stopover regions identi-
fied to date are experiencing high rates of deforestation and are considered hotspots for future
deforestation (Wassenaar et al. 2007). In addition, bird-friendly practices which have the
potential to mitigate forest loss, such as shade-grown coffee, may not be as beneficial to birds
on stopover (Bayly et al. 2016) as they are considered to be for over-wintering migrants
(Komar 2006).
Much of south-eastern Mexico, such as in the state of Veracruz, has been deforested and
although this trend has been reversed in recent years (Ruelas Inzunza et al. 2009), the extremely
low percentage of remaining forest cover likely remains a barrier to a successful migration for many
species. The expansion of cattle pastures in north-western and eastern Honduras and eastern
Nicaragua also represents a major threat to forests that are expected to constitute important
stopover habitats. In particular, the montane forests of north-western Honduras that are used
by the threatened Cerulean Warbler during spring migration (Welton et al. 2012) are under
considerable pressure. Likewise, in Colombia, the Darién bottleneck is a hotspot for deforesta-
tion to make way for cattle pastures. The Sierra Nevada de Santa Marta is also under pressure,
as both coffee cultivation and cattle pastures move up slope (Bayly et al. 2012b). In western
Mexico, loss of dry tropical forest (Trejo and Dirzo 2000) has likely diminished habitat, where
many species have been shown to require primary forest (Hutto 1989).
While populations of most raptor species that migrate through the Neotropics are stable or
increasing (Ruelas Inzunza et al. 2010a), dramatic changes along the main flyway could reverse
current fortunes, particularly for species concentrating in large proportion in narrow corridors.
The development of wind farms, in particular, has been cited as a potential hazard for migrating
raptors but mortality rates reported elsewhere do not support this claim (De Lucas et al. 2008,
Marques et al. 2014). Given our excellent knowledge of the routes taken by raptors, however, any
potentially detrimental projects can be identified rapidly and measures to mitigate their impact
can be proposed based on sound scientific justification.
The next steps: research and conservation needs

The last five years have seen an upsurge in new information on the migratory strategies
of a variety of landbirds, largely due to the innovation of geolocator technology (McKinnon
et al. 2013a), but also as a result of on-the-ground field studies and monitoring programmes in
northern Central America (Martínez Leyva et al. 2005), Colombia (Bayly et al. 2013) and Costa
Rica (Elizondo-Camacho and Ralph 2012). Despite our growing understanding of migratory strate-
gies and stopover site use, we are still only scratching the surface of a topic that will require dec-
ades of dedicated research to determine the needs of most species. To increase our understanding
of migration strategies, and to identify the most important stopover sites and habitats for declin-
ing Nearctic-Neotropical migratory birds, we recommend a coordinated research initiative com-
bining targeted field studies with the latest technologies across key regions of the Neotropics:
including regions 1, 2, 3, 6, 7, and 10 in Figure 3, premontane forests (600–1,500 m) in Panama
and Costa Rica, Pacific dry forests in Costa Rica and Nicaragua, lowland dry forests in northern
Colombia, and a general exploration of regions south of the equator. The goals of this initiative
would be to (1) identify new and previously unknown stopover regions for migratory landbirds;
(2) evaluate sites and habitats within those regions that are most critical for successful migration;
and (3) develop strategies to conserve these sites and habitats.
The deployment of geolocators or GPS tags on a wider range of landbird species, especially
those of conservation concern, is a clear research priority in order to identify new stopover regions,
and studies are already underway on Cerulean, Golden-winged, Canada Cardellina canadensis,
Kirtland’s Setophaga kirtlandii and Prothonotary Protonotaria citrea Warblers. It is also neces-
sary to build on successful geolocator studies by ensuring representative sampling across breed-
ing/wintering ranges, thereby shedding light on how stopover use varies between populations
(Jahn et al. 2013, Fraser et al. 2013, Hobson and Kardynal 2015) or the degree to which popula-
tions converge on major stopover regions, as occurs in Purple Martin (Fraser et al. 2013), Veery
(González-Prieto et al. 2011, Heckscher et al. 2011, Hobson and Kardynal 2015) and Bobolink
(Renfrew et al. 2013).
While geolocators enable us to rapidly identify broad stopover regions, their lack of precision
and inability to tell us about the habitats and resources that birds are using, means that more
precise technology (e.g., GPS tags) and on-the-ground field studies in the Neotropics are essential.
Such studies will not only help to pinpoint the actual location of birds within the broad regions
identified by geolocators (McKinnon et al. 2013b) but also determine what habitats are used and
how their quality varies for birds on stopover (Bayly et al. 2016). We therefore recommend that
geolocator studies be followed up or complemented by standardised surveys within these broad
regions, in combination with multi-year mark-recapture studies, that can: (1) estimate occupancy
and persistence of landbirds by region and habitat during migration; (2) determine the energetic
contribution of a given stopover region to migration strategies; (3) provide site-specific habitat
and resource associations for defining realistic conservation priorities; and (4) provide estimates
of use at the population level for communities of migrants.
The recapture of individually marked birds, for example, is essential to estimate stopover
durations (Schaub et al. 2001), rates of energy accumulation (Schaub and Jenni 2000), and ulti-
mately the contribution that a given stopover site makes to the overall energetic needs of migra-
tion (Bayly et al. 2012a). By comparing rates of mass gain (a surrogate for energy deposition)
between habitats, we can also examine habitat quality (Dunn 2001, Bayly et al. 2016). Mark-
recapture studies can also make use of innovative techniques to gather data on difficult-to-catch
and canopy-dwelling species, including the use of coded nano-tags and tower arrays to determine
stopover durations and subsequent movements of tagged individuals (Gómez et al. 2017, Taylor
et al. 2017).
Once we know where to focus conservation resources, the scale of action required to ensure
adequate habitat availability in major stopover regions throughout the Neotropics is potentially
overwhelming. To be successful, strategies to conserve stopover habitats need to be integrated
within regional conservation efforts, including those focused on Neotropical resident and threat-
ened species, protection of watersheds, reducing deforestation and socio-economic development
(Barlow et al. 2016). Any approach must go above and beyond the strategic purchase of private
reserves, given the considerable areas occupied by migratory birds, and include activities such as
lobbying for changes in detrimental agricultural and land-use policies, community-based conser-
vation efforts, ecotourism activities, promotion of migrant friendly agricultural practices such as
shade-grown coffee and silvopastures, and the use of tools such as Payment for Environmental
Services. Indeed, as we identify new stopover sites, the actions required to ensure that they con-
tinue to function as key links in the lives of migratory birds will have to be tailored to the unique
conditions in each.
As bird conservation plans have focused largely on breeding habitat, and to a lesser extent on
overwintering sites, new information on the specific needs of birds on migration must be incor-
porated into full life-cycle bird conservation plans (Berlanga et al. 2010, Rosenberg et al. 2016).
Only through large-scale and highly collaborative efforts, such as the GoMAMN (https://www.
gomamn.org/, Cohen et al. 2017) or the Neotropical Flyways Project (http://selva.org.co/
research-programs/migratory-species/nfp/) will we be able to understand and conserve the spec-
tacle of migration throughout the Western Hemisphere.
While our suggestions for continued and future research will yield important advances that can
help guide conservation, some habitats and regions are so threatened that we can’t afford to wait
for the next generation of data. Thus, we want to emphasise the importance of continuing ongo-
ing habitat conservation and restoration efforts throughout the region, especially in natural habi-
tats already in decline, as these efforts will undoubtedly benefit Neotropical migrant and resident
species alike.
Acknowledgements
This review grew out of the Migration Stopover Special Session at the Partners in Flight V
Conference and Conservation Workshop in Snowbird, Utah in August 2013, and we thank the
organisers of Partners in Flight V, especially American Bird Conservancy, for including the ses-
sion within the programme. Having begun as an overview of our knowledge prior to the special
session, the discussions during the session highlighted the need for this review. We thank all
participants in the session for their contributions and insights and subsequent discussions, espe-
cially, Rosa M. Vidal, Elisa Peresbarbosa, C. J. Ralph, Melinda Welton and Pablo Elizondo. This is
contribution No. 1 of the Neotropical Flyways Project and Hawk Mountain conservation science
contribution number 285. We thank an anonymous reviewer for suggestions that improved this
manuscript. Attendance at PIFV by NB and KVR was supported by the Cornell Lab of Ornithology,
and travel support for other conference participants was provided by American Bird Conservancy.
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NICHOLAS J. BAYLY*
SELVA: Investigación para la Conservación en el Neotropico, Bogotá D.C., Colombia.
CAMILA GÓMEZ
SELVA: Investigación para la Conservación en el Neotropico, Bogotá D.C., Colombia, and
Laboratorio de Biología Evolutiva de Vertebrados, Universidad de los Andes, Bogotá, Colombia.
KENNETH V. ROSENBERG
Cornell Lab of Ornithology, Ithaca, New York, USA, and American Bird Conservancy,
The Plains, Virginia, USA.
WENDY E. EASTON
Environment and Climate Change Canada, Canadian Wildlife Service, Delta, Canada.
JAY CARLISLE
Intermountain Bird Observatory, Boise State University, Boise, Idaho, USA.
DAVID N. EWERT
The Nature Conservancy, Lansing, Michigan, USA.
ANNA DRAKE
Department of Forest and Conservation Sciences, University of British Columbia, Vancouver, Canada.
LAURIE GOODRICH
Hawk Mountain Sanctuary Association, Orwigsburg, Pennsylvania.
*Author for correspondence; e-mail: [email protected]
Received 25 January 2017; revision accepted 17 July 2017;

Published online 30 October 2017

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Bird Conservation International (2018) 28:1–26.

© BirdLife International, 2017

Major stopover regions and migratory

The needs of migrating landbirds in the Neotropics

Current knowledge of stopover regions and migratory bottlenecks for

Raptors and other diurnal migrants

Findings from geolocators and other tracking technologies

Emerging picture of stopover regions in the Neotropics

Threats facing landbirds on migration in the Neotropics

The next steps: research and conservation needs

Gallinago media: intriguing variations on Moore, F. R. and Buler, J. J. (2011) Migrant-

*Author for correspondence; e-mail: [email protected]

Received 25 January 2017; revision accepted 17 July 2017;

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