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Long-distance bird migration within South America revealed

by light-level geolocators

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Journal:

Manuscript ID:

Manuscript Type:

Complete List of Authors:

AUK-12-077
Research Article
16-Apr-2012

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Date Submitted by the Author:

The Auk

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Jahn, Alex; University of Florida, Biology; Universidad de Buenos Aires,

Ecologia, Genetica y Evolucion
Levey, Douglas; University of Florida, Biology
Cueto, Victor; Universidad de Buenos Aires, Ecologia, Genetica y Evolucion
Pinto Ledezma, Jesus; Museo de Historia Natural Noel Kempff Mercado,
Tuero, Diego; Universidad de Buenos Aires, Ecologia, Genetica y Evolucion
Fox, James; British Antarctic Survey,
Masson, Diego; Universidad de Buenos Aires, Ecologia, Genetica y
Evolucion
Argentina, austral, Fork-tailed flycatcher, Neotropics, Tyrannus savana,
Tyrannidae

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Keywords:

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Fork-tailed Flycatcher Migration

LONG-DISTANCE BIRD MIGRATION WITHIN SOUTH AMERICA REVEALED

BY LIGHT-LEVEL GEOLOCATORS

Alex E. Jahn (1,2), Douglas J. Levey (1), Vctor R. Cueto (2), Jess Pinto Ledezma

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(3), Diego Tuero (2), James W. Fox (4), and Diego Masson (2)

1. Department of Biology, University of Florida, Gainesville, Florida, 326118525 USA

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2. Instituto de Ecologa, Gentica y Evolucin, Facultad de Ciencias Exactas y

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Naturales, Universidad de Buenos Aires, CONICET, Ciudad Universitaria,

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C1428EHA, Buenos Aires, Argentina

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3. Museo de Historia Natural Noel Kempff Mercado, Av. Irala 565, Santa Cruz
de la Sierra, Bolivia

4. British Antarctic Survey, Natural Environment Research Council, High Cross

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Madingley Road, Cambridge, CB3 0ET, United Kingdom

Corresponding author:
Alex Jahn
Instituto de Ecologa, Gentica y Evolucin
Facultad de Ciencias Exactas y Naturales
Universidad de Buenos Aires, Ciudad Universitaria
C1428EHA
Buenos Aires, Argentina
Email: alexjahn77@yahoo.com

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Abstract.- Little is known about the timing of migration, migration routes, and
migratory connectivity of most of the >230 species of birds that breed at south
temperate latitudes of South America, then migrate towards the tropics to overwinter.
We used light-level geolocators to track the migration of six Fork-tailed Flycatchers
(Tyrannus savana) captured on their breeding territories in Argentina. All individuals
(three males and three females) initiated fall migration from late January to late

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February, migrating 44-124 km day-1 in a northwesterly direction through central

South America to either one or two overwintering sites. Five flycatchers first spent
several weeks (primarily April and May) in western Amazonia (Peru, northwestern

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Brazil and southern Colombia), before moving northeast to spend the rest of the nonbreeding season in a region centered on south-central Venezuela. Duration of fall

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migration was three to nine weeks, over a distance of ~2700-3800 km. These results

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represent the first data on the overwintering locations, migration timing and routes of

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individual migrant passerine birds that breed in South America. Given the general lack
of information of such basic data as this for most migratory birds that breed in South

America, geolocator technology has the potential to revolutionize our understanding

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about how passerine birds migrate and the threats they face on South Americas
rapidly changing landscape.

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Key words: Argentina, austral, Fork-tailed Flycatcher, Neotropics, Tyrannus savana,

Tyrannidae.
Every year, billions of birds across the planet migrate between temperate breeding
grounds and tropical wintering grounds. These seasonal and often spectacular
movements have long been studied in the Northern Hemisphere, resulting in a large,

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detailed body of work on the migratory timing, duration, speed, and routes of many
species (reviewed by Newton 2008, Faaborg et al. 2010). Thus, scientists in the
Northern Hemisphere are in a position to evaluate effects of anthropogenic changes on
migratory birds. For example, Marra et al. (2005) demonstrated that birds on spring
migration in North America can adjust their rate of migration in response to changes in
temperature.

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In contrast, the basic natural history of bird migration elsewhere in the world is
poorly understood, largely due to lack of infrastructure, fewer ornithologists, and the
logistical challenges associated with tracking any organism across a vast scale (Jahn et

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al. 2009). Bird species that migrate entirely within the Neotropics represent a
significant proportion of all New World migrants. South America alone hosts >230

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species that migrate entirely within the continent. In terms of species richness, they

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represent the third-largest bird migration system on the planet (Chesser 1994, Jahn et

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al. 2004). Furthermore, bird migration in South America is increasingly viewed as

surprisingly complex and distinct from that in North America. Numerous types of bird

migration exist in South America, including migration between temperate breeding

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grounds and the tropics, migration wholly within tropical latitudes or within temperate
latitudes, altitudinal migration, partial migration, and longitudinal migration (i.e.,

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breeding at one longitude and wintering at another; Areta and Bodrati 2008, 2010;
reviewed by Chesser 1994, Joseph 1997, Faaborg et al. 2010).
Additionally, it is becoming increasingly apparent that theories about migration
developed for migrants that breed in the North America cannot be directly applied to
migrants that breed in much of South America. For example, although seasonality

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ultimately drives migration on both continents, the type of seasonality experienced by

birds is fundamentally different between North America, where seasons are largely
defined by variation in temperature, and South America, where they are largely
defined by variation in rainfall. Thus, the decision to migrate may be based on
different currencies in different hemispheres (Jahn et al. 2010). Even large-scale
patterns of bird migration in South America are generally different than those in North

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America. Migrants in South America generally travel shorter distances and exhibit a
greater incidence of seasonal range overlap than their North American counterparts
(Chesser 1994, Stotz et al. 1996, Dingle 2008).

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The recent and rapidly growing technological revolution in the fields of animal
tracking (reviewed by Robinson et al. 2010, Bowlin et al. 2010, Bridge et al. 2011)

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represents an unprecedented opportunity to understand bird migration at a much larger

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scale than previously possible. We describe the migration and winter distribution of

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individual Fork-tailed Flycatchers (Tyrannus s. savana), one of the most common

migratory birds in South America. The nominate subspecies breeds primarily at south

temperate latitudes (Argentina, Paraguay, Uruguay and southern Brazil) and

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overwinters in northern South America (Ridgely and Tudor 1994, Chesser 1995).
Their migration between south temperate breeding grounds and tropical wintering

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grounds has been observed at the population level for decades (Zimmer 1938, Antas
1987, Capllonch et al. 2009). However, no information has been available on the
migratory behavior of individuals. Our goal was to quantify the timing and speed of
migration, and the location of migratory routes and overwintering sites of individual
Fork-tailed Flycatchers.

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STUDY SITE AND METHODS

We attached light-level archival geolocators (Stutchbury et al. 2009, Robinson et al.
2010) to 43 nesting Fork-tailed Flycatchers at Reserva Privada El Destino, (35.13S,
57.40W), Buenos Aires Province, Argentina, during the breeding seasons (November
- December) of 2009 and 2010. The site is composed of temperate grasslands and
marshes, intersected by woodland tracts generally dominated by Celtis ehrenbergiana

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and Scutia buxifolia. The climate consists of hot summers and mild winters, with a
mean annual precipitation of 885 mm (n=10 years). The rainiest months are January
and February (mid to late summer).

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Flycatchers were captured by placing a predator model (e.g., Chimango

Caracara, Milvago chimango) along with calls of this predator from a portable speaker,

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within two meters of one or two 3X12 m (38 mm mesh) mist nets, at a distance of 2-4

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m from an active nest (except during the incubation stage, to avoid abandonment of

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the nest by the adult), or by placing a Fork-tailed Flycatcher model along with a
conspecific call within two meters of a mist net on a defended territory. Captured birds

were banded with a numbered metal band and a unique combination of up to three

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Darvic color bands, and sexed as described in Pyle (1997). Eight individuals were
fitted with an Mk10S light-level geolocator (1.2 g, Fox 2010) during the 2009

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breeding season, and 35 were fitted with an Mk12S light-level geolocator (0.9 g)
during the 2010 breeding season. Geolocators were attached to birds using a leg loop
backpack harness (Rappole and Tipton 1991) made of Filament Kevlar (500 Tex.,
Saunders Thread Co., Gastonia, North Carolina). All birds flew well upon release. We
recovered nine geolocators in subsequent breeding seasons and were able to recover

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usable data from six. None of the recaptured individuals showed any sign of injury
from the geolocator or harness.
Geolocators were programmed to record light intensity every minute but to
permanently store only the maximum reading every 10 minutes. We estimated
geolocator error by analyzing light data from the first seven days after deployment, a
time period during which flycatchers were still present at the study site, typically with

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nestlings or fledglings. After adjusting for clock drift, we used TransEdit (British
Antarctic Survey [BAS], Cambridge, United Kingdom) to estimate the sun elevation
that corresponded to a light threshold level of 16 (Fox 2010). This calibration resulted

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in a mean error distance of 116.5 (77.1 SD) km for the data from the six birds with
recovered geolocators. Sunrise and sunset transitions with obvious shading events

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were excluded from the analysis, as well as those from 15 days before and after the

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austral fall and spring equinoxes, due to very high uncertainty in latitude values (Fox

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2010).

Latitude/longitude coordinates of location fixes were calculated with

BirdTracker software (BAS) using midnight locations, because we assumed the birds

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were most likely stationary at night (to the best of our knowledge Fork-tailed
Flycatchers migrate primarily during the day; A.E. Jahn, pers. obs.). Thus, midnight

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locations should provide greater accuracy than midday locations, given that the bird
was at the same location overnight (Fox 2010). When constructing maps of daily
locations, we did not include five coordinates that were clearly anomalous: one far
from the regular species range (in southern Chile), one over the Pacific Ocean, one
over the Atlantic Ocean at the same longitude as the study site, and two that would

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have necessitated movement of >1500 km in a single day.

Because light-level geolocation can estimate longitude more accurately than
latitude (since shading events can affect estimates of latitude; Bchler et al. 2010,
Heckscher et al. 2011, Bairlein et al. 2012, Fudickar et al. 2012), we defined the
initiation of fall migration by a change in 1 longitude relative to the initial capture
site (i.e., the breeding territory). We defined the end of fall migration (and beginning

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of the overwintering period) as the first day the bird moved 3 longitude over a
period of at least 10 days, and was at that point located 3 longitude from the
longitude(s) it was at during migration. We used this conservative method to define

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the end of fall migration because most birds continued to move throughout winter (see
below), resulting in an indistinct separation between the fall migration route and the
winter range.

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We calculated distance of fall migration as the straight-line distance between

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the breeding site in Argentina and the first fix obtained in winter, and we calculated
the speed of fall migration by dividing the fall migration distance by duration. We

estimated the location of the fall migration route using the mean of daily fixes during

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fall migration, based on a minimum of six fixes per bird, with Hawth's Analysis Tools
for ArcGIS 9.3 (Beyer 2004). We were unable to analyze fall migration of two birds

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(20811 and 12B011) because we obtained no more than three fixes during fall
migration for each, or for another (20796) because of no fixes far (i.e., >900 km) from
the study site. We were unable to document the timing and route of spring migration
due to broad overlap with the austral spring equinox.
We created a spatially explicit model of the winter distribution of each bird

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using kernel density estimator analysis (Silverman 1986, Terrell and Scott 1992) in the
Spatial Analysis tool of ArcMap, v 9.3 (ESRI, Redlands, California). We set the
search radius at 200 km and grid cell size at 2 km (Bchler et al. 2010, Ryder et al.
2011) and required at least 26 coordinates to estimate the winter distribution of each
bird. For each individual, we present kernel densities encompassing >50%, 50-70%,
and 70-95% of the maximum density. We used the austral spring equinox as the cutoff

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date for the analysis of winter range.

RESULTS AND DISCUSSION
Fork-tailed Flycatchers initiated fall migration from late January to late February and

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migrated ~2700-3800 km during a period of up to twelve weeks (Table 1). No major

differences in migration patterns were evident between the sexes (Table 1). The typical

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route of migration was northwest through Uruguay and northeastern Argentina, then

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across central South America (Paraguay and Bolivia) to northern South America (Fig.

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1). This corroborates observational and museum collection records indicating that the
species is a passage migrant in Bolivia from February to April (Davis 1993, Chesser

1997). The fall migration route taken by the birds in this study skirts central Brazil,

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corroborating the observation that Fork-tailed Flycatchers are rare in that region during
fall migration (Antas 1987, Alves 2007). All three individuals appeared to reverse

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direction and fly south for 1-4 days at a time during fall migration. We are uncertain if
these movements are actual or due to calculation error (e.g., associated with proximity
to the austral fall equinox).
After fall migration, all but one individual (12B060) occupied two sites in
succession. The first was in western Amazonia (i.e., Peru, southern Colombia and

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northwestern Brazil), where they remained from one to 11 weeks (primarily April and
May). In May, they flew approximately 1000 km northeast to south-central Venezuela
and far eastern Colombia, where they stayed 3-4 months until returning to Argentina
(Fig. 1). This pattern may be related to spatial and temporal variation in rainfall across
the Amazon Basin. Rainfall in northwestern Amazonia, where the birds first settled
after migration, peaks from April to June (Espinoza et al. 2009), which corresponds to

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when the birds were there (Fig. 1). Occupying such areas of high rainfall may be
important for securing adequate food after breeding. Fork-tailed Flycatchers consume
both fruit and insects outside of the breeding season (Fitzpatrick 2004, A.E. Jahn pers.

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obs.), and rainfall appears to be important in maintaining adequate food supplies for
migrating and overwintering insectivorous migratory birds in both the Neotropics and

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Paleotropics (Sinclair 1978, Hutto 1985, Sherry and Holmes 1996, Herremans 1998,
Brown and Sherry 2006).

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Another explanation for the use of two wintering areas could be related to postbreeding molt. In North America, a close relative of Fork-tailed Flycatchers, Western

Kingbirds (Tyrannus verticalis), exhibit a molt-migration in fall, in which adults

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migrate to the region of the Mexican monsoon (Arizona/northern Mexico), where they
molt primary flight feathers before proceeding further south to overwinter (Barry et al.

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2009). Presumably, they and other passerine migrants that exhibit this molt-migration
do so to take advantage of a temporal window of food abundance associated with the
Mexican monsoon (Rohwer et al. 2005). Fork-tailed Flycatchers also molt primarily
during the non-breeding season (Zimmer 1938, A.E. Jahn pers. obs.), though the exact
timing of the molt and how it varies across the winter range remains unknown.

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Some Nearctic-Neotropical migrants also appear to occupy sequential winter

sites in the Amazon Basin. Veeries (Catharus fuscescens) use two winter sites in
Amazonia, perhaps following flushes in food abundance associated with river flooding
across the Amazon Basin (Heckscher et al. 2011). Similarly, Northern Black Swifts
(Cypseloides niger borealis) overwinter in western Amazonia and have large winter
distributions, with a tendency to move westwards throughout winter (Beason et al.

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2012). Taken together, these studies, along with our own, suggest that large-scale (i.e.,
hundreds to thousands of kilometers) movements of overwintering passerines in the
tropics may be more common than previously thought.

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Fall migration speed ranged between 44 and 124 km day-1, with males
appearing to migrate faster than females; small sample sizes preclude a statistical

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comparison, however (Table 1). These speeds are similar to that of fall migration

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speeds of Wood Thrushes (Hylocichla mustelina), 27-106 km day-1 (Stutchbury et al.

2009), and Veeries (Catharus fuscescens), which average 65-85 km day-1 (Heckscher

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et al. 2011). They are generally slower than those of Purple Martins (Progne subis), at

>150 km day-1 (Stutchbury et al. 2009), Northern Black Swifts (Cypseloides niger

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borealis), which average 341 km day-1 (Beason et al. 2012), and Northern Wheatears
(Oenanthe oenanthe), at ~160 km day-1 (Bairlein et al. 2012).

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This study represents the first tracking of individual passerine migrants that
breed in South America, and demonstrates that individual migratory Fork-tailed
Flycatchers may occupy all three major watersheds on the continent (i.e., Ro de la
Plata, Amazon, and Orinoco River basins) and up to eight countries throughout the
year (Fig. 1). This study also demonstrates the feasibility of using light-level

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geolocators to better understand the annual cycle of migratory birds within South
America. Among short- to medium-distance migrants, geolocators could potentially be
most useful for the various bird species that migrate wholly at south temperate
latitudes (i.e., Patagonia), because geolocator data is most accurate at higher latitudes.
Additionally, given that location estimation based on longitude offers a relatively good
estimate of movement, even for short-distance migrants (Fudickar et al. 2012),

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geolocators could provide novel information about the annual cycle of short-distance
longitudinal migrants in South America (Areta and Bodrati 2008, 2010).
Threats to Fork-tailed Flycatchers resulting from anthropogenic changes remain

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unknown. However, the ecosystems they inhabit in the Ro de la Plata and Orinoco
basins are undergoing large-scale, rapid anthropogenic land-use change. In the

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Orinoco River Basin, conversion of natural grasslands (i.e., llanos) to agro-industrial

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landscapes and mining are major threats, although the impact of such activities on

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migratory birds is still poorly documented (Lasso et al. 2010). Major threats to the
grasslands (i.e., pampas) of the Ro de la Plata River Basin include conversion to

agriculture, agro-chemical application, and mineral extraction (Krapovickas and Di

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Giacomo 1998, A.E. Jahn, pers. obs.).

Geolocators can be used in South America to address many basic and important

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questions, including: (1) What are the potential carry-over effects between the nonbreeding season in northern South America and the breeding season at south temperate
latitudes? (2) What potential benefits do migratory birds gain from occupying two
sites after fall migration? For example, do they undergo a molt at the first site (as in
molt-migration of other species to the Mexican Monsoon region of North America;

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Rohwer et al. 2005), and/or benefit from higher food abundance there? And, (3) What
is the migratory route, timing, speed, and winter distribution of other migratory bird
populations in South America? Until such basic data are collected, the mechanisms
underlying the evolution and regulation of bird migration in South America cannot be
addressed (Cavalcanti 1990, Jahn et al. 2006, Alves 2007).

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ACKNOWLEDGMENTS

We dedicate this paper to our late friend and colleague Marvin Morales Jacinto. We
thank the Elsa Shaw de Pearson Foundation for providing logistical support, the staff
of Reserva Privada El Destino for their hospitality, and Omar Barroso, Natalia

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Bogado, Vanessa Sandoval, Quilln Vidz, and Emily Williams for assistance in the

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field. This research was funded through a Scientific Research Grant (#8444-08) from
the National Geographic Society, a National Science Foundation International

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Research Fellowship to AEJ (IRFP-0965213), Gatorade Fund of the University of

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Florida, and Optics for the Tropics. VRC acknowledges CONICET and Universidad

de Buenos Aires for supporting his research in Argentina. This research was conducted

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with the authorization of the Departamento de Flora y Fauna, Ministerio de Asuntos

Agrarios, Provincia de Buenos Aires, Argentina (Disposicin 256/11).

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49
50
51
52
53
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LITERATURE CITED

Alves, M. A. S. 2007. Sistemas de migraes de aves em ambientes terrestres no

Brasil: exemplos, lacunas e propostas para o avano do conhecimento. Revista
Brasileira de Ornitologia 15:231-238.
Antas, P. T. Z. 1987. Migrao de aves no Brasil, p. 153-187. Anais do II Encontro
Nacional de Anilhadores de Aves. Rio de Janeiro, Editora UFRJ.

12

The Auk is published for the American Ornithologists' Union by the University of California

Page 13 of 21

Areta, J. I., and A. Bodrati. 2008. Seasonal movements and phylogenetic affinity of
the Shear-tailed Gray-Tyrant (Muscipipra vetula). Ornitologia Neotropical 19:201211.
Areta, J.I., and A. Bodrati. 2010. A longitudinal migratory system within the Atlantic
Forest: seasonal movements and taxonomy of the Golden-rumped Euphonia
(Euphonia cyanocephala) in Misiones (Argentina) and Paraguay. Ornitologia

P
ePr

Neotropical 21:71-86.

Bchler, E., S. Hahn, M. Schaub, R. Arlettaz, L. Jenni, J. W. Fox, V. Afanasyev and F.

Liechti. 2010. Year-round tracking of small trans-saharan migrants using light-

ub

level geolocators. Plos One 5:e9566

Bairlein, F., D. R. Norris, R. Nagel, M. Bulte, C.C. Voigt, J. W. Fox, D. J. T. Hussell,

lic

and H. Schmaljohann. 2012. Cross-hemisphere migration of a 25 g songbird.

at

Biology Letters doi:10.1098/rsbl.2011.1223

io

Barry, J. H., L. K. Butler, S. Rohwer, and V. G. Rohwer. 2009. Documenting moltmigration in Western Kingbird (Tyrannus verticalis) using two measures of

collecting effort. Auk 126:260-267.

Co

Beason, J. P., C. Gunn, K. M. Potter, R. A. Sparks, and J. W. Fox. 2012. The Northern
Black Swift: Migration Path And Wintering Area Revealed. Wilson Journal of

py

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60

PDF for Review

Ornithology 124:1-8.

Beyer, H. L. 2004. Hawth's analysis tools for ArcGIS. Available at http://www.

spatialecology.com/htools, accessed 24 July 2011.
Bowlin M. S., I. -A. Bisson, J. Shamoun-Baranes, J. D. Reichard, N. Sapir, P. P.
Marra, T. H. Kunz, D. S. Wilcove, A. Hedenstrm, C. G. Guglielmo, S. kesson,

13

The Auk is published for the American Ornithologists' Union by the University of California

PDF for Review

M. Ramenofsky, M. Wikelski. 2010. Grand challenges in migration biology.

Integrative and Comparative Biology 50:261-279
Bridge, E. S., K. Thorup, M. S. Bowlin, P. B. Chilson, R. H. Diehl, Flron, R. W., P.
Hartl, R. Kays, J. F. Kelly, W. D. Robinson, and M. Wikelski. 2011. Technology
on the Move: Recent and Forthcoming Innovations for Tracking Migratory Birds.
BioScience 61:689-698.

P
ePr

Brown, D. R., and T. W. Sherry. 2006. Food supply controls the body condition of a
migrant bird wintering in the tropics. Oecologia 149:22-32.
Capllonch, P., D. Ortiz, and K. Soria. 2009. Migraciones de especies de Tyrannidae de

ub

la Argentina: Parte 2. Acta Zoolgica Lilloana 53:55-75.

Cavalcanti, R. B. 1990. Migraes de aves no cerrado Pages 110-116 in Anais do IV

lic

Encontro Nacional de Anilhadores de Aves, Brasil.

at

Chesser, R. T. 1994. Migration in South America: an overview of the austral system.

io

Bird Conservation International 4:91-107.

Chesser, R. T. 1995. Biogeographic, ecological, and evolutionary aspects of South

American austral migration, with special reference to the family Tyrannidae.

Co

Ph.D. Dissertation, Louisiana State University, Baton Rouge.

Chesser, R. T. 1997. Patterns of Seasonal and Geographical Distribution of Austral

py

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
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42
43
44
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46
47
48
49
50
51
52
53
54
55
56
57
58
59
60

Page 14 of 21

Migrant Flycatchers (Tyrannidae) in Bolivia. Ornithological Monographs 48:171204.

Davis, S. E. 1993. Seasonal status, relative abundance, and behavior of the birds of
Concepcin, departamento Santa Cruz, Bolivia. Fieldiana Zoology (new series)
71:1-33.

14

The Auk is published for the American Ornithologists' Union by the University of California

Page 15 of 21

Dingle, H. 2008. Bird migration in the southern hemisphere: a review comparing

continents. Emu 108:341-359.
Espinoza, J. C., J. Ronchail, J. L. Guyot, G. Cochonneau, F. Naziano, W. Lavado, E.
De Oliveira, R. Pombosa, and P. Vauchel. 2009. Spatio-temporal rainfall
variability in the Amazon basin countries (Brazil, Peru, Bolivia, Colombia, and
Ecuador). International Journal of Climatology 29:1574-1594.

P
ePr

Faaborg J., R. T., Holmes, A. D. Anders, K. L. Bildstein, K. M. Dugger, S. A.

Gauthreaux, Jr., P. Heglund, K. A. Hobson, A. E. Jahn, D. H. Johnson, S. C. Latta,
D. J. Levey, P. P. Marra, C. L. Merkord, E. Nol, S. I. Rothstein, T. W. Sherry, T.

ub

S. Sillett, F. R. Thompson, III, and N. Warnock. 2010. Recent Advances in

Understanding Migration Systems of New World Landbirds. Ecological
Monographs 80:3-48.

at

lic

Fitzpatrick, J. 2004. Family Tyrannidae (tyrant-flycatchers). Pages 170-462 in

io

Handbook of birds of the world Volume 9: Cotingas to pipits and wagtails (J. del
Hoyo J., A. Elliott, and D.A. Christie, Eds.). Lynx Editions, Barcelona.

Fudickar, A. M., M. Wikelski, and J. Partecke. 2012. Tracking migratory songbirds:

Co

accuracy of light-level loggers (geolocators) in forest habitats. Methods in Ecology

and Evolution 3:47-52.

py

1
2
3
4
5
6
7
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11
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13
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44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60

PDF for Review

Fox, J. W. 2010 Geolocator manual. Version 8. British Antarctic Survey, Cambridge,

United Kingdom.
Heckscher, C. M., S. M. Taylor, J. W. Fox, and V. Afanasyev. 2011. Veery (Catharus
fuscescens) Wintering locations, migratory connectivity, and a revision of its
winter range using geolocator technology. Auk 128:531-542.

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The Auk is published for the American Ornithologists' Union by the University of California

PDF for Review

Herremans, M. 1998. Strategies, punctuality of arrival and ranges of migrants in the

Kalahari basin, Botswana. Ibis 140:585-590.
Hutto, R. L. 1985. Seasonal Changes in the Habitat Distribution of Transient
Insectivorous Birds in Southeastern Arizona: Competition Mediated? Auk
102:120-132.
Jahn, A. E., D. J. Levey, J. E. Johnson, A. M. Mamani, and S. E. Davis. 2006.

P
ePr

Towards a mechanistic interpretation of bird migration in South America.

Hornero 21:99-108.

Jahn, A. E., D. J. Levey and K. G. Smith. 2004. Reflections across hemispheres: a

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system-wide approach to New World bird migration. Auk 121:1005-1013.

Jahn, A.E., S. Ouly, E. Chiang, D.J.Levey and J.A. Clavijo. 2009. Opportunities

lic

through partnerships for conservation and research of austral migrants. Pages 557-

at

564 in Proceedings of the IV International Partners in Flight Conference, Tundra to

io

the Tropics, McAllen, Texas.

Jahn, A. E., D. J. Levey, J. A. Hostetler, and A. M. Mamani. 2010. Determinants of

partial bird migration in the Amazon Basin. Journal of Animal Ecology 79:983-

Co

992.

Joseph, L. 1997. Towards a broader view of Neotropical migrants: consequences of a

py

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6
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8
9
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12
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15
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re-examination of austral migration. Ornitologia Neotropical 8:31-36.

Krapovickas, S., and A.S. Di Giacomo. 1998. Conservation of pampas and campos
grasslands in Argentina. Parks 8:47-53.
Lasso, C. A., J. S. Usma, F. Trujillo, and A. Rial. 2010. Biodiversidad de la cuenca del
Orinoco: bases cientficas para la identificacin de reas prioritarias para la

16

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Page 17 of 21

conservacin y uso sostenible de la biodiversidad. Instituto de Investigacin de

RecursosBiolgicos Alexander von Humboldt, WWF Colombia,
FundacinOmacha, Fundacin La Salle e Instituto de Estudios de la Orinoquia,
Universidad Nacional de Colombia. Bogot, D.C., Colombia.
Marra P. P., C. M. Francis, R. S. Mulvihill, F. R. Moore. 2005. The influence of
climate on the timing and rate of spring bird migration. Oecologia 142:307-315.

P
ePr

Newton, I. 2008. The migration ecology of birds. Elsevier Press, London.

Pyle, P. 1997. Identification guide to North American birds. Part I. Slate Creek

ub

Press, Bolinas, California.

Rappole, J. H., and A. R. Tipton. 1991. New harness design for attachment of radio

lic

transmitters to small passerines. Journal of Field Ornithology 62:335-337.

at

Ridgely, R. S., and G. Tudor. 1994. The birds of South America. Vol. II. The
Suboscine Passerines. University of Texas Press, Austin, Texas.

io

Robinson, W. D., M. S. Bowlin, I. -A. Bisson, J. Shamoun-Baranes, K. Thorup, R. H.

Diehl, T. H. Kunz, S. Mabey, and D. W. Winkler. 2010. Integrating concepts and

Co

technologies to advance the study of bird migration. Frontiers in Ecology and the
Environment 8:354-361.

py

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2
3
4
5
6
7
8
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49
50
51
52
53
54
55
56
57
58
59
60

PDF for Review

Rohwer, S., L. K. Butler, and D. R. Froehlich. 2005. Ecology and demography of

eastwest differences in molt scheduling of Neotropical migrant passerines. Pages
87-105 in Birds of Two Worlds: The Ecology and Evolution of Migratory Birds
(R. Greenberg and P. P. Marra, Eds.). Johns Hopkins University Press, Baltimore,
Maryland.
Ryder, T. B., J. W. Fox, and P. P. Marra. 2011. Estimating migratory connectivity of
17

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PDF for Review

Gray Catbirds (Dumatella carolinensis) using geolocator and mark-recapture data.

Auk 128:448-453.
Sherry, T. W., and R. T. Holmes. 1996. Winter Habitat Quality, Population Limitation,
and Conservation of Neotropical-Nearctic Migrant Birds. Ecology 77:36-48.
Silverman, B. W. 1986. Density estimation for statistics and data analysis. Chapman
and Hall, New York.

P
ePr

Sinclair, A. R. E. 1978. Factors affecting the food-supply and breeding season of

resident birds and movements of Palaearctic migrants in a tropical African
savanna. Ibis 120:480-497.

ub

Stotz, D. F., F. W. Fitzpatrick, T. A. Parker, III, and D. K. Moskovits. 1996.

Neotropical birds: Ecology and Conservation. University of Chicago Press,
Chicago.

at

lic

Stutchbury, B. J. M, S. A. Tarof, T. Done, E. Gow, P. M. Kramer, J. Tautin, J. W. Fox,

io

and V. Afanasyev. 2009. Tracking long-distance songbird migration by using

geolocators. Science 323:896.

Terrell G. R., and D. W. Scott. 1992. Variable kernel density estimation. The Annals

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of Statistics 20:1236-1265.

Zimmer, J. T. 1938. Notes on migrations of South American birds. Auk 55:405-410.

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Table 1. Initiation, duration, distance and speed of fall migration of three male and
three female Fork-tailed Flycatchers captured at Reserva Privada El Destino,
Argentina. Initiation of fall migration is the first date there was a change of 1
longitude relative to that of the breeding site. Distance is the straight-line distance
between the breeding site and the first fix obtained in winter. Speed represents the
distance of migration divided by duration of migration in days.

ID

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Duration (days) Distance (km) Speed (km day-1)

Sex

Initiation

8-Feb-11

12B043

5-Feb-11

20811

27-Feb-10

12B060

11-Feb-11

20796

24-Jan-10

84

12B011

13-Feb-11

65

12B028

22

2738

124

62

3566

58

41

3755

92

55

3791

69

3770

45

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2891

44

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Fig. 1. Winter range and associated error (>50%, 50-70%, and 70-95% kernel
densities), and fall migration routes of six Fork-tailed Flycatchers initially
captured at Reserva Privada El Destino, Argentina (red circle). The individual
identification number of each is at the top right of each map. Locations and
dates of first and second winter sites are indicated. Dates between these two
periods are those for which we have no data. Fall migration routes from three

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birds are missing due to lack of data during that period. Lines depicting the
migratory routes do not always represent the exact migration route because of
gaps in dates of location fixes due to the austral fall equinox and shading
events (see text).

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