Ecology and Evolution - 2023 - Teitelbaum - Post Migratory Nonbreeding Movements of Birds A Review and Case Study

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Received: 29 October 2022 Revised: 17 February 2023 Accepted: 23 February 2023
DOI: 10.1002/ece3.9893
REVIEW ARTICLE
Post-migratory nonbreeding movements of birds: A review and

case study
Claire S. Teitelbaum | Natalie C. Bachner | Richard J. Hall
Odum School of Ecology, Athens, Georgia,

USA Abstract
Seasonal migrations are fascinating and ecologically important, but many migratory
Correspondence
Claire S. Teitelbaum, Odum School of species are declining as climate change and land-use change alter the habitats used by
Ecology, 140 E Green St, Athens, GA
migrants across the annual cycle. While some migratory birds use a single wintering
30605, USA.
Email: claire.teitelbaum@gmail.com site, others undertake large-scale post-migratory movements during the nonbreed-
ing season. Technological advances that enable tracking individual birds are uncover-
Funding information
National Science Foundation, Grant/ ing more examples of post-migratory nonbreeding movements. Documenting these
Award Number: DEB-1518611 and
movements is important for conservation, which requires understanding when and
Graduate Research Fellowship
where migrants use habitats throughout their range. Here, we reviewed existing lit-
erature and collected information on the post-migratory nonbreeding movements of
92 migratory bird species from 18 orders across six continents. Among these records,
the most commonly reported drivers of movements were resource availability and
climate. This strong dependence of post-migratory nonbreeding movements on birds'
abiotic and biotic environments suggests that environmental change will impact the
patterns of these movements and potentially the fitness of species that undertake
them. We also reviewed post-migratory nonbreeding movements in North American-
breeding thrushes from the genus Catharus to examine the drivers of these move-
ments in five closely related migratory species. We find that species that are less
territorial are more likely to use multiple sites during the nonbreeding season; how-
ever, there is little evidence for dietary, evolutionary, or environmental differences
between thrush species that move during winter and those that are stationary. While
we believe our study represents the most comprehensive list of species exhibiting
post-migratory nonbreeding movements to date, biases in sampling, a lack of common
terminology for these movements, and the still-nascent availability of inexpensive,
lightweight tracking devices mean that there are probably more populations that un-
dertake such movements. Future research into the consequences of post-migratory
nonbreeding movements for individual fitness and ecosystem services would advance
our understanding of their conservation importance and their evolution.
KEYWORDS
avian, intra-tropical migration, migration, movement, nomadism, overwintering, telemetry
This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium,
provided the original work is properly cited.
© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
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2 of 14 TEITELBAUM et al.
TA X O N O M Y C L A S S I F I C AT I O N
Behavioural ecology, Global ecology, Macroecology, Movement ecology
1 | I NTRO D U C TI O N drivers, its potential ecological consequences, and evidence for how
post-migratory nonbreeding movements might respond to global
Animal migration is crucial for survival and reproduction in many environmental change. The results from this literature review rep-
species (Alerstam & Högstedt, 1982) and many migratory animal resent a combination of biological patterns and sampling bias; our
populations are declining (Wilcove & Wikelski, 2008). Migration be- systematic literature search identified only “positive” results (i.e.,
tween breeding and nonbreeding areas is the riskiest time of the those where post-migratory nonbreeding movements have been
annual cycle for some migratory species (Rockwell et al., 2017; Sillett documented), so species and populations not captured by the data
& Holmes, 2002). As a result, a large body of work has sought to set could be true absences (i.e., do not perform post-migratory
understand the patterns, drivers, and consequences of migratory nonbreeding movements) or false absences (i.e., not studied or not
movements, including the evolution of partial migration (Chapman captured by our systematic search). Therefore, we also use a single,
et al., 2011), the ecology of stopover sites (Lindström, 1995; well-studied taxonomic group (North American thrushes from the
Schmaljohann & Eikenaar, 2017), and how migratory movements genus Catharus) as a case study to form a more detailed picture of
are shifting under environmental change (Pulido, 2007; Visser why some species or populations might undertake post-migratory
et al., 2009). During the post-migratory nonbreeding period of movements while others do not. Finally, we synthesize this com-
the annual cycle (i.e., the portion of the nonbreeding period that bined literature to highlight important gaps in our current knowledge
excludes migration; commonly called overwintering), some well- and promising avenues for future research.
studied systems indicate that birds exhibit high site fidelity within
and across years (e.g., Blackburn & Cresswell, 2016; Cuadrado
et al., 1995) and relatively high survival rates (Rockwell et al., 2017). 1.1 | Defining post-
This portion of the nonbreeding period is therefore often considered migratory nonbreeding movements
to be relatively safe and predictable compared with migration itself.
Some migratory animals use two or more distinct sites during the We propose a working definition of post-migratory nonbreeding
nonbreeding season following their seasonal migration, a behavior movements that applies across migratory bird taxa. First, we de-
referred to as within-winter movement, winter itinerancy, seasonal fine seasonal migration based on the characterization by Dingle and
nomadism, intratropical migration, or secondary winter movement Drake (2007):
(Moore, 1976; Stutchbury et al., 2016; Teitelbaum & Mueller, 2019).
We hereafter refer to these movements as “post-migratory non- (1) a type of locomotory activity that is notably per-
breeding movements,” which places them in the context of migration sistent, undistracted, and straightened out; (2) a relo-
and explicitly includes migrants from ecosystems that experience cation of the animal that is on a much greater scale,
wet/dry seasonality; we provide a detailed definition of these move- and involves movement of much longer duration, than
ments, and how they differ from other nonbreeding season move- those arising in its normal daily activities; (3) a sea-
ments, in the section below. Post-migratory nonbreeding movements sonal to-and-fro movement of populations between
have been well described in various groups, including African- and regions where conditions are alternately favorable or
neotropical-wintering songbirds and temperate-wintering waterfowl un- favorable (including one region in which breeding
in North America and Eurasia (Faaborg et al., 2010; Jones, 1995). occurs); and (4) movements leading to redistribution
However, no study to date has synthesized the characteristics and within a spatially extended population.
causes of these movements globally and across species. Such syn-
thesis is especially needed given the large-scale declines of bird Based on the first three of these concepts (the fourth of which
populations (Rosenberg et al., 2019) and the potential importance of is less strongly based on the avian migration literature, Dingle &
overwintering habitats for survival and reproductive success (Norris Drake, 2007), we define migration as an annual, persistent, round-
et al., 2004; Rockwell et al., 2017). trip, and long-distance movement between distinct locations, usually
This review seeks to explore the distributions and drivers of along an elevational, environmental, or latitudinal gradient. We thus
post-migratory nonbreeding movements in migratory birds. To do include partial migration, as long as at least some of the population mi-
so, we first propose a working definition of post-migratory non- grates each year, but exclude year-round nomadism, in which animals
breeding movements. We then use this definition to perform a pre- have little or no fidelity to breeding or wintering regions across years
liminary systematic literature review, through which we characterize (Teitelbaum & Mueller, 2019), and irruptive migrations that occur less
the taxonomic distribution of studies of this behavior, its recorded than annually at the population level (Newton, 2006). Post-migratory
abiotic (e.g., climate) and biotic (e.g., diet, species interactions) nonbreeding movements are large-scale movements that occur after
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TEITELBAUM et al. 3 of 14
the post-breeding migration and before the return migration to breed- This definition is designed to account for the diversity of move-
ing grounds, and connect spatially separated nonbreeding sites. ment strategies within and across taxa, but we acknowledge that
To precisely define post-migratory nonbreeding movements, we movement behavior exists along a spectrum (Nathan et al., 2008;
differentiate post-migratory movements from migratory movements Teitelbaum & Mueller, 2019); therefore, while this definition proved
based on a combination of four characteristics that correspond to functional for our global and preliminary review, it may require crit-
the concepts above: direction, duration, timing, and location (Figure 1). ical consideration for some species or uses. For example, for eleva-
We consider a movement to be migratory if there is continued move- tional migrants, the direction criterion would be more appropriately
ment in the direction of migration after stopping at the site (Hobson & applied to altitude than to latitude, and for pelagic birds, longitude or
Kardynal, 2015), indicating directional persistence (concept 1), and to sea currents might be as important as latitude. In addition, for spe-
be a post-migratory movement if the direction of movement changes cies with large home ranges or where management occurs at a large
substantially from that of the migration (Figure 1a). We also expect spatial scale (e.g., reserves of thousands of hectares), the distance for
migratory movements to be punctuated by stopovers of a relatively sites to be considered separate might be large; conversely, in land-
short duration, whereas post-migratory movements connect sites scapes that vary dramatically at small spatial scales (e.g., urban envi-
used for longer (concept 2); we do not establish a quantitative cutoff, ronments), it may be small. This threshold also interacts with study
but we estimate that a stopover site to be used for the order of days design, since some tracking technologies provide very fine-scale lo-
to weeks, and a post-migratory (i.e., overwintering) site on the order cation data (e.g., GPS error of <10 m), while others are much coarser
of weeks to months (Figure 1b, c). Last, the timing of post-migratory (e.g., Motus networks with broadly spaced towers, geolocators).
nonbreeding movements must be during the nonbreeding season
following migration (i.e., excluding molt migrations that occur prior
to migration), and their location must be outside the extent of the 2 | LITE R AT U R E R E V I E W
normal breeding range of the individual (concept 3, Figure 1d). Finally,
we note that the definition of a “movement” inherently considers dis- 2.1 | Systematic literature search
tance, such that movements must be long enough to be considered
movements between sites, rather than part of one continuously used We searched Web of Science and Google Scholar on 10 February
area. In our literature review (see below), we considered wintering 2021 using the following search string: “migrat* AND (*bird* OR
sites to be separate only if they were at least 5 km apart (though avian) AND (*winter* OR nonbreeding OR non-breeding) AND
this threshold could be larger if identified as such by the authors of (nomad* OR ‘*winter* movement*’ OR ‘multiple *winter* sites’
a paper, who we considered to be experts on each species' biology). OR ‘between*site movement*’ OR ‘multiple non*breeding sites’
F I G U R E 1 Defining post-migratory nonbreeding movements. In each panel, breeding sites are shown as yellow polygons, stopover/
staging sites as orange squares, and post-migratory (i.e., winter) sites as blue circles. Polygon size represents stay duration at a given site. The
distinctions of interest are shown with bold lines and bold font. (a) Post-migratory nonbreeding movements can be primarily longitudinal,
whereas migratory movements are primarily latitudinal. (b) Post-migratory nonbreeding movements can be latitudinal if the first winter site
is used for a relatively long duration but (c) would be considered stopover sites if they are used for the order of days instead of weeks or
months. (d) Winter sites must be used after migration and during the winter season, and outside the spatial extent of an individual's breeding
range. In this example, the first stopover site would otherwise meet the criteria for a winter site but is too close to the breeding site. All
panels display a boreal migrant, but the same principles apply to austral migrants.
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OR itineran*) AND (geolocat* OR *capture* OR track* OR gps OR up in our search (Konno et al., 2020), or because of our choice of
MOTUS OR nanotag OR transmitter OR band* OR ring*).” This string search terms (e.g., we did not include the term “floaters,” which re-
was designed to limit the search to migratory birds only and include fers to nonterritorial birds, some of which could use multiple sites;
known terms to describe post-migratory nonbreeding movements. Brown & Long, 2007, Lenda et al., 2012). Thus, although this data set
We did not search for gray literature or use snowballing techniques. represents a possibly biased sample of species and populations that
This search produced 72 papers from Web of Science; because undertake these movements, it nevertheless suggests interesting
Google Scholar produces so many search results and we found very patterns and directions for research.
few relevant papers beyond the first 70 results, we considered the
first 100 as potential candidates for inclusion. After removing du-
plicates, we read the abstract and/or full text of each paper and ex- 2.2 | Patterns of post-
tracted data if appropriate. The most common reasons for exclusion migratory nonbreeding movements
were the time frame of study (i.e., a study included only the breeding
season or only migration), that it studied between-year site fidel- Our literature search identified 91 bird species across 34 families
ity, and that it studied a non-bird species. From the 69 papers from within 18 orders that undertake post-migratory nonbreeding move-
which we extracted data, we read each discussion section, identi- ments (Figure 2a, see Data Availability Statement: Appendices 1 and
fied any references with potential additional data, and extracted 2). Body masses ranged from 8 g to over 6 kg, with most species
data from these references following the same procedures as for weighing <1 kg. We found evidence for post-migratory nonbreeding
the original search. This secondary search produced an additional movements within all diet types in Elton Traits except frugivores/
54 records from 47 papers; a paper could include multiple records if nectarivores. Post-migratory nonbreeding movements were re-
it studied multiple species. From each paper, we extracted informa- ported from all continents except Oceania, and the most common
tion about the species undertaking the post-migratory nonbreeding continents were Africa and North America.
movements and the characteristics of the movements (e.g., number We recorded the number of wintering sites used per individual
of sites used and distance between sites). We also noted any re- (including the minimum and maximum number, if provided) and the
ported drivers of post-migratory nonbreeding movements (resource distance between sites (including min/max if provided). Birds were
availability, climate, intra- or interspecific competition, predation, observed using up to 21 wintering sites in a single season (Meyburg
age, and sex; see below). We recorded whether the reported driver et al., 1995), but the most common number of wintering sites used
was found to have an effect (positive result) or not (null result), and was two (Figure S1), and in many studies, only part of the popula-
whether the effect was studied analytically (as opposed to being pro- tion undertook post-migratory nonbreeding movements. We also
posed as a hypothesis for further investigation). We also recorded found evidence for intraspecific variation in movement propensity
any proposed or reported consequences of these movements. between populations. For example, Scandinavian-breeding Red-
To understand traits of species undertaking post-migratory necked Phalaropes (Figure 3a) undertake multiple winter move-
nonbreeding movements, we linked each species to data from ments within the Arabian sea, while Scottish and Icelandic breeders
Elton Traits database, which provides body masses and diet classi- overwinter at a single site in the Pacific Ocean (van Bemmelen
fications (frugivore/nectarivore, invertebrate-eater [including both et al., 2019). Reported inter-site distances ranged from our minimum
terrestrial—e.g., flying insects—and aquatic—e.g., crustaceans], om- inclusion distance of 5 km to over 1000 km, with most birds mov-
nivore, plant/seed-eater, or vertebrate-eater/scavenger) (Wilman ing 50–500 km between wintering sites (Figure S1). The number of
et al., 2014). Next, we compared the taxonomic, dietary, and body- wintering sites used, as well as the distance between these sites,
mass traits of species in our data set to those of all migratory birds varied across individuals, species, and years, being highly repeat-
identified in the Global Register of Migratory Species (GROMS), able in some species (e.g., Egyptian Vultures Neophron percnopterus;
which provides a list of migratory species based on handbooks, García-Ripollés et al., 2010) and facultative in others (e.g., White
banding data, and web sites (Riede, 2001). The data and taxonomy Storks Ciconia ciconia; Berthold et al., 2002).
used in GROMS did not always match that in our data set because When we compared the taxonomic, dietary, and body-mass
GROMS was last updated in 2001, so we matched species based on traits of species identified in our search to those of all migratory
both common names and scientific names, but one species in our birds in GROMS, we found that 4.1% of migratory bird species had
data set was missing from GROMS (American Dipper Cinclus mexi- documented post-migratory nonbreeding movements. Although
canus, which is a partial migrant). the prevalence of migration was high in songbirds (Passeriformes)
We note that this review was preliminary and our search missed and they composed a plurality of studies on our data set, they were
some examples of post-breeding nonmigratory movements known to proportionally represented in our search results (3.5% as compared
the authors (e.g., Teitelbaum et al., 2016; Wunderle & Latta, 2000). to 4.6% of non-Passeriformes, χ2(1) = 1.775, p = .182). Among mi-
These papers could have been missed because post-migratory non- gratory bird species, the prevalence of documented post-migratory
breeding movements were mentioned but were not a primary focus nonbreeding movements was highest in Phoenicopteriformes (fla-
of the study (e.g., Teitelbaum et al., 2016); because they were doc- mingoes, 33%), Gaviiformes (loons, 25%), Anseriformes (waterfowl,
umented in non-English language publications and thus not picked 14%), Strigiformes (owls, 13%), and Sphenisciformes (penguins,
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(a)
Proportion of migrants
Number of species
30 0.03
20 0.02
10 0.01
0 0.00
Anseriformes
Accipitriformes
Bucerotiformes
Columbiformes
Galliformes
Passeriformes
Phaethontiformes
Apodiformes
Coraciiformes
Gaviiformes
Gruiformes
Otidiformes
Pelecaniformes
Piciformes
Psittaciformes
Pteroclidiformes
Struthioniformes
Caprimulgiformes
Cuculiformes
Falconiformes
Casuariiformes
Charadriiformes
Ciconiiformes
Sphenisciformes
Strigiformes
Suliformes
Phoenicopteriformes
Podicipediformes
Procellariiformes
(b) (c)
Proportion of migrants
0.05
Number of species
0.04
40
0.4
Density
0.03
20 0.02
0.2
0.01
0 0.00 0.0
1e+01 1e+02 1e+03 1e+04 1e+05
Invertebrate
Omnivore
nectarivore
Plants/
Predatory/
Frugivore/
scavenger
seeds
Body mass (g)
Post−migratory nonbreeding No Yes

movements?
F I G U R E 2 Taxonomic, dietary, and body-mass distributions of species reported to undertake post-migratory nonbreeding movements,
compared with all documented migratory bird species in the Global Register of Migratory Species (GROMS). (a) The taxonomic distribution
of species. Black bars show the number of species from each order that undertook post-migratory movements. Gray bars show the
proportion of migrants in GROMS in each order that were documented to undertake post-migratory movements. Empty bars indicate that
some species in that order were migratory, but none had documented post-migratory nonbreeding movements. (b) Dietary categories
from the Elton Traits database for each species. As in (a), black bars show the number of species in our data set and gray bars show the
proportion of migrants in GROMS from each dietary category that have documented post-migratory movements. (c) Body-mass distributions
of migratory species documented to undertake post-migratory nonbreeding movements (black) and of species with no documented
nonbreeding movements (red). Note the log scale of the x-axis.
12%). No post-migratory nonbreeding movements were docu- comprised only 3.5% of migratory birds identified in GROMS. Note
mented in 11 orders that contained migrants; the most numerous that dietary categories in Elton Traits are defined based on the
of these were Coraciiformes (kingfishers and bee-eaters, 35 mi- majority of a species' diet, so bird species that eat fruit are not de-
gratory species), Columbiformes (doves and pigeons, 20 migra- fined as frugivores unless fruit makes up at least 50% of their diet.
tory species), Piciformes (woodpeckers, 16 migratory species), However, the omnivores in our data set tended to use the fewest
Psittaciformes (parrots, 16 migratory species), and Podicipediformes wintering sites (Figure S1), which suggests that the diet breadth of
(grebes, 15 migratory species). Prevalence of post-migratory non- omnivores might allow them to switch diet items rather than moving
breeding movements differed significantly among orders (Figure 2a, to new sites when conditions change.
χ2(28) = 52.362, p = .003). The average body mass of species with Although these geographic, taxonomic, and trait-related pat-
documented post-migratory nonbreeding movements was higher terns prompt interesting ecological and evolutionary questions,
than that of migrants on average (Figure 2c, ln-transformed body sampling bias inevitably played some role in producing these pat-
mass, χ2(1) = 9.267, p = .002). terns, which highlights the need for more primary research on
Past work on irruptive migration hypothesized that birds with post-migratory nonbreeding movements. For example, the trend
specialized diets might need to move more in response to food short- of larger body masses in species undertaking post-migratory non-
ages compared with dietary generalists (Newton, 2006). We found breeding movements could equally stem from a biological pattern
little evidence that the prevalence of post-migratory nonbreeding (i.e., large species more likely to move during the nonbreeding sea-
movements differed among dietary groups (Figure 2b, χ2 (4) = 8.272, son, possibly due to energetic needs) and from sampling biases (i.e.,
p = .082); there were no frugivores in our data set, but frugivores large species have a longer history of individual tracking because
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F I G U R E 3 Examples of species that undertake post-migratory nonbreeding movements, illustrating the diversity of taxa and geographic
locations represented. Photos are all by R. Hall, with the exception of Wahlberg's Eagle (by Mike's Birds, Wikimedia Commons: https://
commons.wikimedia.org/wiki/File:Wahlberg%27s_Eagle_(6609722249).jpg) and Pallid Swift (by Bogbumper, Wikimedia Commons: https://
commons.wikimedia.org/wiki/File:Apuspallidus_9043.JPG).
(a) (b) F I G U R E 4 Causes and associations of

60 post-migratory nonbreeding movements.
Green bars indicate positive results,
Number of records
Number of records
15 i.e., that a factor was associated with

40 post-migratory nonbreeding movements;
orange bars indicate negative results. Pale
10
bars indicate that an effect was proposed
20 but was not tested analytically. A single
5 study can appear in multiple categories.
(a) Reported relationships between age
0 0
or sex and post-migratory nonbreeding
movements. (b) Reported ecological
competition
resources
causes of post-migratory nonbreeding

predation
age
climate
sex
movements. Note the different y-axis

scales across panels.
Effect detected No effect detected

Effect hypothesized No effect hypothesized
of constraints on tag weight). We also found few studies of austral capacity for scientific research globally, could move towards miti-
migrants or migrants from Oceania, which could stem from either gating this bias (Pettorelli et al., 2021). Comprehensively collating
true phylogeographic patterns and/or from under-representation studies that document post-migratory nonbreeding movements
of authors from the Global South (Maas et al., 2021; Macgregor- is also challenging because of the diversity of terms that refer to
Fors et al., 2021; Stocks et al., 2008). In addition, despite being these movements. We hope that the continued development of
highly mobile and showing evidence of multiple extended station- lightweight and inexpensive tracking devices such as nanotags
ary periods in the nonbreeding season (e.g., Amélineau et al., 2021), (Griffin et al., 2020; Mckinnon et al., 2019) will spur additional
pelagic seabirds were poorly represented in our data set, probably research in under-represented taxa and geographic regions, and
because their movements do not fit traditional definitions of mi- will enable meaningful future comparative analyses of the phylo-
gration developed for terrestrial species. Inclusion of non-English genetic, geographic, and life-history correlates of post-migratory
language papers, alongside political and social actions that build nonbreeding movement across species.
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2.3 | Drivers of post-migratory regions, meaning that temperatures at their wintering sites will fluc-
nonbreeding movements tuate more dramatically, potentially reaching birds' thermal minima
and forcing them to move. Further, these thermal minima might be
Like nomadic and migratory movements (Shaw, 2016; Teitelbaum & more important for temperate water-associated taxa (such as wa-
Mueller, 2019), post-migratory nonbreeding movements could be terfowl), since their habitats freeze at specific temperatures (e.g., in
linked to changes in climate or weather over the nonbreeding season Mallards Anas platyrhynchos and Common Pochards Aythya ferina;
(Sauter et al., 2010); resource tracking (Knight et al., 2019); intraspe- Figure 3b, Keller et al., 2009, Sauter et al., 2010).
cific demographic factors such as age or sex (Fudickar et al., 2013; Although inter- and intraspecies interactions are established
Teitelbaum & Mueller, 2019); and/or species interactions such as drivers of movement behavior (Cohen & Satterfield, 2020; Hopcraft
competition and predation (Smith et al., 2011). In our literature re- et al., 2014), few studies explored competition (N = 6) or predation
view, resource availability, climate, intra- or interspecific competi- (N = 4) as drivers of post-migratory nonbreeding movements, and
tion, predation, age, and sex were all proposed as potential causes exactly half of the studies in each category dismissed each of these
and correlates of post-migratory nonbreeding movements. drivers. Similarly, despite the potential role of pathogens in the evo-
Age and sex could influence the movement propensity of birds lution of migration itself (Altizer et al., 2011; Shaw & Binning, 2016)
because dominance behaviors, territoriality, and energetic consider- and the documented effects of infections on the movement behav-
ations for reproduction differ across sex and age classes (Fudickar ior of migratory birds (van Gils et al., 2007), we found no record of an
et al., 2013; Marques et al., 2009). Our review showed that age and/ association (hypothesized or empirical) between infection and post-
or sex were only sometimes related to post-migratory nonbreeding migratory nonbreeding movements.
movements (Figure 4a), and there was no consensus about whether
adults, juveniles, males, or females were more likely to undertake
post-migratory nonbreeding movements. Of the 12 studies that 2.4 | Methods for studying post-
examined age, seven (58%) found that age was related to post- migratory nonbreeding movements
migratory nonbreeding movements. Juveniles were more likely to
undertake post-migratory nonbreeding movements than adults Advances in tracking technologies over the past 30 years have
in four of these cases (Moore, 1976; Quillfeldt et al., 2010; Sauter transformed the study of post-migratory nonbreeding movements,
et al., 2010; Strandberg et al., 2012), but adults were more likely allowing detailed descriptions of individual movement patterns
than juveniles to move in three cases (Cox & Afton, 2000; Mckinnon across increasingly diverse taxa (McKinnon et al., 2013; Meehan
et al., 2019; Strandberg et al., 2008), indicating that the role of age in et al., 2022; Smith et al., 2022). In our review, post-migratory non-
driving post-migratory nonbreeding movements is far from univer- breeding movements were documented using a wide range of
sal. In the 19 studies that reported migration patterns by sex, only tracking technologies, including (in order of frequency): geoloca-
four (21%) found an association between sex and the use of multi- tors, satellite telemetry, ringing/banding, GPS telemetry, radio te-
ple wintering sites; these studies were evenly split between male lemetry, banding, point counts, and stable isotopes (Figure 5). Even
and female biases in movement (Burgess et al., 2020, Mckinnon before the widespread use of automated tracking devices, work on
et al., 2019, Quillfeldt et al., 2010, Wunderle, 1995).
Climate and resource availability are arguably the most fre-
quently cited drivers of animal movements across taxa (Mueller & 40
Fagan, 2008; Neumann et al., 2015). Accordingly, across all 88 re-

Number of studies
cords that reported potential causes of post-migratory nonbreeding 30
movements, climate, and resource availability were the most common

(n = 47 and n = 57, respectively; Figure 4b), and 25 studies reported 20
a joint influence of climate and resources. Resource-driven move-

ments were most common in invertebrate-eaters and Passeriformes 10
(songbirds), whereas climate-driven movements were more com-

monly reported than resource-driven movements in Anseriformes 0
(waterfowl), Charadriiformes (shorebirds), and Phoenicopteriformes

GPS
telemetry
Satellite
Isotopes
Survey
Band/resight
Geolocator
Radiotelemetry
(flamingoes) (Figure S2). Although it is difficult to differentiate be-

tween climate as a proximate versus an ultimate driver of move-
ment because of its effects on resource availability (e.g., Studds &
Marra, 2007), we hypothesize that this association between taxo-
Tracking method
nomic groups and drivers of movement could be related to winter-
ing habitat and constraints on site choice (Terrill & Ohmart, 1984). F I G U R E 5 Frequency of different tracking technologies used to
Larger-bodied birds (e.g., Anseriformes) and predators might be more document post-migratory nonbreeding movements. Methods are
likely than smaller-bodied insectivores to overwinter in temperate sorted by the potential weight and modernity of the device.
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migratory songbirds in sub-Saharan Africa and waterfowl in Europe 2.5 | Consequences of post-
identified the presence of post-migratory nonbreeding movements migratory nonbreeding movements
(Gätke, 1879; Moreau, 1972), showing that, while high-frequency
and high-precision data are useful, they are not strictly necessary Migratory and nomadic movements are known to affect population
for identifying these movements. dynamics, community composition, and ecosystem processes (Bauer
Early satellite telemetry of large birds identified the number of & Hoye, 2014; Teitelbaum & Mueller, 2019), and arguably the same
nonbreeding sites used and inter-site distances, even though it was could be true for post-migratory nonbreeding movements. Most of
at lower spatial and temporal resolution than many current technol- the studies included in this literature review focused on patterns
ogies; for example, satellite telemetry identified that a Wahlberg's and drivers of post-migratory nonbreeding movements; although
Eagle (Figure 3c), an austral migrant, undertook movements among the consequences of these movements are much less well-explored,
more than 20 sites separated by 10s-100s of km on its Nigerian win- a number of studies suggest the potential for strong effects of post-
tering grounds (Meyburg et al., 1995). More recently, light-level geo- migratory nonbreeding movements on population, community, and
locators, which allow tracking of smaller, lighter birds, have shown ecosystem processes. We recorded hypotheses from each paper in
that birds' winter movements follow local environmental conditions our data set for these consequences qualitatively.
and life-history events (e.g., post-migratory nonbreeding movements At the population level, post-migratory nonbreeding movements
of Bobolinks from Venezuela to Argentina track changes in grassland could influence survival and produce carry-over effects that affect
productivity (Figure 3d, Renfrew et al., 2013)). Tracking methods that future breeding success (Stutchbury et al., 2016), in ways that have
provided automated data at relatively high spatial and temporal res- both benefits and drawbacks for fitness. While theory suggests that
olution were generally better represented in our data set than tech- populations with high movement capacity and those that are spread
niques that required continued field collection (Figure 5), probably over large geographic areas might be buffered from locally harsh
because it is difficult to observe such large-scale movements with conditions (Burgess et al., 2020), others that obligately use multi-
studies at only a single or a few sites; using banding or point counts ple sites could be vulnerable to poor conditions or habitat loss at a
alone, it not possible to measure the movement distances or fates of single site (Smith et al., 2011). For example, in Great Reed Warblers
birds that disappear during the nonbreeding season unless individu- (Acrocephalus arundinaceus) that use two nonbreeding sites, dry
als happen to be recaptured elsewhere. However, creative methods conditions at the first site affected body condition upon arrival at
have occasionally allowed the identification of post-migratory non- the second site, but body condition was similar across years by the
breeding movements without measuring movement directly; among end of the nonbreeding season, suggesting that environmental con-
Common Chiffchaffs wintering in Portugal (Figure 3e), fat reserves ditions at the second site were sufficient to counteract poor rain-
of winter-itinerant birds were more similar to winter residents than fall conditions at the first site (Sorensen et al., 2016). This ability to
to migrants stopping over (Catry et al., 2003), suggesting differences use multiple sites for different functions (e.g., molting vs. fueling for
in physiology between movements during winter compared with migration) could promote future migration and breeding success
those during migration. (Sorensen et al., 2016). Identifying the prevalence of post-migratory
Ongoing technological and methodological developments show nonbreeding movements within a population can also inform esti-
promise for shaping future work on post-migratory nonbreeding mates of annual survival; some species that were thought to have
movements. A recent study using nanotags, which are lightweight low overwinter survival were later discovered to have individuals
and—unlike geolocators—do not require recapture for data retrieval, that undertake post-migratory nonbreeding movements (Blackburn
revealed that Snow Buntings wintering in Ontario move 25–50 km & Cresswell, 2016; Lindberg et al., 2007).
every few days, covering hundreds of square kilometers over the Post-migratory nonbreeding movements could also influence
winter (Figure 3f, Mckinnon et al., 2019). Advances in the statistical metacommunity and ecosystem dynamics at broad spatial scales, for
analysis of movement data can also help identify post-migratory non- example by providing pulses of resources or predators that influence
breeding movements. For example, Pallid Swifts (Figure 3g), which local trophic dynamics; shaping the movement patterns of predators
spend the entire nonbreeding period on the wing, could theoreti- and prey (Cohen & Satterfield, 2020); or providing crucial ecosys-
cally continuously track resources across space, but kernel density tem functions and services such as seed dispersal and pest control
estimation of geolocator tracks showed that, instead, they under- (Lundberg & Moberg, 2003). One of the most commonly hypoth-
take protracted stays in multiple regions punctuated by movements esized consequences of post-migratory nonbreeding movements
(Norevik et al., 2019). Finally, the proliferation of openly accessible for community dynamics in our data set was the potential role of
movement tracks (e.g., MoveBank, Bird Migration Explorer), distri- migrants in dispersing pathogens across their winter range. Multiple
bution data (e.g., eBird), and open-source software packages for con- papers expressed concern about the potential dispersal of avian in-
ducting movement analyses (Joo et al., 2020) provide powerful tools fluenza, especially given the pathogen's high prevalence in water-
for studying post-migratory nonbreeding movements across popula- bird species that frequently undertake post-migratory nonbreeding
tions or taxa. The decreasing cost of tracking devices will also reduce movements (Gourlay-L arour et al., 2012; Guillemain et al., 2010;
current biases in sampling by making these study methods accessible Keller et al., 2009; Vos & Wietses, 2010). For example, Common
to more scientists studying a broader array of species. Teal (Anas crecca), a known host of low pathogenic avian influenza,
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moved up to 640 km in 2 days, well within the shedding period for sites could eventually disappear in this species, potentially linked to
this virus, suggesting their strong potential as a dispersal vector long-term changes in environmental conditions.
(Brochet et al., 2009). However, few studies have explicitly linked Changes in food availability associated with agriculture and ur-
infection data to post-migratory nonbreeding movement patterns banization, along with warmer temperate winters, could also drive
(but see Vos & Wietses, 2010). In addition, although these processes changes in winter site use or promote the adoption of novel win-
were not addressed by any papers in our data set, we hypothesize ter movement patterns. For example, some White Ibis (Eudocimus
that post-migratory nonbreeding movements could be important for albus) wintering in South Florida, USA have recently begun to pref-
seed dispersal, as has been documented for nomadic movements erentially winter in urban parks, where they are hand-fed by people
(Mueller et al., 2014; Teitelbaum & Mueller, 2019). (Teitelbaum et al., 2020); analysis of movement tracks shows that
The roles of migrants as consumers, predators, and resources urban-wintering birds spend longer overwintering and use fewer
could also influence large-scale ecosystem processes and pro- wintering sites than those that overwinter nonurban areas, which
vide ecosystem services or disservices. For example, as Montagu's move nomadically among wetland sites depending on water depth
Harriers (Circus pygargus) track grasshopper densities across the (Kidd-Weaver et al., 2020; Murray et al., 2021). By contrast, changes
Sahel during the winter (Trierweiler et al., 2013), they provide po- to migration patterns in response to land-use and climate change
tential pest control service for people (Mullié & Guéye, 2010). By could trigger novel post-migratory nonbreeding movements in some
contrast, Bobolinks, which use agricultural fields on their South species. Provisioning of ornamental plants and feeders has allowed
American wintering grounds, are considered a pest species because some hummingbirds to overwinter in the United States, a much
of their large flocks that can consume large quantities of crops shorter migration than their historical migration to Central America
(Renfrew et al., 2013). (Greig et al., 2017). While these hummingbirds can remain at a single
feeder for an entire winter, birds wintering at more northerly lati-
tudes might be forced to undertake novel post-migratory nonbreed-
2.6 | Responses to global change ing movements associated with mid-winter cold snaps; this pattern
was observed in a banded Rufous Hummingbird (Selasphorus rufus)
Given that resources and environmental conditions are common in Ohio, USA, which departed its wintering site in mid-December
drivers of post-migratory nonbreeding movements, ongoing cli- but returned to the same feeder the following October (Chartier
mate change, and landcover change could impact populations and et al., 2012).
movement patterns of species undertaking these movements.
Migratory species are generally considered especially suscep-
tible to the consequences of global environmental change (Both 3 | P OS T- M I G R ATO RY N O N B R E E D I N G
et al., 2010), but how migration interacts with post-migratory non- M OV E M E NT S I N C ATH A R U S TH RU S H E S
breeding movements is less well understood. Species with larger
nonbreeding ranges have been documented to have more stable To complement our broad systematic review, we conducted a re-
populations (Gilroy et al., 2016), but the extent to which species view of post-migratory nonbreeding movements in a single taxo-
can adapt their wintering behavior to changing conditions might nomic group, the North American-breeding thrushes in the genus
depend on whether these movements are obligate, or flexible be- Catharus. This focused review allowed us to further explore hypoth-
tween years and individuals. In exceptionally dry years, obligately eses for why some species might display these movements while
migratory Montagu's Harriers in the Sahel used the same number others do not, as well as to more thoroughly document the ways
of wintering sites as in wetter years, but left early winter sites that nonbreeding behavior is studied and described across species.
sooner, departed their final wintering site later, and foraged more Within this genus, two of the five migratory species, Swainson's
intensively (Schlaich et al., 2016), indicating that they adapt their Thrush (C. ustulatus) and Veery (C. fucescens), exhibit post-migratory
movement phenology to environmental conditions rather than nonbreeding movements (Figure 3h, i). We conducted a systematic
using more sites. This adaptation might become especially impor- search of nonbreeding ecology of this genus (Web of Science search
tant because droughts in the Sahel are predicted to become more for “Catharus AND [winter* OR over*winter* OR non*breeding]”)
frequent with ongoing climate warming (Held et al., 2005). By con- and found no evidence for post-migratory nonbreeding movements
trast, in White Storks that employ a variety of wintering strategies, in other three species (Bicknell's Thrush C. bicknelli, Hermit Thrush
a bird wintering in sub-Saharan Africa moved similarly in dry and C. guttatus, and Gray-cheeked Thrush C. minimus).
wet years but did not attempt to breed in the breeding season fol- Studies of nonbreeding ecology of the three stationary winter
lowing the dry year (Berthold et al., 2002), which could indicate species frequently contrasted territorial individuals with “float-
fitness consequences of movement patterns that do not change in ers,” or individuals that did not maintain winter territories (Brown
response to environmental conditions. A larger study found that et al., 2002; Kwit et al., 2004; Townsend et al., 2010). While it might
juvenile White Storks wintering in Europe or North Africa have be tempting to classify floaters as performing post-migratory non-
higher survival than those moving to sub-Saharan Africa (Cheng breeding movements, it is equally possible that they simply use a
et al., 2019), suggesting that storks' use of sub-Saharan wintering larger area during the winter than territorial birds. For example, in
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Bicknell's Thrush, floaters were detected via radiotelemetry at a 4 | CO N C LU S I O N S A N D FU T U R E

single site throughout winter even though they did not appear to D I R EC TI O N S FO R TH E S T U DY O F P OS T-
establish territories (Townsend et al., 2010). Nevertheless, these M I G R ATO RY N O N B R E E D I N G M OV E M E NT S
patterns suggest that territory acquisition and maintenance might
contrast with post-migratory nonbreeding movements, for example, Our literature review identified post-migratory nonbreeding move-
if the investment in territories is high enough that they are worth ments across diverse taxa, geographies, and environments. Overall,
maintaining throughout the winter, even if conditions deteriorate while much of the work synthesized in this study describes pat-
(Brown & Long, 2007). In Swainson's Thrush, stationary populations terns and drivers of post-migratory nonbreeding movements, we
are thought to be territorial (Mack et al., 2020) while transient ones found relatively few studies that address their wider ecological
are not (Inserillo et al., 2020). In Veery, where all tracked individu- consequences. This gap in the literature is especially important
als undertook long-distance nonmigratory movements (Heckscher because understanding the consequences of movement can help
et al., 2015), winter territoriality is not recorded (Heckscher inform predictions for how global change will impact the ecosys-
et al., 2020). Therefore winter-territorial species, populations, or in- tems through which these birds move. This line of research could
dividuals might be less likely to undertake post-migratory nonbreed- draw on our understanding of the consequences of nomadic animal
ing movements than nonterritorial species (although the ultimate movements (e.g., for seed dispersal, Mueller et al., 2014), irruptions
causes of differences in territorial behavior remain an open ques- (Newton, 2006), stopover site use (Schmaljohann & Eikenaar, 2017),
tion; Stutchbury & Morton, 2016). and molt migration (Tonra & Reudink, 2018). For example, year-
Diet, climate, and evolutionary history are all hypothesized round high-resolution tracking of migratory populations in which
as potential drivers of post-migratory nonbreeding movements. some individuals are stationary during the nonbreeding season while
Since all Catharus species consume a mixture of fruits, seeds, and other individuals move could help clarify whether there are carry-
insects and switch seasonally to more frugivory between breed- over effects of these movements for survival and reproduction dur-
ing and post-breeding migration, food seems an unlikely driver of ing migration and the breeding season.
post-migratory nonbreeding movements in this genus. Supporting We also found that, although post-migratory nonbreeding move-
this idea, stationary Hermit Thrushes that defend winter territories ments were often linked to climate and resource availability at the
do not change their territory size when fruit was experimentally level of individuals or populations, our interspecific comparison of
removed, suggesting within-season dietary flexibility (Brown & migratory Catharus thrushes showed little evidence for dietary, geo-
Long, 2006). However, in Ecuador, transient Swainson's Thrushes graphic, or evolutionary patterns in which species undertake these
were hypothesized (but not tested) to be tracking ephemeral fruit movements. Therefore, while the environment certainly plays an
availability (Inserillo et al., 2020), while Veery intratropical migra- important role in driving post-migratory nonbreeding movements
tions are associated with seasonal flooding of the Amazon, poten- of individuals, it interacts with other life-history traits and other ad-
tially reflecting a preference to continue foraging for invertebrates aptations to environmental variability to determine whether these
on the ground (Heckscher et al., 2011, 2015). In the remaining spe- movements occur at the species or population level. A combination
cies, we found no clear climatic or dietary correlates of nonbreed- of large-scale phylogenetic studies (such as those that study the or-
ing movements. igins of migration, (Dufour et al., 2020, Salewski & Bruderer, 2007))
Past work on the evolutionary relationships of Catharus and smaller-scale population studies could help clarify how move-
thrushes and the evolution of migration does not support a ment behavior interacts with other adaptations to environmental
phylogenetic basis to post-migratory nonbreeding movements. variability both within and across taxa.
Swainson's Thrushes were inferred to evolve a southward migra- Our review also uncovered wide variation within the cate-
tion from North America, while Veery and its stationary-wintering gory of “post-migratory nonbreeding movements.” For example,
congeners were thought to independently evolve their north- movement distances ranged from just a few km to over 1000 km.
bound migration from sedentary ancestors in Central America Longer movements can be riskier than shorter-distance move-
(Voelker et al., 2013). The repeated, seemingly obligate long- ments (Cheng et al., 2019), can be associated with differences in
distance intratropical migration of Veery in South America, shown landscape structure or exposure to more diverse environments
by individuals from different breeding populations, has been hy- (Tucker et al., 2019), and pose larger energetic requirements than
pothesized to pre-date the breeding migration into North America do shorter-distance movements (Alerstam & Lindström, 1990).
(Heckscher et al., 2015). Furthermore, neither migration distance Therefore, like all animal behaviors, it is likely that there are
nor breeding/nonbreeding site latitudes are predictive of post- subgroupings within this behavior, which might be linked to dif-
migratory nonbreeding movements, since California-b reeding ferent social or environmental forces (e.g., local competition for
Swainson's Thrushes that undertake relatively short migrations to high-quality territories vs. large-s cale spatiotemporal differences
Mexico (Cormier et al., 2013), and Veeries that undertake long- in resource availability). The same distinctions could be made
distance migrations to South America (Heckscher et al., 2011), across the numbers of sites moved, proportions of a population
both demonstrate these movements. moving, and interannual variation in movement behavior. Future
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comparative studies could address these questions by simultane- ORCID

ously studying the physiological, environmental, and social con- Claire S. Teitelbaum https://orcid.org/0000-0001-5646-3184
texts of post-migratory nonbreeding movements across species
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Ecology and Evolution - 2023 - Teitelbaum - Post Migratory Nonbreeding Movements of Birds A Review and Case Study

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Ecology and Evolution - 2023 - Teitelbaum - Post Migratory Nonbreeding Movements of Birds A Review and Case Study

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Received: 29 October 2022 Revised: 17 February 2023 Accepted: 23 February 2023

Post-migratory nonbreeding movements of birds: A review and

Claire S. Teitelbaum | Natalie C. Bachner | Richard J. Hall

Odum School of Ecology, Athens, Georgia,

Ecology and Evolution. 2023;13:e9893. |

Body mass (g)

Post−migratory nonbreeding No Yes

(a) (b) F I G U R E 4 Causes and associations of

15 i.e., that a factor was associated with

causes of post-migratory nonbreeding

movements. Note the different y-axis

Effect detected No effect detected

Fagan, 2008; Neumann et al., 2015). Accordingly, across all 88 re-

cords that reported potential causes of post-migratory nonbreeding 30

movements, climate, and resource availability were the most common

a joint influence of climate and resources. Resource-driven move-

(songbirds), whereas climate-driven movements were more com-

(waterfowl), Charadriiformes (shorebirds), and Phoenicopteriformes

(flamingoes) (Figure S2). Although it is difficult to differentiate be-

Bicknell's Thrush, floaters were detected via radiotelemetry at a 4 | CO N C LU S I O N S A N D FU T U R E

comparative studies could address these questions by simultane- ORCID

You might also like

Ecology and Evolution - 2023 - Teitelbaum - Post Migratory Nonbreeding Movements of Birds A Review and Case Study

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Document Information

Original Title

Copyright

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Ecology and Evolution - 2023 - Teitelbaum - Post Migratory Nonbreeding Movements of Birds A Review and Case Study

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Copyright:

Available Formats

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Received: 29 October 2022 Revised: 17 February 2023 Accepted: 23 February 2023

Post-­migratory nonbreeding movements of birds: A review and

Claire S. Teitelbaum | Natalie C. Bachner | Richard J. Hall

Odum School of Ecology, Athens, Georgia,

Ecology and Evolution. 2023;13:e9893. ﻿ |

Body mass (g)

Post−migratory nonbreeding No Yes

(a) (b) F I G U R E 4 Causes and associations of

15 i.e., that a factor was associated with

causes of post-­migratory nonbreeding

movements. Note the different y-­axis

Effect detected No effect detected

Fagan, 2008; Neumann et al., 2015). Accordingly, across all 88 re-

cords that reported potential causes of post-­migratory nonbreeding 30

movements, climate, and resource availability were the most common

a joint influence of climate and resources. Resource-­driven move-

(songbirds), whereas climate-­driven movements were more com-

(waterfowl), Charadriiformes (shorebirds), and Phoenicopteriformes

(flamingoes) (Figure S2). Although it is difficult to differentiate be-

Bicknell's Thrush, floaters were detected via radiotelemetry at a 4 | CO N C LU S I O N S A N D FU T U R E

comparative studies could address these questions by simultane- ORCID

You might also like

Post-migratory nonbreeding movements of birds: A review and

Ecology and Evolution. 2023;13:e9893. |

causes of post-migratory nonbreeding

movements. Note the different y-axis

cords that reported potential causes of post-migratory nonbreeding 30

a joint influence of climate and resources. Resource-driven move-

(songbirds), whereas climate-driven movements were more com-