Received: 3 January 2020

| Revised: 31 January 2020 | Accepted: 4 February 2020

DOI: 10.1002/ece3.6126

ORIGINAL RESEARCH

Evolution of altitudinal migration in passerines is linked to diet
Claudie Pageau1
| Mariana M. Vale2,3 | Marcio Argollo de Menezes4,5 |
Luciana Barçante6 | Mateen Shaikh7 | Maria Alice S. Alves8 | Matthew W. Reudink1
1

Department of Biological Sciences, Thompson Rivers University, Kamloops, BC, Canada

2

Ecology Department, Federal University of Rio de Janeiro, Rio de Janeiro, Brazil

3

National Institutes for Science and Technology in Ecology, Evolution and Biodiversity Conservation, Goiás, Brazil

4

Physics Institute, Fluminense Federal University, Niteroi, Brazil

5

National Institute of Science and Technology on Complex Systems, Rio de Janeiro, Brazil

6

Programa de Pós-graduação em Ecologia e Evolução, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil

7

Department of Mathematics & Statistics, Thompson Rivers University, Kamloops, BC, Canada

8

Departamento de Ecologia, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, Brazil

Correspondence
Matthew W. Reudink, Department of
Biological Sciences, Thompson Rivers
University, Kamloops, BC V2C 0C8, Canada.
Email: mreudink@tru.ca
Funding information
Fundação Carlos Chagas Filho de
Amparo à Pesquisa do Estado do Rio
de Janeiro, Grant/Award Number: CNE
E-26/202.835/2018 and FAPERJ E-26/
100.811/2009; Natural Sciences and
Engineering Research Council of Canada;
Thompson Rivers University; Conselho
Nacional de Desenvolvimento Científico e
Tecnológico, Grant/Award Number: CNPq
201297/ 2014-0, CNPq 304309/2018-4
and CNPq-PQ 306.579/2018-9; Fonds
de Recherche du Québec - Nature
et Technologies; Coordenação de
Aperfeiçoamento de Pessoal de Nível
Superior, Grant/Award Number: 001

Abstract
Bird migration is typically associated with a latitudinal movement from north to south
and vice versa. However, many bird species migrate seasonally with an upslope or
downslope movement in a process termed altitudinal migration. Globally, 830 of the
6,579 Passeriformes species are considered altitudinal migrants and this pattern has
emerged multiple times across 77 families of this order. Recent work has indicated an
association between altitudinal migration and diet, but none have looked at diet as a
potential evolutionary driver. Here, we investigated potential evolutionary drivers of
altitudinal migration in passerines around the world by using phylogenetic comparative methods. We tested for evolutionary associations between altitudinal migration
and foraging guild and primary habitat preference in passerines species worldwide.
Our results indicate that foraging guild is evolutionarily associated with altitudinal migration, but this relationship varies across zoogeographical regions. In the Nearctic,
herbivorous and omnivorous species are associated with altitudinal migration, while
only omnivorous species are associated with altitudinal migration in the Palearctic.
Habitat was not strongly linked to the evolution of altitudinal migration. While our
results point to diet as a potentially important driver of altitudinal migration, the evolution of this behavior is complex and certainly driven by multiple factors. Altitudinal
migration varies in its use (for breeding or molting), within a species, population, and
even at the individual level. As such, the evolution of altitudinal migration is likely
driven by an ensemble of factors, but this study provides a beginning framework for
understanding the evolution of this complex behavior.
KEYWORDS

bird movement, evolution, foraging guild, Passeriformes, phylogenetic comparative analysis
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.
© 2020 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Ecology and Evolution. 2020;00:1–8.	

www.ecolevol.org

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PAGEAU et al.

1 | I NTRO D U C TI O N

seasonally in mountainous environments, such as Nepal (Katuwal
et al., 2016).

Altitudinal migration is generally described as a seasonal move-

Altitudinal migration has been observed in every zoogeo-

ment from lower elevations to higher elevations for the breeding

graphical region in the world (Barçante et al., 2017) although some

season and a downslope movement for the nonbreeding season

hotspots seem to host a higher proportion of altitudinal migrants,

(Barçante, Vale, & Alves, 2017; Hayes, 1995; Mackas et al., 2010).

such as the Himalayas and western North America (Boyle, 2017).

Some species also engage in altitudinal movements to reach molt-

It is important to note, however, that some of this variation in

ing grounds (Rohwer, Rohwer, & Barry, 2008; Wiegardt, Wolfe,

the proportion of altitudinal migrants could result from a differ-

Ralph, Stephens, & Alexander, 2017). Altitudinal migration has

ence in sampling efforts across the world (Barçante et al., 2017).

been observed in a broad diversity of bird species; in total, 1,238

Alternatively, environmental conditions in those regions, such as

species across 130 families of birds have been described as alti-

habitat availability and seasonality, may also favor the evolution of

tudinal migrants (Barçante et al., 2017), suggesting repeated in-

altitudinal migration.

dependent evolution of this behavior (Figure 1). There are three

Our goal was to examine potential drivers of the evolution

main advantages ascribed to altitudinal migration: reduction in

of altitudinal migration in passerines. The order Passeriformes

the risk of predation (Boyle, 2008a), avoidance of harsh climatic

represents approximately half of the avifauna and 13% of them

conditions (Boyle, 2008b; Boyle, Norris, & Guglielmo, 2010;

are described as altitudinal migrants, making them a good choice

Hahn, Sockman, Nreuner, & Morton, 2004), and tracking of food

for this study. Of the 6,579 passerines species and subspecies

resources (Chaves-Campos, 2004; Kimura, Yumoto, & Kikuzawa,

recorded in this study, 830 species are considered altitudinal

2001; Levey, 1988; Loiselle & Blake, 1991; Solorzano, Castillo,

migrants and are distributed across 77 of the 137 families of

Valverde, & Avila, 2000).

Passeriformes (Figure 1). Using a speciose and globally distributed

Most studies on altitudinal migration have focused on the

group of birds, we conducted large-scale phylogenetic compara-

food abundance hypothesis rather than predation and climatic

tive analyses to examine evolutionary associations between alti-

conditions, which are extremely challenging to study across a

tudinal migration and diet (foraging guild) and habitat. In addition,

wide range of species and habitats. Some studies on altitudinal

we asked whether these associations differ depending on the zoo-

migration have provided evidence that frugivorous bird abun-

geographic region. We expected that frugivorous and nectivorous

dance is linked to fruit and flower abundance (Chaves-Campos,

species were driven toward altitudinal migration in the Neotropics

2004; Kimura et al., 2001; Levey, 1988; Loiselle & Blake, 1991)

because they were tracking fruit and flower abundance which

while others have shown no evidence of this phenomenon (Boyle,

varies seasonally (Barçante et al., 2017; Chaves-Campo, 2004;

2010; Hart et al., 2011; Papeş, Peterson, & Powell, 2012; Rosselli,

Kimura et al., 2001; Levey, 1988; Loiselle & Blake, 1991). For

1994). Boyle (2017), Chaves-Campos (2004), Kimura et al. (2001)

every other region, invertivorous species would be driven toward

and Pratt, Smith, and Beck (2017) suggested that food abundance

altitudinal migration (Barçante et al., 2017). We also expected al-

drives uphill migration only, but this might depend of the spe-

titudinal migration to be evolutionary associated with forest hab-

cies since Loiselle and Blake (1991) described downhill move-

itats in the Neotropics because altitudinal migrants in Costa Rica

ment for some frugivorous species in Costa Rica when food was

(Stiles, 1988; Stiles & Clarke, 1989) and southeastern Brazil (Stotz,

decreasing.

unpublished—see Stotz, Fitzpatrick, Parker, & Moskovits, 1996),

If altitudinal migration evolved as a strategy to track food resources, we would predict a link between diet (foraging guild) and

for instance, include a high number of restricted-range and forest-dependent species.

altitudinal migration; however, the evidence for this relationship
remains unclear. Frugivory has been suggested as a driver of altitudinal migration, in part because frugivorous altitudinal migrants
have been observed more frequently at higher elevations in Costa
Rica (Blake & Loiselle, 2000; Boyle, Conway, & Bronstein, 2011)

2 | M E TH O DS
2.1 | Ethics statement

and Nepal (Katuwal et al., 2016). However, Barçante et al. (2017)
examined the foraging guild of all altitudinal migrant birds and

No permits were required for this project.

showed that invertivorous altitudinal migrants are most abundant worldwide, except in the Neotropics where frugivores and
nectivores are more abundant. Despite the fact that insect abun-

2.2 | Data collection

dance in temperate regions is often posited as a major driver of
the evolution of long-distance migration, little research has been

We compiled data for species and subspecies of songbirds across

dedicated to the role of insect abundance in the study of altitu-

the world, from supplementary material in Barçante et al. (2017) and

dinal migration even though insect intake might be crucial during

Wilman et al. (2014), and data mining from two online databases:

the breeding season (Chaves-Campos, 2004; Levey, 1988) and

IUCN Red List and BirdLife Data Zone (retrieved in November 2018).

invertivore bird species have been shown to vary in elevation

All entries were checked for nomenclature inconsistencies. Our

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PAGEAU et al.

F I G U R E 1 Phylogeny of all
Passeriformes and occurrences of
altitudinal migration represented by red
circles. Speciose families' (>100 species)
names and silhouettes are shown along
the outside of the phylogeny

universe consists of all 6,579 passerines in the IUCN Red List data-

nectarivores, 547 seed/plant materials, 4,018 invertivores, 1,213

base, downloadable from their website https://www.iucnr​edlist.org/

omnivores, and 20 vertebrates/fish/scavengers. Seventy-one spe-

search after restricting (advanced) searches by taxonomy selecting, in

cies had no information on Willman et al. (2014) and were classi-

the "search filters" option [Kingdom = Animalia; Phylum = Chordata;

fied with information from the Handbook of the Birds of the World

Class = Aves; Order = Passeriformes]. We associated four variables

Alive (del Hoyo, Elliott, Sargatal, Christie, & Kirwan, 2019) (47 spe-

to each species: altitudinal migration status, primary habitat prefer-

cies) or closest related species (24 species).

ence, foraging guild, and zoogeographic region.

To build the zoogeographic region (Newton & Dale, 2001), we

A species was classified in our dataset as altitudinal migrant if

downloaded from IUCN Red List website 13 lists of Passeriformes,

its (common or scientific) name is listed in Barçante et al. (2017) ei-

each with all Passeriformes observed on a specific "Land Region"

ther as altitudinal (238 species) or probable altitudinal migrant (592

(selected in the "search filters" option) and translated those re-

species). BirdLife Data Zone provides, among many other informa-

gions to a reduced set of zoogeographical regions as follows:

tion, the list of preferred breeding and nonbreeding habitats of a

"Caribbean islands" = Neotropical, "Antarctica" = Neotropical,

given species on the webpage http://dataz​one.birdl​ife.org/speci​es/

"East

facts​heet/common_name-scien​tific_name/details (where spaces are

"Mesoamerica" = Neotropical, "North Africa" = Checked indi-

Asia"

=

Indomalayan,

"Europe"

=

Palearctic,

replaced by the character "-" on its common and scientific names).

vidually; "North America" = Neartic. "North Asia" = Palearctic.

Considering the great variety of habitats, we only used the major

"Oceania" = Australian. "South America" = Neotropical, "South and

natural breeding habitat for each species and collapsed habitats into

Southeast Asia" = Indomalayan, "Sub-Saharan Africa" = Afrotropical,

four major categories: dense habitat (forest + shrubland, 4,635 spe-

"West and Central Asia" = Checked individually. Species resid-

cies), open habitat (grassland + savanna + open woodland + rocky

ing on more than one zoogeographical region were classified as

areas, 563 species), water habitat (wetland + marine, 164 species),

"Widespread" after manual investigation of their breeding distribution

and generalist (species that occupied two or more major categories,

maps in the IUCN website. Our dataset consists of 1,298 Afrotropical

1,217 species). A total of 1,217 species occupied two or more major

(11% migrant), 816 Australasian (6% migrant), 1,422 Indomalayan

categories and were classified as generalists.

(17% migrants), 288 Nearctic (31% migrant), 2,387 Neotropical (10%

Foraging guild data were fetched from Willman et al. (2014),
with species distributed among five categories: 754 frugivores/

migrant), 342 Palearctic (20% migrant), and 26 Widespread (42% migrant) species.

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PAGEAU et al.

2.3 | Phylogeny

terms in the top-ranked model, we found strong effects of foraging

We downloaded the first 1,000 trees from Hackett backbone phy-

Figure 2c), and a foraging guild:region interaction (F12 = 10.05,

logenetic trees (Hackett et al., 2008). Hackett backbone phyloge-

p < .0001). The interaction model revealed that herbivore/wide-

netic trees are available from https://BirdT​ree.org (Jetz, Thomas,

spread (t = 4.75, p < .0001), omnivore/Palearctic (t = 3.26, p = .0011),

guild (F2 = 6.48, p = .0016; Figure 2a), region (F6 = 23.77, p < .0001;

Joy, Hartmann, & Mooers, 2012). The trees were read in Rstudio

and omnivore and herbivore/Nearctic (t = 7.43, p < .0001, t = 4.43,

(RStudio Team, 2016) using the ape package (Paradis & Schliep,

p < .0001) species were more likely to exhibit altitudinal migration

2018). We trimmed 4,105 species to only keep Passeriformes spe-

(Table 2).

cies using the drop.tip function in the phytools package (Revell,
2012). Using TreeAnnotator (Rambaut & Drummond, ), a maximum
clade credibility tree was created with 1% burn-in and mean heights.

4 | D I S CU S S I O N

The final tree used in the analysis consisted of 5,888 species and 691
subspecies. Most subspecies are considered full species by IUCN

We explored two potential drivers of the evolution of altitudinal

(2019), but are not included in Birdt​ree.org phylogenies (Jetz et al.,

migration in passerines by conducting large-scale phylogenetic

2012). Since they were absent from the Hackett backbone phylog-

comparative analyses. Our results indicate that foraging guild is evo-

eny, subspecies were added to the tree by matching the genus and

lutionarily associated with altitudinal migration, but this relationship

species names of the sister species (e.g., Acrocephalus luscinius hiwae

varies across zoogeographic regions. Habitat did not appear to be

matched with Acrocephalus luscinius) which created polytomies in-

strongly linked to the evolution of altitudinal migration.

side the phylogeny.

Globally, species eating fruit/nectar or seed/plant material
were more likely to exhibit altitudinal migration than omnivores

2.4 | Statistical analysis

and invertivores, despite the fact that most (61%) passerine
birds are insectivorous. This observation follows most of the literature, which emphasizes that frugivorous altitudinal migrants

To examine evolutionary associations between altitudinal migra-

should track fruit and flower abundance seasonally, particularly in

tion and life history characteristics, we used phylogenetic general-

Costa Rica (Blake & Loiselle, 2000; Boyle et al., 2011) and Nepal

ized least squares (pgls) analyses from the packages ape (Paradis &

(Katuwal et al., 2016). Note that Barçante et al. (2017) in a study

Schliep, 2018) and nlme (Pinheiro, Bates, DebRoy, & Sarkar, 2019).

including all bird orders (not only Passeriformes) showed that in-

Brownian correlation and the maximum likelihood method were

vertivorous altitudinal migrants were more abundant around the

applied to each model. The models consisted of the response vari-

world. Indeed, the number of invertivore species that are altitu-

able (altitudinal migration) coupled with each predictor individually

dinal migrants is higher than any other foraging guild; however,

(diet, habitat, and region), predictors paired together, or all predic-

most Passeriformes eat invertebrates as their main diet and that

tors together. Two models also included an interaction; one be-

foraging guild is by far the most speciose (4,018 of 6,579 species).

tween diet and region and one between habitat and region. The

However, the proportion of invertivorous altitudinal migrants was

interaction was included to test whether the patterns of guild vary

relatively low and we found no evolutionary association between

from one zoogeographical region to another as shown by Barçante

altitudinal migration and invertivory for passerines, neither glob-

et al. (2017); the same was applied to habitat. For the models with

ally nor within regions. Note, however, that the classification of

the interaction, we had to merge frugivore/nectarivore with seed/

each species to one foraging guild is tricky because diet can vary

plant material and vertebrate/fish/scavenger with invertivore, re-

through the seasons. Some birds might rely heavily on the intake

sulting in three diet categories: herbivore, omnivore, and inverti-

of insects during the breeding season, but switch to fruits during

vore. We ranked the models using Akaike's information criterion

the nonbreeding season. If food abundance is driving altitudinal

(AIC). We considered the top models competitive if they differed

migration, such a species may respond to insect abundance during

by <4 AIC units.

the breeding season and fruit during the nonbreeding season.
This situation likely reduced the effect of the patterns that we

3 | R E S U LT S

observed as we only considered the primary foraging guild (e.g.,
the main guild for each species depending on the distribution of
the percentage among the diet categories, according to Wilman

The best phylogenetic generalized least square model that pre-

et al., 2014).

dicted altitudinal migration included diet, region, an interaction

The regions that revealed an evolutionary association be-

between diet and region (Table 1 and Figure 2). The addition of

tween altitudinal migration and foraging guild were the Nearctic,

habitat as a predictor did not improve the model's AIC. However,

Palearctic, and Widespread. However, the foraging guilds asso-

habitat was still associated with altitudinal migration (F3 = 3.98,

ciated with altitudinal migration differed between these regions.

p = .0076; Figure 2b), with more altitudinal migrants in open habitat

In the Nearctic, herbivore and omnivore species were more likely

than dense habitat, water, and generalist. When we examined the

to be altitudinal migrants, a finding consistent with Boyle (2017).

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PAGEAU et al.

TA B L E 1 AIC results for each pgls
model. The models are ranked from best
to worst

F I G U R E 2 Mosaic plots representing
the proportion of passerine species that
are altitudinal migrant (black) or not (gray)
for each foraging guild (a), habitat (b), and
zoogeographical region when considering
only breeding distribution (c). The width
of the bars of the x-axis indicates the
proportion of species in each category

Rank

Model

DF

AIC

ΔAIC

1

Diet + Region + Diet:Region

22

7,880.136

0

2

Diet + Habitat + Region

15

7,961.286

81.15

3

Diet + Region

12

7,975.992

95.856

4

Habitat + Region

11

7,980.389

100.253

5

Region

8

7,984.467

104.331

6

Diet + Habitat + Diet:Habitat

13

8,086.023

205.887

7

Diet

6

8,093.052

212.916

8

Diet + Habitat

9

8,097.665

217.529

9

Habitat

5

8,105.545

225.409

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PAGEAU et al.

TA B L E 2 T-values for each variable included in the top-ranked
model Diet + Region + Diet:Region

2000; Guillaumet, Kuntz, Samuel, & Paxton, 2017; Katuwal et al.,
2016; Kimura et al., 2001; but see Barçante et al., 2017). However,
only 26 species are considered to have a widespread breeding

Value

Standard error

t-Value

p-Value

Intercept

0.077

0.53

0.14

.88

Omnivore

−0.0082

0.026

−0.32

.75

Herbivore

0.033

0.031

1.05

.29

Australasian

0.0091

0.033

0.27

.78

Indomalayan

0.021

0.026

0.80

.42

Nearctic

−0.069

0.054

−1.28

.20

model, providing no additional information beyond what diet and

Neotropical

−0.088

0.052

−1.70

.090

region already provided. The proportion of altitudinal migrants

Palearctic

−0.051

0.035

−1.45

.15

present in each habitat were extremely similar (12%–13%); and

Widespread

0.32

0.092

3.43

.0006

no habitat had a disproportionate number of altitudinal migrants.

Omnivore:
Australasian

0.032

0.043

0.74

.46

However, habitat was still significant in the model with habitat only,

Herbivore:
Australasian

−0.062

0.051

−1.21

.22

Omnivore:
Indomalayan

−0.040

0.035

−1.16

.24

in our analysis.

Herbivore:
Indomalayan

−0.013

0.040

−0.32

.75

of the variability in the expression of the behavior. For instance,

Omnivore:
Nearctic

0.40

0.054

7.43

<.0001

Herbivore:
Nearctic

0.27

0.061

4.43

<.0001

Omnivore:
Neotropical

−0.017

0.037

−0.47

.64

Herbivore:
Neotropical

0.054

0.045

1.19

.23

Omnivore:
Palearctic

0.19

0.058

3.26

.001

up or downslope to reach molting grounds (Rohwer et al., 2008;

Herbivore:
Palearctic

0.073

0.061

1.20

.23

difficult to generalize and categorize birds as altitudinal migrants.

Omnivore:
Widespread

0.13

0.26

0.50

.62

Herbivore:
Widespread

0.59

0.12

4.75

<.0001

distribution, so this interpretation should be taken with caution.
For the other regions (Neotropical, Indomalayan, Afrotropical, and
Australasian), foraging guild was not directly associated with altitudinal migration, potentially due to the vast complexity of tropical ecosystems.
Habitat was not associated with altitudinal migration in the top

with open habitats evolutionary associated with altitudinal migration. Thus, habitat may have played a role in the evolution of altitudinal migration, but foraging guild remains the main factor driving
Altitudinal migration is challenging to study in part because
some populations within the same species are altitudinal migrants
while the others are resident (Boyle, 2017; Green, Whitehorne,
Middleton, & Morrissey, 2015). There is also variation in the propensity to engage in altitudinal migration among individuals within
a population (Boyle, 2008b, 2017; Pratt et al., 2017; Rohwer et al.,
2008) and within individuals across time (Hahn et al., 2004). In addition, most studies focus on the importance of altitudinal migration
to birds moving to reach breeding grounds, but birds may also move
Wiegardt et al., 2017). As such, this variation makes it extremely
We suggest that more studies are needed about specifics of altitudinal migration encompassing species not yet studied and these
should begin to formalize distinctions between different types of
altitudinal migration (e.g., facultative, breeding, and molting) to better understand this behavior and the drivers behind it (sensu Tonra
& Reudink, 2018, formalization of molt-migration). Molting and
breeding are both energetically demanding and could both lead to

However, it is interesting that omnivorous species appear to be

strong selection for altitudinal movements. However, there are still

linked with altitudinal migration. This might support Chaves-

some major differences between molting and breeding and those

Campos (2004) and Levey (1988), who suggest that birds should

differences could be crucial in explaining the evolution of altitudinal

follow fruit abundance during the nonbreeding season and in-

migration.

sect abundance during the breeding season. Altitudinal migration

Another limitation in our study is the lack of information for

would then be beneficial for omnivorous species. Omnivorous

some regions (Barçante et al., 2017). We have confidence in the

species are also linked to altitudinal migration in the Palearctic.

Nearctic since it has been well sampled and documented; approxi-

Barçante et al. (2017) indicated that the proportion of frugivore/

mately 31% species are altitudinal migrant which is the highest pro-

nectarivore species that are altitudinal migrant in the Palearctic

portion within passerines (exception for Widespread). Otherwise,

was lower than expected; our results demonstrating a dispropor-

most studies in the Neotropics are concentrated in Costa Rica and

tionate number of omnivorous species agrees with their findings.

there is limited research on altitudinal migration in the Afrotropical,

For the species with a widespread distribution, herbivorous spe-

Indomalayan, and Australasian regions (Barçante et al., 2017). Even

cies were associated with altitudinal migration. This finding agrees

the Palearctic, which is rich on research in avifauna, lacks data on

with previous studies where herbivorous species have been indi-

altitudinal migration. This could mean either that altitudinal migra-

cated as altitudinal migrants all around the world (Blake & Loiselle,

tion is rare in the Palearctic or that it has not been studied in depth.

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PAGEAU et al.

The present study is the first to examine potential large-scale drivers of the evolution of altitudinal migration in passerines. Altitudinal
migration has evolved independently in different regions of the world
under the different environmental pressures coupled with varying life
history characteristics. Our results have reinforced the idea that diet
(foraging guild) played a major role in the evolution of altitudinal migration. However, the relationship between diet and altitudinal migration
is complex and varies across different regions in the world. Given the
prevalence of this behavior across foraging guilds, diet is clearly not the
only factor that drove the evolution of altitudinal migration, but rather
the evolution of this trait was likely driven by an ensemble of factors.
AC K N OW L E D G M E N T S
We would like to thank S. Joly for providing drawing of birds used
in Figure 1 and F. Pouw and V. Adams-Parsons with their assistance
with assembling the dataset. This research was funded by a Natural
Sciences and Engineering Research Council of Canada Discovery
Grant; a British Columbia Graduate Scholarship; a Master's research scholarship by the Fond de Recherche Nature et technologies, doctoral scholarship from Carlos Chagas Filho Foundation for
Research Support (FAPERJ E-26/100.811/2009), a National Council
for Scientific and Technological Development postdoctoral scholarship (CNPq 201297/2014-0), and productivity fellowships (CNPq
304309/2018-4 and CNPq-PQ 306.579/2018-9). This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal
de Nível Superior—Brasil (CAPES)—Finance Code 001. Additionally,
we thank a grant Research of FAPERJ (CNE E-26/202.835/2018).
C O N FL I C T O F I N T E R E S T S
None declared.
AU T H O R S C O N T R I B U T I O N
M.W. Reudink, M.M. Vale, and C. Pageau conceived the project. C.
Pageau wrote the manuscript, and all the authors edited it. C. Pageau,
M. Shaikh, and M.W. Reudink wrote the code and conducted the
statistical analysis. L. Barcante, M.A.S. Alves, M.A. de Menezes, and
M.M. Vale aggregated the data.
OPEN RESEARCH BADGES

This article has earned an Open Data Badge for making publicly
available the digitally-shareable data necessary to reproduce the
reported results. The data is available at https://doi.org/10.5061/
dryad.jwstq​jq5n.
DATA AVA I L A B I L I T Y S TAT E M E N T
Data are accessible on Dryad (https://doi.org/10.5061/dryad.jwstq​
jq5n).
ORCID
Claudie Pageau

https://orcid.org/0000-0003-0371-5602

Matthew W. Reudink

https://orcid.org/0000-0001-8956-5849

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birds: A review of the literature and a comprehensive list of species.
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How to cite this article: Pageau C, Vale MM, de Menezes MA,
et al. Evolution of altitudinal migration in passerines is linked
to diet. Ecol Evol. 2020;00:1–8. https://doi.org/10.1002/
ece3.6126