J. Field Ornithol. 89(3):193–206, 2018 DOI: 10.1111/jofo.

12256

Endemic birds of the Atlantic Forest: traits, conservation

status, and patterns of biodiversity
Mariana M. Vale,1,6 Luara Tourinho,2,3 Maria Lucia Lorini,4 Henrique Raj~ao,5 and
Marcos S. L. Figueiredo4
1
Departamento de Ecologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Ilha do Fund~ a o,
Rio de Janeiro, Rio de Janeiro C.P. 68020, Brazil
2
Programa de Po s Graduacß~a o em Ecologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro, Ilha do
Fund~a o, Rio de Janeiro, Rio de Janeiro C.P. 68020, Brazil
3
Programa de Po s-graduacß~a o em Ecologia e Evolucß~a o, Instituto de Biologia Roberto Alc^ a ntara Gomes, Universidade
do Estado do Rio de Janeiro, Rua S~a o Francisco Xavier, 524, Maracan~ a , Rio de Janeiro, Rio de Janeiro
CEP: 20550-900, Brazil
4
Departamento de Ci^e ncias Naturais, Instituto de Bioci^e ncias, Universidade Federal do Estado do Rio de Janeiro,
Av. Pasteur, 458, Urca, Rio de Janeiro, Rio de Janeiro CEP: 22290-040, Brazil
5
Departamento de Biologia, Pontifıcia Universidade Cato lica do Rio de Janeiro, Rua Marqu^e s de S~ a o Vicente 225,
Ga vea, Rio de Janeiro, Rio de Janeiro CEP 22451-900, Brazil
Received 4 April 2018; accepted 25 May 2018

ABSTRACT. The bird fauna of the Brazilian Atlantic Forest is exceptionally diverse and threatened, with
high levels of endemism. Available lists of the endemic birds of the Atlantic Forest were generated before
recent taxonomic revisions lumped or split species and before the recent increase in species occurrence records.
Our objective, therefore, was to compile a new list of the endemic birds of the Atlantic Forest, characterize
these species in terms of conservation status and natural history traits, and map remaining vegetation and
protected areas. We combined GIS analysis with a literature search to compile a list of endemic species and,
based on the phylogeny and distribution of these species, characterized areas in terms of species richness,
phylogenetic diversity, and endemism. We identified 223 species of birds endemic to the Atlantic Forest,
including 12 species not included in previous lists. In addition, 14 species included in previous lists were not
considered endemic, either because they occur outside the Atlantic Forest biome or because they are not
considered valid species. The typical Atlantic Forest endemic bird is a small forest-dependent invertivore. Of
the species on our list, 31% are considered threatened or extinct. Only ~ 34% of the spatial analysis units had
> 10% forest cover, and protected area coverage was consistently low (< 1%). In addition, we found spatial
incongruity among the different measures of biodiversity (species richness, relative phylogenetic diversity,
restricted-range species, and irreplaceability). Each of these measures provides information concerning different
aspects of biological diversity. However, regardless of which aspect(s) of biodiversity might be considered most
important, preservation of the remaining areas of remnant vegetation and further expansion of protected areas
are essential if we are to conserve the many endemic species of birds in the Atlantic Forest.

RESUMEN. Aves endemicas del Bosque Atlantico: caracterısticas, estatus de conservaci on y

patrones de biodiversidad
La avifauna del Bosque Atlantico de Brasil es excepcionalmente diversa y amenazada, con altos niveles de
endemismo. Los listados disponibles de especies endemicas al Bosque del Atlantico fueron generados antes de
las recientes revisiones taxonomicas que agregan o separan especies y antes del reciente incremento de especies
con registros de presencia. Por ello, nuestro objetivo fue compilar una nueva lista de especies endemicas de
aves del Bosque Atlantico, caracterizar el estatus de conservacion de estas especies, caracterısticas de su historia
natural, y mapear la vegetacion remanente y areas protegidas. Combinamos un analisis de SIG con b usquedas
de literatura para recopilar una lista de especies endemicas, y, basados en la filogenia y distribuci on de estas
especies, caracterizar areas en terminos de riqueza de especies, diversidad filogenetica y endemismo.
Identificamos 223 especies de aves endemicas el Bosque Atlantico, incluidas 12 especies que no habıan sido
incluidas en listados previos. Ademas, 14 especies incluidas en listados previos no fueron consideradas
endemicas, porque se encuentran fuera del bioma del Bosque Atlantico o porque no son consideradas especies
validas. La tıpica ave endemica del Bosque Atlantico es un peque~ no invertıvoro que depende de habitats
boscosos. De las especies en nuestra lista, 31% estan consideradas como endemicas o extintas. Solo un ~ 34%
de las unidades de analisis espacial tenıan > 10% de cobertura de bosque y la cobertura de areas protegidas fue
consistentemente baja (< 1%). Adicionalmente, encontramos incongruencia espacial entre las diferentes
medidas de biodiversidad (riqueza de especies, diversidad filogenetica relativa, especies de rangos restringidos e
irremplazabilidad). Cada una de estas metricas provee informacion respecto a diferentes aspectos de la
diversidad biologica. Sin embargo, independientemente de que aspectos de la biodiversidad puedan ser
6
Corresponding author. Email: mvale.eco@gmail.com
© 2018 Association of Field Ornithologists
193
194 M.M. Vale et al.
considerados mas importantes, la preservacion de las areas de vegetacion remanentes y una mayor expansion de
las areas protegidas son esenciales si nuestra intencion es conservar las muchas especies de aves endemicas del
Bosque Atlantico.
Key words: Biodiversity, Brazil, habitat conservation, phylogenetic diversity, spatial patterns
INTRODUCTION
The Brazilian Atlantic Forest is among the
most biodiverse and threatened biomes in the
world (Myers et al. 2000), and is well known
for its exceptionally high bird diversity and
endemism (Stattersfield et al. 1998, Myers
et al. 2000, Silva et al. 2004, Jenkins et al.
2015). The Atlantic Forest originally covered
~ 1200 million km2 along the Atlantic coast
of South America (Olson et al. 2001), pri-
marily in Brazil, but also extending into Para-
guay and Argentina. However, only ~ 12% of
the original forest cover remains, mostly in
small and isolated forest fragments (Ribeiro
et al. 2009). As a result, the Atlantic Forest
has a high concentration of threatened bird
species (Jenkins et al. 2015).
An important step in conserving biodiver-
sity is identifying the species endemic to a
biome. For example, the most effective means
of identifying conservation priorities among
Brazilian ecoregions involved determining the
number and identity of endemic species in
those ecoregions (Loyola et al. 2007). In
addition, a study of the endemic birds of the
Atlantic Forest was used to show that the
extent of deforestation accurately predicted
the threat of extinction (Brooks et al. 1999),
and to determine the threshold range size at
which forest-dependent species become threat-
ened with extinction (Harris and Pimm
2007).
Haffer (1974) was the first to attempt to
identify the endemic avifauna of the Atlantic
Forest region and compiled a list of 160 spe-
cies. Later, Parker et al. (1996) identified 202
species of birds found only in the Atlantic For-
est zoogeographic region of the Neotropics.
Other investigators have provided estimates of
181 endemic species (Brooks et al. 1999) and
218 endemic species (Bencke et al. 2006) of
birds in the Atlantic Forest. Since publication
of the last estimate in 2006, however, there
have been a number of taxonomic revisions
where species found in the Atlantic Forest have
either been lumped or split (e.g., Batalha-Filho
et al. 2013, Buzzetti et al. 2013, Barnett and
J. Field Ornithol.
Buzzetti 2014, Ferreira et al. 2017, Garcıa
et al. 2018). There has also been an increase in
species occurrence records associated with new
online databases such as WikiAves (http://
www.wikiaves.com.br/), a Brazilian citizen-
science project that compiles bird records with
associated photos and song recordings. These
changes in distributional maps and taxonomy
can impact the observed patterns of species
diversity (Blackburn and Gaston 1998, Black-
burn et al. 2004).
Most studies focusing on spatial patterns
of diversity are based on species richness,
which does not take into account species
ecological or evolutionary singularities
because one of the assumptions of these
indexes is that all species should be consid-
ered biologically equivalent (Magurran
2004). Because quantifying and interpreting
species richness is relatively easy, this biodi-
versity descriptor has been the main focus of
macroecological studies, and also the princi-
pal information used to determine conserva-
tion priorities (Cadotte and Davies 2010,
Zupan et al. 2014). Measures of phyloge-
netic diversity have been developed as a
promising way to account for differences
between species in terms of evolutionary
characteristics (Devictor et al. 2010, Pavoine
and Bonsall 2011). Unlike spatial patterns of
species richness, which has been a central
topic of ecology for many years (Hawkins
2001), patterns of phylogenetic diversity are
mostly unknown due to, until recently, the
lack of adequate data for the analyses (Vos-
kamp et al. 2017). Phylogenetic diversity is
expected to be positively related to species
richness (Morlon et al. 2011, Rodrigues
et al. 2011, Campos et al. 2017), but this
relationship can vary spatially (e.g.,
Gonzalez–Maya et al. 2016). Examination
and comparison of these patterns could pro-
vide new clues about the processes that
shaped the diversification of a biota (Figueir-
edo and Grelle 2018).
Phylogenetic diversity is a measure that can
provide information about the history of
diversification of a lineage (i.e., speciation
Vol. 89, No. 3 Endemic Birds of the Atlantic Forest
minus extinctions). For example, if the phylo-
genetic diversity of a community reflects the
number of lineages it contains, then commu-
nities with low phylogenetic diversity relative
to the number of species present apparently
results from a large radiation of a few lin-
eages. This suggests that the diversity in this
region has an autochthonous origin and that
other lineages of the same taxon were not
successful in colonizing it or became extinct
before they could diversify. Likewise, species-
rich regions with high phylogenetic diversity
may have many different phylogenetic lin-
eages, with colonization of multiple lineages
followed by diversification (Davies and Buck-
ley 2011, Fritz and Rahbek 2012).
Given the aforementioned taxonomic revi-
sions and sources of new information, our
primary objective was to compile an updated
list of birds endemic to the Atlantic Forest.
Additional objectives were to characterize
these species of birds in terms of their conser-
vation status and natural history traits, and
characterize areas of the Atlantic Forest based
on the presence of remnant vegetation and
protected areas, species richness, and phyloge-
netic diversity.
METHODS
Compiling a list of endemic bird
species. We combined GIS analysis with a
literature search to compile a list of bird species
endemic to the Atlantic Forest, focusing in the
geographic limits of the biome as the main cri-
terion, independently of the vegetation type
where species occurred. Therefore, the list is
not restricted to forest specialists. First, we used
the digital maps of BirdLife Interna-
tional (http://www.iucnredlist.org/technical-
documents/spatial-data), and created a list of
species that have at least 80% of their distribu-
tion within the original extent of the Atlantic
Forest biome, as defined by the ecoregions
(Olson et al. 2001, http://www.worldwildlife.
org/publications/terrestrial-ecoregions-of-the-
world) and the Brazilian Ministry of the
Environment (http://www.mma.gov.br/biomas/
mata-atlantica/mapa-da-area-de-aplicacao).
In addition to the maps, we also conducted
a non-systematic search of the literature for
newly described species. That preliminary list
was then compared to the lists of Atlantic
Forest endemics provided by Stotz et al.
195
(1996) and Bencke et al. (2006). Finally, we
examined occurrence records provided by two
standard online databases, GBIF (http://www.
gbif.org) and SpeciesLink (http://splink.cria.
org.br), plus WikiAves (http://www.wikiaves.
com.br/).
There are important differences between
the taxonomy used by BirdLife International
(BirdLife Taxonomic Checklist v7.0, http://
www.birdlife.org/datazone/info/taxonomy) and
the Brazilian Committee on Ornithological
Records (CBRO) (Piacentini et al. 2015). To
accommodate these differences, we compiled
two lists of endemics, one incorporating all
recent taxonomic splits by either BirdLife
International or CBRO, and a more conserva-
tive list that only included species recognized
by BirdLife International, the international
authority responsible for establishing the con-
servation status of birds and priorities under
the IUCN RedList.
Conservation, natural history, and
diversity of Atlantic Forest birds. We
characterized all Atlantic Forest endemic spe-
cies in terms of their natural history and con-
servation-relevant traits. Natural history traits
included foraging guild (Wilman et al. 2014),
body mass (Dunning 2008, and the Hand-
book of the Birds of the World Alive; https://
www.hbw.com/species), and conservation sta-
tus based on the International Union for
Conservation of Nature (IUCN 2018). Two
additional conservation-relevant traits we con-
sidered were geographic range size and forest-
dependency level (Birdlife Data Zone; http://
www.birdlife.org/datazone/species). We searched
additional literature when the information we
needed was not available from these sources.
For species on our conservative list (i.e.,
recognized by BirdLife International), we
aimed to describe the spatial patterns of spe-
cies richness, phylogenetic diversity, and
endemism. We projected the digital distribu-
tion maps from IUCN (2016) over an equal-
area hexagonal grid (~ 500-km² cells) that
encompassed the entire Atlantic Forest, and
obtained a list of all species present in each
cell. Phylogenetic information regarding these
species was extracted from a recently pub-
lished super tree (Jetz et al. 2012). Those spe-
cies included on the conservative list, but
absent from the supertree, were added based
on criteria proposed by Schipper et al.
(2008), i.e., species added to a polytypic
196 M.M. Vale et al.
genus were included at the most basal node
to the genus and, in the case of a species
added to a monotypic genus, the genus was
split into two or more species and the divi-
sion was set at half of the branch length.
To describe the spatial structuring of ende-
mic bird diversity in the Atlantic Forest, we
mapped the patterns of five measurements
estimated for each cell of the hexagonal grid,
including (1) species richness, (2) phyloge-
netic diversity (PD), estimated based on the
PD index (Faith 1992) that represents the
sum of all branch lengths of a phylogenetic
tree comprised of all species in a given cell,
(3) relative phylogenetic diversity (rPD),
which is the phylogenetic diversity after the
effect of richness is removed (Davies et al.
2007), (4) restricted-range richness, estimated
as the sum of the inverse of the geographic
range size of all species present in a cell
because species with restricted geographic
ranges contributed more to this index than
those with large geographic ranges, and
(5) irreplaceability, estimated as the average
of the inverse of the number of cells of all
species present in a cell, so cells with higher
irreplaceability values would have more spe-
cies found in a smaller number of cells. Irre-
placeability scores for each cell describe the
potential cost of losing unique faunas (Dulvy
et al. 2014), which, in our study, would be
unique endemic bird faunas. In addition, we
also estimated the percent cover of (6) rem-
nant vegetation and (7) protected areas for
each hexagonal cell. Remnant vegetation data
were obtained from the Atlantic Forest frag-
ment maps for 2014 (SOS Mata Atl^antica
and INPE 2015). Data summarizing the cur-
rent network of protected areas were retrieved
from online databases (accessed on November
2017), including the World Database on Pro-
tected Area (WDPA, http://protectedplanet.ne
t), Brazil’s Ministry of Environment Database
(http://mapas.mma.gov.br), SIMRPPN (http://
sistemas.icmbio.gov.br/simrppn/publico), and
the WWF-Brazil Protected Areas Observatory
(http://observatorio.wwf.org.br). We only con-
sidered protected areas that were established to
protect biodiversity, including categories I to
IV from IUCN PAs System, plus two addi-
tional categories of Brazilian Protected Area
System (SNUC), the Area of Relevant Ecologi-
cal Interest (ARIE) and the Private Natural
Heritage Reserve (RPPN).
J. Field Ornithol.
In addition, we mapped species richness
using the list of bird species endemic to
the Atlantic Forest compiled by Stotz et al.
(1996) because several authors have used
that list as a reference. We compared that
map with our updated map of species rich-
ness to verify whether the known spatial
pattern of endemism has changed due to
recent taxonomic reviews and newly
described species.
We used Pearson’s coefficient to assess the
correlation and spatial match between the
seven measurements estimated for each and
every cell of the hexagonal grid, including spe-
cies richness, restricted range species (weighted
endemism), irreplaceability, phylogenetic diver-
sity, relative phylogenetic diversity, remnant
vegetation coverage and protected area cover-
age. Phylogenetic diversity PD indices were
calculated using the package “picante” (Kembel
et al. 2010), and the other analyses were per-
formed using the basic package of the R envi-
ronment 3.4.3 (R Core team 2017).
RESULTS
We compiled a list of 223 species of birds
endemic to the Atlantic Forest (Table 1),
with 215 of these species recognized by Bird-
life International taxonomy (Table S1). Six
species were only on our liberal list, recog-
nized as valid species by CBRO, but not
Birdlife International (Table S1). These are
essentially subspecies that have been elevated
to the species level, including Pernambuco
Woodcreeper (Dendrocincla taunayi), Mar-
garetta’s Hermit (Phaethornis margarettae),
Atlantic Woodpecker (Piculus polyzonus),
Bahia Piculet (Picumnus exilis (exilis)), Per-
nambuco Piculet (Picumnus pernambucensis),
and Diamantina Tapaculo (Scytalopus diaman-
tinensis). In addition, several species on our
conservative list are not recognized by CBRO,
including Cryptic Treehunter (Cichlocolaptes
mazarbarnetti), Atlantic Black-breasted
Woodpecker (Celeus tinnunculus), Festive
Coquette (Lophornis chalybeus), and Northern
Surucua Trogon (Trogon aurantius).
Some species with small ranges occur in
the contact zone between the Atlantic Forest
and other biomes, e.g., Hooded Visorbearers
(Augastes lumachella) and Sincora Antwrens
(Formicivora grantsaui) at the border of the
Caatinga xeric shrublands, and Hyacinth
Vol. 89, No. 3 Endemic Birds of the Atlantic Forest
Table 1. Species of birds endemic to the Atlantic
Forest, following the taxonomy of the South Ameri-
can Classification Committee (SACC).
TINAMIFORMES
Tinamidae
Tinamus solitarius (Solitary Tinamou)
GALLIFORMES
Cracidae
Pipile jacutinga (Black-fronted Piping-guan)
Ortalis araucuan (East Brazilian Chachalaca)
Ortalis squamata (Scaled Chachalaca)
Crax blumenbachii (Red-billed Curassow)
Mitu mitu (Alagoas Curassow)
GALLIFORMES
Odontophoridae
Odontophorus capueira (Spot-winged Wood-
Quail)
COLUMBIFORMES
Columbidae
Claravis geoffroyi (Purple-winged Ground-dove)
CAPRIMULGIFORMES
Caprimulgidae
Macropsalis forcipata (Long-trained Nightjar)
APODIFORMES
Trochilidae
Ramphodon naevius (Saw-billed Hermit)
Glaucis dohrnii (Hook-billed Hermit)
Phaethornis squalidus (Dusky-throated
Hermit)
Phaethornis idaliae (Minute Hermit)
Phaethornis eurynome (Scale-throated Hermit)
Phaethornis margarettae (Margaretta’s Hermit)
Lophornis chalybeus (Festive Coquette)
Clytolaema rubricauda (Brazilian Ruby)
Stephanoxis lalandi (Green-crowned
Plovercrest)
Stephanoxis loddigesii (Violet-crowned
Plovercrest)
Thalurania watertonii (Long-tailed
Woodnymph)
Thalurania glaucopis (Violet-capped
Woodnymph)
GRUIFORMES
Rallidae
Aramides saracura (Slaty-breasted Wood-rail)
ACCIPITRIFORMES
Accipitridae
Leptodon forbesi (White-collared Kite)
Buteogallus lacernulatus (White-necked Hawk)
Pseudastur polionotus (Mantled Hawk)
STRIGIFORMES
Strigidae
Megascops sanctaecatarinae (Long-tufted
Screech-Owl)
Megascops atricapilla (Black-capped Screech-
Owl)
Pulsatrix koeniswaldiana (Tawny-browed
Owl)
197
Strix hylophila (Rusty-barred Owl)
Glaucidium mooreorum (Pernambuco Pygmy-
Owl)
Glaucidium minutissimum (Least Pygmy-Owl)
TROGONIFORMES
Trogonidae
Trogon aurantius (Northern Surucua Trogon)
CORACIIFORMES
Momotidae
Baryphthengus ruficapillus (Rufous-capped
Motmot)
GALBULIFORMES
Galbulidae
Jacamaralcyon tridactyla (Three-toed Jacamar)
Bucconidae
Notharchus swainsoni (Buff-bellied Puffbird)
Malacoptila striata (Crescent-chested Puffbird)
PICIFORMES
Ramphastidae
Ramphastos dicolorus (Red-breasted Toucan)
Selenidera maculirostris (Spot-billed Toucanet)
Pteroglossus bailloni (Saffron Toucanet)
Picidae
Picumnus exilis (Golden-spangled Piculet)
Picumnus pernambucensis (Pernambuco
Piculet)
Picumnus temminckii (Ochre-collared Piculet)
Melanerpes flavifrons (Yellow-fronted
Woodpecker)
Veniliornis maculifrons (Yellow-eared
Woodpecker)
Campephilus robustus (Robust Woodpecker)
Celeus tinnunculus (Atlantic Black-breasted
Woodpecker)
Hylatomus galeatus (Helmeted Woodpecker)
Piculus polyzonus (Golden-green Woodpecker)
Piculus aurulentus (Yellow-browed
Woodpecker)
PSITTACIFORMES
Psittacidae
Touit melanonotus (Brown-backed Parrotlet)
Touit surdus (Golden-tailed Parrotlet)
Brotogeris tirica (Plain Parakeet)
Pionopsitta pileata (Pileated Parrot)
Triclaria malachitacea (Blue-bellied Parrot)
Pionus reichenowi (Blue-breasted Parrot)
Amazona vinacea (Vinaceous-breasted
Amazon)
Amazona pretrei (Red-spectacled Amazon)
Amazona rhodocorytha (Red-browed Amazon)
Amazona brasiliensis (Red-tailed Amazon)
Pyrrhura cruentata (Ochre-marked Parakeet)
Pyrrhura frontalis (Maroon-bellied Parakeet)
Pyrrhura griseipectus (Grey-breasted Parakeet)
Pyrrhura leucotis (White-eared Parakeet)
PASSERIFORMES
Thamnophilidae
Hypoedaleus guttatus (Spot-backed Antshrike)
198 M.M. Vale et al.
Mackenziaena leachii (Large-tailed Antshrike)
Mackenziaena severa (Tufted Antshrike)
Biatas nigropectus (White-bearded Antshrike)
Thamnophilus ambiguous (Sooretama Slaty-
antshrike)
Rhopias gularis (Star-throated Antwren)
Dysithamnus stictothorax (Spot-breasted
Antvireo)
Dysithamnus xanthopterus (Rufous-backed
Antvireo)
Dysithamnus plumbeus (Plumbeous Antvireo)
Myrmotherula luctuosa (Silvery-flanked
Antwren)
Myrmotherula minor (Salvadori’s Antwren)
Myrmotherula urosticta (Band-tailed Antwren)
Myrmotherula unicolor (Unicolored Antwren)
Myrmotherula snowi (Alagoas Antwren)
Herpsilochmus pileatus (Bahia Antwren)
Formicivora erythronotos (Black-hooded
Antwren)
Formicivora serrana (Serra Antwren)
Formicivora paludicola (Marsh Antwren)
Formicivora acutirostris (Parana Antwren)
Drymophila ferruginea (Ferruginous Antbird)
Drymophila rubricollis (Bertoni’s Antbird)
Drymophila genei (Rufous-tailed Antbird)
Drymophila ochropyga (Ochre-rumped
Antbird)
Drymophila malura (Dusky-tailed Antbird)
Drymophila squamata (Scaled Antbird)
Terenura sicki (Orange-bellied Antwren)
Terenura maculata (Streak-capped Antwren)
Cercomacra brasiliana (Rio de Janeiro
Antbird)
Pyriglena atra (Fringe-backed Fire-eye)
Pyriglena leucoptera (White-shouldered Fire-
eye)
Rhopornis ardesiacus (Slender Antbird)
Myrmoderus ruficauda (Scalloped Antbird)
Myrmoderus loricatus (White-bibbed Antbird)
Myrmoderus squamosus (Squamate Antbird)
Conopophagidae
Conopophaga melanops (Black-cheeked
Gnateater)
Conopophaga cearae (Ceara Gnateater)
Grallariidae
Hylopezus nattereri (Speckle-breasted Antpitta)
Rhinocryptidae
Psilorhamphus guttatus (Spotted Bamboowren)
Merulaxis ater (Slaty Bristlefront)
Merulaxis stresemanni (Stresemann’s
Bristlefront)
Eleoscytalopus psychopompus (Bahia Tapaculo)
Eleoscytalopus indigoticus (White-breasted
Tapaculo)
Scytalopus gonzagai (Boa Nova Tapaculo)
Scytalopus speluncae (Mouse-colored
Tapaculo)
J. Field Ornithol.
Scytalopus diamantinenis (Diamantina
Tapaculo)
Scytalopus petrophilus (Rock Tapaculo)
Scytalopus pachecoi (Planalto Tapaculo)
Scytalopus iraiensis (Marsh Tapaculo)
Formicariidae
Chamaeza meruloides (Cryptic Antthrush)
Chamaeza ruficauda (Rufous-tailed
Antthrush)
Furnariidae
Sclerurus cearensis (Ceara Leaftosser)
Sclerurus scansor (Rufous-breasted Leaftosser)
Dendrocincla taunayi (Pernambuco
Woodcreeper)
Dendrocincla turdina (Plain-winged
Woodcreeper)
Xyphorhynchus atlanticus (Atlantic
Woodcreeper)
Xiphorhynchus fuscus (Lesser Woodcreeper)
Campylorhamphus falcularius (Black-billed
Scythebill)
Lepidocolaptes squamatus (Scaled
Woodcreeper)
Lepidocolaptes falcinellus (Scalloped
Woodcreeper)
Cinclodes pabsti (Long-tailed Cinclodes)
Anabazenops fuscus (White-collared Foliage-
gleaner)
Cichlocolaptes mazarbarnetti (Cryptic
Treehunter)
Cichlocolaptes leucophrus (Pale-browed
Treehunter)
Heliobletus contaminates (Sharp-billed
Treehunter)
Philydor novaesi (Alagoas Foliage-gleaner)
Philydor atricapillus (Black-capped Foliage-
gleaner)
Anabacerthia amaurotis (White-browed
Foliage-gleaner)
Anabacerthia lichtensteini (Ochre-breasted
Foliage-gleaner)
Clibanornis dendrocolaptoides (Canebrake
Groundcreeper)
Automolus lammi (Pernambuco Foliage-
gleaner)
Automolus leucophthalmus (White-eyed
Foliage-gleaner)
Leptasthenura striolata (Striolated Tit-
spinetail)
Leptasthenura setaria (Araucaria Tit-spinetail)
Phacellodomus erythrophthalmus (Orange-eyed
Thornbird)
Phacellodomus ferrugineigula (Orange-breasted
Thornbird)
Asthenes moreirae (Itatiaia Spinetail)
Acrobatornis fonsecai (Pink-legged Graveteiro)
Thripophaga macroura (Striated Softtail)
Cranioleuca obsoleta (Olive Spinetail)
Vol. 89, No. 3 Endemic Birds of the Atlantic Forest
Cranioleuca pallida (Pallid Spinetail)
Synallaxis cinerea (Bahia Spinetail)
Synallaxis infuscate (Pinto’s Spinetail)
Synallaxis ruficapilla (Rufous-capped Spinetail)
Tyrannidae
Phyllomyias virescens (Greenish Tyrannulet)
Phyllomyias griseocapilla (Grey-capped
Tyrannulet)
Phylloscartes kronei (Restinga Tyrannulet)
Phylloscartes beckeri (Bahia Tyrannulet)
Phylloscartes ceciliae (Alagoas Tyrannulet)
Phylloscartes paulista (Sao Paulo Tyrannulet)
Phylloscartes oustaleti (Oustalet’s Tyrannulet)
Phylloscartes difficilis (Serra do Mar
Tyrannulet)
Phylloscartes sylviolus (Bay-ringed Tyrannulet)
Myiornis auricularis (Eared Pygmy-Tyrant)
Hemitriccus diops (Drab-breasted Pygmy-
Tyrant)
Hemitriccus obsoletus (Brown-breasted Pygmy-
Tyrant)
Hemitriccus orbitatus (Eye-ringed Tody-
Tyrant)
Hemitriccus nidipendulus (Hangnest Tody-
Tyrant)
Hemitriccus mirandae (Buff-breasted Tody-
Tyrant)
Hemitriccus kaempferi (Kaempfer’s Tody-
Tyrant)
Hemitriccus furcatus (Fork-tailed Tody-Tyrant)
Todirostrum poliocephalum (Yellow-lored
Tody-Flycatcher)
Calyptura cristata (Kinglet Calyptura)
Platyrinchus leucoryphus (Russet-winged
Spadebill)
Onychorhynchus swainsoni (Atlantic Royal
Flycatcher)
Knipolegus nigerrimus (Velvet Black-Tyrant)
Muscipipra vetula (Shear-tailed Grey Tyrant)
Attila rufus (Grey-hooded Attila)
Piprites pileata (Black-capped Piprites)
Cotingidae
Carpornis cucullata (Hooded Berryeater)
Carpornis melanocephala (Black-headed
Berryeater)
Phibalura flavirostris (Swallow-tailed Cotinga)
Cotinga maculata (Banded Cotinga)
Lipaugus lanioides (Cinnamon-vented Piha)
Lipaugus ater (Black-and-gold Cotinga)
Lipaugus conditus (Grey-winged Cotinga)
Procnias nudicollis (Bare-throated Bellbird)
Xipholena atropurpurea (White-winged Cotinga)
Pipridae
Neopelma aurifrons (Wied’s Tyrant-Manakin)
Neopelma chrysolophum (Serra do Mar
Tyrant-Manakin)
Antilophia bokermanni (Araripe Manakin)
Chiroxiphia caudata (Blue Manakin)
199
Ilicura militaris (Pin-tailed Manakin)
Machaeropterus regulus (Eastern Striped
Manakin)
Tityridae
Schiffornis virescens (Greenish Schiffornis)
Iodopleura pipra (Buff-throated Purpletuft)
Laniisoma elegans (Shrike-like Cotinga)
Vireonidae
Hylophilus poicilotis (Rufous-crowned
Greenlet)
Corvidae
Cyanocorax coeruleus (Azure Jay)
Polioptilidae
Polioptila lactea (Creamy-bellied Gnatcatcher)
Thraupidae
Nemosia rourei (Cherry-throated Tanager)
Orchesticus abeillei (Brown Tanager)
Hemithraupis ruficapilla (Rufous-headed
Tanager)
Haplospiza unicolor (Uniform Finch)
Tachyphonus coronatus (Ruby-crowned
Tanager)
Ramphocelus bresilius (Brazilian Tanager)
Dacnis nigripes (Black-legged Dacnis)
Sporophila falcirostris (Temminck’s Seedeater)
Sporophila frontalis (Buffy-fronted Seedeater)
Saltator maxillosus (Thick-billed Saltator)
Saltator fuliginosus (Black-throated Grosbeak)
Castanozoster thoracicus (Bay-chested
Warbling-Finch)
Thlypopsis pyrrhocoma (Chestnut-headed
Tanager)
Microspingus lateralis (Buff-breasted
Warbling-Finch)
Tangara cyanoptera (Black-headed Tanager)
Tangara brasiliensis (White-bellied Tanager)
Tangara peruviana (Black-backed Tanager)
Tangara cyanomelas (Silvery-breasted Tanager)
Tangara seledon (Green-headed Tanager)
Tangara fastuosa (Seven-colored Tanager)
Tangara cyanocephala (Red-necked Tanager)
Tangara desmaresti (Brassy-breasted Tanager)
Tangara cyanoventris (Gilt-edged Tanager)
Tangara ornata (Golden-chevroned Tanager)
Mitrospingidae
Orthogonys chloricterus (Olive-green Tanager)
Passerellidae
Arremon semitorquatus (Half-collared Sparrow)
Icteridae
Anumara forbesi (Forbes’s Blackbird)
Fringillidae
Euphonia chalybea (Green-throated Euphonia)
Euphonia pectoralis (Chestnut-bellied
Euphonia)
Tyrannidae
Pogonotriccus eximius (Southern Bristle-
Tyrant)
Mionectes rufiventris (Grey-hooded Flycatcher)
200 M.M. Vale et al. J. Field Ornithol.

Fig. 1. Habitats and foraging guilds of birds endemic to the Atlantic Forest. (A) Dependence on forest
habitat, and (B) Foraging guild. Invert = invertivores (consume mainly invertebrates, including insects),
Frug/Nect = frugivores/nectarivores (consume mainly fruits and/or nectar), Omniv = omnivores (con-
sume both animal and plant matter), Plant = plantivores (consume primarily plant material other than
fruit or nectar, e.g., granivores and folivores), and Carniv = carnivores (consume mainly vertebrates and/
or carrion).

Fig. 2. Conservation status, body mass, and geographic ranges of birds endemic to the Atlantic Forest.
(A) Number of species, (B) Mean body mass, and (C) Box plot showing median geographic range sizes
of species in each conservation-status category, with circles indicating outliers. Conservation status: LC,
Least Concern; NT, Near Threatened; VU, Vulnerable; EN, Endangered; CR, Critically Endangered,
and EX, Extinct or Extinct in the wild.

Visorbearers (Augastes scutatus), Cipo Canas-
teros (Asthenes luizae), Cipo Cinclodes (Cin-
clodes espinhacensis), Pale-throated Pampa-
finches (Embernagra longicauda), and Dia-
mantina Tapaculos (Scytalopus diamantinensis)
at the border of the Cerrado savannas. In
such cases, the decision about whether to
include or not was based on the preferred
habitat of each species. For example, Hya-
cinth Visorbearers, Hooded Visorbearers, and
Sincora Antwrens are all associated with xero-
phitic vegetation in mountain regions and,
therefore, were not included (Handbook
of the Birds of the World Alive; https://www.
hbw.com/species). Diamantina Tapaculos,
however, inhabit forests (Bornschein et al.
2007) and were included.
Most species of birds endemic to the Atlan-
tic Forest are small (N = 217, X = 103.2 g,
median = 20.5 g, S = 342.3 g) forest-
dependent invertivores (Fig. 1, Table S1).
The IUCN recognizes 31% of the Atlantic
Forest endemic birds as threatened or extinct
and another 19% as near threatened. Threat-
ened or extinct species tend to be larger and
have smaller geographic ranges (Fig. 2).
Except for richness and phylogenetic diver-
sity, which were highly correlated (r = 0.98,
P < 0.001), biodiversity measurements
revealed a spatial mismatch (Table S2,
Fig. 3). The spatial pattern of diversity of
birds endemic to the Atlantic Forest, as mea-
sured by species richness and phylogenetic
diversity, followed a well-known pattern of
greater diversity at the core of the biome at
the Serra do Mar coastal forest ecoregion,
with a gradual decrease toward its extremes
(Fig. 3A). The spatial pattern of species rich-
ness did not change when using the species
list compiled by Stotz et al. (1996) (Fig. S1).
Vol. 89, No. 3 Endemic Birds of the Atlantic Forest 201

Fig. 3. Distribution patterns of remnant vegetation, protected areas, and bird species endemic to the Atlantic
Forest. For all figures: dark gray color = areas not analyzed, light gray = 0, remaining color coding varies from
dark blue (minimum value) to red (maximum value). (A) Remnant vegetation area (minimum = 0.0001, maxi-
mum = 97.18), (B) Protected area (minimum = 0.0001, maximum = 100), (C) Species richness (mini-
mum = 1, maximum = 149), (D) Relative phylogenetic diversity (minimum = 3.0, maximum = 1.9), (E)
Restricted-range weighted richness (minimum = 0.0001, maximum = 0.35), and (F) Irreplaceability (mini-
mum = 0.0001, maximum = 0.11). [Color figure can be viewed at wileyonlinelibrary.com]
202 M.M. Vale et al.
Once the effect of species richness over phylo-
genetic diversity was removed, the rPD
revealed a different spatial pattern, with the
southeastern Brazilian mountain ranges hav-
ing lower values of phylogenetic diversity than
expected based on the observed species rich-
ness, and with the highest values at the
Parana and Paulista plateaus and the southern
Bahia (Fig. 3B).
Restricted-range species were more concen-
trated in the southeastern and southern
Brazilian mountain ranges along the Atlantic
coast, but with some found in the northeast-
ern region. In contrast, bird communities in
the interior forests consisted primarily of
wide-ranging species (Fig. 3C). Irreplaceabil-
ity presented a somewhat different pattern,
with the highest values in the northeastern
Atlantic Forest and intermediate values in the
southeastern Brazilian mountain ranges and
along the Bahia coastal forests. The lowest
values of irreplaceability were in the interior
Atlantic Forest (Fig. 3D).
Only ~ 34% of grid cells had remnant veg-
etation coverage greater than 10%. Highest
values of vegetation remnant coverage
(> 50%) were largely restricted to southeast-
ern and southern Brazilian mountain ranges
and the Atlantic coast, whereas the interior
and northeastern Atlantic Forest had the low-
est values (< 25%; Fig. 3). Protected-area
coverage was consistently low (< 1%) in most
cells, especially in the northeastern Atlantic
Forest (Fig. 3). Values of protected areas
> 25% were limited to a few cells in the
southeastern and southern Brazilian mountain
ranges and the interior Atlantic Forest
(Fig. 3).
DISCUSSION
Atlantic Forest endemic bird listing and
conservation. Our compilation of 223
Atlantic Forest endemics includes some new
species and excludes a number of species pre-
viously considered endemic. Our list includes
12 species not listed by either Stotz et al.
(1996) or Bencke et al. (2006), including the
Araripe Manakin (Antilophia bokermanni),
Atlantic Black-breasted Woodpecker (Celeus
tinnunculus), Cryptic Treehunter (Cichloco-
laptes mazarbarnetti), Pernambuco Wood-
creeper (Dendrocincla taunayi), Festive
Coquette (Lophornis chalybeus), Swallow-tailed
J. Field Ornithol.
Cotinga (Phibalura flavirostris), Blue-breasted
Parrot (Pionus reichenowi), Buff-breasted War-
bling-finch (Microspingus lateralis), Boa Nova
Tapaculo (Scytalopus gonzagai), Northern
Surucua Trogon (Trogon aurantius), Marsh
Antwren (Formicivora paludicola), and Rock
Tapaculo (Scytalopus petrophilus).
Many new species on our list were due to
taxonomic reviews that elevated subspecies
endemic to the Atlantic Forest to the species
level, but some are newly described species,
such as the Rock Tapaculo, Marsh Antwren,
and Cryptic Treehunter (Whitney et al.
2010, Buzzetti et al. 2013, Barnett and Buz-
zetti 2014), with the latter already considered
Critically Endangered (BirdLife International
[online] 2017a,b). In addition, we did not
consider 14 species listed by Stotz et al.
(1996) and/or Bencke et al. (2006) as ende-
mic because their occurrence records revealed
distributions well outside the Atlantic Forest
biome, including the Sombre Hummingbird
(Aphantochroa cirrochloris), Yellow-legged
Tinamou (Crypturellus noctivagus), Black Jaco-
bin (Florisuga fusca), Southern Surucua Tro-
gon (Trogon surrucura), Eastern Slaty Thrush
(Turdus subalaris), White-rimmed Warbler
(Myiothlypis leucoblephara), White-throated
Hummingbird (Leucochloris albicollis),
Dubois’s Seedeater (Sporophila ardesiaca),
Red-ruffed Fruitcrow (Pyroderus scutatus),
White-spotted Woodpecker (Veniliornis spilo-
gaster), Blue Seedeater (Amaurospiza moesta),
Black-bellied Seedeater (Sporophila melanoga-
ster), and two species found on Fernando de
Noronha oceanic islands (Noronha Elaenia,
Elaenia ridleyana, and Noronha Vireo, Vireo
gracilirostris). The number of Atlantic forest
endemics might increase considerably if taxo-
nomic reviews reveal that some subspecies
endemic to the Atlantic Forest deserve to be
elevated to species status, such as Pyroderus
scutatus scutatus, Conopophaga lineata vulgaris,
Knipolegus nigerrimus nigerrimus, and Cryp-
turellus noctivagus noctivagus.
Threatened and extinct species tend to be
larger in size and have smaller geographic
ranges (Gaston and Blackburn 1995, Harris
and Pimm 2007, Lee and Jetz 2010), a pat-
tern confirmed for Atlantic Forest endemic
birds. About one-third of the endemics are
already either threatened or extinct according
to the IUCN Redlist. This estimate, however,
is likely conservative. One species, the Alagoas
Vol. 89, No. 3 Endemic Birds of the Atlantic Forest
Curassow (Mitu mitu), is considered extinct
in the wild by the International Union for
Conservation of Nature (IUCN), and three
other species considered Critically Endangered
by IUCN are already considered Extinct by
Brazil’s Ministry of Environment and other
authorities (Cryptic Treehunter, Cichlocolaptes
mazarbarnetti; Alagoas Foliage-gleaner, Phily-
dor novaesi; and Pernambuco Pygmy-owl,
Glaucidium mooreorum) (Brasil 2014, Pereira
et al. 2014). In addition, the conservation sta-
tus of the seven species that are not recog-
nized as valid species by BirdLife
International is unknown. Many of them,
however, have small geographic ranges and
are endemic to the northern portion of the
Atlantic Forest that is particularly deforested,
fragmented, and unprotected (Ribeiro et al.
2009, Fig. 3). Not surprisingly, four of these
species are already on the official Brazilian list
of threatened species, including Pernambuco
Woodcreeper (Dendrocincla taunayi), Mar-
garetta’s Hermit (Phaethornis margarettae),
Atlantic Woodpecker (Piculus polyzonus), and
Diamantina Tapaculo (Scytalopus diamanti-
nensis) (Brasil 2014). On the brighter side, 67
species including in our list are included in
the National Plan for the Conservation of
Atlantic Forest Birds (Table S1, MMA 2018).
Biodiversity patterns and threats. The
pattern of greater species richness and phylo-
genetic diversity in the southeastern Atlantic
Forest has previously been shown for birds
(Davies et al. 2007, Souza et al. 2011, Jenk-
ins et al. 2015, Vale et al. 2018), amphibians
(Fritz and Rahbek 2012, Jenkins et al. 2015,
Campos et al. 2017), and marsupials (Fig-
ueiredo and Grelle 2018). This pattern may
be due to the high topographic and climatic
variability (Fine 2015) in the Serra do Mar
region or may just be an artifact caused by
geometric constraints, the mid-domain effect
(Colwell and Lees 2000).
Relative phylogenetic diversity increased
toward the Parana and Paulista plateaus and
the southern Bahia, and was lower in the
southeastern mountain ranges. Low values of
relative phylogenetic diversity apparently
result from the large radiation of a few lin-
eages and have previously been reported in
mountainous regions (Fritz and Rahbek
2012, Voskamp et al. 2017, Figueiredo and
Grelle 2018), due to multiple cases of vicari-
ance in these regions (Davies and Buckley
203
2011). High values of relative phylogenetic
diversity, on the other hand, suggest the pres-
ence of phylogenetic lineages of various ori-
gins and are usually found in contact zones
among ecoregions with different species pools
(Davies and Buckley 2011). As such, it is not
surprising that its highest values are observed
in contact zones with Pantanal and Cerrado
(Parana and Paulista plateaus), or in southern
Bahia, where the avifaunas of the southern
and northern Atlantic Forest meets.
Of the five biodiversity measurements we
examined, irreplaceability is the most worri-
some. The area with the most exclusive set of
species – the northeastern Atlantic Forest – is
also the area most affected by human impacts
(Ribeiro et al. 2009). This region also had
the lowest coverage of remnant vegetation
and protected areas. The northeastern Atlantic
Forest is a center of endemism for birds in
the biome (Silva et al. 2004) and has a large
number of threatened species (Pereira et al.
2014). Many species of birds found in the
northeastern Atlantic Forest do not occur
below the S~ao Francisco River and, therefore,
loss of forest cover in this region could lead
to the disappearance of many species not
found elsewhere; a great amount of diversity
will be lost even if related phylogenetic lin-
eages remain in other parts of the Atlantic
Forest. Indeed, three species officially consid-
ered as globally extinct by Brazil’s Ministry of
Environment (Brasil 2014) – Cryptic Tree-
hunter (Cichlocolaptes mazarbarnetti), Alagoas
Foliage-gleaner (Philydor novaesi), and Per-
nambuco Pygmy-owl (Glaucidium mooreo-
rum) (Pereira et al. 2014) – previously
occurred in the northeastern Atlantic Forest.
The spatial incongruence among the four
measures of biodiversity in our study (bird
richness, relative phylogenetic diversity,
restricted-range species, and irreplaceability)
corroborates and extends the results of previ-
ous studies and highlights the importance of
considering multiple measures of biodiversity
when formulating conservation strategies
because each of these measures provides
valuable information regarding either evolu-
tionary (phylogenetic diversity) or biogeo-
graphical (weighted endemism and
irreplaceability) processes. Biological diversity
is a complex and multifaceted phenomenon,
and different biodiversity measures capture
different aspects of this diversity (Magurran
204 M.M. Vale et al.
2004). However, regardless of which aspect
(s) might be considered most important, our
results clearly indicate that preservation of
the remaining areas of remnant vegetation
and further expansion of protected areas are
essential if we are to conserve the many
endemic species of birds in the Atlantic For-
est. This is particularly the case for the
northeastern Atlantic Forest, the area with
the least coverage of remnant vegetation and
fewest protected areas, but the center of
endemism for birds in the biome. Any fur-
ther loss of forest cover in this region would
likely lead to the extinction of many of the
endemic species of birds found in the Atlan-
tic Forest.
ACKNOWLEDGMENTS
We thank Thiago V. Souza for analytical help, J.
Niemeyer for help in trait compilation, and G. Ritchi-
son and two anonymous reviewers for valuable sugges-
tions for improving our manuscript. This paper is a
contribution of the Brazilian Network on Global Cli-
mate Change Research (Rede CLIMA) and the
National Institutes for Science and Technology in
Ecology, Evolution and Conservation of Biodiversity
(INCT EECBio). We thank the Coordination of Supe-
rior Level Staff Improvement (CAPES) for graduate
fellowships to LT and the National Council for Scien-
tific and Technological Development (CNPq) for a
postdoctoral fellowship to MSLF.
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SUPPORTING INFORMATION
Additional Supporting Information may be
found in the online version of this article at
the publisher’s web-site.
Fig. S1. Richness of bird species endemic
to the Atlantic Forest. (A) Richness using the
species list compiled in this study (maximum
of 148 species per hexagon), (B) richness
using species list by Stotz et al. (1996) (maxi-
mum of 128 species per hexagon), and
(C) difference in richness between maps.
Table S1. Species database.
Table S2. Pearson’s correlation between
biodiversity measurements, coverage of pro-
tected area, and vegetation remnant area
(P < 0.001).