Global Ecology and Biogeography, (Global Ecol. Biogeogr.

) (2004) 13, 85 –92

Areas of endemism for passerine birds in

Blackwell Publishing, Ltd.

RESEARCH
PAPER
the Atlantic forest, South America
José Maria Cardoso da Silva*†‡, Marcelo Cardoso de Sousa†§ and
Carlos H. M. Castelletti†

†Universidade Federal de Pernambuco, Centro ABSTRACT

de Ciências Biológicas, Departamento de
Zoologia, Recife, Pernambuco, Brazil. E-mail:
j.silva@conservation.org.br, ‡Conservation Inter-
national do Brazil, Avenida Nazaré 541/Sala
310, 66035 –170, Belém, Pará, Brazil and
§Universidade Tiradentes, Centro de Ciências
Biológicas e da Saúde, Avenida Murilo Dantas
300, 49032– 490, Aracaju, Sergipe, Brazil
*Corresponding author
Aim To use the method of parsimony analysis of endemism to identify areas of
endemism for passerine birds in the Atlantic Forest, South America, and to compare
the locations of these areas with areas previously identified for birds as well as other taxa.
Location The Atlantic Forest, eastern South America.
Methods We analysed a matrix composed of the presence (1) or absence (0) of 140
endemic species in 24 quadrats of 1 × 1 degree distributed along the Atlantic Forest
to find the most parsimonious area cladogram.
Results Fourteen most parsimonious cladograms were found and then summarized
in a single consensus tree. Four areas of endemism were identified: Pernambuco,
Central Bahia, Coastal Bahia, and Serra do Mar.
Main conclusions Avian areas of endemism in the Atlantic Forest have significant
generality, as they are highly nonrandom and congruent with those of other groups
of organisms. A first hypothesis about the historical relationships among the four
areas of avian endemism in the Atlantic Forest is delineated. There is a basal dichot-
omy among areas of endemism in the Atlantic Forest, with Pernambuco forming a
northern cluster and Coastal Bahia, Central Bahia and Serra do Mar comprising a
southern cluster. Within the southern cluster, Central Bahia and Serra do Mar are
more closely related to each other than to Coastal Bahia.
Keywords
Areas of endemism, Atlantic Forest, biogeography, Brazil, evolution, Neotropics,
passerine birds, phylogeny.

endemism harbour unique biotas and therefore are considered

INTRODUCTION
Two basic patterns of geographical distribution are recognized in
biogeography (Brown & Lomolino, 1998). The first is that geo-
graphical distributions of organisms are limited by ecological or
historical factors; thus no taxon is completely cosmopolitan but
many are endemic to restricted regions. The second pattern is
that endemic species are not distributed randomly but they tend
to be concentrated in some regions of the world, constituting a
phenomenon called provincialism.
Regions with at least two endemic taxa are named areas of
endemism (Cracraft, 1985; Platnick, 1991). They are important
for two reasons. The first is that they represent the smallest geo-
graphical units of analysis in historical biogeography, forming
the base for postulating hypotheses about the history of geo-
graphical units and their biotas (Cracraft, 1985, 1994; Morrone,
1994; Morrone & Crisci, 1995). The second is that areas of
as priority targets for conservation action (Terborgh & Winter,
1982; Fjeldså, 1993).
There are several methods used to identify areas of endemism
at a continental scale. The method more commonly used consists
of superimposing distribution maps for a number of taxa to
identify areas characterized by high concentrations of overlap-
ping ranges (Müller, 1973). However, the overlap among species’
ranges may not be perfect and arbitrary decisions are taken when
a great number of species are analysed (Platnick, 1991; Silva &
Oren, 1996).
Recently, a new method was proposed to identify areas of
endemism (Morrone, 1994). The method applies a parsimony
algorithm (commonly used in phylogenetic analyses) on matri-
ces with the raw distributions of organisms to identify areas or
sets of areas that share a unique group of endemic taxa. This
method presents several advantages compared to the traditional

© 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/geb 85

J. M. C. da Silva et al.
method. It is clearer, direct and can easily be improved when
more information on geographical distribution becomes avail-
able (Posadas & Miranda-Esquivel, 1999).
The Atlantic Forest is one of the best defined biogeographical
regions in South America. From a continental perspective, the
Atlantic Forest can be considered an island, because it is isolated
from other large blocks of South American forests (Amazonian
and Andean forests) by a corridor of open to semiopen forma-
tions, comprising Caatinga, Cerrado and Chaco (Ab’Saber, 1977).
Due to this isolation, the Atlantic Forest harbours a unique biota
with many endemic genera and species (Myers et al., 2000).
The Atlantic Forest, however, is not homogeneous. Several fac-
tors contribute to the environmental diversity of this region. One
is the latitudinal range, of about 25 degrees, thus including dif-
ferent climatic regimes (Nimer, 1979). Another relevant factor is
altitude, as forests cover areas from sea level to 1700 m a.s.l.
(Rizzini, 1997). Latitude and altitude together produce condi-
tions for the development of distinctive types of vegetation
(Hueck, 1972; Rizzini, 1997). This, in turn, influences species’
distributions (Brown & Lomolino, 1998).
Several attempts have been made to identify avian areas of
endemism in the Atlantic Forest. Although using the traditional
methods, their results differed in several aspects. Müller (1973)
analysed several terrestrial vertebrates’ ranges (including several
bird species) and considered the Atlantic Forest as a single area of
endemism. However, he identified at least three subareas of
endemism: Pernambuco, Bahia and Paulista. Cracraft (1985)
identified two areas of endemism for birds along the Atlantic
Forest: Serra do Mar, which stretches from Pernambuco to Santa
Catarina along the coastal zone; and Paraná, which encompasses
the region covered by Araucaria forests in southern Brazil. Haffer
(1985) regarded the Atlantic Forest as a single area of endemism,
named South-eastern Brazil. Stattersfield et al. (1998) identified
five avian areas of endemism in eastern Brazil: the Atlantic Slope
of Alagoas and Pernambuco; Deciduous forests of Bahia; Decid-
uous forests of Minas Gerais and Goiás; Atlantic Forest lowlands;
and Atlantic Forest mountains.
This paper has three main goals. The first is to use Morrone’s
methodology to identify areas of endemism along the Atlantic
Forest using passerine birds as a study group. The second is to
evaluate the generality of the areas identified using this method
by comparing them with those identified for other organisms.
The final goal is to use the area cladogram generated in the
analysis to propose, for the first time, a general hypothesis of
historical relationships among areas of endemism of the
Atlantic Forest.
METHODS
Study area
About 95% of the Atlantic Forest lies in Brazil, the remainder in
Argentina and Paraguay. The core region of Atlantic Forest corre-
sponds to an almost continuous zone composed of several forest
types along the Brazilian coast, from Rio Grande do Norte to Rio
Grande do Sul (Fig. 1). In addition, other disjunct regions are
86 Global Ecology and Biogeography, 13, 85 –92, © 2004 Blackwell Publishing Ltd
currently also considered as Atlantic Forest (Conservation
International et al., 2000): (a) evergreen to semideciduous forest
islands (regionally known as ‘brejos’) located in the slopes of
residual plateaus in the Caatinga region; (b) deciduous and sem-
isolated forests along the middle section of the São Francisco
river and in southern Piauí; and (c) deciduous and semidecidu-
ous forests isolated along the Serra da Bodoquena, Mato Grosso
do Sul. In total, this area corresponds to about 1,400,000 km2.
Identification of areas of endemism
The method proposed by Morrone (1994) comprises the follow-
ing steps: (a) define operational geographical unites (OGU); (b)
construct a data matrix; (c) perform a parsimony analysis of the
data matrix; (d) delimit the OGU or groups of OGUs defined by
at least two endemic species; and (e) map the species endemic to
each OGU or groups of OGUs to delineate the boundaries of
each area of endemism.
Sample localities are the ideal OGUs for the parsimony analy-
sis of distribution data (Rosen, 1988), however, our knowledge of
the geographical distribution of bird species along the Atlantic
Forest is incomplete as several places have not been adequately
sampled for birds (Conservation International et al., 2000). To
circumvent this problem, several authors have used as OGUs a
previous regionalization scheme, such as provinces, districts,
interfluvial regions, or ecoregions (Cracraft, 1991, 1994; Silva &
Oren, 1996; Morrone & Escalante, 2002). However, Silva & Oren
(1995) pointed out that this strategy is based on a critical
assumption, namely that the biotic difference within an OGU is
smaller than among OGUs. We used a different approach. We
selected 24 quadrats of 1 × 1 degree, which together cover most
of the Atlantic Forest (Fig. 1). These quadrats were selected based
on two criteria. The first (‘sampling criterion’) is that each quad-
rat must have at least one or a set of localities that have been rea-
sonably sampled for birds. The second criterion (‘representation
criterion’) is that these quadrats must collectively encompass the
major forest types (according to the IBGE, 1988) and latitudinal
belts of the Atlantic Forest.
The endemic passerine species of the Atlantic Forest were
selected from a list presented by Pacheco & Bauer (1999). Then,
the general distribution and ecological requirements of each
species were separately re-evaluated to confirm that species were
endemic to the region and also restricted to forest environments.
As a result, some species (Formicivora serrana, Oreophylax morei-
rae, Elaenia ridleyana, Vireo gracilirostris, Hylophilus poicilotis,
Euphonia pectoralis, Sporophila melanogaster) listed as endemic
by Pacheco & Bauer (1999) were excluded from the final list of
species, whereas others (Lepidocolaptes wagleri, Lepidocolaptes
squamatus, Arremon franciscanus and Phylloscartes roquettei)
were included. Species’ ranges were obtained from several
sources, such as books (Pinto et al. 1978; Ridgely & Tudor, 1994,
1997; Sick, 1997), papers on systematics and distribution
(Bornschein et al., 1995, 1998, 2001; Gonzaga & Pacheco, 1995;
Gonzaga et al., 1995; Pacheco & Gonzaga, 1995; Whitney et al.,
1995a,b; Silva & Straube, 1996; Isler et al., 1997; Pacheco, 1997;
Raposo, 1997; Whitney & Pacheco, 1997; Willis, 1988), regional/

Figure 1 The Atlantic Forest in eastern South
America. A set of 24 quadrats was selected to
cover most of the environmental variation that
there is in this biome.
site lists (Pinto, 1952; Willis, 1979; Willis & Oniki, 1981; Scott
& Brooke, 1985; Belton, 1994; Rosário, 1996; Straube et al.,
1996; Aleixo & Galetti, 1997; Anjos & Schuchmann, 1997;
Magalhães, 1999; Parrini et al., 1999; Lins, 2001), unpublished
field observations by JMCS and MCS, and specimens of the
ornithological collections of the Universidade Federal de
Pernambuco (UFPE), Universidade Federal de Minas Gerais
(UFMG) and Museu de Zoologia da Universidade de São Paulo
(MZUSP).
A data matrix was built in which the columns represent species
(analogous to characters in a phylogenetic analysis) and rows
represent the quadrats (analogous to taxa in a phylogenetic
analysis). In the matrix, the absence of species is coded as ‘0’ and
the presence as ‘1.’ A hypothetical quadrat coded ‘0’ for all
columns is added to root the final cladogram. We used the
commands mh*; bb* of the computer program Hennig 86 (Farris,
1989) to find the most parsimonious cladograms. We used the
command ‘nelsen’ to generate a strict consensus tree.
The most basal quadrats or clusters of quadrats that have at
least two endemic species were considered as areas of endemism
(Morrone, 1994). Then, the ranges of endemic species that
delimit each area of endemism were mapped by using site
records processed by ARCVIEW 3.2. Finally, limits of each area of
endemism were proposed based on the degree of overlap among
their endemic species’ ranges.
Global Ecology and Biogeography, 13, 85– 92, © 2004 Blackwell Publishing Ltd
Bird areas of endemism in the Atlantic Forest
RESULTS
One hundred and forty species of endemic passerines were
included in this analysis (Appendix 1). Fourteen most parsimo-
nious cladograms were found. Thus, a strict consensus cladog-
ram was generated to identify unambiguous clusters of OGUs
(Fig. 2). Four areas of endemism may be identified from this
analysis. The first area is composed of quadrat A, defined by six
species (Myrmotherula snowi, Terenura sicki, Synallaxis infuscata,
Philydor novaesi, Phylloscartes ceciliae and Tangara fastuosa). The
second area is composed of quadrats D and E, and is defined by
four endemic species (Synallaxis cinerea, Phylloscartes beckeri,
Rhopornis ardesiaca and Formicivora iheringi). The third area is
composed only of quadrat F, and is defined by two species (Acro-
batornis fonsecai and Scytalopus psychopompus). The fourth area
of endemism is composed of quadrats Q, U, J, V, S, R, P, M, and
N. This area is defined by two species (Orthogonys chloricterus
and Dacnis nigripes). When compared with all other areas of
endemism identified, the fourth area is the only one in which
nested subareas of endemism may be also recognized. Thus, the
subcluster composed of quadrats U, J, V, S, R, P, M and N is
defined by two species (Myrmotherula gularis and Tangara des-
maresti); the subcluster S, R, P, M, and N is defined by two species
(Biatas nigropectus and Drymophila rubricollis); and quadrat N is
defined by four species (Myrmotherula fluminensis, Formicivora
87
J. M. C. da Silva et al.
Figure 2 Cladogram obtained from the parsimony analysis of the
data matrix of Appendix 1. Numbers above the branches are the
number of endemic species.
littoralis, Tijuca condita and Calyptura cristata). By combining
the ranges of the endemic species listed above, it is possible to
delimit four areas of endemism (Fig. 3): Pernambuco (A),
Central Bahia (D and E), Coastal Bahia (F), and Serra do Mar
(Q, U, J, V, S, R, P, M, and N).
Pernambuco includes all evergreen, semideciduous and decid-
uous forests that are orientated as a belt following the Atlantic
coast, north to the river São Francisco. It comprises the coastal
zone of the states of Paraíba, Pernambuco and Alagoas. Central
Bahia includes a large and topographically complex area that
harbours a mosaic of evergreen, semideciduous and deciduous
forests, which cover both slopes and adjacent areas of the
Chapada da Diamantina and all surrounding plateaus in central
Bahia and northern Minas Gerais. Coastal Bahia encompasses a
small area dominated by lowland evergreen forests roughly
between the rivers Jiquiriça and Jequitinhonha, in the state of
Bahia. Serra do Mar includes a very heterogeneous habitat gradi-
ent that ranges from coastal restingas at sea level to forests at
1700 m a.s.l. This area follows roughly the belt of high-elevation
ranges that is generally named ‘Serra do Mar’ and that extends
from central Espirito Santo to Santa Catarina.
88 Global Ecology and Biogeography, 13, 85 –92, © 2004 Blackwell Publishing Ltd
DISCUSSION
Avian areas of endemism
The analysis indicates that there are distinct areas of passerine
bird endemism in the Atlantic Forest. Four distinctive and basal
areas of endemism were identified and they form the base upon
which a systematic research program on avian biogeography in
this region may be developed. This result supports the prediction
by Cracraft (1985) that his ‘Serra do Mar Centre’ would be sub-
divided into smaller areas of endemism when bird distributions
were examined in more detail.
The identification of the four areas of endemism differs from
previous analyses. In contrast with Müller (1973), the area of
endemism Pernambuco does not include both banks of the river
São Francisco, because all endemic bird species that delimit this
region are restricted to the northern bank. Also, Müller’s Bahia
subcentre does not strictly follow the Atlantic coast. Finally,
although Müller’s Paulista subcentre is largely congruent with
the area of endemism Serra do Mar, it does not extend up to
Espírito Santo.
In contrast with Cracraft (1985), we could not find any
support for the Paraná Centre (encompassing central Paraná,
western Santa Catarina and northern Rio Grande do Sul) as an
area of endemism. Cracraft listed 12 bird taxa as supporting
the Paraná Centre, including six passerine species: five forest
specialists (Clibanornis dendrocolaptoides, Leptasthenura setaria,
Leptasthenura striolata, Hemitriccus kaempferi, Cyanocorax
caeruleus), and one grassland specialist (Anthus nattereri). The
ranges of these species are not restricted to Paraná Centre (see
Appendix 1) and include also other areas of endemism. In fact,
our analysis suggests that at least the eastern portion of Cracraft’s
Paraná Centre (quadrat Q in Fig. 2) is part of the area of ende-
mism Serra do Mar, whilst other sectors (quadrats T, X and Z)
comprise a transition zone with other biomes.
Stattersfield et al. (1998) recognized only one of the areas of
endemism that we identified: Pernambuco (they named this area
as ‘Atlantic slope of Alagoas and Pernambuco’). They also identi-
fied an area in central Bahia (‘Deciduous forests of Bahia’), but
their area is smaller and does not encompass the ranges of all
species that we suggest are endemic to this region. Bird species
that define the area of endemism Central Bahia inhabit different
habitats. Two species (Synallaxis cinerea and Phylloscartes beckeri)
are restricted to evergreen forests along the slopes of some high
plateaus while Rhopornis ardesiaca and Formicivora iheringi are
restricted to dry forests that surround these plateaus. Cracraft
(1985) stressed that there is no reason to expect that endemic
species defining an area of endemism (a primarily historical
entity) should have the same ecological requirements. In
addition, although these four species are not syntopic, they are
sympatric at the spatial scale analysed herein and their ranges
coincide spatially to define an area of endemism. Stattersfield
et al. (1998) recognized one large area of endemism named
‘Atlantic Forest lowlands’, which is largely congruent with the
areas Coastal Bahia and Serra do Mar identified herein. They also
identified an area of endemism named ‘Atlantic Forest mountains’,

Figure 3 The boundaries of the four areas of
endemism identified for passerine birds in the
Atlantic Forest. Boundaries are based on the
percentage of overlap among ranges of the
endemic species. (a) Pernambuco (b) Central
Bahia and Coastal Bahia, and (c) Serra do Mar.
pointing out that there are differences between lowland and
highland avifaunas along the Serra do Mar. We could not find
any evidence for this area of endemism and it is included in the
Serra do Mar or in Central Bahia. We found no support to recog-
nize Stattersfield’s area of endemism ‘Deciduous forests of Minas
Gerais and Goiás’. This area of endemism is composed of two dis-
junct patches of dry forests (one in the São Francisco valley and
another in the Paranã valley) separated by a large plateau covered
mostly by cerrado. The ranges of the two species that they used to
recognize this area of endemism are not fully coincident, because
Knipolegus franciscanus occurs in both São Francisco and Paranã
valleys, whilst Phylloscartes roquettei occurs only in the São
Francisco valley.
Comparison with areas of endemism of other
organisms
The four areas of endemism exhibit considerable congruence
with those identified for other organisms in the Atlantic Forest.
Prance (1982) identified three areas of endemism for woody
plants (Pernambuco, Bahia and Rio de Janeiro) that are largely
coincident with the areas identified herein (Pernambuco, Costal
Bahia and Serra do Mar, respectively). Erwin & Pogue (1988)
suggested that all coastal Atlantic Forest south of the river São
Francisco is a single biogeographical unity that they named
‘South Atlantic Coast’. This scheme is not contradictory with the
one suggested herein. Soderstrom et al. (1988) found two areas
Global Ecology and Biogeography, 13, 85– 92, © 2004 Blackwell Publishing Ltd
Bird areas of endemism in the Atlantic Forest
of endemism for bamboos along the Atlantic Forest. One (Bahia)
is largely coincident with Coastal Bahia and another (Serra do
Mar) is coincident with our area Serra do Mar. Tyler et al. (1994)
identified four areas of endemism along the Atlantic Forest.
Their area of endemism Pernambuco overlaps 100% with the
Pernambuco area identified herein. Tyler et al.’s Bahia area
includes a large extension of Coastal Bahia and a small portion of
Central Bahia. Tyler et al.’s Rio de Janeiro and Santa Catarina
areas are mostly included within the area of endemism Serra do
Mar.
These comparisons indicate that passerine areas of endemism
have significant generality, because endemic passerine species
along the Atlantic Forest tend to cluster in the same regions as
other groups of organisms. According to Cracraft (1985), this
generality implies that there is a pattern to be investigated,
namely, the origin and development of biotas as represented by
these areas of endemism.
Historical relationships among areas of endemism
Although the analysis of species’ raw distributions using parsi-
mony is effectively one good method to identify the areas of
endemism at intracontinental scales (Silva & Oren, 1996; Posadas
et al., 1999; Garcia-Barros et al., 2002; Morrone & Escalante,
2002), there are doubts if this method may be used to estimate
historical relationships among areas of endemism (Brooks, 1981;
McLennan & Brooks, 2002). Some authors claim that the results
89
J. M. C. da Silva et al.
from this method are congruent with those obtained using more
sophisticated biogeographical analyses based on cladistic bio-
geography methods and on the phylogenies of different groups
of species (e.g. Cracraft, 1991, 1994; Silva & Oren, 1995; Ron,
2000). Cracraft (1991) provided a rationale for this claim, by
stating that shared taxa among areas may be evidence of histo-
rical relatedness, because they are the consequence of failure to
differentiate once the areas became segregated. Cracraft (1994)
also suggested that if biotic dispersion is historically constrained,
it should be expected that the distribution of natural taxa will
exhibit hierarchical congruence when examined cladistically.
Silva & Oren (1995) suggested that the analysis of raw distribu-
tions of species using parsimony should be a useful tool to iden-
tify preliminary hypotheses of historical relationships among
areas of endemism that may later be evaluated by the methods of
cladistic biogeography.
Several authors have investigated the historical relationships of
the Atlantic Forest with other lowland forest regions in South
America (Cracraft & Prum, 1988; Bates, 1998; Costa et al., 2000).
In all these broad-scale analyses, the hypothesis that the Atlantic
Forest is a composite of at least two or more areas of endemism is
well supported. The realization that the Atlantic Forest is com-
posed of at least four areas of avian endemism will help refine
those analyses and shed light on the complex issue of historical
relationships among the areas of endemism in South America
(Bates et al., 1998).
Recent studies indicate that most biotas are mosaics due to
vicariance, peripheral isolates speciation, postspeciation disper-
sal, nonresponse to vicariance and extinction and that the same
areas often have reticulated histories with respect to the species
that live in them (McLennan & Brooks, 2002). Reticulated histo-
ries are in fact sets of different dichotomic relationships, each one
representing a piece of the complex evolutionary history of a
biota. The results obtained herein may be used to advance a first
(but by no means the only) set of historical dichotomic relation-
ships among areas of endemism for birds in the Atlantic Forest.
The hypotheses derived from Fig. 2 are as follows: (a) there is a
basal dichotomy among areas of endemism in the Atlantic Forest,
with Pernambuco constituting a northern cluster, and Coastal
Bahia, Central Bahia and Serra do Mar comprising a southern
cluster; and (b) within the southern cluster, Central Bahia and
Serra do Mar are more closely related to each other than to
Coastal Bahia. This hypothesis may be tested by evaluating the
phylogeny of monophyletic groups that have taxa (species or
subspecies) endemic to at least three of these four areas of ende-
mism. Three passerine groups could be good candidates for these
tests. The first is Lepidocolaptes fuscus, because it is largely distrib-
uted in the Atlantic Forest and has a subspecies in each one of the
areas of endemism: Pernambuco (atlanticus), Central Bahia (bre-
virostris), Coastal Bahia (tenuirostris), and Serra do Mar ( fuscus).
The two other groups are also informative (Gonzaga & Pacheco,
1995; Pacheco & Gonzaga, 1995), but they lack species in Coastal
Bahia: one is comprised of Synallaxis infuscata (Pernambuco), S.
cinerea (Central Bahia), and S. ruficapilla (Serra do Mar); and
the other is formed by Phylloscartes ceciliae (Pernambuco),
P. beckeri (Central Bahia), and P. v. virescens (Serra do Mar).
90 Global Ecology and Biogeography, 13, 85 –92, © 2004 Blackwell Publishing Ltd
ACKNOWLEDGEMENTS
This project was conducted with support from FACEPE, CNPq,
Universidade Tiradentes, Universidade Federal de Pernambuco
(Programa de Pós-Graduação em Biologia Animal), and Conser-
vation International do Brazil. We thank José Fernando Pacheco
and Sonia A. Roda for data about some critical species, and John
Bates and an anonymous referee for very useful criticisms. We
particularly wish to thank Luis Barbosa for preparing the figures.
SUPPLEMENTARY MATERIAL
The following material is available from http://
www.blackwellpublishing.com/products/journals/suppmat/
GEB/GEB077/GEB077sm.htm
Appendix S1. Presence (1) and absence (0) of 140 endemic
passerine endemic species in 24 quadrats in the Atlantic Forest.
Lines are quadrats and columns are species. The line ‘root’ is an
area with no taxon at all that is used to root the most parsimoni-
ous cladograms.
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BIOSKETCHES
José Maria Cardoso da Silva is an ornithologist
interested in the systematics, evolution, ecology,
biogeography and conservation of Neotropical birds.
Marcelo Cardoso de Sousa is an ornithologist
interested in the species distributions and ecology of
Atlantic Forest and Caatinga birds.
Carlos Henrique Madeiros Casttelleti is a biologist
interested in biogeography, landscape ecology and
conservation.