Revista Brasileira de Ornitologia - Sociedade Brasileira de Ornitologia

Revista Brasileira 

de Ornitologia 

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Volume 21 / Issue 21 

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Março 2013 / March 2013 

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Sociedade Brasileira de Ornitologia / Brazilian Ornithological Society 

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Revista Brasileira de Ornitologia 

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Front cover: Four poorly-known bird species photographed in the Floresta Nacional do Tapajós, Estado do Pará; an inventory of the avifauna of the wider Santarém 

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Revista Brasileira 

de Ornitologia 

ISSN (impresso/printed) 0103-5657 

ISSN (on-line) 2178-7875 

www.ararajuba.org.br/sbo/ararajuba/revbrasorn.htm 

Volume 21 / Issue 21 

Número 1 / Number 1 

Março 2013 / March 2013 

Publicada pela / Published by the 

Sociedade Brasileira de Ornitologia / Brazilian Ornithological Society 

Belém - PA


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ISSN: 0103-5657 (impresso) 

ISSN: 2178-7875 (on-line) 

1. Ornitologia. I. Sociedade Brasileira de Ornitologia.


Volume 21 – Número 1 – Março 2013 / Issue 21 – Number 1 – March 2013 

SUMÁRIO / CONTENTS 

artigos / papers 

Plumage variation in the Planalto Woodcreeper (Dendrocolaptes platyrostris) and the melanocortin-1 receptor gene 

(MC1R) 

Josmael Corso, Gustavo Sebastián Cabanne, Fernando Mendonça d’Horta, Élgion Lucio da Silva Loreto and Cristina Yumi 

Miyaki............................................................................................................................................................................. 

Early singing onset in the black-cheeked gnateater (Conopophaga melanops) 

André Magnani Xavier de Lima and James Joseph Roper...................................................................................................... 

Sazonalidade na assembleia de aves aquáticas em uma lagoa marginal do rio Mogi Guaçu, estado de São Paulo, Brasil 

Paulo Rubim..................................................................................................................................................................... 

One hundred and thirty-five years of avifaunal surveys around Santarém, central Brazilian Amazon 

Alexander C. Lees, Nárgila G. de Moura, Christian Borges Andretti, Bradley J. W. Davis, Edson V. Lopes, L. Magalli Pinto 

Henriques, Alexandre Aleixo, Jos Barlow, Joice Ferreira and Toby A. Gardner........................................................................ 

Las aves de la isla Lobos de Tierra, Perú: revisión bibliográfica y nuevos registros (1684-2011) 

Judith Figueroa.................................................................................................................................................................. 

Aves da Serra de Maracaju, Mato Grosso do Sul, Brasil 

Alessandro Pacheco Nunes, Maurício Neves Godoi, Maria Antonietta Castro Pivatto, José Carlos Morante Filho, Eduardo 

Weffort Patrial, Paulo Antonio Silva, Vanessa Katherinne Stavis, Daniel De Granville Manço, Marco de Barros Costacurta, 

Caroline Leuchtenberger e Carlos Rodrigo Lehn.................................................................................................................. 

1 

5 

10 

16 

58 

75 

NotaS / Short-CommunicationS 

On the nest, eggs and nestlings of the Short-tailed Antthrush (Chamaeza campanisona) 

Ismael Franz..................................................................................................................................................................... 

Registros documentados da gaivota-de-franklin Leucophaeus pipixcan (Wagler, 1831) no Nordeste do Brasil e da gaivota-decabeça-cinza 

Chroicocephalus cirrocephalus (Vieillot, 1817) nos Estados de Alagoas e Pernambuco 

Sergio Leal, Lúcio Cláudio H. Serapião e Glauco Alves Pereira............................................................................................. 

101 

105 

Instructions to Authors

Revista Brasileira de Ornitologia, 21(1), 1-4 

March 2013 

article 

Plumage variation in the Planalto Woodcreeper 

(Dendrocolaptes platyrostris) and the 

melanocortin-1 receptor gene (MC1R) 

Josmael Corso 1,6 , Gustavo Sebastián Cabanne 2 , Fernando Mendonça d’Horta 3 , Élgion Lucio da Silva Loreto 4 

and Cristina Yumi Miyaki 5 

1 

Programa de Pós-Graduação em Genética e Biologia Molecular, Departamento de Genética, Universidade Federal do Rio Grande do Sul, Porto 

Alegre, RS, Brazil - josmaelcorso@gmail.com 

2 

Museo Argentino de Ciencias Naturales, CONICET, Buenos Aires, Argentina - gscabanne@yahoo.com 

3 

Departamento de Genética e Biologia Evolutiva, Instituto de Biociências, Universidade de São Paulo, São Paulo, SP, Brazil - fmhorta@ig.com.br 

4 

Departamento de Biologia, Universidade Federal de Santa Maria, Santa Maria, RS, Brazil - elgion.loreto@pq.cnpq.br 

5 

Departamento de Genética e Biologia Evolutiva, Instituto de Biociências, Universidade de São Paulo, São Paulo, SP, Brazil - cymiyaki@ib.usp.br 

6 

Correspondence: Josmael Corso – Programa de Pós-Graduação em Genética e Biologia Molecular, Departamento de Genética, Universidade 

Federal do Rio Grande do Sul, Porto Alegre, RS, Brazil – CEP:91501-970 - Tel.: 51 3308.6726. E-mail: josmaelcorso@gmail.com 

Received on 7 August 2011. Accepted on 6 January 2013. 

ABSTRACT: The Planalto Woodcreeper (Dendrocolaptes platyrostris) presents “pale” and “dark” plumage variants, which are 

distributed throughout the Cerrado and Caatinga, and throughout the Atlantic Forest, respectively. To understand the genetic 

nature of the plumage variation in the species, we partially sequenced the melanocortin-1 receptor (MC1R) gene, which is associated 

with melanic phenotypes in vertebrates. We found no correlation between variation at MC1R sequences and plumage color in D. 

platyrostris. Aminoacid sites that were correlated with variation in melanic plumage in other bird species were monomorphic in D. 

platyrostris. Our results suggested that MC1R seems not to be involved in controlling plumage variation in D. platyrostris. 

KEY-WORDS: plumage color polymorphism; melanocortin-1 receptor, Dendrocolaptes 

INTRODUCTION 

The genetic basis of phenotypic diversity within 

species is of great interest to evolutionary biologists because 

adaptive evolution depends on selection of genetic variants 

(Theron et al. 2001). Genetic changes resulting in color and 

pigmentation variation among closely related taxa might 

represent important evolutionary events. However, the 

molecular basis and developmental pathways responsible 

for phenotypic difference are unknown in most cases 

(Cheviron et al. 2006). Recently, it has been suggested that 

the locus encoding the melanocortin-1 receptor (MC1R) 

may cause color polymorphisms in wild populations 

(Mundy 2005). This gene encodes the MC1R protein, 

which is expressed in the melanocytes of developing 

feathers and hair follicles and plays a critical role in the 

control of melanin synthesis (Theron et al. 2001, Baião 

et al. 2007). Point mutations in this locus were associated 

with color polymorphisms based on melanin and can 

cause changes from light to dark color all over the body in 

a variety of taxa (revision in Corso et al. 2012). 

The planalto woodcreeper (Dendrocolaptes 

platyrostris) is an endemic bird of the Atlantic forest 

that inhabits forest enclaves and gallery forests within 

the Cerrado, Chaco, and Caatinga (Figure 1). As 

many other woodcreepers, D. platyrostris does not have 

plumage sexual dimorphism. It has two parapatrically 

distributed subspecies: D. p. platyrostris – dark plumage 

morph - individuals are predominantly streaked buff, 

have blackish crown, dull chestnut wings and tail, and 

brown underparts, and inhabit the Atlantic Forest 

domain in southeastern Brazil, eastern Paraguay and 

extreme northeastern Argentina; D. p. intermedius – 

pale plumage morph - individuals have browner crown, 

almost no streaking on mantle, paler and brighter rufous 

wings and tail, and slightly paler underparts, and inhabit 

gallery forests of Cerrado / Caatinga domains (Ridgely 

& Tudor 1994, Willis & Oniki 2001, Cabanne et al. 

2011; Figure 1). The study of Cabanne et al. (2011) 

indicated that plumage type in D. platyrostris (i.e., dark 

and pale morphs) was not correlated with neutral genetic 

divergence (mitochondrial DNA) but confirmed that it

2 

Plumage variation in the Planalto Woodcreeper (Dendrocolaptes platyrostris) and the melanocortin-1 receptor gene (MC1R) 

Josmael Corso, Gustavo Sebastián Cabanne, Fernando Mendonça d’Horta, Élgion Lucio da Silva Loreto and Cristina Yumi Miyaki 

was correlated to different types of habitats, with dark 

and pale morphs occurring respectively in more humid 

(Atlantic Forest) and dry (Cerrado / Caatinga) habitats. 

In this study we examined the MC1R sequence 

variation of D. p platyrostris and D. p. intermedius. 

Specifically, we looked for fixed non-synonymous 

differences in MC1R between plumage morphs. Our 

main question was whether the light and dark phenotypes 

of this species could be related to changes in the MC1R 

coding region. 

Figure 1. Distribution of biomes and of the two subspecies of Dendrocolaptes platyrostris. Modified from Cabanne et al. (2011). 

MATERIAL AND METHODS 

We sampled five adult individuals of D. platyrostris 

intermedius and five D. p. platyrostris, from localities that are 

1600 km apart (Table 1). Total genomic DNA was isolated 

from blood or muscle samples by a standard phenol/ 

chloroform extraction protocol (Sambrook et al. 2001). 

A fragment of approximately 500 bp of the avian MC1R 

gene was amplified by PCR using the following primers: 

lcorMSHR72 – 5’ AYGCCAGYGAGGGCAACCA 3’ 

(Cheviron et al. 2006) and MC1RIntRev – 

5’ AACATGTGRATGTAGAGCACC 3’. PCR 

Revista Brasileira de Ornitologia, 21(1), 2013



3 

conditions were: initial denaturation for 3 min at 94°C, 

followed by 45 cycles (denaturation at 94°C for 45 s, 

annealing at 50-60°C for 60 s and extension at 72°C 

for 90 s) and a final extension at 72°C for 5 min. This 

MC1R fragment includes main sites previously shown to 

be associated with melanic phenotypes in birds (Cheviron 

et al. 2006). Each specimen was submitted to at least two 

independent amplification and sequencing reactions 

(using the same primers used in the PCR) to confirm 

the sequences. Consensus sequences were obtained and 

deposited in GenBank under the accession numbers: 

FJ985683–FJ985688, JN224986-JN224989. 

DNA sequences were aligned and their nucleotides 

and deduced amino acids were compared to those 

from bananaquits (Coereba flaveola; GenBank number 

AF362605 and AF362598) using BIOEDIT v. 7 (Hall, 

1999; Table 2). We tested for evidence of selection at 

MC1R by calculating Tajima’s D statistic using MEGA 5 

(Tamura et al. 2011). 

Table 1. Specimens sequenced, sampling localities, sample identification, and voucher identification of tissue samples. 

Taxon a Locality (habitat) Tissue identification b Vouchers c 

D. p. platyrostris 

Pinhalão, Paraná (PR). 23°46’S; 50°3’W 

(Atlantic forest) 

Wenceslau Braz, PR. 22°5’S; 48°47’W 


Morro Grande State Park, São Paulo (SP). 

23°42’S; 46°59’W (Atlantic forest) 

Barreiro Rico, SP. 22°38’S; 48°13’W 


LGEMA P885 MZUSP 75622 

LGEMA P957 MZUSP 75690 

LGEMA P2480, P2482 - 

LGEMA P1696 - 

D. p. intermedius 

National Park of Serra das Confusões, 

Piauí. 9°40’S; 44°8’W (Caatinga) 

LGEMA P2277, P2278, 

P2329, P2379, P2418 

MZUSP 77719, 77720, 

77721, 77722 

a D. p.= Dendrocolaptes platyrostris. 

b Samples are deposited at the Laboratório de Genética e Evolução Molecular de Aves (LGEMA), Universidade de São Paulo, São 

Paulo, Brazil. 

c Voucher specimens housed at MZUSP: Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil. A dash indicates that 

a voucher specimen does not exist. 

Table 2. MC1R sequences of Dendrocolaptes platyrostris. Sites are numbered based on the alignment with bananaquit (Coereba flaveola) MC1R 

sequences (see methods). Sites in bold are associated with color polymorphism in other avian taxa (amino acid replacement), all D. platyrostris 

individuals are monomorphic at these sites. Variable sites in D. platyrostris that resulted in putative aminoacid substitution are underlined. Asterisks 

show sites that are identical to those of Coereba flaveola. 

Taxa Nucleotide site Aminoacid position Plumage 

1 1 2 2 5 1 3 6 8 9 1 1 1 1 2 

1 2 0 7 1 6 8 8 5 2 1 5 6 7 0 

3 0 3 4 4 1 7 6 2 7 

C. flaveola M5 T C A A G A I H V K V Q V V H melanic 

C. flaveola Y24 * * * G * * * * * E * * * * * yellow 

D. p. platyrostris P885; P2480; P2482; P1696 * T * G * . * * * E * * * * * dark 

D. p. platyrostris P957 * T/C * G * . * * * E * * * * * dark 

D. p. intermedius P2278; P2379; P2418 * T * G A . * * * E * * * I * pale 

D. p. intermedius P2329 T/A T * G A . * * * E * * * I * pale 

D. p. intermedius P2277 * T G G * . * R * E * * * * * pale 


4 



RESULTS 

The alignment matrix presented 525 characters 

encompassing sites 100 to 624 of the MC1R gene. It had 

96–97% nucleotide identity with sequences from other 

passerine birds such as Coereba flaveola (Theron et al. 

2001), Phylloscopus warblers (MacDougall-Shackleton 

et al. 2003), and Lepidothrix coronata (Cheviron et 

al. 2006). Two D. platyrostris individuals presented 

heterozygous sites at positions 113 and 120 (Table 2). 

There were three variable sites that resulted in nonsynonymous 

substitutions (amino acid positions 38, 68, 

and 172), which allowed us to identify three polymorphic 

DNA sites in pale morph birds: c.113 T > A , c.203 A > 

G, and c.514 G > A (Table 2). These changes could not be 

correlated with phenotype differences, once those changes 

maintain the same polarities of original aminoacids. As 

the number of variable sites observed was very low, our 

statistical power to detect selection was also low (Tajima’s 

D: -0.78, not significant at P > 0.10). 

DISCUSSION 

The results obtained are in contrast with some 

studies of vertebrate taxa, including birds (reviewed in 

Mundy 2005), which observed a linkage between MC1R 

mutations and the appearance of melanic phenotype. In 

Coereba flaveola, a non-synonymous substitution (E92K) 

was associated with melanic plumage (Mundy et al. 2004). 

In the red-footed booby (Sula sula), the white/melanic 

polymorphism observed was associated to two point 

substitutions, V85M and H207R (Baião et al. 2007). These 

three sites were monomorphic in D. platyrostris (Table 2). 

However, these results should be interpreted with caution, 

as our sample size is small and it may not have allowed 

the identification of association between phenotypic and 

genotypic variations. However, even though we did not 

completely sequence the MC1R, the segment studied 

contained the majority of sites previously shown to be 

correlated with plumage differences in birds (Mundy et al. 

2004, Mundy 2005, Cheviron et al. 2006). Therefore, D. 

platyrostris seems to be another instance of a bird species 

that does not present any evidence that MC1R is correlated 

with pigmentation differences as documented previously 

in Phylloscopus warblers (MacDougall-Shackleton et al. 

2003) and Lepidothrix coronata (Cheviron et al. 2006), 

the latter belonging to the same Passeriformes suborder 

(Tyranni) as D. platyrostris. 

ACKNOWLEDGMENTS 

This work was funded by Conselho Nacional de 

Desenvolvimento Científico e Tecnológico (CNPq) 

Fundação de Amparo à Pesquisa do Estado de São Paulo 

(FAPESP) Brazil. We are also grateful to Fabio Raposo 

and Luis Fabio Silveira (MZUSP), and the members 

of Laboratório de Biologia Molecular (LabDros) for 

their help. 

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Ridgely, R. S. & Tudor, G. 1994. The Birds of South America: The 

Suboscine Passerines, v. 2. Austin: University of Texas Press. 

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Cloning: a Laboratory Manual. New York: Cold Spring Harbor 

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notes. Wilson Bulletin, 113: 231-233. 

Associate Editor: Camila C. Ribas 



March 2013 

article 

Early singing onset in the black-cheeked gnateater 

(Conopophaga melanops) 

André Magnani Xavier de Lima a, c and James Joseph Roper a, b 

a 

b 

c 

Programa de Pós-Graduação em Ecologia e Conservação, Universidade Federal do Paraná, Curitiba, PR, Brazil 

Departamento de Ecologia de Ecossistemas, Universidade de Vila Velha, Vila Velha, ES, Brazil 

Corresponding author: andremxlima@gmail.com 

Received on 7 August 2012. Accepted on 19 January 2013. 

ABSTRACT: Early singing onset in the black-cheeked gnateater (Conopophaga melanops). Song learning in birds has been 

intensively studied, mostly in the suborder Oscines. While studying the black-cheeked gnateater Conopophaga melanops (Suboscine) 

in southern Brazil nestling vocalizations were observed. Adult and nestlings vocalizations were described and compared to examine 

age-specific variation in song structure. Nestling vocalization was more similar to adult song than to alarm calls. However, nestling 

and adults songs differed primarily in maximum frequency and total length of the song. Nestling age was also correlated with some 

nestling song features. These patterns suggest that while vocalization at an early age may be innate, nonetheless, there may also be 

a stage during which individuals could learn the adult song. Because this early song is not the same as the adult song, we suggest 

that some Suboscines may achieve their song only after practice, although not necessarily with an imitative-learning basis, such as 

in Oscines. Further study will be required, with observations beginning with young birds on the nests, to better understand the 

ontogeny of song and individual variability in adults Suboscines. 

KEY-WORDS: Conopophagidae; innate behavior; learning; Suboscine; vocalization. 

INTRODUCTION 

Questions about the development of avian song 

learning have a long history of debate (Marler & Tamura 

1964, Nottebohm 1970, Immelmann 1975). One 

general pattern is clear in which some songbirds, parrots, 

and hummingbirds learn from imitation (Gaunt et al. 

1994, Farabaugh et al. 1994, Berg et al. 2011) while other 

birds inherit song templates (Kroodsma 1984, Kroodsma 

& Konishi 1991). Oscines (songbirds, suborder Passeri), 

the best studied group, may learn and modify their 

songs over time (Marler & Tamura 1962, Chaiken et al. 

1994, Brenowitz & Beecher 2005, Kirn 2010), while the 

Suboscines (the other passerines, suborder Tyranni) are 

poorly studied and thought to have innate songs that 

are relatively unmodified over time (Kroodsma 1984). 

Although this pattern is widely accepted, more empirical 

and experimental data are available for those species that 

“learn” than there is for the non-learners, the Suboscines 

(and others). However, learning may be more common 

than thought, even among the Suboscines (Leger 2005, 

Saranathan et al. 2007). 

In those species that learn songs, young birds pass 

through a sensitive period during which they acquire 

information that will shape song development. In some 

species, song learning occurs at a certain age, while others 

may learn at any age (Nottebohm 1970, Farabaugh et 

al. 1994, Tramontin & Brenowitz 1999). The learning 

stage (involving auditory and recording mechanisms 

prior to song production) usually begins the first month 

after hatching and may continue through the first year 

(Kroodsma & Konishi 1991, Marler 1997, Brenowitz & 

Beecher 2005). In most species within the Suboscines, 

when singing begins and when the song acquires the 

adult form are unknown. Although they are not expected 

to go through a typical song-learning phase, it is unlikely 

that the first vocalizations of a young bird will be that of 

an adult song (Kroodsma 1984). 

The Atlantic Forest endemic black-cheeked gnateater 

Conopophaga melanops (South American bird family 

Conopophagidae) is a tracheophone Suboscines (Irestedt 

et al. 2002). This gnateater is a socially monogamous 

and territorial species, and in general has life history 

traits more related to slow pace species (Wiersma et al. 

2007). Such traits would include an extended nest cycle, 

low clutch size and extended parental care. As example, 

the nest cycle may take more than 50 days since the nest 

building (Lima & Roper 2009a). 

While studying the population dynamics of the 

black-cheeked gnateater in southern Brazil (Lima &

6 


André Magnani Xavier de Lima and James Joseph Roper 

Roper 2009a), an unusual nestling singing behavior was 

observed. While based on few and sporadic observations, 

the lack of knowledge on song ontogeny within the 

Suboscines suggests that our information may be useful 

to better understand such process. Thus, we describe and 

compare adult and nestling vocalizations to discuss a first 

approach on the song development in this Suboscine 

passerine. 

METHODS 

The black-cheeked gnateater Conopophaga melanops 

was studied at a Private Reserve located in the municipality 

of Guaraqueçaba, state of Paraná, southern Brazil 

(25°13´S, 48°17´W). The reserve is at the lowland and 

mountain tropical forest, comprising approximately 2300 

ha, in a larger matrix of mixed successional forest stages 

(> 400 000 ha) contiguous with the largest area of wellpreserved 

Atlantic Forest. We captured and monitored 

gnateaters in a 25 ha plot (Lima & Roper 2009a). 

Nestlings were observed and monitored at five of 

18 nesting attempts. Some nestlings (three individuals in 

two nests) were observed as they occasionally vocalized. 

In eight observations, vocalizations were digitally 

recorded (Tascam DR08 with an external microphone 

Sennheiser ME66). Also, digital videos were recorded 

twice of singing nestlings on the nest (Sony DSC/H2). 

Songs of the three nestlings were recorded from 2-4 times 

on different days (thus, diverse ages). Nestling age was 

estimated based on the last date eggs were found and in 

comparison with nests found before hatching. Songs of 

adult males were recorded in the field (N = 10). One song 

per adult was used. 

Sound was analyzed in Raven Lite 1.0 (Cornell Lab of 

Ornithology, USA). Audio spectrograms were compared 

among nestlings and adults. Acoustic parameters used 

for comparison include total length (s), note rate (s - 

1 

), and fundamental frequency range (kHz), measured 

as the maximum and the minimum frequencies at two 

moments of each song: after the first second (f1) and at 

the beginning of the last second (f2) (i.e., a song that last 

8.3 s had frequencies measured at 1.0 s, as f1, and f2 at 7.3 

s). Adult and nestling song features were compared using 

t tests and regression analysis. We carried out regression 

analysis for checking the correlation between nestling 

age and song features. Tests were considered statistically 

significant when P < 0.05. 

RESULTS 

Adults sing a long and multi-note song (mean 

+ SD = 9.3 + 0.8 s, with 12.2 + 0.2 notes s -1 , n = 10, 

Figure 1). Adult song had usually presented two or three 

harmonics, with the lower frequency harmonic ranging 

around 2.7 – 4.2 kHz (f1 max 

+ SD = 3.5 + 0.2 kHz, and 

f2 max 

+ SD = 4.0 + 0.3 kHz, n = 10). The two succeeding 

harmonics ranges around 4.7 – 5.7 and 6.2 – 7.7 kHz. 

A unique recording presented four evident harmonics, 

which was the only one including the harmonic with 

the lower and fundamental frequency (1.5 – 1.9 kHz), 

with a second and dominant harmonic around 2.8 – 3.9 

kHz. Comparisons of initial and final frequencies were 

possible for the dominant harmonic, which increased the 

modulation toward the end of the song (f1 = 2.7 – 

min – max 

3.7 kHz, f2 = 3.3 – 4.2 kHz, r = 0.49, df = 9, P < 

min – max 

0.05). The alarm call is quite different than the song (Fig. 

1) and it is a single short pulse (~ 0.2 s) with a high and 

broad frequency (3.2 – 16 kHz). 

The nestling emits a similar but shorter song (mean 

+ SD = 3.9 + 0.3 s, 12.2 + 0.6 notes s -1 , n = 8; Figure 1). 

Nestling songs start and finish at lower frequencies than 

those of adults (f1 max 

+ SD = 2.8 + 0.2 kHz, t = 4.08, P < 

0.05, and f2 max 

+ SD = 3.4 + 0.2 kHz, t = 6.5, P < 0.05), 

and there is an increase in the modulation throughout the 

song for nestlings (r = 0.84, df = 7, P < 0.05). Only one 

harmonic, at 4.7-5.9 kHz, were found in one recording 

(from the older nestling in the sample, with the higher 

maximum frequency in figure 2). The emission of the 

sound was in apparent response to approach, during 

which the nestling stayed in the usual nestling posture. 

During sound emission, the body of the nestling vibrated 

in synchrony with the sound (we can send the video file 

for those interested). Begging was never seemed during 

observations of parental care at five nests where nestlings 

had survived (Lima e Roper 2009a). No other sounds 

were noted, even when nestlings were manipulated (for 

banding). 

The age of nestlings was estimated for one nest 

found on 10 December 2006 with two eggs that by 

12 December had hatched. The first juvenile observed 

singing in that nest was on 22 December (at 10-11 days 

after hatching). Another nest was found with two chicks 

on 12 January 2007, and when compared to nestlings of 

known age, they were estimated to be 8 days old. One 

nestling were recorded singing when the nest was found, 

and then, both nestlings “sang” in three (13, 18 and 23 

January) of six observations before they fledged on 25 

January, a date that corroborate with the estimated age 

(Lima & Roper 2009a). Thus, the age of nestlings at this 

nest was 8 and 9 days after hatching when was it found. 

Nestling age was correlated with some characteristics 

of the song. Maximum frequency at the end of the song 

(f2) and nestling age were correlated (r = 0.83, df = 7, 

P < 0.05, Figure 3). Total song length and age were 

correlated (r = 0.64) but at n = 9, P = 0.07. Minimum 

and maximum frequencies at the beginning of the song 

(f1) were independent of nestling age. 




7 

20 

15 

Adult 

Song 

Calls 

10 

5 

0 

0 1 2 3 4 5 6 7 8 9 10 

20 Nestlings 

15 

10 

5 

0 

0 1 2 3 4 5 6 7 8 9 10 

Time (s) 

Figure 1. Song and call spectrogram of the adult black-cheeked gnateater Conopophaga melanops, and the audio spectrogram of two nestlings. 

4.2 

4.0 

18 

3.8 

16 

3.6 

3.4 

14 

3.2 

12 

3.0 

2.8 

10 

Notes s -1 Frequency (kHz) 

Adult Nestling 

Max frequency (kHz) 

Figure 2. Song features points for adults and nestlings of black-cheeked gnateater Conopophaga melanops: black dots show note rate and white 

circles show maximum frequency (kHz). 


8 



Maximun frequency of f2 (kHz) 

3.2 3.3 3.4 3.5 3.6 3.7 3.8 

10 

12 14 16 18 

Age (days) 

Figure 3. Correlation between maximum frequency (kHz) in the end of the song (f2) and the age (in days) of nestlings of black-cheeked gnateater 

Conopophaga melanops. 

DISCUSSION 

Nestling vocalizations always more closely resembled 

adult songs than they did begging or alarm calls. This was 

somewhat surprising because vocalizations seemed to be 

in response to our approaching the nest, and so might have 

been in alarm. Or, perhaps the noise we made suggested 

to the bird that adults were returning with food. Despite 

of the reason they sang, the similarity between adult and 

young songs suggests that a song template is inherited, 

as expected for Suboscines (Kroodsma 1984, Kroodsma 

& Konishi 1991), but which must be improved as the 

nestling grows. 

Nestling black-cheeked gnateaters begin to sing 

earlier than other not related species (Kroodsma & 

Pickert 1984, Podos et al. 2004). Furthermore, some 

features of the song seem age-dependent. The maximum 

frequency of the dominant harmonic increased with age 

approaching that of the adult song, while young birds 

remained on the nest (Figure. 3). A similar process goes 

through the total length of their song (Figure 1). This 

appears to be learning on the part of the young blackcheeked 

gnateater and which would aim to result in an 

adult song with respect to frequency and song length. 

Thus, while nestling black-cheeked gnateaters inherit 

a song template, apparently the song is being practiced 

since early age such that learning may be involved to 

“fine-tuned” the song into that of an adult (such as longer 

duration and higher frequencies). 

Such fine-tuning may be due to the young bird 

comparing its own song with that of the adult singing 

nearby (Leger 2005, Saranathan et al. 2007). Or also, it 

may be due to the young bird comparing its own song 

with some innate “template”, because nestlings can sing 

similar to an adult prior to leaving the nest. This could 

be also somehow associated to the fitness or the body 

developmental rate of each individual, such as an agedependent 

tonus of the syringe´s muscles, or the body 

(and vocal tract) size (Marler 1997, Podos et al. 2004, 

Cardoso 2010). 

Nevertheless, even if the song ontogeny does not 

include any auditory feedback-imitation, the song of 

young birds did not entirely fit as an adult song. As adult’ 

songs presents some variation in the maximum frequency, 

and the song has an ascendant modulation, potential 

song individual variability would likely appears in adults 

by differences in the frequency level in the end of the 

song, and perhaps either in the song duration. Thus, it is 

likely that grown-up individuals will not achieve exactly 

the same song features when adults (Lampe & Espmark 

1994, Nelson & Marler 2005, Berg et al. 2011), even 

being a Suboscine species (Ippi et al. 2011). 

We suspect that learning may be more broadly 

correlated with song ontogeny here than previously 

thought. This could be potentially important for 

this species, due to the role of the song as a territorymaintenance 

device (Lima & Roper 2009a, b). If so, then 

social context, phenotypic and environmental variation 

may also influence song development in this and perhaps 

several others Suboscine species, but not essentially as it 

does in the Oscines (Mace 1987, Hoi-Leitner et al. 1995, 

Nowicki et al. 2002, O’Loghlen & Rothstein 2002, 

Kojima & Doupe 2011). 




9 


We thank the Fundação o Boticário de Proteção à 

Natureza and CAPES for the financial support. We thank 

Donald E. Kroodsma and anonymous reviewers for their 

throughout revision of early version of the manuscript. 

The authors were licensed by the Brazilian Agencies 

SISBIO/CEMAVE. 

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Associate Editor: Luiz Antônio Pedreira Gonzaga 



March 2013 

article 

Sazonalidade na assembleia de aves aquáticas 

em uma lagoa marginal do rio Mogi Guaçu, 

estado de São Paulo, Brasil 

Departamento de Botânica, Universidade Federal de São Carlos, 

CP 676, CEP 13565-905, São Carlos, SP, Brazil 

E-mail: rubimpaulo@gmail.com 

Paulo Rubim 

Recebido em 15 de julho de 2010. Aceito em 20 de janeiro de 2013. 

ABSTRACT: Seasonality in an assemblage of waterbirds in a marginal lake of the Mogi Guaçu River, São Paulo state, Brazil. I 

analyzed the seasonality of a non-passerine waterbird assemblage in a marginal lake of the Mogi Guaçu River during monthly visits 

from March 2003 to February 2004. I found a total of 15 species belonging to eight families. Ardeidae (five species) and Anatidae 

(four) were the families with the highest number of species. There was no significant variation in the number of species between the 

rainy and dry seasons. However, the number of recorded individuals approximately doubled during the rainy season. There was high 

rate of temporal succession among species throughout the year, because more than half of the species were recorded only once during 

the year. Vanellus chilensis (Molina, 1782) and Amazonetta brasiliensis (Gmelin, 1789) were the commonest species, but they were not 

considered residents. Apparently the occurrence of waterbirds at the study sites may be primarily related to their foraging strategies, 

so I suggest that for any individual species there are periods of higher probability of occurrence according to both its foraging strategy 

and the local climate seasonality. 

KEY-WORDS: feeding guilds, foraging strategy, temporal succession. 

Introdução 

Aves aquáticas não-Passeriformes possuem 

adaptações biológicas e ecológicas que propiciaram uma 

grande radiação adaptativa e, consequentemente, grande 

proporção das espécies desse grupo é cosmopolita (Sick 

2001). Devido à grande capacidade de locomoção, esse 

grupo de aves realiza deslocamentos regionais e algumas 

espécies fazem migrações continentais (Sick 2001). Dessa 

forma, pequenos açudes e lagoas propiciam ambiente 

de descanso e alimentação ao longo de suas rotas de 

deslocamentos e migrações (Sick 2001). 

O deslocamento das aves aquáticas em geral pode 

ser influenciado por diversas razões, como por exemplo, 

competição (Cox 1968), abundância de presas em 

potencial como insetos e macro-invertebrados (Wolda 

1988, Marques & Vicente 1999), alterações no nível 

de água (tanto excesso quanto falta), procura de locais 

para descanso e muda de penas (Sick 2001), variações 

climáticas (Alves & Pereira 1998), sazonalidade na 

disponibilidade de hábitat em geral (Hayes & Fox 1991) e 

provavelmente a poluição. Além desses fatores que podem 

promover deslocamento espaço-temporal, é de esperar 

que as aves aquáticas com semelhantes estratégias de 

forrageamento e guildas de alimentação tendam a possuir 

períodos semelhantes de deslocamentos para os mesmos 

hábitats por fins característicos, como disponibilidade dos 

respectivos hábitats apropriados, alimentos, início e fim 

do período reprodutivo propício para cada espécie. 

Dessa forma, esse estudo enfoca uma assembleia de 

aves aquáticas em relação à variação temporal entre grupos 

com semelhantes estratégias de forrageamento e guildas de 

alimentação, e se atenta a responder as seguintes questões: 

[1] A riqueza e abundância de aves aquáticas durante o 

ano têm relação com a sazonalidade climática do período 

de estudo? Meu prognóstico é que o maior número 

de indivíduos ocorra durante a estação seca devido ao 

menor volume de água aumentar a chance de captura 

de alimentos (Alves & Ferreira 1998, Ishikawa-Ferreira 

et al. 1999, Rodrigues & Michelin 2005); [2] Caso haja 

sazonalidade, as diferentes estratégias de forrageamento 

e guildas de alimentação auxiliam no entendimento do 

padrão de distribuição encontrado? Meu prognóstico 

é que piscívoros pernaltas (Ardeidae) e piscívoros 

subaquáticos (Phalacrocoracidae e Alceadinidae) utilizem 

preferencialmente a estação seca, provavelmente devido


Paulo Rubim 

11 

à diminuição do nível de água que facilita a captura de 

alimento (Alves & Ferreira 1998), enquanto onívoros 

flutuantes (Anatidae) podem aumentar em densidade em 

relação ao aumento das chuvas e nível da água e fim do 

período reprodutivo (Sick 2001). 

Métodos 

Realizei esse estudo em uma lagoa marginal do rio 

Mogi Guaçu localizada dentro da Reserva Particular do 

Patrimônio Natural (RPPN) São Marcelo (22˚21’50”S; 

46˚58’47”W) e pertencente à empresa de papel e celulose 

International Paper do Brasil. Essa lagoa está situada à 

margem esquerda do rio Mogi Guaçu (a 20 m do rio), 

distando cerca de 20 km à montante da Estação Ecológica 

Mogi Guaçu (Fazenda Campininha) e a 15 km à jusante 

do Reservatório Usina Mogi Guaçu. Em seu entorno 

(cerca de 500 metros) há talhões de reflorestamento de 

espécies nativas em diferentes estágios de sucessão, um 

talhão de eucalipto formando assim um mosaico com 

diferentes tipos de cobertura vegetacional. 

A lagoa apresentou formato circular com cerca de 608 

m 2 de lâmina d’ água na estação chuvosa e cerca de 510 

m 2 durante a estação seca, com variação de 20 cm do nível 

da água durante o ano. Suas margens estavam desprovidas 

de macrófitas e árvores, havendo pouca vegetação rasteira 

e substrato composto por manchas de argila e seixos em 

baixa inclinação, caracterizando um ambiente bastante 

homogêneo durante o período de estudo. 

O clima para a região de Mogi Guaçu tem precipitação 

média anual de 1365 mm e temperatura média anual de 

20,6ºC entre o período de 1971 a 2003. A precipitação 

do período de estudo foi de 1411 mm e a temperatura 

média de 20,5°C. Em geral o clima para a região de Mogi 

Guaçu é caracterizado por uma estação chuvosa e quente 

(verão) de outubro a abril, com precipitação superior a 

100 mm e uma estação seca e fria (inverno) de maio a 

setembro, com precipitação em torno de 60 mm e déficit 

hídrico entre os meses de julho e agosto. Todos os dados 

climáticos foram cedidos pela estação meteorológica da 

Estação Ecológica Mogi Guaçu, que dista cerca de 20 km 

da lagoa estudada. 

Durante um ano (março de 2003 a fevereiro de 2004) 

realizei visitas mensais de cinco dias consecutivos à lagoa ao 

final de cada mês. Fiz três visitas diárias com duração de 30 

minutos cada: amanhecer (6 h às 6 h30 min), entardecer 

(18 h às 18 h 30 min) e noturna (21 h às 21 h 30 min). 

Fiz as observações com um binóculo (8 x 42) e utilizei uma 

lanterna nas observações noturnas. O maior número de 

indivíduos encontrados entre os cinco dias de cada mês foi 

considerado como a população mensal. 

Agrupei as espécies da mesma família que possuem 

estratégias de forrageamento e guildas tróficas semelhantes, 

resultando em quatro grupos de aves: Piscívoros Pernaltas 

(PP): aves pernaltas da família Ardeidae que se alimentam 

preferencialmente de peixes às margens do corpo d’água, 

sem flutuar sobre a mesma; Piscívoros Subaquáticos (PS): 

espécies das famílias Phalacrocoracidae e Alcedinidae que 

pairam ou voam sobre os corpos d’água, mergulhando e 

perseguindo ativamente peixes como principal fonte de 

alimentação; Onívoros Flutuantes (OF): aves da família 

Anatidae que flutuam sobre a água e filtram sedimentos, 

com alimentação preferencialmente baseada em algas, 

pequenos insetos e organismos que se fixam em seu aparato 

bucal adaptado à alimentação por filtração; e Insetívoros das 

Margens (IM): aves das famílias Jacanidae, Charadriidae, 

Scolopacidae e Recurvirostridae, preferencialmente 

insetívoras que forrageiam as margens dos corpos d’água, 

se alimentando geralmente de larvas, lagartas e pupas de 

insetos aquáticos que utilizam as margens de corpos d’água 

em algum período de seu ciclo de vida. 

O critério para separação entre estação seca e chuvosa 

foi baseada na pluviosidade mensal, onde a estação 

chuvosa e quente (verão) de outubro a abril (precipitação 

> 100 mm) e a estação seca e fria (inverno) de maio a 

setembro (precipitação < 60 mm). O número de espécies 

e indivíduos foram considerados nas respectivas estações 

e devido a baixa recorrência amostral não foi necessário 

realizar uma estatística robusta para visualização da 

sazonalidade, somente fenogramas foram suficientes. Para 

comparar o número de espécies e indivíduos entre estações 

utilizei o teste de Mann-Whitney (U) (Zar 1999). 

Para uma análise com toda amostragem da 

assembleia de aves aquáticas, utilizei correlação linear 

simples de Spearman (Zar 1999) onde o número total 

de espécies e indivíduos quantificados mensalmente 

(variáveis dependentes) foi correlacionado com a 

precipitação, temperatura e comprimento do dia 

(variáveis independentes) do mesmo período (mês). Cada 

variável ambiental foi correlacionada individualmente 

com cada variável dependente. Utilizei o mesmo 

procedimento entre os grupos com diferentes estratégias 

de forrageamento (PP, PS, OF e IM) e as mesmas variáveis 

independentes. Construí fenogramas com o número de 

indivíduos durante os meses do ano para visualização da 

variação temporal segundo cada categoria de estratégia 

de forrageamento. Considerei espécie residente as que 

permaneceram no lago por 10 ou mais meses do ano 

segundo Alves & Pereira (1998). 

Resultados 

Não houve diferença entre o número de espécies 

entre as estações (U = 11,5; P = 0,357), dessa forma cinco 

espécies foram encontradas somente na estação chuvosa, 

cinco na estação seca e cinco espécies permaneceram ao 

menos um mês na estação chuvosa e seca. Porém, apesar 

do teste de Mann-Whitney não indicar significância (U 


12 


Paulo Rubim 

= 22,5; P = 0, 460), quando são considerados números 

de indivíduos os meses da estação chuvosa apresentaram 

média de 10.4 aves/mês, enquanto que na estação seca foi 

de 6 aves/mês. (Tabela 1). Das 15 espécies pertencentes 

a oito famílias que utilizaram a lagoa marginal durante 

o ano de estudo (Tabela 1), mais da metade (53,3%) 

foi avistada apenas um mês durante o ano de coleta, 

caracterizando sucessivas trocas entre espécies ao longo do 

ano (Figura 1). A espécie que utilizou a lagoa com maior 

freqüência foi Vanellus chilensis (oito meses), seguida por 

Amazonetta brasiliensis (sete) e Ardea alba Linnaeus, 1758 

e Dendrocygna autumnalis (Linnaeus, 1758) (quatro meses 

cada). Actitis macularius (Linnaeus, 1766) foi a única 

espécie migratória setentrional e permaneceu três meses 

consecutivos durante a estação chuvosa. Não encontrei 

nenhuma espécie residente ou nidificando próximo ao 

Tabela 1: Composição das espécies, estratégias de forrageamento, número de indivíduos e freqüência relativa para cada espécie de ave aquática 

encontrada na lagoa marginal do rio Mogi Guaçu, Mogi Guaçu, São Paulo. PS = Piscívoro Subaquático, PP = Piscívoro Pernalta, OF = Onívoro 

Flutuante e IM = Insetívoro das Margens. Área em cinza = precipitação média superior a 100 mm. 

Table 1: Species composition, foraging strategy, number of individuals and relative frequency of encounter for each species of waterbird found in 

the marginal lake of Mogi Guaçu river, Mogi Guaçu, São Paulo. PS = Piscivorous underwater, PP = piscivorous waders, OF = Floating omnivorous 

and IM = insectivorous of margins. Gray area = average rainfall exceeding 100 mm. 

FAMÍLIA/Espécie 

Estratégias 

de captura 

mar abr mai jun jul ago set out nov dez jan fev 

freq. 

(%) 

ANATIDAE Leach, 1820 

Dendrocygna viduata (Linnaeus, 1766) OF 0 7 0 0 0 0 0 0 0 0 0 0 8.33 

Dendrocygna autumnalis (Linnaeus, 1758) OF 0 7 0 0 2 0 0 0 4 0 2 0 33.33 

Cairina moschata (Linnaeus, 1758) OF 0 0 1 0 0 0 0 0 0 0 0 0 8.33 

Amazonetta brasiliensis (Gmelin, 1789) OF 4 10 2 2 2 0 0 0 0 0 10 5 58.33 

PHALACROCORACIDAE Reichenbach, 1849 

Phalacrocorax brasilianus (Gmelin, 1789) PS 0 0 0 0 2 2 0 0 0 0 0 0 16.67 

ARDEIDAE Leach, 1820 

Butorides striata (Linnaeus, 1758) PP 0 0 0 0 0 0 0 0 0 0 0 2 8.33 

Bubulcus íbis (Linnaeus, 1758) PP 0 0 0 0 0 1 0 0 0 0 0 0 8.33 

Ardea alba Linnaeus, 1758 PP 0 0 0 1 1 1 0 0 0 0 2 0 33.33 

Ardea cocoi Linnaeus, 1776 PP 0 0 0 1 1 0 0 0 0 0 1 0 25.00 

Egretta thula (Molina, 1782) PP 0 0 0 0 0 1 0 0 0 0 0 0 8.33 

CHARADRIIDAE Leach, 1820 

Vanellus chilensis (Molina, 1782) IM 2 1 2 0 2 1 4 2 0 4 0 0 66.67 

RECURVIROSTRIDAE Bonaparte, 1831 

Himantopus melanurus Vieillot, 1817 IM 0 2 0 0 0 0 0 0 0 0 0 0 8.33 

SCOLOPACIDAE Rafinesque, 1815 

Actitis macularius (Linnaeus, 1766) IM 0 0 0 0 0 0 0 0 3 1 3 0 25.00 

JACANIDAE Chenu & Des Murs, 1854 

Jacana jacana (Linnaeus, 1766) IM 0 1 0 0 0 0 0 0 0 0 0 0 8.33 

ALCEDINIDAE Rafinesque, 1815 

Megaceryle torquata (Linnaeus, 1766) PS 0 0 0 0 0 1 0 0 0 0 0 0 8.33 

Total de espécies 2 6 3 3 6 6 1 1 2 2 5 2 

Total de indivíduos 6 28 5 4 10 7 4 2 7 5 18 7 



Paulo Rubim 

13 

lago. Não houve correlação significativa entre o número 

de espécies e indivíduos totais em relação à precipitação, 

temperatura e o comprimento do dia. 

Em relação às estratégias de forrageamento, 

somente piscívoros subaquáticos apresentaram correlação 

negativa significativa com a temperatura (r = - 0,591; 

P = 0,043). Piscívoros pernaltas pertencentes à família 

Ardeidae (cinco espécies) ocorreram com no máximo três 

indivíduos tanto durante a estação seca quanto na estação 

chuvosa, porém de forma descontínua (Figura 2a). 

Piscívoros subaquáticos de duas espécies pertencentes a 

duas famílias ocorreram somente durante a estação seca, 

com no máximo três indivíduos no mês de agosto (Figura 

2b). Onívoros flutuantes da família Anatidae (quatro 

espécies) foram os mais abundantes durante o período 

chuvoso, sendo que o máximo de indivíduos ocorreu 

durante o mês abril, com 24 indivíduos (Figura 2c). 

Insetívoros das margens (quatro espécies pertencentes a 

quatro famílias) ocorreram praticamente durante todo o 

ano, sendo ausentes somente durante os meses de junho e 

fevereiro (Figura 2d). 

Número meroÊdeÊespecies 

especies 

8 

7 

6 

5 

4 

3 

2 

1 

0 

1 2 3 4 5 6 7 8 

tempoÊdeÊpermanÿnciaÊ(meses) 

permanência FIGURA 1. Distribuição das espécies segundo o tempo de permanência 

(em meses) para a assembléia de aves aquáticas observadas na lagoa 

marginal da RPPN São Marcelo, Mogi Guaçu, São Paulo, Brasil. 

FIGURE 1. Species distributions according to occurrence at the study 

site (in number of months) for waterbird species in a marginal lake of 

the Mogi Guaçu River, São Paulo state, Brazil 

25 

20 

15 

(a) 

piscívoros pernaltas 

10 

5 

0 

número de indivíduos 

25 

20 

15 

10 

5 

0 

25 

20 

15 

10 

(b) 

(c) 

piscívoros subaquáticos 

onívoros flutuantes 

5 

0 

25 

20 

15 

10 

5 

0 

(d) 

insetívoros das margens 

mar abr mai jun jul ago set out nov dez jan fev 

2002 2003 

FIGURA 2. Distribuição anual de indivíduos de aves aquáticas agrupadas segundo sua estratégia e guilda de alimentação na lagoa marginal do Rio 

Mogi Guaçu, RPPN São Marcelo, São Paulo, Brasil. Área em cinza = precipitação média superior a 100 mm. 

FIGURE 2. Annual distribution of waterbirds following according to feeding guild in a marginal lake of the Mogi Guaçu River, São Paulo state, 

Brazil. Gray area= average rainfall exceeding 100 mm. 


14 


Paulo Rubim 

Discussão 

Pequenas variações na riqueza de espécies durante o 

ano também foram encontradas em outros dois estudos 

em ambiente lóticos no estado de São Paulo (Ishikawa- 

Ferreira et al. 1999) e Rio de Janeiro (Alves & Pereira 

1998), Minas Gerais (Rodrigues & Michelin 2005) 

e estuarino de Santa Catarina (Branco 2007). Porém 

esses mesmos estudos evidenciaram maior número de 

indivíduos durante a estação seca, discordando com 

meus dados e prognóstico inicial. Bandos de Anatídeos, 

principalmente Amazonetta brasiliensis ocorreram durante 

abril de 2003 e janeiro de 2004 foram responsáveis pelo 

aumento significativo no número de indivíduos na estação 

chuvosa. Essa espécie é uma das mais abundantes na 

Estação Ecológica Mogi Guaçu (Willis & Oniki 1981), 

que dista cerca de 20 km da lagoa do presente estudo e 

responsável pelo padrão encontrado. 

A diversidade e sazonalidade de aves aquáticas 

podem ser fortemente influenciadas de maneira geral 

pela heterogeneidade temporal de hábitat (Hayes & 

Fox 1991), configuração da paisagem (Guadagnin et al. 

2009) e mais especificamente por diferentes tipos de 

vegetação (Anderson et al. 1983). A lagoa estudada não 

possui vegetação aquática, troncos, rochas e macrófitas, 

propiciando um ambiente com pouca diversidade 

estrutural em suas margens e conseqüentemente não 

oferecem local para esconderijo ou nidificação. Portanto, 

o número de espécies encontradas pode ser reflexo da 

homogeneidade de ambientes na lagoa estudada. Por 

exemplo, os martins-pescadores preferencialmente 

empoleiram num galho a margem do corpo d’água 

antes mergulhar a perseguição da presa (Skutch 1957). 

A pequena taxa de encontro com os martins-pescadores 

provavelmente foi influenciada pela escassez de poleiros 

de pouso ou também devido à escassez de alimento. 

Actitis macularius foi a única espécie migratória 

encontrada durante o verão do período de estudo. No 

interior do estado de São Paulo, essa espécie é encontrada 

esporadicamente, sendo que a principal rota de migração 

no estado ocorre no litoral durante verão (Willis & 

Oniki 2003). Actitis macularius e Himantopus melanurus 

Vieillot, 1817 não foram encontrados em nenhum dos 

dois estudos realizados em reservatórios no estado de São 

Paulo (Ishikawa-Ferreira et al. 1999) e Rio de Janeiro 

(Alves & Pereira 1998), indicando que essas espécies não 

necessariamente utilizam grandes corpos d’água como 

área de alimentação e rotas de migrações e deslocamentos. 

Porém, Rodrigues & Michelin (2005) classificaram 

Himantopus melanurus como provável residente em lago 

em Minas Gerais. 

Apesar da baixa abundância de aves aquáticas 

encontrada, houve um significativo número de espécies 

utilizando a lagoa em pequenos períodos sucessionais, 

provavelmente para intervalos de descansos durante rotas 

maiores de deslocamentos reprodutivos e de alimentação 

das aves aquáticas da região. Piscívoros pernaltas e 

subaquáticos utilizaram a lagoa preferencialmente durante 

a estação seca, resultado similar ao encontrado por Alves 

& Pereira (1998), e concordando com meu prognóstico 

inicial. Esse resultado provavelmente é devido à menor 

quantidade de água na lagoa, facilitando a captura de 

presas. A procriação das garças geralmente ocorre no início 

ou fim da estação seca, quando o alimento para estas aves 

aquáticas é normalmente mais abundante (Sick 2001). Não 

foi observado nenhum ninhal de ardeídeo nos arredores ao 

lago, e parece que sua presença nesse período se deveu à 

diminuição do volume de água durante a estação seca. 

Onívoros flutuantes ocorreram em maior número 

de indivíduos durante a estação chuvosa, concordando 

com meu prognóstico inicial. A maioria das espécies de 

aves se reproduz durante a estação chuvosa (Sick 2001), 

e a maioria dos bandos encontrados de anatídeos durante 

o verão possuía filhotes e jovens, o que influenciou na 

abundância desse grupo durante esse período. Insetívoros 

das margens foram observados durante quase todo o ano 

devido à permanência de Vanellus chilensis por oito meses 

não consecutivos. Jacana jacana e Himantopus melanurus 

ocorreram apenas um mês ao final da estação chuvosa, 

provavelmente em movimentação à procura de sítio de 

alimentação. Actitis macularius ocorreu provavelmente 

devido a seu período de migração. Houve pouca 

coexistência temporal entre as espécies dessa categoria, 

provavelmente devido a diferentes períodos de migração 

e também possivelmente para diminuir a competição, 

já que a lagoa possui pequena área de forrageio para 

muitos indivíduos em suas margens. Riveros et al. (1981) 

encontraram diferentes áreas de forrageio entre espécies 

simpátricas em lago no Chile. Provavelmente essas 

espécies insetívoras possuem territórios diferentes para 

forrageio enquanto utilizam a lagoa no mesmo período. 

Estudos populacionais podem detalhar melhor os 

padrões de sazonalidade inter e intra-anual de um grupo 

específico, como o exemplo de Phalacrocorax brasilianus em 

ambiente estuarino (Branco 2002), que ficam obscuros em 

estudos em nível de comunidade. Os resultados indicam 

que aparentemente há alternância temporal no período de 

permanência das aves aquáticas segundo as suas respectivas 

estratégias de forrageamento e guildas tróficas, por esse 

motivo, sugiro existir períodos com maior probabilidade 

de encontro com determinada espécie focal. 

Agradecimentos 

Agradeço à International Paper do Brasil pela 

ajuda logística e autorização para trabalhar na RPPN 

São Marcelo. À Leslie Calderon pelo envio de uma das 

referências. Agradeço aos comentários de dois revisores 

anônimos que contribuíram a esse manuscrito. 



Paulo Rubim 

15 

Referências 

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seasonality of birds species in a lagoon of an urban area (Lagoa 

Rodrigo de Freitas) of Rio de Janeiro, Brazil. Ararajuba, 6: 110- 

116. 

Anderson, B. W.; R. D. Ohmart & Rice, J. 1983. Avian and 

vegetation community structure and this seasonal relationship in 

the lower Colorado river valley. Condor, 85: 392-405. 

Branco, J. O. 2002. Flutuações sazonais na abundância de 

Phalacrocorax brasilianus (Gmelin) no estuário do Saco da Fazenda, 

Itajaí, Santa Catarina, Brasil. Revista Brasileira de Zoologia, 19: 

1057-1062. 

Branco, J. O. 2007. Avifauna aquática do Saco da Fazenda (Itajaí, 

Santa Catarina, Brasil): uma década de monitoramento. Revista 

Brasileira de Zoologia, 24: 873-882. 

Cox, G. 1968. The role of competition in the evolution of migration. 

Evolution, 22: 180-192. 

Guadagnin, D. L.; Maltchik, L. & Fonseca, C. R. 2009. Speciesarea 

relationship of Neotropical waterbird assemblages in remnant 

wetlands: looking at the mechanisms. Diversity and Distributions, 

15: 319-327. 

Hayes, F. & Fox J. A. 1991. Seasonality, habitat use, and flock sizes 

of shorebirds at the Bahía de Asunción, Paraguay. The Wilson 

Bulletin, 103: 637-649. 

Ishikawa-Ferreira, L.; Ribeiro-Neto, F. B. & Höfling, J. C.. 

1999. Avifauna aquática do reservatório Salto Grande e Varjão 

de Paulínia, Bacia do Rio Piracicaba, São Paulo, Brasil: espécies 

principais e variação temporal. Bioikos, 13(1/2): 7-18. 

Marques, P. A. M. & Vicente, L. 1999. Seasonal variation of 

waterbird prey abundance in the sado estuary rice fields. Ardeola, 

46: 231-234. 

Riveros, G. G.; Serey, E. I. & Drouilly, L. P. 1981. Estructura y 

diversidad de la comunidad de aves acuaticas de la laguna el Peral, 

Chile central. Anales del Museo de Historia Natural, 14: 189-196. 

Rodrigues, M. & Michelin, V.B. 2005. Riqueza e diversidade de 

aves aquáticas de uma lagoa natural no sudeste do Brasil. Revista 

Brasileira de Zoologia, 22: 928-935. 

Sick, H. 2001. Ornitologia Brasileira. Rio de Janeiro: Editora Nova 

Fronteira. 

Skutch, A. F. 1957. Life history of the amazon kingfisher. Condor, 

59: 217-229. 

Willis, E. O. & Oniki, Y. 1981. Levantamento preliminar de aves em 

treze áreas do Estado de São Paulo. Revista Brasileira de Biologia, 

41: 121-135. 

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Claro: Editora Divisa. 

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Hall. 

Editor Associado: Luís Fábio Silveira 



March 2013 

article 

One hundred and thirty-five years of avifaunal surveys 

around Santarém, central Brazilian Amazon 

Alexander C. Lees 1,9 , Nárgila G. de Moura 1 , Christian Borges Andretti 2 , Bradley J. W. Davis 3 , Edson V. Lopes 4 , 

L. Magalli Pinto Henriques 5 , Alexandre Aleixo 1 , Jos Barlow 1,6 , Joice Ferreira 7 and Toby A. Gardner 1,6,8 

1 

Museu Paraense Emílio Goeldi, CP 399, Belém, Pará, Brazil 

2 

Coleção de Aves, Instituto Instituto Nacional de Pesquisas da Amazônia (INPA), Avenida André Araújo 2936, Bairro Petrópolis, Manaus, Amazonas 

69060, Brazil 

3 

Rua São Cosme e Damião, 247, CEP 78580-000. Alta Floresta, Mato Grosso, Brazil 

4 

Universidade Federal do Oeste do Pará CEP 68005-100 Santarém, Pará, Brazil. 

5 

INPA, Avenida André Araújo 2936, Bairro Petrópolis, Manaus, Amazonas 69060, Brazil 

6 

Lancaster Environment Centre, Lancaster University, Lancaster LA1 4YW, UK 

7 

Embrapa Amazônia Oriental Trav. Dr. Enéas Pinheiro s/nº Caixa Postal, 48 Belém, CEP 66095-100, Pará, Brazil 

8 

Department of Zoology, University of Cambridge, Cambridge CB2 3EJ, UK 

9 

Corresponding author. Email: alexanderlees@btopenworld.com 

Received on 16 July 2012. Accepted on 22 November 2012 

ABSTRACT: We present an updated annotated avifaunal checklist for the Santarém region of central Pará state, Brazil, 

an area that has one of the oldest histories of ornithological exploration in South America. We combine data from a fivemonth 

quantitative survey of the birds of the municipalities of Santarém and Belterra (east of the Tapajós River) between 

2010 and 2011 with an exhaustive search of material in museum collections worldwide and digital vouchers deposited 

online. Our own survey sampled habitats across a gradient of disturbance ranging from ‘undisturbed’ primary forest, 

through logged and burnt forest, patches of secondary forest, cattle pastures and intensive mechanized agriculture. 

Given the potential for species misidentifications in avian inventories, we paid special attention to obtaining voucher 

documentation. Here we present a collection of publicly accessible digital vouchers for all of the new species, in addition 

to providing museum catalogue numbers for all old records. We added 24 species to the regional list, principally species 

associated with anthropogenic land-uses, but also including seven species restricted to primary forest habitats which were 

missed from both recent published inventories and over the course of two centuries of intensive collecting efforts. The regional 

list now stands at 583 species for which voucher documentation is available, with an additional 26 undocumented species. Many 

of the species reported here are poorly known or represent notable range extensions, and we present new data on their status and 

distribution. 

KEY-WORDS: bird survey, Amazonia, conservation, range extension, digital voucher. 

INTRODUCTION 

The compilation of accurate biodiversity inventories 

represents a critical first step for understanding natural 

patterns of environmental heterogeneity and speciesspecific 

responses to human-induced environmental 

change. Even for birds, perhaps the best studied of the 

Neotropical biota, such inventories remain a labor 

intensive and error prone task, particularly in extremely 

diverse tropical forest regions such as the Amazon basin 

(Remsen 1994, Cohn-Haft et al. 1997). 

The Santarém region of central Pará (PA) state, 

south of the Amazon and east of the Tapajós Rivers, is 

one of the ornithologically best-studied landscapes in 

Amazonian Brazil, with a history of specimen collection 

starting from at least 1834 (Pelzeln 1871) and avian 

inventories spanning over 135 years (e.g. Allen 1876, 

Sclater & Salvin 1878, Riker 1891, Griscom & Greenway 

1941, Henriques et al. 2003). Intensive sampling effort in 

the 19 th and early 20 th centuries saw many thousands of 

specimens collected in the region, but this data has never 

been synthesized in one place. The fruits of this labour 

during this period included the discovery of several new 

birds to science including Klage’s Antwren Myrmotherula 

klagesi, Bare-eyed Antbird Rhegmatorhina gymnops and 

Point-tailed Palmcreeper Berlepschia rikeri. 

The most exhaustive contemporary inventory 

undertaken in the region - Henriques et al. (2003) -

One hundred and thirty-five years of avifaunal surveys around Santarém, central brazilian amazon 

Alexander C. Lees, Nárgila G. de Moura, Christian Borges Andretti, Bradley J. W. Davis, Edson V. Lopes, L. Magalli Pinto Henriques, Alexandre Aleixo, 

Jos Barlow, Joice Ferreira and Toby A. Gardner 

17 

focused on the terra firme forest avifauna in the Floresta 

Nacional do Tapajós (Tapajós National Forest, hereafter 

FLONA), a 560,000-ha protected area managed by the 

Instituto Chico Mendes de Conservação da Biodiversidade 

- ICMBio. Subsequent studies in the FLONA have 

investigated avian response to forest gaps (Wunderle et 

al. 2005) and reduced impact logging (Wunderle et al. 

2006, Henriques et al. 2008). Elsewhere, the savannah 

enclave of Alter do Chão has been the subject of several 

quantitative avian studies (see e.g. Sanaiotti & Cintra 

2001 and Cintra & Sanaiotti 2005). However, beyond 

the FLONA and Alter do Chão, the region has been 

relatively poorly inventoried, especially in non-forest 

landscapes. 

We carried out a five month survey of the birds 

of the municipalities of Santarém and Belterra under 

the auspices of the ‘Rede Amazônia Sustentável’ (RAS: 

www.redeamazoniasustentavel.org), a collaborative 

research initiative focused on the study of land-use 

sustainability in eastern Amazonia, involving more than 

30 institutional partners from Brazil, the UK, Australia 

and US. Coordinating institutions are the Goeldi 

Museum and Embrapa Amazônia Oriental (Belém), 

and the Universities of Cambridge and Lancaster in the 

United Kingdom. The overall aim of RAS is to contribute 

towards an improved understanding of the long-term 

environmental and socio-economic consequences of 

current land-use and land-use change processes in the 

eastern Brazilian Amazon (Gardner et al. in press). In 

this paper we present an updated and annotated species 

list derived from the avian component of RAS study 

region in the municipalities of Santarém/Belterra, our 

incidental observations from surrounding non-study 

landscapes e.g. Alter do Chão, and a critical review of old 

records, including a search of global museum holdings 

from the region. 

METHODS 

Study Landscape: climate and biophysical conditions 

Santarém has a mean annual temperature of 25°C 

and a mean relative humidity of 86%, with annual 

rainfall averaging 1920 mm and a short dry season of 2–3 

months, usually between August and October with severe 

droughts in El Niño years (Parrotta et al. 1995, Nepstad 

et al. 2002). Canopy heights of undisturbed terra firme 

forests are typically in the range of 30 to 40 m, with 

occasional emergent species up to 50 m tall. Most of the 

survey landscape is situated on a flat terrace of Tertiary 

sediments capped by the Belterra Clay Formation 

(Clapperton 1993), at least 90 m above the water level 

of the adjacent Tapajós and Amazon rivers. Regional soils 

are predominantly oxisols dominated by kaolinite clay 

minerals and free of hardpan or iron oxide concretions 

in the upper 12 m (Nepstad et al. 2002). Originally the 

survey region was entirely covered by lowland tropical 

forest. By 2008 approximately one third had been 

deforested with much of the forest outside the FLONA 

having been degraded from the impacts of logging and 

fire (RAS unpubl. data). 

At the extreme north-western point of the region 

(Figure 1), there is an enclave of about 10,000 ha of 

savannah habitat on a peninsula beyond the town of 

Alter do Chão. The vegetation here is dominated by 

an herbaceous stratum composed principally of tuftforming 

grasses (e.g. Paspalum carinatum and Trachypogon 

plumosus) and sedges (e.g. Rhyncospora hirsute) interspersed 

with patches of trees and shrubs (principally the families 

Myrtaceae and Rubiaceae) (Miranda 1993, Magnusson et 

al. 1999, Magnusson et al. 2008) and some larger forest 

fragments. The trees are short in stature, often with 

tortuous trunks, a thick cortex and leathery leaves, and 

do not form a continuous canopy. Regular semi-annual 

burning can significantly reduce the area covered by the 

common shrub species, which then become dominated 

by the grass P. carinatum (Sanaiotti & Magnusson 

1995). Such savannah formations were formerly more 

widespread; Griscom & Greenway (1941) states of the 

environs of the city: ‘the built-up part is surrounded by 

savannahs for a distance of about two kilometres, beyond 

which the dense vegetation, high and savage, begins.’ 

The northern border of the region is delimited by the 

várzea forests and associated series of sedimentary islands 

and channels resulting from constant fluvial action. 

Behind these, on clay soils, lie savannahs and open lakes, 

both of which flood seasonally. The lakes swell and retract 

according to the flood cycle, sometimes covering tens of 

square kilometers. Large grasses found on the flooded 

savannah include Echinochloa polystachya, E. spectabilis, 

Hymenachne amplexicaulis and Leersia hexandra, in 

addition to sedges such as Scirpus cubensis, Cyperus 

luzulae and Scleria geniculata. At the ecotone between the 

savannah and forest habitats dominant shrubs and small 

vines include Artemisia artemisiifolia, Ipomoea fistulosa, 

Polygonum punctatusm, Mimosa pigra, Montrichardia 

linifolia, Rhabdadenia macrostoma and Clitonia triquetum 

(Pires & Prance 1985, Daly & Mitchell 2000). 

2010-2011 survey experimental design 

To develop our sampling approach the municipalities 

of Santarém-Belterra in the region between the Tapajós 

and Curuá-Una rivers, bordered to the north by the 

Amazon river and extending approximately 140 km 

south along the BR-163 highway (Figure 1), were divided 

up into catchments of 5,000 – 6,000 ha, which were 

delineated using a digital elevation model and SWAT 

(Soil and Water Assessment Tool) for ARCGIS 9.3 


18 




(ESRI 2008). We then selected a subset of 18 catchments 

(Table 1, Figure 1) to represent a gradient of accumulated 

forest loss from 78% (28% remaining forest cover) to 

0% (100% remaining forest cover) (Figure 1). Total 

deforestation extent is correlated with many other factors 

including age of occupation, types of historical landuse 

change, road access as well as biophysical variables 

(such as topography). Once a set of candidate catchments 

was identified to capture the full deforestation gradient, 

a final selection of 18 catchments was made to ensure 

satisfactory representation of current land-use practices, 

the spatial distribution of the rural population, and major 

soil types. All landowners in each catchment were visited 

prior to any fieldwork to introduce the RAS project 

and secure permissions for surveys in private properties 

(Gardner et al in press). 

Within each catchment, we used a stratifiedrandom 

sampling design that helped ensure that sample 

data provide a representative assessment of the overall 

environmental condition. In each catchment a standard 

density (1 per 400 ha) of 300 m study transects was 

distributed across the landscape in proportion to the 

percent cover of forest (including primary and secondary 

forests) and production areas (including agriculture, 

Figure 1. A map of the municipality of Santarém illustrating major land-use types and the locations (and numbers) of the 18 study catchments. 





19 

pasture, fruticulture and silviculture) – such that if half of 

the catchment is covered by forest then it receives only half 

of the study transects. Within each of these major landuse 

categories sample transects were distributed randomly 

to increase the likelihood that we captured important 

internal heterogeneities in forest and/or production 

systems. A minimum separation distance rule of 1,500 m 

between transects was employed to minimize dependence 

between points. Where forest cover fell below 1,200 ha, 

we maintained a minimum of three sample transects in 

forest (ensuring we captured a reasonable sample of the 

state of the forest in that catchment). 

Table 1. Co-ordinates, total area and percentage forest cover (using using a 2008 Landsat-Palsar classified image courtesy of The Nature 

Conservancy) of the 18 catchments sampled during the study. 

Catchment code Latitude and Longitude of catchment centroid Catchment size (ha) % forest cover 

69 2°32’53”S; 54°40’35”W 4299 46 

81 2°37’45”S; 54°31’23”W 4659 57 

99 2°40’28”S; 54°38’44”W 4546 47 

103 2°40’30”S; 54°54’33”W 4105 39 

112 2°42’37”S; 54°28’55”W 4795 38 

125 2°45’21”S; 54°36’32”W 4852 39 

129 2°44’17”S; 54°45’57”W 4963 52 

157 2°49’8”S; 54°28’48”W 4321 81 

160 2°47’0”S; 54°51’5”W 4841 60 

165 2°49’44”S; 54°59’51”W 3447 99 

199 2°51’52”S; 54°47’58”W 3228 28 

236 2°57’50”S; 54°44’1”W 3681 63 

260 3°1’7”S; 54°52’55”W 4219 59 

261 3°1’7”S; 55°0’12”W 4654 100 

307 3°9’14”S; 54°51’27”W 3451 87 

357 3°16’50”S; 54°52’41”W 3518 67 

363 3°19’1”S; 54°58’12”W 5166 100 

399 3°27’40”S; 54°50’17”W 5215 77 

Avian Sampling 

Fieldwork by A. C. L., N. G. M., C. B. A., B. J. W. 

D. and E. V. L. was conducted from 16 October 2010 to 

8 February 2011. We conducted two repetitions of three 

fixed width (75 m) 15-minute point counts per transect 

situated at 150 m intervals along a 300 m transect. All 

point counts (PCs) were conducted by principal observers 

A. C. L., N. G. M., C. B. A. and B. J. W. D. with the 

exception of two transects carried out independently by 

E. V. L. in Catchment 236 (see Figure 1 for numbering of 

study catchments). Surveys were not carried out on days 

with persistent rain and/or strong winds. Any systematic 

effect of seasonality (presence/absence of austral/boreal 

migrants and peaks and troughs in vocalization activity) 

was minimized by systematically rotating surveys between 

catchments of varying total forest cover and between 

habitat types. 

Digital Vouchers 

We have archived digital vouchers (photo and 

sound-recording e-vouchers) on the internet to provide 

documentary evidence for all species recorded (Appendix 

1). Such vouchers are not intended to supplant traditional 

specimen vouchers (cf. Monk & Baker 2001), although 

even these can be wrongly identified, but instead are aimed 

at providing the opportunity for general peer-review, 

which is not possible if documentary vouchers such as 

archived museum skins, photographs or sound recordings 

are not also made electronically available. Minimum 

criteria for inclusion on the list include multiple sight 

records by multiple observers, of species easy to identify 

and considered to be biogeographically likely in the 

region (i.e. there are documented records at other sites 

close to the study region). Our images have been archived 

on the Brazilian avian photo archive Wikiaves (www. 


20 




wikiaves.com.br) and our sound-recordings are archived 

on the global avian sound library Xeno-canto (www.xenocanto.org). 

Recordings on both sites are searchable by the 

catalogue number provided in Appendix 1, in addition 

we also provide catalogue numbers for ‘background 

species’ on Xeno-canto recordings. Where we are unable 

to provide a voucher (4% of species) we moved the species 

to Appendix II and also provide observer(s) names and 

date and details of the sighting. 

Historical Analysis 

We provide accession numbers for voucher 

specimens of species previously collected in the region in 

Appendix 1. We compiled a list of specimens collected by 

previous fieldworkers from the Museu Paraense Emílio 

Goeldi, Belém, Brazil (MPEG) and were provided with 

digital data for the holdings of the Carnegie Museum of 

Natural History, Pittsburgh, USA (CM) and partial data 

(only non-passerines available) for the Museu de Zoologia 

Universidade de São Paulo, São Paulo, Brazil (MZUSP). 

We used the digital database Ornis http://www.ornisnet. 

org/ to search for historically-collected specimens and 

retrieved records from the American Museum of Natural 

History, New York, NY, USA (AMNH), the Academy 

of Natural Sciences, Philadelphia, PA, USA (ANSP), the 

Field Museum of Natural History, Chicago, IL, USA 

(FMNH), the Los Angeles County Museum of Natural 

History, Los Angeles, CA, USA (LACM), the Louisiana 

State University, Baton Rouge, LA, USA (LSU), the 

University of Michigan, Museum of Zoology, Ann Arbor, 

MI, USA (UMMZ) and the United States National 

Museum, Washington, D.C., USA (USNM). Collecting 

localities were located using Paynter & Traylor (1991). 

We critically reviewed specimens and solicited 

photographic documentation of any specimens deemed 

by us and independent collaborators (Curtis Marantz 

& Bret Whitney) to be biogeographically unlikely. This 

search of museum holdings was accompanied by a review 

of previous published ornithological inventories from 

the region and we also include digital vouchers of images 

and sound-recordings archived on Wikiaves and Xeno- 

Canto by non-authors separately, coupled with voucher 

numbers for sound-recordings archived at the Macaulay 

Library http://macaulaylibrary.org/ (principally by Curtis 

Marantz) of species listed in Henriques et al. (2003). 

Our taxonomy follows the checklist of Brazilian 

birds compiled by the Comitê Brasileiro de Registros 

Ornitológicos (CBRO 2011). 

RESULTS 

During our 100 days of fieldwork we recorded 427 

species in 70 families (Appendix I), of these we provide our 

own digital vouchers for 375 species (88%, 250 species 

represented by images and 266 by sound-recordings). 

Historical collecting effort in Santarém was intense; we 

located records of over 10,000 specimens of 531 species 

in 10 collections. This in addition to a significant number 

of early skins deposited at the British Museum, Tring, UK 

which are as yet undigitalised. By totaling these historical 

records (and other contemporary records supported by 

digital vouchers) we can add a further 156 species to the 

total giving a total of 583 species in 70 families. Species 

recorded by us and missed by all previous inventories 

included the expected transient or scarce resident 

waterbirds (e.g. Snowy Egret Egretta thula), potentially 

colonizing non-forest species (e.g. Plain-breasted Grounddove 

Columbina minuta), the poorly sampled nocturnal 

avifauna (e.g. Long-tailed Potoo Nyctibius aethereus but 

also that would be considered core members of the terra 

firme forest community such as Brown-banded Puffbird 

Notharcus ordii and Grey Elaenia Myiopagis caniceps. 

These latter species represent surprising omissions, but 

their canopy lifestyles probably put them ‘beyond the 

shotgun reach’ of many earlier collectors and may have 

been missed in contemporary surveys by a combination 

of local rarity and their unobtrusive habits. We retained 

one unvouchered species: Para Gnatcatcher Polioptila 

paraensis on the main list given multiple detections by 

our and past inventories; the presence of this species in 

the region is also supported by documented records from 

adjacent municipalities. 

A number of species from recent inventories or 

unpublished observations (including our own) did not 

meet our minimum criteria for inclusion in the main 

list and these records (of 26 species) are summarized 

in Appendix II. In most cases we simply consider these 

records to be unproven and are not inferring necessarily 

that an identification is certainly in error. However, in the 

case of the report of Green-barred Woodpeckers Colaptes 

melanochloros from Alter do Chão listed in Sanaiotti 

& Cintra (2001) we consider it highly likely that these 

were misidentified Spot-breasted Woodpeckers Colaptes 

punctigula which are a common resident in that region and 

absent from the list of Sanaiotti & Cintra (2001). Likewise, 

the records of Rufous-capped Motmots Baryphthengus 

ruficapillus listed in Henriques et al. (2008) appeared in 

error and referred to Rufous Motmots B. martii. 

We follow Silveira et al. (2005) in considering the 

presence of Sulphur-breasted Parakeet Aratinga maculata 

in the region as unproven. There are two specimen records 

from Santarém - one collected by E. Garbe in 1920 

(MZUSP 10644) and the other by A. M. Olalla in 1935 

(MZUSP 18451). The former is suspected as having come 

from Monte Alegre and the latter was apparently of captive 

origin (Silveira et al. 2005). In addition to these two 

specimen records, Silva & Willis (1986) reported a series 

of sight records of this species from Santarém – groups of 





21 

3, 5 and 6 in várzea forest at Maicá on 16 January 1984, 

2 feeding on small melastomataceous fruits in seasonally 

flooded forest at Rodagém, Santarém on 18 October 1984 

and groups of 3 and 5 in secondary forest at Urumari, in 

February 1985, all considered unproven by Silveira et al. 

(2005). Given that this species’ distribution has recently 

been found to be far more extensive than previously 

thought, extending east to Amapá (da Costa et al. 2011) 

and north into Suriname (Mittermeier et al. 2010), then a 

confirmed record from the south bank of the Amazon river 

seems less far-fetched than was previously considered. 

We paid particular attention to trying to validate 

historical records that were not supported by recent 

field observations and those which appeared to be 

biogeographically unlikely. At the top of this list was a 

record of Brown Tanager Orchesticus abeillei (UMMZ 

22269) collected by Joseph Steere. We were unable to 

obtain images of the specimen but this record of an Atlantic 

Forest endemic is entirely unlikely and presumably either 

refers to a misidentified or mislabeled specimen. A number 

of skins collected by A. M. Olalla from the region were 

adjudged to be likely misidentified and this proved to be 

the case on examining images of the original skins. These 

included a specimen of Semipalmated Sandpiper Calidris 

pusilla which we re-identified as Least Sandpiper Calidris 

minutilla (MCZ 173283 see separate species account 

below); a specimen of Black-bellied Antwren Formicivora 

melanogaster (MCZ 174889) which we reidentified as a 

female Rusty-backed Antwren F. rufa; and a specimen of 

Black-necked Red-cotinga Phoenicircus nigricollis (MCZ 

171158) which we reidentified as Guianan Red-cotinga 

P. carnifex. In addition we consider the identification of a 

female Thick-billed Euphonia Euphonia laniirostris (MCZ 

176604) to be improbable by range and more likely to 

relate to a Violaceous Euphonia E. violacea, separation of 

females of these two replacement species is very difficult. 

Riker & Chapman (1891) list a record of an unidentified 

Attila sp. that they considered ‘may be the as yet undescribed 

female of A. citriniventris’ [Citron-bellied Attila]. The 

specimen is deposited in the collection of the National 

Museum of Natural History (USNM 121134) and until 

recently was labeled as A. citriniventris. However, this 

would be biogeographically unlikely considering that this 

species is restricted in Brazil to the western Guianas. T. 

Chesser (in litt.) examined the specimen on our behalf 

and found the plumage to be in poor condition, stained 

by some unknown chemical, but noted that plumage 

coloration (to the extent that it can be discerned) and bill 

morphology and coloration match those of Dull-capped 

Attila A. bolivianus. Moreover, “yellow iris” is noted on 

the back of the original collector’s label; a yellowish-white 

iris is found among species of Attila only in bolivianus. An 

old specimen record of Peruvian Recurvebill Simoxenops 

ucayalae (MPEG 32018), purportedly from Santarém has 

proven rather controversial. Novaes (1978) considered 

the specimen likely mislabeled, as at the time there were 

no records from the eastern Amazon, but the species has 

subsequently been found at various disjunct locations 

in eastern Amazonia, including as close as Altamira 

(230 km south-east), so although there have been no 

subsequent records from the region this species may 

occur in (or close to) the region (Aleixo et al. 2000). 

These exceptions aside we are confident that specimens 

labeled as ‘Santarém’ were taken from our study region 

south of the Amazon River and east of the Tapajós given 

the absence of specimens of common replacement terra 

firme forest species from adjacent areas of endemism 

(such as the west bank of the Tapajós, or north of the 

Amazon). However, an element of doubt remains over 

records of the following generalist and edge species 

which are typically widespread in anthropogenic habitat 

elsewhere in Amazonia: Rusty-fronted Tody-Flcatcher 

Poecilotriccus latirostris, Euler’s Flycatcher Lathrotriccus 

euleri and Chalk-browed Mockingbird Mimus saturninus 

but which are only represented by historic specimens 

(and no contemporary observations). There remains the 

possibility that these species might have been collected 

from river-islands closer to the north than the south bank 

of the river Amazon or have simply failed to colonize terra 

firme habitats in the region. 

Our own fieldwork produced several unconfirmed 

records (Appendix II). The most notable of these were 

the multiple detections of Spix’s Guan Penelope jacquacu, 

which most contemporary distribution maps indicating 

that this species does not occur north of the Serra do 

Cachimbo (a significant faunal and floral barrier 600 km 

south of the region) in the Tapajós-Xingu interfluvium. 

However, this species was reported north of the Serra 

do Cachimbo, in Novo Progresso by Pacheco & Olmos 

(2005), has been collected 200 km SW of our region at 

Fazenda Jamanxim, Altamira, PA on 24 November 2005 

by A. A., E. Portes and M. Silva (MPEG 59303) where 

the species was also recently recorded by C. B. A. and 

A. Whittaker, suggesting that our records may not be in 

error, despite the lack of previous reports of this large and 

generally conspicuous species. 

Although not listed in Appendix II, a possible aural 

contact of Black-chested Tyrant Taeniotriccus andrei from 

secondary forest in catchment 112 is worthy of mention 

here given the lack of previous reports from the western 

half of the Tapajós-Xingu interfluvium. The distant and 

poorly heard single note contact call was only detected on 

revision of the point count recording, and therefore cannot 

be confirmed. Although Zimmer & Whittaker (2004) list 

a specimen (MPEG 49278) from ‘Novo Fazenda, Jaburu, 

Santarém, PA’ this actually refers to a bird collected at 

Fazenda Jaburu, Novo Santarém; confusion owing to a 

slightly ambiguous specimen label. Novo Santarém lies 

east of Belém, a region where T. andrei is reasonably 

common (cf. Lees & Moura 2011). 


22 




Selected species accounts for taxa of significant 

biogeographic or conservation interest recorded 

during RAS fieldwork 

Brown Tinamou Crypturellus obsoletus 

N. G. M. sound recorded several vocalising 

individuals in river-edge forest in catchment 165 

on 14 December 2010 (Moura 2010a). This species 

was unrecorded by Henriques et al. (2003), but has 

previously been collected from the region by S. M. Klages 

who obtained three individuals at ‘Colônia do Mojuy’ 

(=Mojuí dos Campos) in November 1919 (Blake 1961). 

These birds pertain to the subspecies griseiventris which 

is significantly vocally and morphologically distinct from 

other Amazonian and Atlantic Forest populations and 

might be better considered a separate species. 

Crested Eagle Morphnus guianensis 

Although recorded from the first inventory, we 

include an account for this species given the collection of 

data on the species’ breeding biology. João Batista Ferreira, 

a local landowner on whose property we had a transect 

(catchment 103), took us to see a nest of an ‘eagle’, which 

transpired to be the active nest of a pair of Morphnus 

guianensis with a dependent (circa 7 month old) juvenile 

(Andretti 2010a). The nest (Figure 2, Lees 2010a) was 

located within a patch of old secondary forest on the edge 

of the town of Belterra. The structure was quite small, 

120 cm x 105 cm and 62 cm deep, positioned 30 m 

up in a ‘morototó’ tree, family Araliaceae (Programa de 

Conservação do Gavião-real in litt. 2011). This is the first 

report of a suburban pair of Morphnus from anywhere in 

the world and only the 7 th nest of this species recorded 

from Brazil. This discovery parallels that of a suburban 

pair of Harpy Eagles Harpia harpyja in Alta Floresta, 

Mato Grosso (MT), which bred successfully for at least 

three consecutive years in a 270 ha forest fragment (Lees 

2006). These two examples illustrate how large forest 

eagles may not be prey-limited in small forest fragments, 

but are probably extremely susceptible to being hunted 

should they become accustomed to prey upon small 

livestock (Trinca et al. 2008). 

Aplomado Falcon Falco femoralis 

We first recorded this falcon in catchment 260 

where A. C. L. observed a single adult hunting over 

soy bean fields on 6 December 2010 (Lees 2010b). 

We subsequently recorded this species on a further five 

occasions including an additional two catchments (99 

and 125), all hunting over open farmland. In addition, E. 

V. L. photographed a juvenile (Lopes 2011a) at Alter do 

Chão on 6 March 2011; a location where this species has 

previously been reported by Sanaiotti & Cintra (2001), 

who suspected on the basis of a single July record that 

this species may be a migrant in the region. Considering 

our records in the austral summer, we assume this species 

to be a rare resident in the region. There is one historical 

record from the region: one (MCZ 173143) collected by 

A. M. Olalla from ‘Santarém, Tapajós river’. These records 

are apparently the only ones from central Amazonia, with 

the closest records coming from the southern savannahs 

of Guyana and Roraima (RO), 650 km NW (Robbins et 

al. 2004, Santos & Silva 2007), Vila Nova, AP, 520 km 

NE (Schunk et al. 2011), and Alta Floresta, MT, 815 km 

south (Mahood et al. 2012, Lees et al. 2013). 

Plain-breasted Ground-dove Columbina minuta 

We recorded this species on two occasions: single 

individuals photographed (Moura 2011a), and soundrecorded 

(Moura 2011b) by N. G. M. from cattle 

pasture in catchment 69 on 8 January 2011, and from a 

smallholder’s fruit farm in catchment 112 on 31 January 

2011. We are only aware of two previous reports from 

central Amazonia – an individual collected from the 

savannahs of Monte Alegre, PA (Vasconcelos et al. 2011) 

and sight records from the Juruti region, PA (Santos et 

al. 2011) but this species has been reported from several 

peri-Amazonian sites (e.g. Schunk et al. 2011, Somenzari 

et al. 2011). Our records probably relate to individuals 

colonizing anthropogenic habitats from these savannah 

enclaves rather than individuals spreading in from peri- 

Amazonian areas. We predict that this species will prove 

to be considerably more widespread in Amazonia than 

these scant records indicate. 

Hyacinth Macaw Anodorhynchus hyacinthinus 

We encountered this threatened macaw on two 

occasions from two different catchments; C. B. A. 

observed a single individual flying overhead on 17 

October 2010 in catchment 261, and B. J. W. D. and A. 

C. L. independently heard and sound-recorded a single 

passing over the canopy in catchment 363 on 23 January 

2011 (Davis 2011a). We assume that these pertain to 

wandering individuals from populations further south 

along the BR-163 (e.g. Pacheco & Olmos 2005) and 

highlight the current local rarity of the species. The 

species was formerly more widespread in the Santarém 

region; Riker (1891) obtained three specimens ‘twentyfive 

miles back from Santarém’ on 10 June 1887. 

Long-tailed Potoo Nyctibius aethereus 

We recorded this enigmatic potoo on two occasions, 

the first records from the Santarém region. C. B. A. 

sound-recorded one singing distantly (Andretti 2010b) 





23 

from catchment 261 on 20 October 2010 and B. J. 

W. D. sound-recorded one in catchment 363 on 24 

January 2011. Despite regular night-time searches (and 

fairly regular aural contacts with White-winged Potoos 

Nyctibius leucopterus) we were unable to find Rufous 

Potoo N. bracteatus in the region. The closest records of 

this latter species are one sound-recorded 200 km south 

of the region from Trairão on 7 June 2008 by C. B. A. and 

on the west bank of the lower Tapajós at Juruti (Santos et 

al. 2011) and the Reserva Extrativista Tapajós-Arapiuns 

(MPEG 72300 and 72301). 

Great Horned Owl Bubo virginianus 

E. V. L. photographed a single individual dayroosting 

on the campus of the Universidade Federal do 

Oeste do Pará on 13 October 2011 (Lopes 2011b). There 

are few records of this species from the central Amazon, 

although this species is present on savannahs in Roraima 

(Naka et al. 2006) and Suriname (Mittermeier et al. 2010). 

Streak-throated Hermit Phaethornis rupurumii 

We recorded the amazonicus subspecies of this 

hermit on eight occasions from three different (although 

geographically adjacent) catchments (99, 125 and 129); 

most of these were secondary forest sites although we also 

encountered this species in logged and burnt primary 

forest. A. C. L. located two different leks – one each in 

125 and 129 where the birds were photographed (Figure 

3, Lees 2011a) and sound recorded (Lees 2011b). This 

taxon is typically considered to be restricted to the várzeas 

of the river Amazon and its major tributaries. However our 

observations, of leks in secondary forest over 25 km from 

a major river, mirror those of Schunck et al. (2011) from 

Vila Nova, Amapá, who found this species ‘in woodlots 

and narrow riverine forest within the mosaic of savannistic 

formations of Vila Nova, distant from the widest rivers’. 

This confirms that this species has a broader tolerance of 

forest habitats than previously suspected but we cannot 

rule out that this expansion into non-riparian habitats 

may be a recent phenomenon following land-use change. 

We may have overlooked this species if present at a low 

density elsewhere in the region owing to the sympatric 

presence of as many as six species of Phaethornis hermits 

(and Glaucis hirsutus), which made identification of flythrough 

individuals at times difficult or impossible. 

Tapajós Hermit Phaethornis aethopyga 

This species, recently re-elevated to species status 

(Piacentini et al. 2009) is endemic to the Tapajós-Xingu 

interfluvium, occurring between the river Teles Pires 

and the river Amazon and was listed as Phaethornis 

Figure 2. Nest of Crested Eagle Morphnus guianensis at catchment 103 in Belterra (A. C. L.). 


24 




Figure 3. Streak-throated Hermit Phaethornis rupurumii at lek in a fragment of secondary forest (A. C. L.). 

longuemareus in Henriques et al. (2003). We found it 

to be the most common Phaethornis hermit within the 

FLONA, but to be uncommon or absent from most 

of the catchments outside of the reserve where it was 

largely replaced by Reddish Hermit Phaethornis ruber 

and P. rupurumii, although S. M. Klages collected one 

individual at Colônia do Mojuy on 27 October 1919. 

Whether this current distribution is potentially related to 

topographically-determined micro-habitat preferences or 

direct replacement by these more ruderal hermit species 

remains unclear, but on current evidence this species 

appears to be quite disturbance intolerant cf. Henriques 

et al. (2008) although also see Piacentini et al. (2009). 

Brown-banded Puffbird Notharchus ordii 

We recorded this poorly known puffbird on two 

occasions: C. B. A. tape-recorded (Andretti 2010c) one 

in catchment 399 on 1 November 2010 and saw a second 

individual in catchment 261 on 19 October 2010. C. B. 

A. also recorded this species from the region of Trairão 

where the species was recorded on four dates in September 

2009 on the Transamazônica 80 km NE of Itaituba and 

on the river Cupariri 92 km east of Itaituba (PA). This 

species is often reported as being associated with stunted 

forest on white sandy soils e.g. in Acre (Guilherme & 

Borges 2011), north-eastern Peru (Alonso & Whitney 

2003), southwestern Venezuela, and the upper river Negro 

region of northern Brazil (Zimmer & Hilty 1997) and in 

dept Pando, Bolivia (Tobias & Seddon 2007). This record 

however, coupled with others from Alta Floresta (Zimmer 

et al. 1997), Novo Progresso (Aleixo et al. 2008), the 

Juruti region of Pará (Santos et al. 2011) and Tambopata, 

south-eastern Peru (A. C. L. & A. Whittaker) reinforces 

the notion that this species may be under-recorded in tall 

stature central Amazonian terra firme forests. Vasconcelos 

et al. (2011) lists a record from the opposite bank of the 

river Amazon at Monte Alegre, PA - a female (MPEG 

4405) collected by A. Costa on 17 November which 

would be the first record of N. ordii east of the river Negro 

and north of the river Amazon. However, there is some 

uncertainty surrounding the locations of some Costa 

specimens from the region, which may have been taken 

on the south bank (F. Lima in litt.). Costa collected a 

second N. ordii specimen from Monte Cuçari on the south 

bank, seven days before collecting MPEG 4405 allegedly 

from Monte Alegre, this specimen is held in Berlin (ZMB 

311582). Given these doubts and a lack of subsequent 

records, we consider the presence of N. ordii north of the 

Amazon and east of the Negro to be unproven. 





25 

Purple-throated Cotinga Cotinga cotinga 

This spectacular cotinga was recorded on just 

two occasions: A. C. L. photographed (Lees 2010c) a 

single adult male from the LBA Tower at KM-67 on 5 

December 2010; and observed a female in the canopy of 

old secondary forest in catchment 160 on 18 December 

2010. The only other record for the region we managed 

to trace were two (USNM 120921 and USNM 120922) 

collected by C. Riker at Diamantina, one mentioned in 

Riker & Chapman (1891) as collected on 4 July 1887, 

the other listed as ‘1886’. 

Pale-breasted Spinetail Synallaxis albescens 

We recorded this non-forest spinetail from cattle 

pasture in just two transects (e.g. Lees 2011c) in two 

different catchments (129 and 157), this in sharp contrast 

to its abundance in our sister landscape in Paragominas 

where the species was a near-ubiquitous inhabitant of 

agropastoral landscapes (Lees et al. 2012). Both landscapes 

contain catchments with similar deforestation histories 

and abut areas where the species ancestrally occurred, 

so it remains unclear why the species has proliferated in 

Paragominas and not in Santarém. Aleixo et al. (2008) 

reported this species from disturbed habitats between 

Moraes de Almeida (50 km north of Novo Progresso) 

and Santarém on 11 December 2005. The only historical 

record we were able to find for the region concern a pair 

collected by S. M. Klages in April 1919, the male of 

which was later designated as the type of S. a. griseonota 

by Todd (1948). This proposed race was described as 

having a paler crown and wing-coverts and more greyish 

underparts than inaequalis, but has subsequently been 

synonymised with the latter (Remsen 2003). 

Fiery-capped Manakin Machaeropterus pyrocephalus 

We encountered this unobtrusive manakin twice: 

from catchment 157 on 2 February 2011 (A. C. L.), and 

from catchment 125 on 7 February 2011 (Davis 2011b). 

This species had been collected three times previously 

from the region: a male collected from the ‘right bank 

of the Tapajós at Santarém’ by A. M. Olalla on 19 

June 1934; and two males collected by J. M. Cardoso 

da Silva at Urumari on 10 January and 2 February 

1984. These scant records do not permit a confident 

appraisal of whether or not the lack of previous records 

from the FLONA (Henriques et al. 2003, our data) 

reflects a genuine absence from this site and other areas 

lacking sandy soils along the main Tapajós riverbank 

or difficulties in detecting the species on account of its 

relatively cryptic vocalisations and mist-net avoidance 

combined with its local rarity. 

Yellow-crowned Elaenia Myiopagis flavivertex 

We detected this flycatcher from three transects in 

two different catchments (69 and 81) between 12 and 17 

January 2011 (e.g. Lees 2011d). Myiopagis flavivertex is 

widely considered to be a specialist of várzea forests, but 

all of our records come from logged and burnt terra firme 

forest sites on the plateau, although in all cases never more 

than 5 km from the river Amazon. These records might 

either represent wandering males which have been unable 

to secure ‘high quality’ territories in adjacent várzea forests 

or alternatively indicate a potentially new trend towards 

colonisation of moderately disturbed terra firme forests. 

Gray Elaenia Myiopagis caniceps 

This canopy flycatcher was found to be an apparently 

rare member of canopy mixed-species flocks and was 

detected just six times from five different catchments 

in addition to a pair regularly present at the LBA Tower 

at KM-67 (Figure 4). This species was missed by both 

historic and recent inventories owing to its unobtrusive 

canopy habits. The taxonomy of this species is under 

investigation by C. B. Andretti and collaborators, birds 

from Santarém are of the same vocal type as other eastern 

Amazonian and Atlantic Forest populations (although 

morphologically distinct from the latter) but are very 

different from populations in south-west Amazonia and 

northern Amazonia. 

Bank Swallow Riparia riparia 

A. C. L. photographed two individuals (Lees 2011e) 

within a migrating flock of c.1000 Barn Swallows Hirundo 

rustica hawking over cattle pasture in catchment 125 on 5 

February 2011. This species is apparently rare in centraleastern 

and eastern Amazonia (Stotz et al. 1992), with 

no records from extensive surveys in the Belém centre of 

endemism (e.g. Novaes & Lima 1998, Portes et al. 2011) 

and only a single record from the Alta Floresta region 

(Lees et al. 2013), although the species was reported by 

Fávaro & Flores (2009) from the Estação Ecológica Terra 

do Meio, PA. This rarity should reinforce the notion 

that Neotropical migrant swallows are not uniformly 

distributed across the South American continent as 

illustrated in many published distribution maps and may 

be very spatiotemporally localised (cf. Remsen 2001). 

Cocoa Thrush Turdus fumigatus 

We include a species account for this taxon as it 

seems a rather odd omission from the Henriques et al. 

(2003) inventory, as it ought to be a ‘core terra firme’ 

species. However, we only recorded this species from 

three different transects in three different catchments in 


26 




Figure 4. Gray Elaenia Myiopagis caniceps photographed from the tower at KM-67 in the FLONA (A.C. L.). 

addition to a relatively confiding pair that frequented the 

LBA Base at KM-83 (Figure 5, Lees 2010d). S. M. Klages 

collected four individuals in 1919, one from ‘Colônia do 

Mojuy’ and three from ‘Santarém (Tapajós river; Right 

Bank) and Riker & Chapman (1890) collected three 

specimens and described the species as ‘common in semipalm 

growths’. 

Red-crested Finch Lanio cucullatus 

We recorded this species on two occasions from 

catchment 369, two different singing males (3 km 

apart) located on 3 December 2010 by A. C. L (e.g. Lees 

2010e). The first was singing from the edge of primary 

forest, bordered by a ploughed field and the second from 

scrubby second growth bordering primary forest. Further 

afield, C. B. A. photographed and sound-recorded two 

individuals of this species from the town of Trairão 220 

km south-west of the region on 8 and 15 June 2008. 

These records represent substantial range extensions from 

the nearest sites in Alta Floresta (Lees et al. 2013) and 

Paragominas (Portes et al. 2011, Lees et al. 2012), we 

cannot eliminate the possibility that such records might 

relate to local introductions, but considering the speed 

at which open country species have colonized much of 

the Amazon, natural colonization seems more likely (cf. 

Mahood et al. 2012). 

Historical Records 

Sharp-shinned Hawk Accipiter striatus 

Whilst searching through the catalogue of birds 

collected by S. M. Klages from the region, we came 

across a record of a female Accipiter striatus (CM 72339) 

collected at Santarém (Tapajós river; Right Bank) on 2 

May 1919 and assigned to the subspecies erythronemius. 

Accipiter striatus is unrecorded from the Brazilian 

Amazon, or indeed anywhere in lowland Amazonia, so 

given the importance of the record we solicited images of 

the original skin from S. Rogers at the Carnegie Museum. 

The images (Figure 6) confirm that the specimen pertains 

to A. striatus and can be further aged as a subadult female 

by the retained (streaked) juvenile feathers on the throat. 

This record represents the first confirmed record from 

the Brazilian Amazon. Subsequently M. Cohn-Haft (in 

litt.) collected an immature plumaged bird in savannah 

woodland on 7 May 2007 in Amazonas (AM) in the 

Madeira-Purus interfluvium on the Ramal do Mucuim, 

50 km west of Porto Velho at 8° 40’ S; 64° 25’ W. Other 





27 

Figure 5. Cocoa Thrush Turdus fumigatus at the LBA Base KM-83, FLONA forest (A. C. L.). 

Figure 6. Composite image of the first Brazilian Amazonian record of Sharp-shinned Hawk Accipiter striatus (S. Rogers copyright Carnegie 

Museum). 


28 




sight records include two undocumented sight records 

from Manaus, AM in Cohn-Haft et al. (1997) and two 

sight records from Alter do Chão on 11 and 29 November 

2000 (R. Cintra in litt.). 

Least Sandpiper (Calidris minutilla) 

A record of a ‘Semipalmated Sandpiper Calidris 

pusilla’ collected by A. M. Olalla on 18 November 1932 

(MCZ 173283) from ‘Santarém’ (Griscom & Greenway 

1941, Stotz et al. 1992) was to our knowledge the only 

documented record of this species in the interior of the 

Brazilian Amazon. We examined digital images (Figure 7) 

of the original specimen and reidentified the individual 

as a Least Sandpiper C. minutilla based on the thin, 

slightly decurved beak, extensive dark-centers to the 

mantle feathers and yellowish legs. Least Sandpiper is an 

uncommon vagrant/scarce passage migrant to the interior 

of Amazonia with documented records from MT, PA, RO 

and AM (Stotz et al. 1992). We consider Semipalmated 

Sandpiper to be an unproven vagrant to Amazonia and 

any future reports should preferably be documented with 

high quality digital images. 

Gull-billed Tern Gelochelidon nilotica 

The only record that we can trace for the region 

concerns a single breeding-plumaged adult photographed 

by Kurazo Okada (Aguiar 2010) at the Lago do Maicá on 

31 July 2010. The status of this species in the interior of 

the Amazon basin is unclear, but circumstantial evidence 

suggests that this species maybe a regular seasonal visitor 

(breeder?) along the river Amazon. For instance, Kirwan 

et al. (2012) recorded four individuals of Gelochelidon 

nilotica associating with a mixed colony of Large-billed 

Terns Phaetusa simplex and Black Skimmers Rhynchops 

niger and exhibiting indications of breeding on the Ilha 

da Benta, Itacoatiara, Amazonas state (c.400 km WSW 

of Santarém) on 21–22 November 2011. Closer to the 

study region, G. M. Kirwan and C. F. Collins observed 

one midstream in the river Amazon c.20 km west of 

Monte Alegre, Pará, on 8 December 2005 (Kirwan et al. 

2012). Further afield, this species has been collected from 

Marajó Island (Henriques & Oren 1997) and we (A. C. 

L. and N. G. M.) have recorded flocks of this species 

on the Pará coast at Salinópolis, Bragança and Augusto 

Corrêa (e.g. Lees 2011f). 

Figure 7. Composite image of Least Sandpiper Calidris minutilla originally identified as Semipalmated Sandpiper Calidris pusilla (J. Trimble, 

copyright Museum of Comparative Zoology, Harvard University). 





29 

Scaled Ground-cuckoo Neomorphus squamiger 

The type series of the micro-endemic Neomorphus 

squamiger comes from Colônia do Mojuy by S. M. Klages 

– four individuals (two males and two females) collected 

on three dates in October and November 1919. Klages, 

in Todd (1926) remarked of the habitat preferences of 

this taxon: “It lives on or near the ground in the dense forest, 

where it accompanies the hunting ants, and is rare so far 

as my experience goes. It was never met with in the littoral 

area, nor yet in the contiguous forested mesa, but only upon 

penetrating back into the more elevated Mojuy district. We 

sought for it in vain along the Tapajós.” Subsequently A. M. 

Olalla collected two (MCZ 173562 and MCZ 173563) 

at Tauary, 39 km south-west of Santarém and alongside 

the Tapajós. We know of no subsequent reports for the 

region. Although we have no evidence for its continued 

persistence within the FLONA, we assume that the 

species is likely still extant there in more isolated regions 

and likely also persists in extensive areas of unsurveyed 

upland forest in the east of the region. Elsewhere, C. B. 

A. briefly observed one at Trairão (PA) on 14 September 

2009 following a large understorey mixed species bird 

flock in selectively-logged forest. The absence of a breast 

band was noted and the bird was observed removing loose 

bark from a decomposing fallen tree. 

Pavonine Quetzal Pharomachrus pavoninus 

One (MCZ 173835) was collected by A. M. Olalla at 

Tauary and has apparently been overlooked in subsequent 

publications. The nearest records from the Tapajós-Xingu 

interfluvium were made by Pacheco & Olmos (2005) at 

Vicinal Progresso (07°10’S; 55°06’W), 30 km SSE from 

Novo Progresso, PA (440 km south of Santarém) on 16 May 

2002 and Aleixo et al. (2008) recorded this species from 

the Floresta Nacional de Altamira, near Moraes Almeida 

(PA) in December 2005 (370 km south of Santarém). The 

south-central FLONA probably represents the northern 

limit of the range for a species which generally occurs at 

low density throughout its range. 

Red-billed Scythebill Campylorhamphus trochilirostris 

Two specimens collected by S. M. Klages from 

Santarém (Tapajós river; Right Bank) in “swamp forest” 

on 26 March (CM 71504) and 13 June (73210) 1919 

were originally identified as C. procurvoides multostriatus 

by Todd (1948), but later re-identified as C. trochilirostris 

snethlageae by A. A. upon direct examination of the 

specimens involved and comparison with dozens of 

Campylorhamphus specimens from several collections. 

Both specimens from Santarém possess the typical brickreddish 

hue on the underparts distinguishing the várzea 

specialist C. t. snethlageae (Zimmer 1934), rather than 

the distinct brownish olivaceous, which characterizes the 

underparts of C. procurvoides populations of Santarém 

found exclusively in upland terra firme forest. Despite 

Todd’s misidentification, Klages himself had noticed 

that those two Santarém specimens collected in várzea 

belonged to a different taxon than the Campylorhamphus 

found in nearby upland terra firme forest as shown by 

his field notes, transcribed as follows: “The birds with the 

serial number 2436 were collected in the upland forest. I 

consider this series to be different from series 2401”. Both 

Santarém specimens mentioned above belong to Klages’ 

series 2401, whereas all 2436 series birds included only 

specimens of two C. procurvoides taxa associated with 

terra firme: multostriatus and notabilis (A. A. pers. obs). 

Klages could distinguish those two sympatric (but not 

syntopic) species of Campylorhamphus from Santarém 

mainly by their bill color, still well preserved shortly after 

collection, as indicated by his field notes: “This form with 

the redder h. (unreadable) and less deeply curved bill seems 

to be restricted to the swampy-forest”. 

Zimmer’s Woodcreeper Dendroplex kienerii 

S. M. Klages collected four individuals of this 

seasonally-flooded forest (várzea and igapó) specialist 

between 24 March and 8 April 1919 from Santarém 

(Tapajós river; Right Bank) and A. A. and J. D. Weckstein 

collected two females and one male on 22 July 2000 11 

km south east of Santarém, in tall forest at Lago do Maicá 

(MPEG 55159, 55160, 55290). The distribution of this 

woodcreeper seems confined mostly to western Amazonia 

and the Negro river basin, with the easternmost records 

coming from the vicinity of Santarém. 

White-eyed Tody-tyrant Hemitriccus griseipectus 

S. M. Klages collected one male (Figure 8, CM 

74717) as ‘Hemitriccus zosterops’ at Colônia do Mojuy on 

1 November 1919. At the suggestion of B. M. Whitney 

we solicited images of the skin to check the identification 

and on comparison with skins of all Amazonian 

Hemitriccus and Lophotriccus species can confirm that the 

identification is correct (identification also independently 

checked by M. Cohn-Haft) and we have no reason to 

doubt the provenance of the skin. We do not believe we 

missed H. griseipectus during our own surveys, the voice 

of which all observers are familiar, and suggest that this 

species may be restricted to tall terra firme only in the east 

of the region and its distribution may be associated with 

as yet undiagnosed topographical factors. The nearest 

records of this species come from the FLONA do Trairão 

90 km east of Itaituba (C. B. A. unpubl. data). There are no 

confirmed records of Snethlage’s Tody-tyrant Hemitriccus 

minor from any sites in the Tapajós-Xingu interfluvium 

north of the Teles Pires river (Cohn-Haft 2000). 


30 




Figure 8. Composite image of the only regional record of White-belied Tody-tyrant Hemmitriccus grisepectis (S. Rogers copyright Carnegie 

Museum). 

‘Trail’s Flycatcher’ Empidonax traillii/alnorum 

An Empidonax flycatcher (Figure 9) was collected by 

G. P. Silva at Vila Mojuí dos Campos, Estrada do Palhal 

km 5 on 24 February 1978. This individual (MPEG 

32320), was identified as Willow Flycatcher Empidonax 

trailli by E. Eisenmann and A. R. Phillips (Sick 1985), the 

first and only Brazilian record of this species. However, 

without comment the same record is listed as Alder 

Flycatcher Empidonax alnorum in Stotz et al. (1992) and 

again in Vasconcelos et al. (2008). This has created some 

confusion in the subsequent literature – for instance 

Silva (2011) lists February records for both species for 

Santarém based on different sources. We re-examined 

the specimen (aged as a first winter based on prominent 

growth-bars on the tail) but unfortunately its biometrics 

fell within the range of overlap in the discriminant 

formulas of Pyle (1997) so robust identification will have 

to await molecular testing (A. C. L., A. A, G. Thom in 

prep.). Vasconcelos et al. (2008) list just three records 

of Empidonax alnorum, the aforementioned Santarém 

record, a singing bird at Manaus, AM on 15 December 

1984 (Stotz et al. 1992) and an unsexed individual 

(DZUFMG 4580) collected by M. F. Vasconcelos on 19 

November 2005 in the Pantanal at Fazenda Figueirinha 

(Corumbá municipality) MS. Additional records include 

an individual seen and sound-recorded (ML 117234) by 

Curtis Marantz at Igarape Crajari, AM on 5 April 1997, 

a female sound-recorded and collected by M. Cohn-Haft 

at Igrapé Craiata, 9 km ESE of Benjamin Constant AM 

on 5 April 1991, a male collected at Feijó, Envira river, 

Locality Novo Porto, Fóz do Igarapé Paraná do Ouro, 

AC by E. Guilherme and N. S. Brígida on 20 November 

2011 and one collected by E. Guilherme and P. Maurício 

at Manoel Urbano, BR 364, Seringal “Sardinha”, AC on 

10 November 2004. 

Gray-cheeked Thrush Catharus minimus 

G. P. Silva collected one specimen (MPEG 47943, 

Figure 10) at km-84 of the BR-163 on 15 December 

1972 and LMPH captured one individual in the FLONA 

on 20 March 2000. Stotz et al. (1992) considered this 

species to be ‘almost completely unknown from south of the 

Amazon’. The 1972 record is the first from the southern 

Brazilian Amazon. Outside of our region, subsequent 

southern Amazonian records include one collected by G. 

P. Silva from the Sena Madureira (AC) on 4 November 

1976 (Novaes 1978), and a sight record from Alta Floresta 

(MT) by A. Lang on 12 December 2002 (Lees et al. 2013). 

DISCUSSION 

This updated checklist provides a solid baseline for 

future quantitative studies and we believe that the list 

covers all core members of the regional avifauna. However, 

we anticipate that the list will continue to increase in size 

as new open-habitat colonizers, migrants and vagrants are 

added, especially considering the colonization possibilities 





31 

Figure 9. Composite image of ‘Trail’s Flycatcher’ Empidonxa trailli/alnorum (A. C. L. copyright Museu Paraense Emilio Goeldi). 

Figure 10. Gray-cheeked Thrush Catharus miniimus collected on 15 December 1972 (A. C. L. copyright Museu Paraense Emilio Goeldi) 


32 




afforded for non-forest species following extensive habitat 

conversion (Lees & Peres 2006, Mahood et al. 2012) 

and even the periodic incursion of pelagic vagrants 

into Amazonia (cf. Teixeira et al. 1986). The region is 

particularly rich in boreal migrant and vagrant passerines 

for a central Amazonian site with 12 species recorded, 

perhaps indicating that the Tapajós may function as a 

migration corridor for boreal migrants. However, species 

richness for shorebirds is quite low, with notable omissions 

including Greater Yellowlegs Tringa melanoleuca and 

White-rumped Sandpiper Calidris fuscicollis, more intense 

surveys of suitable habitats at peak migration times will 

no doubt plug these gaps in the pool of expected species. 

Our own fieldwork did not focus on river island and 

várzea habitats which are regionally of high conservation 

importance, recognized in the Important Bird Area PA04 

‘Várzeas de Monte Alegre’ which includes parts of the 

municipalities of both Santarém and Belterra (De Luca 

et al. 2009), although historical collecting effort in these 

areas was quite intense. 

A quantitative analysis of regional beta diversity is 

beyond the scope of this paper, but it is evident that even 

among least disturbed terra firme forests of the region 

there is considerable heterogeneity, probably driven by 

topographic and edaphic factors and resulting in a patchy 

distribution for many species (cf. Alonso & Whitney 2003). 

Nearly two hundred years of fieldwork have failed to find 

within the study region many terra firme forest bird species 

known from the Tapajós-Xingu interfluvium as close as 

Trairão 200 km SW of the region. These apparently absent 

species include Collared Trogon Trogon collaris, Whitebrowed 

Antbird Myrmoborus leucophrys, Black-throated 

Antbird Myrmeciza atrothorax and Striped Woodhaunter 

Hyloctistes subulatus, which probably reflects different 

forest physiognomies between these adjacent regions. 

This turnover is also reflected in the absence of records of 

Golden Parakeet Guaruba guarouba (Laranjeiras & Cohn- 

Haft 2009) and documented records of both Band-tailed 

Antbird Hypocnemoides maculicauda and Speckled Spinetail 

Cranioleuca gutturata (B. Whitney in litt.) from the southern 

boundary of the FLONA, but outside of our study region. 

These absences also illustrate that published distribution 

maps for many Amazonian bird species are very liberal, as 

they are frequently based on the extent of occurrence, while 

the actual area of occupancy for many species is far smaller 

as they are extremely patchily distributed even with the 

same interfluvium (cf. Gaston & Fuller 2009). 

Santarém has one of the longest histories of 

ornithological fieldwork in the Brazilian Amazon; that our 

own fieldwork added core terra firme birds to the regional 

list is testament to the low population density and patchy 

distribution of many rarer taxa, and the importance of 

thorough familiarity with vocalizations of such species 

which may be easily missed in rapid inventories or by 

inexperienced observers Modern avian surveys (sensu 

Aleixo 2009) are an invaluable tool for uncovering true 

biogeographic patterns, and forming robust baselines 

for conservation policies, and should include as much 

accessible documentary evidence as possible to allow for 

general peer review (Lees et al. 2012). 

ACKNOWLEDGEMENTS 

We are grateful to the following for financial 

support; Instituto Nacional de Ciência e Tecnologia 

– Biodiversidade e Uso da Terra na Amazônia (CNPq 

574008/2008-0), Empresa Brasileira de Pesquisa 

Agropecuária – Embrapa (SEG: 02.08.06.005.00), 

the Coordenação de Aperfeiçoamento de Pessoal de 

Nível Superior-Capes, the UK government Darwin 

Initiative (17-023), The Nature Conservancy, and 

Natural Environment Research Council (NERC) (NE/ 

F01614X/1 and NE/G000816/1). We also thank the 

farmers and workers unions of Santarém, Belterra and 

Paragominas and all collaborating private landowners 

for their support and The Nature Conservancy for access 

to detailed land-use maps of the municipality. We thank 

the Large-scale Biosphere-Atmosphere Experiment in 

Amazonia (LBA) for logistic support. We are indebted to 

support from our field team including Renilson Menezes 

de Freitas, Eucielde Pantoja de Oliveira, Gilson de Jesus 

Oliveira, Jony Martins Oliveira, and Manoel Aviz do 

Nascimento. We thank Fatima Lima (MPEG), Jeremiah 

Trimble (MCZ), Luís Fábio Silveira (MZUSP), Stephen 

Rogers (CM) Stephen Rogers, Terry Chesser (USNM) 

and Thomas Trombone (AMNH) for supplying the 

digitised data and/or images from their respective museum 

collections. We thank Mario Cohn-Haft, Curtis Marantz, 

Jeremy Minns, Bret Whitney, Andrew Whittaker for 

details of unpublished records. A. A. is supported by a 

productivity fellowship from CNPq. This paper is #13 in 

the Rede Amazônia Sustentável publication series. 

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Associate Editor: Marcos Pérsio Dantas Santos 


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Appendix 1 

List of 583 species recorded from the Santarém-Belterra region, south of the Amazon and east of the Tapajós (PA, Brazil). Inventories are as follows: 1= this study (* 

denotes if recorded during quantitative fieldwork), 2 = Henriques et al. 2003, 3 = Sanaiotti and Cintra (2001). Photo reference and sound reference numbers are searchable 

in the online databases of www.wikiaves.com.br (WA), www.xeno-canto.org (XC) and the Macaulay Library http://macaulaylibrary.org/ (ML). Initials given after online 

voucher numbers are those of non-author contributors, photographers: DO = D. Oliveira, DLF = Diogo Lagroteria Faria, FG = Felipe Gomes, FS = Francisco Sérgio, FP 

= Frederico Pereira, GL = Gilmar Leal, HGS = Helena G. Salgado, IT = Ian Thompson, IM = Ingrid Macedo, IG = Ivo Ghizoni-Jr, JAA = J Augusto Alves, KO = Kurazo 

Okada, LATB = Luiz Álvaro Toledo Barros, RC = Robson Czaban, TD = Tulio Dornas, VH = Valdir Hobus and sound-recordists: CM = Curtis Marantz, JM = Jeremy 

Minns, PI = Phyllis Isler, Sidnei Dantas. Accession numbers are presented for species previously collected in the region and housed at the American Museum of Natural 

History, New York City, USA (AMNH), the Academy of Natural Sciences, Philadelphia, USA (ANSP), the Carnegie Museum of Natural History, Pittsburgh, USA (CM), 

the Field Museum of Natural History, Chicago, USA (FMNH), the Los Angeles County Museum of Natural History, Los Angeles, USA (LACM), the Louisiana State 

University Museum of Natural Science, Baton Rouge, USA (LSUMZ), the Museu Paraense Emilio Goeldi, Belém, Brazil (MPEG), the Museu de Zoologia Universidade de 

São Paulo, São Paulo, Brazil (MZUSP), the University of Michigan Museum of Zoology, Ann Arbor, USA (UMMZ) and the United States National Museum, Washington, 

USA (USNM). Taxonomy and nomenclature follows CBRO (2011). 

Family / species Inventories This study Previous fieldwork 

XC foreground XC background Wikiaves Specimen Wikiaves Photographer Sound Recordist 

TINAMIDAE 

Tinamus tao 1*,2 XC91214 MZUSP 10583 ML114917 CM 

Tinamus guttatus 1*,2 XC94649 CM 74874 ML115028 CM 

Crypturellus cinereus 1*,2 XC90693 XC91205 

Crypturellus soui 1*,2 XC90703 XC90764 CM 72221 ML117119 CM 

Crypturellus obsoletus 1 XC94679 CM 74876 

Crypturellus undulatus 1,3 XC94878 CM 78240 

Crypturellus strigulosus 1*,2 XC91207 XC91203 CM 78199 

Crypturellus variegatus 1*,2 XC90705 XC94871 MPEG 56038 

Crypturellus parvirostris 1*,2 XC94650 XC94670 MPEG 47652 

ANHIMIDAE 

Anhima cornuta CM 73737 

ANANTIDAE 

Sarkidiornis sylvicola CM 73268 

Cairina moschata 1 WA580720 UMMZ 27966 WA189071 KO 

Amazoneta brasiliensis 1* WA426586 MZUSP 20920 WA559786 VH 

Dendrocygna autumnalis 1,3 WA429940 CM 73634 WA576641 IT 





37 


CRACIDAE 

Ortalis motmot 1*,2,3 XC94608 WA340078 MZUSP 46267 

Penelope superciliaris 1*,2,3 CM 75036 

Penelope pileata 1* XC91206 MZUSP 21058 

Aburria cujubi 1*,2 WA500190 MZUSP 20832 

Pauxi tuberosum 1*,2 XC91214 WA675633 MZUSP 20467 

ODONTOPHORIDAE 

Odontophorus gujanensis 1*,2 XC94805 MZUSP 10602 

PODICIPEDIDAE 

Tachybaptus dominicus 1 WA500150 MCZ 173025 

CICONIIDAE 

Ciconia maguari MCZ 23047 

PHALACROCORACIDAE 

Phalacrocorax brasilianus 1 MZUSP 21925 WA185783 KO 

ANHINGIDAE 

Anhinga anhinga 1 WA580721 MCZ 173021 WA98813 JAA 

ARDEIDAE 

Tigrisoma lineatum 1 CM 72000 WA100655 JAA 

Agamia agami MZUSP 35885 

Cochlearius cochlearius 1 WA359482 MZUSP 35886 

Zebrilus undulatus 1 CM 75076 

Botaurus pinnatus MCZ 173069 

Ixobrychus exilis CM 72388 

Nycticorax nycticorax CM 78113 WA183342 KO 

Butorides striata 1,2 WA580731 MZUSP 61789 WA77559 LATB 

Bulbucus ibis 1,3 WA372477 MPEG 36473 WA185772 KO 

Ardea cocoi 1 MCZ 23190 WA74313 LATB 

Ardea alba 1* WA329322 LACM 34344 WA557916 VH 

Pilherodius pileatus 1,2 MZUSP 46199 

Egretta thula 1 WA329325 

Egretta caerulea WA675594 WA183310 KO 


38 





THRESKIORNITHIDAE 

Mesembrinibis cayennensis 1* WA588367 MCZ 173072 

Theristicus caudatus 1* WA366370 WA205442 KO 

CATHARTIDAE 

Cathartes aura 1*2,3 WA505835 CM 78110 WA242292 IT 

Cathartes burrovianus 1*,3 WA359441 WA189075 KO 

Cathartes melambrotus 1*2,3 WA333384 

Coragyps atratus 1*2,3 WA333385 CM 78109 WA242294 IT 

Sarcoramphus papa 1,2 AMNH 285739 

PANDIONIDAE 

Pandion haileautus 1,3 WA357402 MCZ 173117 WA549592 VH 

ACCIPITRIDAE 

Leptodon cayanensis 1,2 MCZ 173091 WA918207 RC ML115074 CM 

Chondrohierax uncinatus 1*,2 WA435547 MCZ 173092 

Elanoides forficatus 1*2,3 WA429985 CM 73057 

Gampsonyx swainsonii 1 WA629547 MPEG 34430 

Harpagus bidentatus 1*,2 MPEG 15342 WA320216 FG 

Harpagus didon MPEG 35598 

Accipter superciliosus 1*2 WA361613 CM 72934 

Accipter striatus 3 CM 72517 

Accipter bicolor CM 72339 

Ictinia plumbea 1,2,3 WA936127 

Busarellus nigricollis 1 FMNH 257783 WA185781 KO 

Rostrhamus sociabilis WA435213 FMNH 257787 

Geranospiza caerulescens WA645500 FMNH 257800 

Buteogallus schistaceus FMNH 101510 

Heterospizias meridionalis 1*,3 WA431330 WA180997 KO 

Urubutinga urubitinga 1*,2,3 WA514779 FMNH 257765 

Rupornis magnirostris 1*,2,3 XC94809 WA329171 MZUSP 10134 WA206722 KO ML117158 CM 

Geranoaetus albicaudatus 1*,3 WA443906 MCZ 173102 

Pseudastur albicollis 1*,2 WA432803 MPEG 13772 

Leucopternis melanops MZUSP 46240 





39 


Leucopternis kuhli 1*,2 XC92080 XC94851 WA514724 FMNH 101120 WA320489 FG 

Buteo nitidus 1*,2,3 XC95086 XC94874 WA491528 MPEG 35598 

Buteo brachyurus 1*,3 WA499991 

Buteo swainsoni USNM 121073 

Morphnus guianensis 1 WA356485 

Harpia harpyja 1 WA616225 MPEG 1855 

Spizaetus tyrannus 1*,2 XC96328 XC96343 WA329317 FMNH 101130 WA320256 FG XC85417 JM 

Spizastur melanoleucus 1*,2 WA467097 MCZ 173114 

Spizaetus ornatus 1*,2 XC96376 MCZ 173115 XC85417 JM 

FALCONIDAE 

Daptrius ater 1*,2 WA347314 CM 74791 

Ibycter americanus 1*,2 XC95591 WA356727 CM 72788 ML115015 CM 

Caracara plancus 1*,2 WA580755 ANSP 76478 

Milvago chimachima 1*,2,3 WA500112 MZUSP 35888 WA552031 VH 

Herpetotheres cachinnans 1*,2,3 WA516301 

Micrastur ruficollis 1*,2 XC90680 XC90687 

Micrastur mintoni 1*,2 XC95106 XC90680 MZUSP 18030 

Micrastur mirandollei 1* XC94623 MZUSP 10862 

Micrastur semitorquatus 1*,2 WA346345 CM 74614 ML114997 CM 

Falco rufigularis 1*,2,3 WA447466 CM 75002 

Falco deiroleucus CM 73801 WA632317 IG 

Falco femoralis 1*,3 WA500144 MCZ173143 

Falco peregrinus WA325212 FP 

EURYPYGIDAE 

Eurypyga helias WA517296 CM 72364 

ARAMIDAE 

Aramus guarauna WA583437 CM 73676 WA185149 KO 

PSOPHIDAE 

Psophia dextralis XC96488 WA359490 CM 75034 

RALLIDAE 

Aramides cajanea 1*,2 XC94871 CM 72145 

Amaurolimnas concolor CM 71647 


40 





Laterallus viridis 1*,2 XC94670 MZUSP 35891 ML117040 CM 

Laterallus exilis 1* MCZ 173214 

Neocrex erythrops 1* XC91474 MPEG 74208 

Gallinula galeata MZUSP 22636 

Porphyrio martinica 1*,2 WA500120 CM 71555 WA104023 GL 

Porphyrio flavirostris CM 71615 

HELIORNITHIDAE 

Heliornis fulica WA362963 MZUSP 35892 

CHARADRIIDAE 

Vanellus cayanus 1 WA333899 CM 73189 WA99261 JAA 

Vanellus chilensis 1* WA583439 CM 73677 WA182115 KO 

Pluvialis dominica LACM 34401 WA757451 HGS 

Charadrius collaris 1 WA467107 MZUSP 35894 WA546923 VH 

RECURVIROSTRIDAE 

Himantopus mexicanus WA431336 WA183311 KO 

SCOLOPACIDAE 

Gallinago paraguaiae MPEG 36472 WA205424 KO 

Bartramia longicauda MZUSP 35895 

Actitis macularius WA357318 CM 74312 

Tringa solitaria 1 WA357320 MZUSP 35896 WA242305 IT 

Tringa flavipes WA508864 CM 73689 WA182114 KO 

Calidris melanotos MCZ 173293 WA189072 KO 

Calidris minutilla MCZ 173283 

JACANIDAE 

Jacana jacana WA511911 MZUSP 3376 WA205425 KO 

STERNIDAE 

Sternula superciliaris WA435379 CM 78510 

Phaetusa simplex WA432695 CM 73739 WA549214 VH ML47954 PI 

Gelochelidon nilotica WA176659 KO 

RYNCHOPIDAE 

Rynchops niger WA357316 MCZ 23042 WA559241 VH 





41 


COLUMBIDAE 

Columbina passerina 1*,2,3 XC94650 WA500208 MPEG 17611 WA550785 VH 

Columbina minuta 1* XC94621 XC94956 WA441603 

Columbina talpacoti 1*,2 WA333907 CM 73312 WA319722 FG 

Claravis pretiosa 3 MPEG 47665 ML117176 CM 

Columba livia 1* WA333902 

Patagioenas speciosa 1*,3 MZUSP 10607 

Patagioenas cayennensis 1*,3 WA372475 MZUSP 35897 WA205427 KO 

Patagioenas plumbea 1*,2 XC94779 XC94851 AMNH 285541 ML115068 CM 

Patagioenas subvinacea 1*,2 XC95107 CM 74472 XC87137 JM 

Zenaida auriculata 1*,3 WA359445 MPEG 17612 WA547583 VH 

Leptotila verreauxi 1*,2 XC94620 WA505858 CM 72540 

Leptotila rufaxilla 1*,2,3 XC95111 XC92089 CM 73078 

Geotrygon montana 1*,2,3 XC95572 MZUSP 10606 WA320511 FG 

PSITTACIDAE 

Anodorhynchus hyacinthinus 1* XC91202 MCZ 173413 

Ara ararauna CM 72105 

Ara macao 1*,2 WA522295 ML115115 CM 

Ara chloropterus 1*,2,3 XC95108 MCZ 173415 

Ara severus 1*,2 XC90773 XC90776 WA444684 MZUSP 11834 WA185148 KO 

Orthopsittaca manilata 1* XC94856 CM 72174 

Aratinga leucophthalma 1*,2 XC95676 XC96344 WA426594 CM 74387 WA319711 FG 

Aratinga aurea 1*,3 XC94618 WA357380 MPEG 28147 WA180979 KO ML117043 CM 

Pyrrhura amazonum 1*,2 XC94954 WA356522 MZUSP 3416 XC85381 JM 

Forpus passerinus 1* WA467169 MPEG 2330 WA205438 KO 

Brotogeris versicolurus 1*,3 XC94874 XC96344 WA351745 MZUSP 3410 

Brototeris chrysoptera 1*,2 XC94955 XC87290 WA351744 MPEG 8890 

Brotogeris sanctithomae WA872402 MZUSP 35909 WA183291 KO XC84943 JM 

Touit huetii 1* ML115198 CM 

Pionites leucogaster 1*,2 XC95118 CM 74836 

Pyrilia vulturina 1*,2 XC95120 MZUSP 10630 ML114929 CM 

Graydidascalus brachyurus CM 72417 


42 





Pionus menstruus 1*,2,3 XC95117 XC94832 WA352463 MZUSP 10624 

Pionus fuscus 1*,2 XC95125 WA500133 CM 74545 ML115035 CM 

Amazona festiva CM 72900 

Amazona farinosa 1*,2 XC95112 XC90706 WA356731 CM 74734 ML115064 CM 

Amazona amazonica 1*,2 XC95122 XC90773 LACM 34501 

Amazona ochrocephala 1*,2 XC94682 CM 73608 

Deroptyus accipitrinus 1*,2 XC95123 WA500189 MZUSP 10618 ML114902 CM 

OPISTTHOCOMIDAE 

Opisthocomus hoazin WA432121 MZUSP 35889 WA183343 KO 

CUCULIDAE 

Coccycua minuta 1* XC94622 WA567160 CM 72868 

Piaya cayana 1*,2,3 XC96333 WA500202 MZUSP 61865 WA247315 IT 

Piaya melanogaster 1*,2 XC96382 WA432789 MPEG 56039 

Coccyzus melacoryphus CM 73549 WA182095 KO 

Coccyzus euleri CM 72739 WA552668 VH 

Crotophaga major 1*,3 WA337955 MZUSP 35904 WA189087 KO 

Crothophaga ani 1*,2,3 XC94607 XC94648 WA500149 MPEG 17617 WA189086 KO 

Tapera naevia 1,2 MPEG 47671 

Dromococcyx phasianellus 1* XC87287 XC95171 LACM 34519 

Neomorphus squamiger CM 74616 

TYTONIDAE 

Tyto alba WA436255 MCZ 173144 

STRIGIDAE 

Megascops choliba 1*,2,3 XC94800 WA432016 CM 73578 

Megascops usta 1*,2 XC94645 MPEG 53840 

Lophostrix cristata 1*,2 CM 72585 ML114946 CM 

Pulsatrix perspicillata 1*,2 XC90764 CM 72854 

Bubo virginianus WA481116 

Strix virgata 1* XC94713 MCZ 173158 

Strix huhula 1* XC94712 

Glaucidium hardyi 1*,2 XC94683 XC94710 ML114944 CM 

Athene cunicularia 1* WA509541 





43 


Asio clamator WA357321 

Asio stygius WA583443 MCZ 173148 

NYCTIBIIDAE 

Nyctibius grandis 1,2,3 MZUSP 35913 

Nyctibius aethereus 1* XC94710 

Nyctibius griseus 1*,2,3 WA567100 CM 72237 WA49272 IM 

Nyctibius leucopterus 1*,2 XC94711 ML114943 CM 

CAPRIMULGIDAE 

Nyctiphrynus ocellatus 1*,2 XC95113 MPEG 54302 ML115108 CM 

Antrostomus rufus 1*,3 MZUSP 10894 WA631559 IG 

Antrostomus sericocaudatus 1* XC86600 MPEG 56042 ML114945 CM 

Lurocalis semitorquatus 1*,2 XC90702 

Hydropsalis leucopyga MZUSP 35916 WA756417 HGS 

Hydropsalis nigrescens 1*,2 WA576637 CM 71585 ML115079 CM 

Hydropsalis albicollis 1*,2 XC94800 CM 73263 ML115038 CM 

Hydropsalis parvula CM 73674 

Hydropsalis maculicauda CM 73816 

Hydropsalis climacocerca WA431349 CM 71658 WA98817 JAA 

Hydropsalis torquata MPEG 27368 WA25992 IM 

Chordeiles nacunda CM 73153 WA178884 KO 

Chordeiles rupestris MCZ 173600 

Chordeiles acutipennis 1,3 MPEG 37761 WA316733 DLF 

APODIDAE 

Chaetura spinicauda 1*,2 CM 74412 ML115092 CM 

Chaetura chapmanii 1* WA360041 

Chaetura brachyura 1*,2 XC94831 WA320213 FG 

Tachornis squamata 1*,2 WA573688 CM 73577 

Panyptila cayennensis 1,2 MPEG 37764 

THROCHILIDAE 

Glaucis hirsutus 1*,2,3 WA360047 MPEG 53832 

Phaethornis rupurumii 1* XC84327 WA360065 MPEG 8869 

Phaethornis aethopyga 1*,2,3 XC90519 WA358701 CM 74518 


44 





Phaethornis ruber 1* XC94882 AMNH 148269 WA206708 KO 

Phaethornis bourcieri 1*,2 MPEG 56041 

Phaethornis superciliosus 1*,2,3 XC91212 CM 74606 

Campylopterus largipennis 1,2 ML114922 CM 

Eupetonema macroura CM 78361 WA634736 IT 

Florisuga mellivora 1*,2 MPEG 53839 

Anthracothorax viridigula CM 73471 

Anthracothorax nigricollis 1*,2,3 CM 73265 

Avocettula recurvirostris 2 MZUSP 3409 ML115199 CM 

Topaza pella 1,2 XC5725 SD 

Chlorostilbon notatus 3 MPEG 8881 

Thalurania furcata 1*,2,3 WA567110 MPEG 53837 

Hylocharis sapphirina 1,2,3 WA645522 CM 72123 

Polytmus theresiae 1,3 MCZ 173823 WA185793 KO 

Amazilia versicolor MCZ 173755 

Amazilia fimbriata 1* WA584520, MPEG 35617 WA185769 KO 

Heliothryx auritus 1*,2 CM 78631 

Heliomaster longirostris 1,2 WA624861 MZUSP 3404 WA319712 FG 

Caliphlox amethystina WA625025 WA183292 KO 

TROGONIDAE 

Trogon melanurus 1*,2 XC94717 WA583441 CM 72885 ML115062 CM 

Trogon viridis 1*,2,3 XC95314 XC94623 WA500148 MZUSP 35920 WA78768 LATB ML115159 CM 

Trogon ramonianus 1*,2 WA522329 CM 74432 

Trogon rufus 1*,2 XC95308 XC91207 WA676347 MPEG 53841 ML114920 CM 

Pharomachrus pavoninus MCZ 173835 

ALCEDINIDAE 

Megaceryle torquata 1*,2 WA583444 MPEG 27312 WA325211 FP 

Chloroceryle amazona 1,2,3 WA366364 MZUSP 35922 WA98818 JAA 

Chloroceryle aenea 2,3 MZUSP 15947 

Chloroceryle americana 1*,2,3 WA583449 MZUSP 46551 WA185167 KO 

MOMOTIDAE 

Baryphthengus martii 1*,2 XC90680 WA356477 CM 75042 





45 


Momotus momota 1*,2 XC94679 WA442693 CM 74832 

GALBULIDAE 

Galbula cyanicollis 1*,2 XC95109 XC95110 CM 74550 XC4883 SD 

Galbula ruficauda WA936123 CM 71853 

Galbula dea 1*,2 WA573667 CM 75062 ML115189 CM 

Jacamerops aureus 1*,2 XC94777 XC87290 WA676330 CM 75073 ML115013 CM 

BUCONNIDAE 

Notharchus hyperrhynchus 1*,2 XC91203 WA500142 MZUSP 10683 WA320253 FG ML117113 CM 

Notharus ordii 1* XC94707 

Notharchus tectus 1*,2,3 XC91203 WA363562 MZUSP 10688 WA101535 JAA ML117107 CM 

Bucco tamatia 1*,2 WA544924 CM 71967 WA553066 VH 

Bucco capensis 1*,2 XC94709 XC94871 CM 72995 ML114990 CM 

Nystalus maculatus 1*,2,3 XC94618 WA500146 MPEG 17614 WA551718 VH ML117140 CM 

Malacoptila rufa 1*,2 XC90772 WA567157 MPEG 56044 

Monasa nigrifrons 1 WA583455 MZUSP 35926 

Monasa morphoeus 1*,2 XC95269 XC94679 WA500134 MPEG 40577 ML115175 CM 

Chelidoptera tenebrosa 1*,2,3 WA428002 MZUSP 35928 

RAMPHASTIDAE 

Ramphastos toco 1,3 WA435202 CM 74281 

Ramphatos tucanus 1*,2 XC90703 WA472581 MZUSP 82495 

Ramphastos vitellinus 1*,2,3 XC90774 WA352476 MPEG 14851 ML114981 CM 

Selenidera gouldii 1*,2 XC94803 WA871417 MZUSP 10671 

Pteroglossus inscriptus 1*,2,3 WA352331 MZUSP 3424 WA320254 FG 

Pteroglossus bitorquatus 1*,2 XC95110 WA500203 MZUSP 10659 WA49283 IM ML117120 CM 

Pteroglossus aracari 1*,2,3 XC90777 WA467112 MZUSP 10665 WA319710 FG ML114949 CM 

PICIDAE 

Picumnus aurifrons 1*,2,3 XC90709 WA349052 MPEG 53843 

Picumnus cirratus WA351754 CM 78190 

Melanerpes candidus CM 73144 ML47952 PI 

Melanerpes cruentatus 1*,2 XC95086 WA573656 CM 73063 

Veniliornis affinis 1*,2 XC95304 MPEG 36697 WA320545 FG ML114951 CM 

Veniliornis passerinus CM 72952 


46 





Piculus flavigula 1*,2,3 XC94957 WA675088 CM 75072 ML114956 CM 

Piculus chrysochloros 1* WA356507 CM 72647 

Colaptes punctigula 1,3 WA366369 MZUSP 3420 WA183309 KO 

Celeus grammicus 1*,2,3 XC91204 WA674439 MZUSP 3419 

Celeus elegans 1*,2,3 WA356077 MPEG 56045 

Celeus flavescens CM 73169 

Celeus flavus 1*,2 XC95305 WA359486 MZUSP 10708 ML115217 CM 

Celeus torquatus 1*,2 XC96148 XC96151 CM 74743 WA919233 RC ML114909 CM 

Dryocopus lineatus 1*,2,3 XC94959 WA349050 MZUSP 10716 XC87455 JM 

Campephilus rubricollis 1*2 XC95102 XC95109 WA675642 CM 72866 WA319721 FG ML114957 CM 

Campephilus melanoleucos 1*,3 XC95103 WA442184 MZUSP 35932 WA629817 IT 

THAMNOPHILIDAE 

Myrmornis torquata 2 MPEG 53917 

Pygiptila stellaris 1*2 XC95310 XC90772 CM 74493 

Microrhopias quixensis 1*2 XC94851 MPEG 53900 XC88940 JM 

Myrmeciza hemimelaena 1*2 XC95315 XC90760 WA352334 MPEG 56086 ML115203 CM 

Epinecrophylla leucophthalma 1*2 XC96451 WA356470 MPEG 56078 ML114994 CM 

Epinecrophylla ornata 1*2 MPEG 53893 

Myrmotherula brachyura 1*2 XC94887 XC90774 WA359432 MPEG 56084 

Myrmotherula sclateri 1*2 XC95307 XC90760 CM 74937 ML114962 CM 

Myrmotherula klagesi CM 78427 

Myrmotherula hauxwelli 1*2 XC90707 MPEG 56072 

Myrmotherula axillaris 1*2 XC95311 XC90704 WA621984 MPEG 53897 WA320531 FG 

Myrmotherula longipennis 1*2 XC95317 XC96303 MPEG 56071 ML115032 CM 

Myrmotherula menetriesii 1*2 XC95316 XC96455 MPEG 56074 XC88775 JM 

Myrmotherula assimilis CM 73136 ML47950 PI 

Formicivora grisea 1*,3 XC94648 XC94670 WA583458 MPEG 35616 ML117125 CM 

Formicivora rufa 3 MPEG 37766 WA639485 IT 

Thamnomanes caesius 1*2 XC94719 XC94851 MPEG 56068 ML115122 CM 

Dichrozona cincta 2 MPEG 53904 XC88789 JM 

Herpsilochmus rufimarginatus 1*2 XC95402 XC90760 WA357413 CM 74645 ML114913 CM 

Sakesphorus luctuosos WA583465 CM 72794 WA185174 KO XC87606 JM 





47 


Thamnophilus doliatus 1,3 MPEG 26699 WA634314 FS 

Thamnophilus schistaceus 1*2 XC90697 XC87290 MPEG 56062 

Thamnophilus nigrocinereus CM 72219 

Thamnophilus stictocephalus 1*,3 XC94611 WA619262 MPEG 26710 ML117126 CM 

Thamnophilus aethiops 1*2 XC95318 XC95466 MPEG 53872 

Thamnophilus amazonicus 1* CM 74955 

Cymbilaimus lineatus 1*2 XC94888 XC91205 WA356488 MPEG 56060 ML115024 CM 

Taraba major 1*2 XC94620 CM 72511 

Sclateria naevia 1*2 CM 74856 WA206694 KO 

Schistocichla rufifacies 2 CM 72556 ML115023 CM 

Hypocnemoides melanopogon 1 CM 72350 

Hylophylax naevius 1*2 XC90776 XC91214 MPEG 56093 ML115186 CM 

Hylophylax punctulatus 1*2 XC94780 XC95401 CM 74463 

Pyriglena leuconota 1*2 XC96494 MPEG 40590 ML115047 CM 

Myrmoborus lugubris CM 72226 ML47948 PI 

Myrmoborus myotherinus 1*2 XC90747 XC90773 WA675602 MPEG 56244 

Cercomacra cinerascens 1*2 XC91216 XC91204 MPEG 56064 WA185151 KO 

Cercomacra nigrescens 1*2 XC95465 WA447471 CM 74773 ML114996 CM 

Hypocnemis striata 1*2 XC96355 XC90709 WA356468 MPEG 56096 WA320252 FG ML114911 CM 

Hypocnemis hypoxantha 1*2 XC87289 XC94623 WA356117 CM 74732 ML115172 CM 

Willisornis poecilinotus 1*2 XC91222 CM 75079 

Phlegopsis nigromaculata 1*2 XC90744 XC91206 MPEG 56104 ML114942 CM 

Rhegmatorhina gymnops 1*2 XC96150 XC94872 MPEG 56102 XC90272 JM 

CONOPOPHAGIDAE 

Conopophaga aurita 1*2 XC94952 WA357416 MPEG 56105 ML114979 CM 

GRALLARIDAE 

Grallaria varia 1*2 XC94645 CM 72858 

Hylopezus macularius 1*2 XC86599 XC90705 WA357411 MPEG 56099 ML115081 CM 

Hylopezus berlepschi 1*2 XC94723 XC96341 CM 78386 XC6519 SD 

Myrmothera campanisona 1*2 XC94889 XC91202 CM 74656 ML114910 CM 

FORMICARIIDAE 

Chameaza nobilis CM 75049 


48 





Formicarius colma 1*2 XC95312 MPEG 53920 

Formicarius analis 1*2 XC95313 WA500201 MPEG 53921 

SCLERURIDAE 

Sclerurus mexicanus 1*2 XC96534 MPEG 53866 

Sclerurus rufigularis 1*2 XC96380 XC90707 MPEG 53869 

Sclerurus caudacutus 1*2 XC94774 MPEG 36465 

DENDROCOLAPTIDAE 

Dendrocincla fuliginosa 1*,2 XC94830 XC95269 MPEG 56046 ML114930 CM 

Dendrocincla merula 1*,2,3 XC94829 XC94831 MPEG 53850 ML115052 CM 

Deconychura longicauda 1*,2 XC95571 MPEG 53852 ML115073 CM 

Certhiasomus stictolaemus 1*,2 MPEG 53851 

Sittasomus griseicapillus 1*,2 XC96151 WA356030 MPEG 47735 

Glyphorynchus spirurus 1*,2 XC95678 XC90709 MPEG 56054 WA320215 FG 

Xiphorhynchus spixii 1*,2 XC94876 XC96376 MPEG 56051 ML115123 CM 

Xiphorhynchus obsoletus 1 MPEG 55293 

Xiphorhynchus guttatus 1*,2,3 XC95467 XC90740 WA676332 MPEG 56049 ML114900 CM 

Campylorhamphus procurvoides 1*,2 XC90761 MPEG 56093 

Campylorhamphus trochilirostris CM 71504 

Dendroplex picus 1*,2,3 XC94885 XC90697 WA337960 MPEG 55291 WA189088 KO 

Dendroplex kienerii MPEG 55160 

Lepidocolaptes angustirostris 1,3 WA337961 MPEG 19701 WA550043 VH 

Lepidocolaptes albolineatus 1*,2 XC96153 XC96154 WA675589 MPEG 56055 ML117115 CM 

Nasica longirostris 1 CM 73175 

Dendrexetastes rufigula 1* XC87286 XC96496 

Dendrocolaptes certhia 1*,2 XC90769 XC94679 MPEG 53855 ML114916 CM 

Dendrocolaptes picumnus 1*,2 XC90767 XC90703 WA500147 MPEG 53859 ML115181 CM 

Xiphocolaptes promeropirhynchus 1*,2 XC94715 XC95467 MPEG 47698 ML114924 CM 

Hylexetastes uniformis 1*,2 XC90740 XC90705 WA360070 MPEG 53857 ML117106 CM 

FURNARIIDAE 

Xenops minutus 1*,2 XC96457 MPEG 56057 WA320530 FG ML115050 CM 

Berlepschia rikeri USNM 109221 

Furnarius figulus 1* WA359474 CM 72394 





49 


Furnarius minor CM 72014 WA180980 KO 

Ancistrops strigilatus 1* XC95119 CM 74882 

Automolus ochrolaemus 1*,2 XC94886 XC90693 WA360092 CM 74600 

Automolus paraensis 1*,2 XC95303 MPEG 53864 ML115019 CM 

Automolus rufipileatus 1*,2 XC95468 XC96457 MPEG 53863 

Philydor ruficaudatum 1*,2 XC95121 CM 74983 

Philydor erythrocercum 1*,2 XC96455 WA500204 MPEG 56056 

Philydor pyrrhodes 1*,2 XC95124 MPEG 47737 

Certhiaxis cinnamomeus MPEG 36471 WA98816 JAA 

Certhiaxis mustelinus CM 72392 

Synallaxis albescens 1* XC87288 XC94798 CM 72311 

Synallaxis rutilans 1*,2 XC94606 CM 74663 ML114952 CM 

Synallaxis gujanensis 1*,2 XC94806 CM 72131 XC6583 SD 

Cranioleuca vulpina CM 72181 XC91265 JM 

Cranioleuca muelleri CM 71831 

PIPRIDAE 

Neopelma pallescens 1,3 WA432109 CM 78332 ML117165 CM 

Tyranneutes stolzmanni 1*,2 XC94831 XC94951 WA357339 CM 74778 ML115137 CM 

Pipra aureola CM 73444 

Pipra rubrocapilla 1*,2 XC95466 XC90774 WA360056 CM 72985 

Lepidothrix iris 1*,2 XC95469 XC94872 WA350948 CM 74351 ML115232 CM 

Manacus manacus 1*,2,3 XC95470 XC94611 CM 71803 WA552475 VH ML117132 CM 

Heterocercus linteatus 1 CM 74422 

Machaeropterus pyrocephalus 1* XC91205 MPEG 35610 

Chiroxiphia pareola 1*,2,3 XC90695 XC95100 WA621992 MPEG 27125 ML117138 CM 

TITYRIDAE 

Onychorhynchus coronatus 1*,2 XC94724 MPEG 56122 ML114935 CM 

Terenotriccus erythrurus 1*,2 XC96314 MPEG 53924 WA320528 FG 

Myiobius barbatus 1*,2 MPEG 53929 

Myiobius atricaudus MCZ 175749 

Schiffornis major CM 78261 

Schiffornis turdina 1*,2 XC90687 XC96303 MPEG 56123 


50 





Laniocera hypopyrra 1*,2 XC94719 CM 74551 

Iodopleura isabellae 1*,2 WA500136 

Tityra inquisitor 1*,2,3 CM 74683 

Tityra cayana 1*,2 CM 72152 

Tityra semifasciata 1*,2 WA356510 CM 73293 ML115041 CM 

Pachyramphus rufus 1*,2 XC96341 WA363556 CM 73309 WA634740 IT ML115089 CM 

Pachyramphus castaneus CM 73462 

Pachyramphus polychopterus 1*,3 CM 72347 

Pachyramphus marginatus 1*,2 XC90680 WA500164 CM 74892 

Pachyramphus minor 1*,2 XC94849 WA674422 CM 74712 ML115056 CM 

Pachyramphus validus MPEG 56120 

COTINGIDAE 

Lipaugus vociferans 1*,2 XC95589 XC90679 CM 74728 ML114921 CM 

Gymnoderus foetidus 1,3 CM 74417 

Xipholena lamellipennis 1*,2 XC96154 WA467154 CM 78385 WA320546 FG 

Cotinga cotinga 1* WA356517 USNM 120922 

Cotinga cayana 1*,2 WA467112 CM 74452 WA320488 FG 

Querula purpurata 1*,2 XC91212 WA585264 CM 72789 WA320527 FG ML115077 CM 

Phoenicircus carnifex 1*,2 XC90519 WA358706 MZUSP 10781 ML115046 CM 

RHYNCHOCYCLIDAE 

Platyrinchus saturatus 1*,2 MPEG 56112 ML114961 CM 

Platyrinchus coronatus 2 MPEG 47907 ML114912 CM 

Platyrinchus platyrhynchos 1*,2 XC96149 MPEG 56114 ML114975 CM 

Piprites chloris 1*,2 XC94949 XC95466 MPEG 53943 

Mionectes oleagineus 1*,2 XC95577 CM 74861 

Mionectes macconnelli 1*,2 XC95582 XC95589 WA358691 MPEG 56109 ML115011 CM 

Corythopis torquata 1*,2 XC94832 CM 74611 

Rhynchocyclus olivaceus 1*,2 XC90704 XC90703 MPEG 56119 

Tolmomyias assimilis 1*,2 XC90760 XC94647 CM 75085 ML115022 CM 

Tolmomyias poliocephalus 1*,2 XC94953 WA573666 CM 73047 ML115009 CM 

Tolmomyias flaviventris 1*,3 XC94813 XC87286 WA584504 MPEG 47911 

Todirostrum maculatum 1,3 WA357387 MPEG 15446 WA181010 KO 





51 


Todirostrum cinereum 1,3 WA363019 CM 78277 

Todirostrum chrysocrotaphum 1* XC95679 WA359478 CM 73630 

Poecilotriccus latirostris CM 73669 

Myiornis ecaudatus 1*,2 XC94889 WA360064 CM 73468 ML115043 CM 

Hemitriccus griseipectus CM 74717 

Hemitriccus striaticollis 1*,2,3 XC94612 WA361101 MPEG 50976 ML115017 CM 

Hemitriccus minimus 1*,2 XC86601 XC95119 WA357409 CM 78150 ML114970 CM 

Lophotriccus galeatus 1*,2 XC95590 XC94953 MPEG 56106 ML117099 CM 

TYRANNIDAE 

Zimmerius acer 1*,2 XC95352 XC90774 WA357348 CM 78409 ML114932 CM 

Inezia subflava MCZ 175873 

Ornithion inerme 1*,2 XC95592 WA357363 CM 78584 ML117101 CM 

Camptostoma obsoletum 1*,2,3 XC94885 MPEG 25716 

Elaenia flavogaster 1*,2,3 XC94776 WA359465 MPEG 35603 ML117175 CM 

Elaenia parvirostris 3 CM 73503 

Elaenia cristata 1*,3 WA357381 MPEG 17659 WA206691 KO 

Elaenia pelzelni CM 73687 

Elaenia chiriquensis 1,3 WA361082 MPEG 32449 ML117144 CM 

Suiriri suiriri 3 MPEG 26415 

Myiopagis gaimardii 1*,2,3 XC90775 XC90697 WA444713 CM 74859 ML115069 CM 

Myiopagis caniceps 1* XC104022 WA357356 

Myiopagis flavivertex 1* XC87285 CM 71927 

Myiopagis viridicata 1* CM 74642 

Tyrannulus elatus 1*,2,3 XC94803 WA363020 MPEG 47920 

Capsiempis flaveola 1* XC94620 CM 78368 

Phaeomyias murina 1*,3 XC94648 WA361751 MPEG 40568 ML117159 CM 

Serpophaga hypoleuca CM 72406 

Attila cinnamomeus 1*,2 XC94611 WA352332 MPEG 35605 

Attila spadiceus 1*,2 XC90702 WA676334 MPEG 53931 ML114928 CM 

Attila bolivianus CM 72909 

Legatus leucophaius 1*,2,3 XC94953 WA515389 CM 73516 

Ramphotrigon ruficauda 1*,2 XC94680 WA357403 MPEG 8627 


52 





Myiarchus tuberculifer 1*,2 WA472584 MPEG 40565 

Myiarchus swainsonii 3 CM 72386 

Myiarchus ferox 1*,2,3 XC94723 WA361742 MPEG 47899 ML115096 CM 

Myiarchus tyrannulus 1*,3 WA357382 MPEG 25539 ML117042 CM 

Rhytipterna simplex 1*,2 XC95588 XC95590 MPEG 56121 

Rhytipterna immunda CM 78626 

Casiornis fuscus CM 73783 

Pitangus sulphuratus 1*,2,3 XC94775 WA363015 CM 72043 

Philohydor lictor 1*,2 WA584507 CM 73735 WA100649 JAA 

Myiodinastes maculatus 1*,2,3 XC94723 MPEG 47896 WA446708 TD ML117079 CM 

Tyrannopsis sulphurea 1*,3 WA359435 CM 73243 

Megarhynchus pitangua 1*,2,3 XC94956 WA361104 CM 73775 ML47949 PI 

Myiozetetes similis 1 WA361748 CM 78404 

Myiozetetes cayanensis 1*,2,3 XC94650 WA361745 MPEG 40564 WA205426 KO 

Myiozetetes luteiventris 1*,2 XC96496 WA515439 CM 74488 ML114934 CM 

Tyrannus albogularis 1,3 WA590808 MPEG 26016 WA421557 DO 

Tyrannus melancholicus 1*,2,3 XC94812 WA143667 MPEG 47892 WA446713 TD 

Tyrannus savana 1*,3 WA436252 CM 72317 WA446706 TD 

Griseotyrannus aurantioatrocristatus 1 WA361093 MPEG 26478 

Empidonomus varius 1*,2,3 XC94874 WA361089 MPEG 47895 ML115083 CM 

Conopias trivirgata 1*,2 XC87290 XC94680 WA357332 CM 72771 

Colonia colonus 1* XC95681 

Myiophobus fasciatus 1* XC94776 

Sublegatus obscurior MPEG 56107 

Sublegatus modestus MCZ 175958 

Pyrocephalus rubinus CM 72875 

Fluvicola albiventer 1 CM 71619 WA185168 KO 

Arundinicola leucocephala 1 MPEG 36694 

Cnemotriccus fuscatus 1* XC94878 CM 78450 

Lathrotriccus euleri CM 78471 

Empidonax traillii MPEG 32320 

Contopus nigrescens ML114941 CM 





53 


Knipolegus poecilocercus CM 78169 

VIREONIDAE 

Cyclarhis gujanensis 1*,2,3 XC94878 WA363554 MPEG 40572 

Vireolanius leucotis 1*,2 XC90679 XC87290 WA357364 CM 74926 ML114963 CM 

Vireo olivaceus 1*,2,3 XC90697 XC94648 WA363560 MPEG 54788 WA552752 VH 

Vireo altiloquus 1 MCZ 176304 

Hylophilus semicinereus 1*,2 XC92089 XC94620 MCZ 176308 

Hylophilus pectoralis 1*,2,3 XC94620 WA363555 MPEG 47949 ML117123 CM 

Hylophilus hypoxanthus 1*,2 XC94847 XC94680 WA360087 MPEG 47950 XC6510 SD 

Hylophilus ochraceiceps 1*,2 XC96312 MPEG 36470 ML115235 CM 

HIRUNDINIDAE 

Atticora fasciata 1,2 WA584510 

Stelgidopteryx ruficollis 1*,2,3 WA472579 CM 72747 WA100657 JAA 

Progne tapera 1,3 WA462582 MPEG 54787 

Progne subis 1 WA573665 LACM 38905 

Progne chalybea 1*,2 XC96378 XC95572 WA348548 CM 74309 WA205428 KO 

Tachycineta albiventer 1*,2,3 WA348554 CM 72712 WA206709 KO 

Hirundo rustica 1*,3 WA360057 MPEG 47927 

Riparia riparia 1* WA360063 

TROGLODYTIDAE 

Microcerculus marginatus 1*,2 XC94706 MPEG 56138 XC6568 SD 

Odontorchilus cinereus 1*,2 XC90774 XC94623 WA335247 CM 74980 ML114936 CM 

Troglodytes musculus 1*,2,3 XC94798 WA358650 MPEG 47936 

Campylorhynchus turdinus 1*,2 XC96294 CM 75082 

Pheugopedius coraya 1*,2 XC90737 XC86601 WA360049 MPEG 53947 ML115045 CM 

Canthorchilus leucotis 1*,2,3 XC94806 XC94620 WA142405 MPEG 47935 ML117127 CM 

Cyphorhinus arada 1*,2 XC96732 MPEG 56136 ML117074 CM 

DONACOBIIDAE 

Donacobius atricapilla 1* XC94812 CM 71507 

POLIOPTILIDAE 

Ramphocaenus melanurus 1*,2 XC94882 CM 74871 ML114958 CM 

Polioptila plumbea 1 WA352469 CM 78424 


54 





Polioptila paraensis 1*,2 

TURDIDAE 

Catharus fuscescens MPEG 54844 

Catharus mimus MPEG 47943 

Turdus nudigenis CM 72988 

Turdus leucomelas 1*,3 XC90695 XC94670 WA443928 MPEG 35602 WA446714 TD 

Turdus fumigatus 1* WA358714 CM 74475 

Turdus albicollis 1*,2 XC96492 MPEG 56139 

MIMIDAE 

Mimus saturninus MPEG 08546 

MOTACILIDAE 

Anthus lutescens 1* XC96368 CM 73185 

COEREBIDAE 

Coereba flaveola 1*,2,3 WA333900 MPEG 53952 ML115037 CM 

THRAUPIDAE 

Saltator grossus 1*,2 XC90772 XC92089 WA515530 MPEG 56142 ML114919 CM 

Saltator maximus 1*,2 XC96344 MPEG 23662 

Saltator coerulescens 1* WA610324 CM 72178 

Parkerthraustes humeralis 1*,2 XC104858 WA357371 

Lamprospiza melanoleuca 1*,2 XC94951 XC94949 WA500186 CM 74850 

Nemosia pileata 1,3 WA357328 CM 72632 ML117145 CM 

Tachyphonus rufus 1*,2,3 XC96330 WA358064 FMNH 258333 WA509899 IT 

Ramphocelus carbo 1*,2,3 XC94611 WA358061 MPEG 22794 WA100650 JAA ML115208 CM 

Ramphocelus nigrogularis CM 72702 

Lanio luctuosus 1*,2 CM 75084 

Lanio cristatus 1*,2 WA467147 CM 74707 

Lanio cuculatus 1 XC94890 WA435508 

Lanio versicolor 1*,2 XC96152 MPEG 53955 XC91264 JM 

Lanio surinamus 1*,2 CM 75078 ML115051 CM 

Lanio penicillatus CM 73592 

Tangara mexicana 1*,2,3 XC96313 CM 72207 XC5981 SD 

Tangara velia 1*,2 WA357353 





55 


Tangara varia 1* XC96295 

Tangara punctata 1*,2 WA467140 

Tangara episcopus 1*,2,3 XC94878 WA358065 MPEG 17778 WA182124 KO 

Tangara palmarum 1*,2,3 XC94648 WA500207 CM 72052 WA182125 KO 

Tangara cayana 1,3 WA444716 CM 78198 WA509900 IT 

Schistoclamis melanops WA551855 MPEG 37767 

Paroaria gularis 1,2,3 WA340077 CM 73727 WA559505 VH 

Dacnis lineata 1*,2 WA356513 

Dacnis flaviventer CM 72799 

Dacnis cayana 1*,2,3 WA467126 MPEG 23826 

Cyanerpes caeruleus 1*,2 WA487618 CM 74612 

Cyanerpes cyaneus 1*,2,3 WA356513 CM 72808 ML117142 CM 

Chlorophanes spiza 1* WA443043 MCZ 22928 ML115025 CM 

Hemithraupis guira 1*,2 WA500180 CM 74941 

Conirostrum bicolor CM 73679 

EMBEREZIDAE 

Amnodramus humeralis 1,3 WA544922 MPEG 23449 ML117039 CM 

Ammodramus aurifrons 1 WA583468 CM 73732 WA183290 KO 

Sicalis colombiana 1 WA144015 MPEG 36695 WA446718 TD ML47953 PI 

Sicalis luteola CM 73513 

Volatinia jacarina 1*,2,3 XC94618 WA467122 CM 71954 ML117171 CM 

Sporophila schistacea MPEG 47983 

Sporophila americana 1* XC94776 WA444715 CM 71800 

Sporophila lineola 1* WA500123 CM 72651 

Sporophila nigricollis 1* WA347325 WA576640 IT 

Sporophila caerulescens 2 MCZ 176848 

Sporophila minuta 1* WA467116 CM 72072 

Sporophila castaneiventris WA340079 CM 71617 

Sporophila angolensis 1*,2,3 XC94874 WA514803 CM 72521 

Arremon taciturnus 1*,2 XC94778 MPEG 53961 ML115061 CM 

CARDINALIDAE 

Piranga flava 3 USNM 276980 


56 





Piranga rubra WA924652 MZUSP 47382 

Habia rubica 1*,2 XC96312 MPEG 35338 ML88362 CM 

Granatellus pelzelni 1* XC92090 CM 74460 

Periporphyrus erythromelas 1*,2 XC104023 

Cyanoloxia cyanoides 1*,2 XC94734 MPEG 35608 ML115042 CM 

PARULIDAE 

Phaeothlypis rivularis MPEG 53957 ML114985 CM 

Dendroica striata 3 MPEG 50977 ML117141 CM 

Geothlypis aequinoctialis CM 78459 

ICTERIDAE 

Psarocolius viridis 1*,2 XC91202 CM 75037 ML115067 CM 

Psarocolius decumanus 1*,2,3 CM 71975 ML115059 CM 

Psarocolius bifasciatus 1*,2 XC94714 CM 73313 

Procacicus solitarius CM 71999 

Cacicus haemorrhous 1*,2 CM 74580 

Cacicus cela 1*,2,3 XC94775 WA441607 MPEG 23351 ML115210 CM 

Icterus cayanensis 1,2 WA675100 CM 72081 

Icterus croconotus CM 72609 XC91267 JM 

Gymnomystax mexicanus 1 WA348555 CM 71607 WA550297 VH 

Chrysomus icterocephalus CM 71939 

Molothrus oryzivorus 1*,2,3 WA348559 MPEG 15252 WA240665 IT 

Molothrus bonariensis 1*,2 WA348556 MPEG 36693 WA550669 VH 

Sturnella militaris 1*,3 WA467176 MPEG 35614 

FRINGILIDAE 

Euphonia chlorotica 1,3 WA357326 CM 72623 WA370312 IT 

Euphonia violacea 1*,2 WA584513 CM 72853 ML115106 CM 

Euphonia minuta 1*,2 CM 73799 

Euphonia xanthogaster CM 74535 

Euphonia rufiventris 1*,2 XC94738 WA514793 ML115140 CM 

PASSERIDAE 

Passer domesticus 1 WA349047 





57 

Appendix 2 

List of 26 species reported from the Santarém-Belterra region, south of the Amazon and east of the Tapajós (PA, Brazil) but without any permanent vouchering material. 

Species Details of sighting 

Penelope jacquacu Sight records, C. B. A., A. C. L., B. J. W. D., Catchments: 69, 81, 99, 103, 157, 165, 236, 260, 261, 307, 399 

Egretta tricolor Sight record, A. Whittaker 14/11/1988, Alter do Chão 

Ictinia mississippiensis Sight record, G. M. Kirwan & C. F. Collins, 19 Alter do Chão 4/12/2005, listed in Whittaker et al. (2008) 

Helicolestes hamatus Sight record listed in Henriques et al. (2003) 

Buteo albonotatus Sight record listed in Sanaiotti & Cintra (2001) 

Falco columbarius Sight record, E. L., 30/11/2011, campus of the Universidade Federal do Oeste do Pará 

Aratinga maculata Sight records by E. Willis: Maicá 16/011984, Rodagém, 18/10/1984, Urumari, in Feb,1985, listed in Willis & Silva (1986) 

Pyrrhura lepida Aural records, E. L, 15/02/2012, Rio Curuauna 

Cypseloides sp. Sight record, A. C. L., 27/01/2011, Catchment 129 

Threnetes leucurus Mist net captures listed by Henriques et al. (2003) 

Phaethornis hispidus Sight record, B. J. W. D. 31/01/2011, Catchment 112 

Lophornis ornatus Sight record Henriques et al. (2003) 

Chrysolampis mosquituus Sight record B. Whitney Km 21 on road to Alter do Chão, 19 June and again 7 July 1995) 

Trogon curucui Aural records C. B. A., A. C. L., B. J. W. D., Catchments: 81, 112 

Brachygalba lugubris Sight records in Henriques et al. (2003) 

Xenops rutilans Sight records C. B. A., Catchment: 157 

Microxenops milleri Sight records C. Marantz, 23/8/1999, 18/09/1999, 09/10/1999, Base de Sucupira, FLONA 

Dixiphia pipra Mist-net capture, Henriques et al. (2003) 

Tolmomyias sulphurescens Mist-net capture, reported in Henriques et al. (2003) 

Sirystes sibilator Sight record listed in Sanaiotti & Cintra (2001) 

Contopus cooperi Sight record, C. Marantz, 26/09/1999, Base de Sucupira, FLONA 

Petrochelidon pyrrhonota Sight record, A. Whittaker, 14/111988, Alter do Chão (in Stotz et al. 1992) 

Atticora tibialis Sight record, C. B. A.,18/11/2010, Catchment 307 

Cyanocorax chrysops Sight record, B. Whitney, 06/071995, Maicá 

Tersina viridis Sight record in Sanaiotti & Cintra (2001) 

Cissopis leverianus Sight record, C. B. A. 03/11/2010, Catchment 399 



March 2013 

article 

Las aves de la isla Lobos de Tierra, Perú: revisión 

bibliográfica y nuevos registros (1684-2011) 

Judith Figueroa 

Asociación para la Investigación y Conservación de la Biodiversidad - AICB. Lima, Perú. E-mail: aicb.peru@gmail.com 

Enviado en 30 Octubre 2011. Acceptado en 7 Junio 2012. 

ABSTRACT: The birds of Lobos de Tierra island, Peru: a review and new records (1684-2011). The present work is based 

on a detailed review of bird observations made on Lobos de Tierra island (06°26’S; 80°51’W) by various authors since 1684, as 

well as interviews with island rangers and other researchers, and field assessments carried out in 1999, 2000, 2004 and 2011. This 

compilation came to a total of 43 species (11 breeding), Sula nebouxii and Pelecanus thagus being the most abundant. The high 

number of records compared with other islands in Peru are related to the island’s proximity to the coast and its location on the 

convergence zone of the cold waters of the Peruvian Current and the warm waters of the Equatorial Countercurrent. Of all the 

species, 17 are listed in a conservation category. 

KEY-WORDS: birds, Lobos de Tierra island, Peru. 

INTRODUCCIÓN 

MÉTODOS 

Área de estudio 

La isla Lobos de Tierra se ubica al norte del Perú, 

entre las regiones de Piura y Lambayeque. Es una isla 

deshabitada y un área reservada para la explotación del 

guano de aves marinas desde inicios del siglo pasado, 

formando parte de la recientemente creada Reserva 

Nacional Sistema de Islas, Islotes y Puntas Guaneras 

(MINAM 2010). Además, está considerada como un área 

importante para la conservación y reproducción de las 

aves marinas en el Perú (IBA PE012; Franke et al. 2005, 

Franke 2006). 

Los primeros alcances sobre las especies de aves y 

su abundancia en esta isla fueron realizados por Forbes 

(1914), Coker (1919), Murphy (1925a, 1936) y Tovar 

(1968). Posteriormente, se desarrollaron numerosas 

investigaciones principalmente orientadas a la ecología 

y dieta de las aves guaneras (Phalacrocorax bougainvillii, 

Sula variegata y Pelecanus thagus) y Sula nebouxii (Duffy 

1987, Jahncke & Goya 1997a, 1998a, Jahncke et al. 

1997, Zavalaga et al. 2007, 2008, 2009a, 2010a, 2010b, 

2011). 

El objetivo del presente trabajo es sistematizar la 

información existente relacionada a la avifauna de la 

isla Lobos de Tierra, así como dar a conocer los nuevos 

registros, incluyendo en algunos casos su número 

poblacional y distribución. 

La isla Lobos de Tierra (LT) se ubica frente al límite 

de las regiones Lambayeque y Piura (06°26’S; 80°51’O) 

(Figura 1). Es una de las islas más grandes del litoral 

peruano, con un área de 14.3 km 2 (10.5 km de largo por 

4.0 km en su parte más ancha). Está orientada en dirección 

norte-sur, a 11.4 km de la costa, con una altura máxima 

de 92 m. Presenta una geografía accidentada, con laderas 

pedregosas de roca granítica y playas arenosas (DHN 2003), 

casi desprovista de vegetación, a excepción de algunos 

pequeños parches de verdolaga (Sesuvium portulacastrum) 

en las orillas arenosas del lado sureste de la isla (Figuras 2a 

y 2b). Como ocurre a lo largo del todo el litoral peruano, 

las mayores temperaturas del aire se presentan en el primer 

trimestre del año (23.4°C), disminuyendo en los siguientes 

trimestres (entre 17.7°C y 20.7°C) y aumentando en el 

último (18.0°C; Schweigger 1931, BCAG 1934-1941). 

En el caso de la temperatura superficial del mar también 

se presenta este mismo comportamiento (Carbajal et al. 

2003, 2004a, 2004b, 2005b, 2006, De la Cruz et al. 2006). 

En base a la presencia de indicadores biológicos, 

como las diatomeas Lithodesmium undulatum, Skeletonema 

costatum, Chaetoceros debilis, Thalassiosira subtilis, entre 

otras, se conoce que LT presenta activos procesos de 

afloramientos costeros (Carbajal et al. 2004a, 2004b). 

Asimismo, los datos oceanográficos (temperatura, salinidad



59 

FIGURA 1. Mapa de la isla Lobos de Tierra y lugares donde fueron observadas las aves: Pc - Phoenicopterus chilensis, Sh - Spheniscus humboldti, 

Pi - Phoebastria irrorata, Pc - Pterodroma cookii, Om - Oceanodroma markhami, Oh - Oceanodroma hornbyi, Pg - Pelecanoides garnotii, Fm - Fregata 

magnificens, Sn - Sula nebouxii, Sv - Sula variegata, Sg - Sula granti, Sle - Sula leucogaster, Pb - Phalacrocorax brasilianus, Pg - Phalacrocorax gaimardi, 

Pbo - Phalacrocorax bougainvillii, Pt - Pelecanus thagus, Et - Egretta thula, Cau - Cathartes aura, Fp - Falco peregrinus, Cha - Charadrius alexandrinus, 

Ha - Haematopus ater, Np - Numenius phaeopus, Cal - Calidris alba, Cc - Chroicocephalus cirrocephalus, Lm - Leucophaeus modestus, Lb - Larus belcheri, 

Ld - Larus dominicanus, Sl - Sternula lorata, Li - Larosterna inca. 

FIGURE 1. Map of the Lobos de Tierra island and places where birds were observed: Pc - Phoenicopterus chilensis, Sh - Spheniscus humboldti, Pi 

- Phoebastria irrorata, Pc - Pterodroma cookii, Om - Oceanodroma markhami, Oh - Oceanodroma hornbyi, Pg - Pelecanoides garnotii, Fm - Fregata 

magnificens, Sn - Sula nebouxii, Sv - Sula variegata, Sg - Sula granti, Sle - Sula leucogaster, Pb - Phalacrocorax brasilianus, Pg - Phalacrocorax gaimardi, 

Pbo - Phalacrocorax bougainvillii, Pt - Pelecanus thagus, Et - Egretta thula, Fp - Falco peregrinus, Cha - Charadrius alexandrinus, Ha - Haematopus 

ater, Np - Numenius phaeopus, Cal - Calidris alba, Cc - Chroicocephalus cirrocephalus, Lm - Leucophaeus modestus, Lb - Larus belcheri, Ld - Larus 

dominicanus, Sl - Sternula lorata, Li - Larosterna inca. 


60 



a 

b 

FIGURA 2. (a) Áreas planas al noreste de la isla, con colonias de Pelecanus thagus. Foto: M. Stucchi (b) Áreas accidentadas al centro-sur de la isla, 

con presencia de Sula nebouxii y S. granti. Foto: JF. 

FIGURE 2. (a) Flat areas in the northeast of the island, with colonies of Pelecanus thagus. Photo: M. Stucchi. (b) Hilly areas in the center-south of 

the island, with presence of Sula nebouxii and S. granti. Photo: JF. 




61 

y oxígeno disuelto) y la presencia del dinoflagelado 

Protoperidinium obtusum, señalan que esta isla se caracteriza 

por la predominancia de aguas costeras frías, cuya salinidad 

puede variar entre los 35.01 y 35.14 ups (Carbajal et al. 

2004a, 2004b, De la Cruz et al. 2006). A inicios de cada 

año presenta una ligera influencia de las aguas subtropicales 

superficiales (Carbajal et al. 2004a, 2005b). Alrededor de 

la isla existen praderas de algas macrofíticas, especialmente 

del género Caulerpa sp. (Carbajal et al. 2003, 2005b, De 

la Cruz et al. 2006), las cuales están asociadas con los altos 

valores de oxígeno disuelto en el mar. 

En el lado centro-oeste de la isla, frente a los islotes 

León, se encuentra un apostadero de lobo marino chusco 

(Otaria flavescens). 

Procedimiento y colecta de datos 

La lista de aves de la isla LT se ha compilado en base 

a cinco fuentes de información: (1) Observaciones hechas 

por la autora el 27 y 28 de marzo de 1999, 9 y 10 de junio 

de 1999, 10 y 11 de agosto de 1999, 24 y 25 de febrero de 

2000, del 6 al 8 de diciembre de 2004 y del 28 de febrero 

al 3 de marzo de 2011. Estas se realizaron de forma 

oportunista, en el desarrollo de los censos del lobo marino 

chusco al oeste de la isla (1999), así como los censos y 

búsqueda de nuevas áreas de reproducción del piquero 

de Nazca (Sula granti) y la gaviota dominicana (Larus 

dominicanus) (2000, 2004 y 2011). Para estos últimos, los 

recorridos se realizaron en dos grupos, tratando de cubrir 

toda la isla, entre las 08 h y 17 h. Se tomaron registros 

fotográficos de algunas especies. 

(2) Observaciones y registros fotográficos realizados 

por otros investigadores. 

(3) Registros entre enero de 2009 a febrero de 2011, 

proporcionados por tres guarda-islas de AGRORURAL 

(Programa de Desarrollo Productivo Agrario Rural - 

Ministerio de Agricultura), quienes se encargan del 

monitoreo de las aves guaneras, por lo que una vez al mes 

realizan un censo general de toda el área. 

(4) Búsqueda de colectas en la base de datos 

del American Museum of Natural History (AMNH) 

(http://entheros.amnh.org/db/emuwebamnh/Query. 

php) y Natural History Museum of London (NHM) 

(http://www.nhm.ac.uk/jdsml/research-curation/research/ 

projects/birdtype/search.dsml). 

(5) Artículos publicados anteriormente sobre la 

avifauna de LT. 

RESULTADOS Y DISCUSIÓN 

Diversidad 

Se registraron 43 especies de aves entre marinas 

y terrestres, 11 de ellas anidando en la isla (Tabla 1). 

Estas pertenecieron a 18 familias, siendo la más diversa 

Laridae con ocho especies, seguida de Procellariidae, 

Hydrobatidae, Sulidae y Scolopacidae, con cuatro especies 

cada una. Del total de especies, 35 fueron registradas 

anteriormente en la bibliografía, y ocho correspondieron 

a nuevos registros para LT: Phoenicopterus chilensis, Sula 

leucogaster, Egretta thula, Falco peregrinus, Charadrius 

alexandrinus, Numenius phaeopus, Chroicocephalus 

cirrocephalus y Leucophaeus modestus. 

Esta diversidad y grandes números poblaciones 

de algunas especies (Sula nebouxii y Pelecanus thagus), 

podrían guardar relación con el conjunto de las siguientes 

condiciones: a) gran área de extensión (segunda isla 

más grande del Perú), b) cercanía a la costa, c) clima y 

fisiografía (Murphy 1925a), d) su ubicación entre las 

aguas de surgencia fría de la Corriente Peruana y la baja 

productividad de las aguas cálidas de la Contracorriente 

Ecuatorial (Duffy et al. 1984) y, e) características 

oceanográficas, que hacen propicio el desarrollo de la 

anchoveta Engraulis ringens, principal especie ictiológica 

de la dieta de las aves guaneras y otras especies, como S. 

nebouxii y S. granti, todas habitantes de LT (Jahncke & 

Goya 1997a, Jahncke et al. 1997, Jahncke 1998, Jahncke 

& Goya 1998a,). 

En comparación con otras islas del Perú el número 

de especies en LT es mayor [isla Foca (Piura) con 34 

especies (Novoa et al. 2010, Rivas 2010, Figueroa & 

Stucchi 2012), Lobos de Afuera (Lambayeque) con 33 

especies (Stucchi et al. 2011), El Frontón (Callao) con 20 

especies (Rivadeneira et al. 1986) y San Lorenzo (Callao) 

con 17 especies (Gonzáles et al. 2005)]. Además de las 

condiciones arriba descritas, esto también podría deberse 

a que LT es una de las islas más estudiadas del país. 

Sin embargo, a pesar que la isla se encuentra 

cercana a la costa (11.4 km), no se ha registrado un 

mayor número de especies terrestres. Esto podría estar 

relacionado a la casi total falta de vegetación en el área. 

Asimismo, llama la atención la ausencia de Progne 

murphyi, la cual es endémica de la costa peruana (Plenge 

2011). Esta ha sido observada en islas cercanas al litoral, 

ubicadas hasta 5 km de la costa: Santa y Ferrol (Ancash) 

(Valverde et al. 2007), Chao y Corcovado (La Libertad) 

(Balta et al. 2005) y Foca (Piura) (Novoa et al. 2010), 

pero principalmente en áreas costeras (Valverde et al. 

2007). Parece existir una relación entre la presencia 

de las colonias de esta especie y la escasa perturbación 

de las áreas, principalmente por la ausencia de la 

extracción del guano (Balta et al. 2005, Valverde et al. 

2007), condición que no se da en LT debido a que esta 

actividad se realizó de forma continua entre 1986 y 

1999 (Proabonos 2008), y después de 11, años en el 

2010. Por otro lado, existe la posibilidad de que la falta 

del registro de P. murphyi y de otras aves terrestres, sean 

debido a que las observaciones fueron realizadas solo de 

forma oportunista. 


62 



TABLA 1. Lista de aves registradas en la isla Lobos de Tierra (1684-2011) 

Registros previos: (1) AMNH (2011), (2) Ayala (2006), (3) Coker (1919), (4) Dampier (1729), (5) Duffy (1987), (6) Duffy et al. (1984), (7) 

Figueroa (2010), (8) Figueroa et al. (2011b), (9) Forbes (1914), (10) Guillén (1991), (11) Jahncke & Goya (1997a), (12) Jahncke & Goya (1997b), 

(13) Jahncke & Goya (1998a), (14) Jahncke & Goya (1998b), (15) Jahncke & Paz-Soldán (1998), (16) Jahncke et al. (1997), (17) Mills (1968), 

(18) Murphy (1925a), (19) Murphy (1925b), (20) Murphy (1936), (21) NHM (2011), (22) Nelson (1978), (23) Schulenberg & Parker (1981), 

(24) Schweigger (1947), (25) Taylor et al. (2010), (26) Tovar (1968), (27) Vogt (1942), (28) Wetmore (1923), (29) Zavalaga (2003), (30) Zavalaga 

et al. (2002), (31) Zavalaga et al. (2007), (32) Zavalaga et al. (2008), (33) Zavalaga et al. (2009a), (34) Zavalaga et al. (2009b), (35) Zavalaga et al. 

(2010a), (36) Zavalaga et al. (2010b), (37) Zavalaga et al. (2011). 

Evidencia: O - Observado, C - Colectado, F - Fotografiado. 

Cantidad: ME - Muy Escaso: menos de diez individuos, E - Escaso: decenas de individuos, M - Medio: centenares de individuos, A - Abundante: 

miles individuos. 

Frecuencia: Cm - Común: observada en toda la isla, L - Localizado: habita en zonas específicas, O - Ocasional: llegan a la isla accidental o eventualmente. 

Estacionalidad: Re - Residente, M - Migratorio, R - Reproductivo, I - Indeterminado. 

a 

Ministerio de Agricultura: Categorización de especies amenazadas de fauna silvestre del Perú. 

b 

IUCN: CR - En peligro crítico, EN - En peligro, VU - Vulnerable, NT- Casi amenazado. 

c 

CITES: Apéndice I: Especie en peligro de extinción, Apéndice II: Especie cuyo comercio debe controlarse a fin de evitar una utilización incompatible 

con su supervivencia. 

d 

CMS: Apéndice I: Especie migratoria en peligro, Apéndice II: Especie migratoria cuyo estado de conservación es desfavorable. 

TABLE 1. List of birds recorded on the Lobos de Tierra island (1684-2011). 

Previous records: (1) AMNH (2011), (2) Ayala (2006), (3) Coker (1919), (4) Dampier (1729), (5) Duffy (1987), (6) Duffy et al. (1984), (7) 

Figueroa (2010), (8) Figueroa et al. (2011b), (9) Forbes (1914), (10) Guillén (1991), (11) Jahncke & Goya (1997a), (12) Jahncke & Goya (1997b), 

(13) Jahncke & Goya (1998a), (14) Jahncke & Goya (1998b), (15) Jahncke & Paz-Soldán (1998), (16) Jahncke et al. (1997), (17) Mills (1968), 

(18) Murphy (1925a), (19) Murphy (1925b), (20) Murphy (1936), (21) NHM (2011), (22) Nelson (1978), (23) Schulenberg & Parker (1981), 

(24) Schweigger (1947), (25) Taylor et al. (2010), (26) Tovar (1968), (27) Vogt (1942), (28) Wetmore (1923), (29) Zavalaga (2003), (30) Zavalaga 

et al. (2002), (31) Zavalaga et al. (2007), (32) Zavalaga et al. (2008), (33) Zavalaga et al. (2009a), (34) Zavalaga et al. (2009b), (35) Zavalaga et al. 

(2010a), (36) Zavalaga et al. (2010b), (37) Zavalaga et al. (2011). 

Evidence: O - Observed, C - Collected, F - Photographed. 

Quantity: ME - Very low: less than ten individuals, E - Low: dozens of individuals, M - Medium: hundreds of individuals, A - Abundant: thousands 

individuals. 

Frequency: Cm - Common: seen throughout the island, L - Located: living in specific areas, O - Occasional: arriving on the island accidentally or 

eventually. 

Seasonality: Re - Resident, M - Migratory, R - Reproductive, I - Indeterminate. 

a 

Ministry of Agriculture: Categorization of endangered species of wildlife of Peru. 

b 

IUCN: CR - Critically Endangered, EN - Endangered, VU - Vulnerable, NT- Near Threatened. 

c 

CITES: Appendix I: Includes species threatened with extinction, Appendix II: Includes species not necessarily threatened with extinction, but in 

which trade must be controlled in order to avoid utilization incompatible with their survival. 

d 

CMS: Appendix I: Endangered migratory species, Appendix II: Migratory species conserved through agreements. 

Categoría de Amenaza 

FAMILIA / Especie Reportes previos Evidencia, año del registro 

Cantidad / 

Frecuencia 

Estacionalidad 

MINAG 2004 a 

IUCN 2011 b 

CITES 2011 c 

CMS 2009 d 

Phoenicopteridae (1) 

Phoenicopterus chilensis O2011 ME / O NT NT II 

Spheniscidae (1) 

Spheniscus humboldti 4, 6, 20, 26 O1684, 1920, 1979, 1982; 

F2011 

Diomedeidae (3) 

E / L Re, R EN VU I I 

Phoebastria irrorata 3, 20 C1912; O1908, 1920, 2002 ME / O M VU CR II 

Thalassarche melanophris 20 C1912, 1920 ME / O M EN II 

Thalassarche salvini 20, 21 C1912 ME / O M VU 




63 



Cantidad / 

Frecuencia 


MINAG 2004 a 

IUCN 2011 b 

CITES 2011 c 

CMS 2009 d 

Procellariidae (4) 

Daption capense 24 O1939 ME / O M 

Pterodroma cookii 20 C1922 ME / O M VU 

Pachyptila belcheri 23 O1978 ME / O M 

Puffinus griseus 18 O1920 ME / O M NT 

Hydrobatidae (4) 

Oceanites gracilis 20 C1920 ME / O I 

Oceanodroma markhami 20 O1920 ME / O I VU 

Oceanodroma hornbyi 17, 28 O1922, 1965 ME / O I 

Oceanodroma melania 28 O1922 ME / O I 

Pelecanoididae (1) 

Pelecanoides garnotii 9, 14, 20 O1913, 1996 I I CR EN I 

Fregatidae (1) 

Fregata magnificens 3, 20 O1907, 1920, 2011; F2006 ME / O 

Sulidae (4) 

Sula nebouxii 2, 3, 5, 10, 15, 16, 

20, 25, 26, 31, 32 

Sula variegata 2, 5, 9, 11, 13, 15, 

16, 20, 22, 25, 33, 35 

O1907, 1920, 1963, 1978, 

2002, 2003; F2011 

A / Cm 

Re, R 

O1913, 1920, 1978; F2011 A / L Re, R EN 

Sula granti 12, 16 O1996; F1999, 2004, 2011 E / L Re, R EN 

Sula leucogaster F2011 ME / O 

Phalacrocoridae (3) 

Phalacrocorax brasilianus 3, 9 O1907, 1913; F2011 E / Cm Re, R 

Phalacrocorax gaimardi 9, 26, 30 O1913, 1963, 1999, 2010 E / L Re, R EN NT 

Phalacrocorax bougainvillii 3, 9, 11, 13, 15, 

16, 20, 22, 27 

Pelecanidae (1) 

O1907, 1913, 1920; F2011 A / L Re, R EN NT 

Pelecanus thagus 3, 20, 22, 27, 36, 37 O1907, 2010; C1920; 

F2011 

Ardeidae (1) 

A / L Re, R EN NT 

Egretta thula O2010 ME / O 

Cathartidae (1) 

Cathartes aura jota 3, 8, 9, 29 O1907, 1913, 2000, 2002, 

1999, 2004; F2011 

Falconidae (1) 

M / Cm 

Re, R 

Falco peregrinus F2011 ME / O M NT I 


64 





Cantidad / 

Frecuencia 


MINAG 2004 a 

IUCN 2011 b 

CITES 2011 c 

CMS 2009 d 

Charadriidae (2) 

Charadrius alexandrinus O2010 ME / O I 

Oreopholus ruficollis pallidus 1 C1912 ME / O M 

Haematopodidae (2) 

Haematopus palliatus pitanay 19, 20 O1920 ME / O I 

Haematopus ater 3 O1907, 2010 ME / O I 

Scolopacidae (4) 

Numenius phaeopus O2011 ME / O M 

Tringa incana 3 O1907 ME / O M 

Calidris alba 3 O1907; F2011 ME / O M 

Calidris mauri 3 O1907 ME / O M 

Stercorariidae (1) 

Stercorarius chilensis 20 O1913 ME / O 

Laridae (8) 

Creagrus furcatus 20 O1920 ME / O M 

Chroicocephalus cirrocephalus F2011 ME / O M 

Leucophaeus modestus O1999; F2011 ME / O M 

Leucophaeus pipixcan 20 O1920 E / O M 

Larus belcheri 20 O1920; F2011 E / L R 

Larus dominicanus 3, 6, 7, 20, 26 O1907, 1920, 1963, 1979; M / L Re, R 

F2004, 2011 

Sternula lorata 20, 34 O1920, 2006, 2010 E / O I VU EN I 

Larosterna inca 26 O1963, 2011 E / L Re, R? VU NT 

Especies registradas 

Phoenicopterus chilensis 

A principios de diciembre de 2010, A. Oliva (com. 

pers.) observó un individuo en la playa El Ñopo, que se 

encontraba en malas condiciones de salud, muriendo 

al día siguiente de ser visto. Este podría tratarse de un 

individuo perdido que formó parte de los grandes grupos 

de P. chilensis (~7000 individuos) que migraron a la laguna 

de Ñapique (Piura) a fines del 2010 (F. Angulo com. pers.). 

Spheniscus humboldti (Figura 3a) 

Es una especie endémica de la Corriente Peruana, 

actualmente muy escasa en la isla LT. Dampier (1729), 

a mediados de noviembre de 1684 arribó al lado noreste 

de LT encontrando una gran cantidad de individuos de 

esta especie. Posteriormente, esta población decreció 

considerablemente. Murphy (1936) no observó 

individuos vivos en la isla; en enero de 1920 solo 

encontró la piel de un pingüino que recientemente 

había sido cazado cerca de los campamentos de pesca. Al 




65 

respecto, el administrador de operaciones de extracción 

del guano, le informó que en años pasados, en los islotes 

adyacentes, grandes números de pingüinos mantenían 

sus madrigueras dentro del guano. En febrero de 1979 

y mayo de 1982, se observaron cinco individuos en cada 

caso (Duffy et al. 1984). Estos mismos autores nombran 

otros conteos realizados en LT por Pesca Perú, 1000 

individuos en diciembre de 1980 y 900 individuos en 

marzo de 1981, sin embargo, señalan que podrían estar 

sobrevalorados. El 24 de febrero de 1999, se observaron 

seis pingüinos en muda en la bahía Canevaro. En marzo 

de 2011 se encontró en esta misma bahía, una colonia 

conformada por 38 individuos (36 adultos en muda y 

dos jóvenes), así como un individuo joven descansando 

en una punta al norte de la isla. 

Phoebastria irrorata (Figura 3b) 

Fue observado en 1908, entre LT y Eten, volando en 

grupos de 30 individuos (Coker 1919). Posteriormente, 

en mayo de 1912 y enero de 1920 se registraron pequeños 

grupos nadando cerca a LT (Murphy 1936). En julio 

de 2002, G. Mori (com. pers.) encontró un individuo 

descansando en un área plana en la parte central de LT, 

y el 24 de junio de 2006, I. García-Godos (com. pers.) 

observó otro individuo adulto descansando en la isla, a 

200 m al sur de la guardianía de La Grama. 

Thalassarche melanophris 

H. O. Forbes colectó dos especímenes en los 

alrededores de LT en junio de 1912. Murphy cazó otro 

espécimen en 1920 (Murphy 1936). 

Thalassarche salvini 

Cuatro especímenes (tres machos y una hembra) 

fueron colectados por H. O. Forbes en los alrededores 

de LT entre el 28 de mayo y 27 de junio de 1912, y 

depositados en el NHM, donde posteriormente fueron 

identificados como Diomedea cauta salvini (Murphy 

1936, Hellmayr & Conover 1948). Además, en esta 

colección, se tiene el registro de un albatros macho 

adulto también colectado por H. O. Forbes cerca a LT 

el 5 de marzo de 1912, e identificado como Diomedia 

cauta peruvia (Mathews 1933, Warren 1966), el cual fue 

identificado como sinónimo de D. c. salvini (Mayr & 

Cottrell 1979). Posteriormente, Diomedea fue cambiado 

al género Thalassarche, y T. salvini fue separado de la 

especie politípica D. cauta (Robertson & Nunn 1998). 

Daption capense 

Schweigger (1947) señaló que esta especie fue 

observada cerca de las islas de Lobos (LT y Lobos de 

Afuera) en septiembre de 1939, e indicó que en años 

posteriores no la volvió a ver. 

Pterodroma cookii 

C. L. Fagan colectó dos especímenes al norte de LT 

el 31 de julio de 1922 (Murphy 1936). 

Pachyptila belcheri 

Entre el 26 de junio y 15 de julio de 1978, M. 

Williams (Schulenberg & Parker 1981) encontró en 

Lambayeque varios individuos muertos: uno en Pimentel, 

tres en San José y uno en la isla LT. 

Puffinus griseus 

Varios individuos de esta especie fueron observados 

por Murphy (1925b) a 3.5 km de LT, en enero de 1920. 

Oceanites gracilis 

Murphy (1936) detalla las medidas morfométricas 

de 40 ejemplares colectados entre Chilca y LT. 

Oceanodroma markhami 

Murphy (1936) observó una bandada en punta 

Sáenz. 

Oceanodroma hornbyi 

Esta especie fue registrada por Wetmore (1923) el 

20 de julio y 24 de noviembre de 1922, a 4.8 km y 8 km, 

respectivamente de la isla LT. Asimismo, Mills (1968) 

observó un individuo al noroeste de esta isla, el 4 de 

octubre de 1965. 

Oceanodroma melania 

Wetmore (1923) observó un individuo hembra el 

24 de noviembre de 1922, a 8 km de LT. 

Pelecanoides garnotii 


actualmente se desconoce su situación en la isla. Forbes 

(1914), en febrero de 1913, observó grandes bandadas 

de esta especie en el islote Unanue (conocido también 

como isla El Colorado o Larrañaga). Murphy (1936) 

señaló a LT como una de las áreas de reproducción de P. 

garnotii anteriormente registrada. Posteriormente, entre 

mayo y noviembre de 1996, Jahncke & Goya (1998b) 

encontraron un individuo muerto en las cercanías de 

la isla. A pesar que en los años 2003, 2004 y 2011 se 


66 



volvió a registrar la presencia de P. garnotii a 55 km al 

sur de LT (Figueroa & Stucchi 2008, Figueroa et al. 

2011a), no se tienen nuevos registros en la isla. Al igual 

que LT, en otras áreas donde se registró la reproducción 

de la especie, como las islas Chinchas, Ballestas, Macabí, 

Guañape, Pescadores, Mazorca y Mongoncillo (Coker 

1919, Murphy 1936, Tovar 1968), esta parece estar 

actualmente ausente. En la última década las únicas 

colonias reproductivas del Perú se encontraron en las 

islas La Vieja, San Gallán y Corcovado (Jahncke & Goya 

1998b, Valverde 2006). Esta pérdida y disminución 

poblacional, está relacionada directamente con la 

actividad extractiva del guano, eliminando el substrato 

en el que P. garnotii anida (Coker 1919, Murphy 1936, 

Tovar 1968). 

Fregata magnificens 

La especie fue registrada en LT en abril de 1907 y en 

enero de 1920. En aquella época, su presencia podría haber 

estado relacionada con los numerosos campamentos de los 

pescadores (Coker 1919, Murphy 1936). Posteriormente, 

G. Mori (com. pers.) observó un ejemplar volando sobre la 

isla el 30 de diciembre de 2006, mientras que M. Bautista 

(com. pers.) observó otro individuo el 26 de febrero de 

2011, sobre el muelle de La Grama. 

Sula nebouxii 

Es la especie de ave marina dominante en la isla. 

Coker (1919), entre el 29 de marzo y el 6 de abril de 

1907, e inicios de diciembre del mismo año, observó 

a esta especie cerca de su campamento, en grandes 

cantidades. También señaló que su época de reproducción 

fue ininterrumpida ya que encontró nidos con huevos, 

pichones recién nacidos, y jóvenes de diversas edades. 

Murphy (1936, 1954) y Tovar (1968) también la 

nombraron como la más abundante en LT, con nidos 

densamente distribuidos, a menudo muy próximos entre 

sí, que incluso rodeaban las casas de las guardianías. 

Posteriormente, Duffy et al. (1984) determinaron la 

presencia de 15,000 parejas en febrero de 1979. Por su 

parte, Guillén (1991) consideró a LT como una de las 

áreas con mayor población de la especie en el Perú, que, 

sin embargo, presenta amplias fluctuaciones poblacionales 

por año y por mes: 1985 (10,000 - 180,000), 1986 

(35,000 - 160,000), 1988 (11,000 - 180,000), 1989 

(2500 - 150,000), 1990 (0 - 90,000). Asimismo, se 

determinó que S. nebouxii ocupaba casi dos tercios de 

la superficie de la isla para su reproducción, con nidos 

uniformemente distribuidos en grandes sub-colonias 

(hasta 10,000 pares), separadas entre sí por colinas 

rocosas, con una estimación de la población reproductiva 

de 75,000 a 100,000 parejas (Zavalaga et al. 2007). En 

marzo de 2011, se observaron algunos pocos huevos, y 

pichones de diferentes edades, pero principalmente de 5 a 

8 semanas de nacidos, de los cuales la mayoría formaban 

grupos. La especie se encontró dispersa por toda la isla, 

pero principalmente en las áreas planas y quebradas del 

sector sur y centro. Valverde y García (2009), registraron 

en 1999, un individuo joven con plumaje aberrante, que 

fue identificado como dilución. 

Sula variegata 


cuyas poblaciones en la isla LT varían drásticamente 

dependiendo de las condiciones ambientales. Murphy 

(1936, 1954), en enero de 1920, encontró siete grandes 

colonias, todas ubicadas en terrenos planos, siendo sin 

embargo, mucho menos numerosa que S. nebouxii. En 

base a una evaluación realizada en la isla en noviembre 

de 1960 y enero de 1962, esta fue considerada como 

una de las áreas con mayor densidad poblacional de S. 

variegata en el Perú, con 400,000 y 470,100 individuos, 

respectivamente (Jordán 1963). Posteriormente, en 

diciembre de 2006 y entre enero y diciembre de 2009, la 

población reproductiva fue estimada entre 2000 y 2500 

parejas y 11,150 individuos, respectivamente (Zavalaga 

et al. 2009a, 2010a, SERNANP & AGRORURAL 

2010). Entre el 28 de febrero y 3 de marzo de 2011, se 

encontraron dos grandes colonias de esta especie: una 

adyacente a la guardianía de La Grama y otra frente a 

Roca Blanca; además se observaron otras cuatro colonias 

pequeñas: dos frente a la bahía Canevaro, una frente a la 

bahía Viveros y otra frente a la bahía Juanchuquita, las 

cuales estuvieron constituidas por pichones de diferentes 

edades, principalmente entre 3 y 5 semanas de nacidos. 

Sula granti (Figura 3c) 

Jahncke & Goya (1997b) observaron por primera 

vez, en los meses de mayo y noviembre de 1996, 10 

parejas reproductivas de S. dactylatra, ubicadas en el lado 

sureste de la isla LT, de los cuales algunos individuos 

fueron anillados. Posteriormente, estas parejas fueron 

reconsideradas como S. granti (Figueroa 2004), ya que 

en base a criterios morfológicos, ecológicos (Pitman & 

Jehl 1998, Roberson 1998) y genéticos (Friesen et al. 

2002), esta especie fue separada de S. dactylatra. En años 

posteriores se han observado individuos reproductivos 

en esta misma área y en otras zonas más hacia el oeste; 

en marzo de 1999, diciembre de 2004 y marzo de 2011, 

se observaron 17, 21 y 12 individuos reproductivos, 

respectivamente; algunos de ellos presentaron anillos de 

marcaje de las islas Galápagos y de LT. M. Bautista (com. 

pers.) observó un individuo descansando en el muelle de 

La Grama, el 27 de diciembre de 2010. 




67 

Sula leucogaster (Figura 3d) 

Si bien el golfo de Guayaquil (Ecuador) representa 

formalmente su límite sur de distribución (Nelson 1978), 

también ha sido reportada en algunas oportunidades en 

el Perú: isla Foca (Rivas 2010), islas Lobos de Afuera 

(Stucchi et al. 2011) e isla Huampanú (Lima) (Valverde 

2007). El 3 de enero de 2011, M. Bautista (com. pers.) 

observó un adulto en el muelle de La Grama. 

- Hibridación de Sulidae 

Entre los años 1997 y 2005, se observaron en las 

islas LT individuos con características de coloración 

compartidas entre S. nebouxii y S. variegata (Ayala 2006), 

que posteriormente fueron genéticamente determinados 

como híbridos (Taylor et al. 2010). En junio de 1999 y 

febrero de 2000, se observaron dos híbridos en la isla, 

frente al islote Roca Blanca. Por otro lado, en marzo de 

1999, se encontró al este del faro de LT una hembra de 

S. nebouxii y un macho de S. granti, con comportamiento 

de cortejo entre ellos. 

Phalacrocorax brasilianus 

Su reproducción en LT ha sido registrada 

anteriormente, entre los meses de diciembre y febrero 

(Forbes 1914, Coker 1919, Tovar 1968). En agosto de 

1999 y marzo de 2011 se observaron varios individuos 

adultos no reproductivos nadando principalmente en 

el muelle de La Grama y en la bahía Canevaro, entre la 

colonia de S. humboldti. 

Phalacrocorax gaimardi 

Coker (1919) en sus viajes de 1907 señaló que no 

observó a esta especie en LT, sin embargo, Forbes (1914) 

encontró varias parejas reproductivas incubando en 

febrero de 1913. Posteriormente, en diciembre de 1963, 

Tovar (1968) solo encontró dos parejas reproductivas 

que anidaban en un acantilado, sobre el apostadero del 

lobo marino. En diciembre 1999, Zavalaga et al. (2002) 

encontraron 10 individuos adultos; un año después, M. 

Bautista (com. pers.) observó dos parejas anidando en los 

acantilados cercanos al apostadero. 

Phalacrocorax bougainvillii 


cuyas poblaciones en LT varían drásticamente 

dependiendo de las condiciones ambientales. Se han 

observado desde grandes poblaciones reproductivas, 

dispersas en agregaciones pequeñas, hasta la ausencia total 

de individuos (Murphy 1936, Jordán 1963). Entre enero 

y diciembre de 2009, la población promedio en LT fue de 

36,083 individuos (SERNANP & AGRORURAL 2010). 

Las áreas de reproducción de esta especie en LT incluyen 

las bahías Canevaro y Viveros, los islotes Albatros, Smith 

y Unanue, y las zonas ventosas de las puntas norte y sur 

(Forbes 1914, Coker 1919, Vogt 1942, Murphy 1954). En 

marzo de 2011, se observaron dos grandes colonias, una 

frente a la bahía Juanchuquita y otra en el islote Unanue. 

Pelecanus thagus 


y al igual que otras aves, sus poblaciones en LT varían 

drásticamente dependiendo de las condiciones 

ambientales. En enero de 1920, Murphy (1936) 

encontró 1600 parejas distribuidas en ocho colonias. 

Posteriormente en noviembre de 1960 y enero de 1962, 

se contabilizaron 20,000 individuos y la ausencia de estos, 

respectivamente (Jordán 1963). Entre enero y diciembre 

de 2009, la población promedio fue de 6644 individuos 

(SERNANP & AGRORURAL 2010), incrementándose 

hasta 160,000 a 210,000 parejas un año después (Zavalaga 

et al. 2011). Entre febrero y marzo de 2011, se observaron 

miles de pichones de diferentes edades (0 a 4 semanas 

de nacidos), de los cuales la mayoría formaban grupos 

mientras esperaban ser alimentados. Estos se encontraban 

asentados principalmente en el lado norte de la isla. 

Egretta thula 

M. Bautista y A. Oliva (com. pers.) observaron un 

individuo en el muelle de La Grama, en diciembre de 2010. 

Cathartes aura jota 

Es una especie común en la isla, la cual fue señalada 

como muy abundante a inicios del siglo XX (Forbes 

1914). Su reproducción en LT ha sido registrada 

en diciembre (Coker 1919). En marzo de 2011, se 

observaron individuos jóvenes y adultos dispersos por 

toda la isla, pero principalmente cerca a las colonias de 

las aves guaneras y S. nebouxii, así como en las áreas más 

altas. Además se encontró un pichón a 500 m al oeste de 

la guardianía de La Grama. Zavalaga (2003) y Figueroa 

et al. (2011b) registraron la presencia de individuos 

con plumaje aberrante de color blanco, observados en 

diversas áreas de la isla entre 1999 y 2011. Estos fueron 

identificados como leucísticos (Figueroa et al. 2011b). 

Falco peregrinus (Figura 3e) 

M. Bautista y S. García (com. pers.) observaron el 

15 y 16 de enero de 2011 a un F. peregrinus cazar a un 

individuo adulto de S. nebouxii y otro de Larus belcheri, 


68 



respectivamente. El primer ataque fue observado en la 

zona sureste de la isla y el segundo en el lado norte, cerca 

a Cabo Cruz. El 28 de febrero de 2011 se encontró una 

hembra adulta parada a unos 10 m de una colonia de S. 

nebouxii, a 500 m de El Ñopo. 

Charadrius nivosus 

En noviembre de 2010, M. Bautista y A. Oliva (com. 

pers.) observaron un grupo de 20 individuos hacia el norte 

de la playa La Grama. 

Oreopholus ruficollis pallidus 

H. O. Forbes colectó una hembra el 19 de junio de 

1912 en LT (AMNH 2011). 

Haematopus palliatus pitanay 

Se colectaron especímenes procedentes de LT 

(Murphy 1925b, 1936). 

Haematopus ater 

Su primer registro fue hecho por Coker (1919) en 

1907. M. Bautista (com. pers.) observó dos individuos en 

unos roqueríos de El Ñopo, a mediados de 2010. 

Numenius phaeopus hudsonicus 

M. Bautista (com. pers.) observó un individuo al 

norte de la playa La Grama el 15 de febrero de 2011. 

Tringa incana 

Fue observada por Coker (1919) el 3 de diciembre 

de 1907. 

Calidris alba (Figura 3f) 

Fue observada por Coker (1919) el 10 de diciembre 

de 1907. El 1 de marzo de 2011, en el área central de esta 

isla, se encontró un individuo muerto. 

Calidris mauri 

El 13 de diciembre de 1907, Coker (1919) observó 

a esta especie en LT. 

Stercorarius chilensis 

Fue registrada por R. H. Beck en LT el 7 de 

septiembre de 1913 (Murphy 1936). 

Creagrus furcatus 

Varias bandadas de esta especie fueron observadas 

por Murphy (1936) entre Lobos de Afuera y LT, el 6 de 

enero de 1920. 

Chroicocephalus cirrocephalus (Figura 3g) 

El 3 de marzo de 2011, desde una embarcación 

artesanal, se observó un individuo adulto volando frente 

a la bahía de Juanchuquita. 

Leucophaeus modestus (Figura 3h) 

Murphy (1936) mencionó su sorpresa sobre la 

ausencia de registros de esta especie en LT, ya que es 

común su observación en otras islas del Perú, y además, al 

estar ligada a las playas de arenas, tendría en LT muchas 

áreas con este hábitat. Se observó un individuo adulto 

en la playa El Ñopo, el 11 de agosto de 1999 y otro el 

28 de febrero de 2011. A pesar de ser un ave endémica 

de la Corriente de Humboldt, es visitante de la costa del 

Pacífico entre Ecuador y Chile, llegando ocasionalmente 

hasta Colombia y Argentina (Harrison 1987), por lo que 

ciertamente es rara su poca presencia en LT. 

Leucophaeus pipixcan 

Fue observada en grandes números, descansando en 

las playas de arena de LT, en diciembre de 1920 (Murphy 

1936). 

Larus belcheri 

Murphy (1936) observó la concentración de esta 

especie en la bahía Canevaro, cerca de una colonia de 

P. bougainvillii, a quienes les robaban sus huevos. En la 

actualidad su presencia es muy escasa. El 28 de febrero 

se observaron dos individuos al sur de la guardianía de 

El Ñopo, y el 2 de marzo de 2011 se encontraron seis 

individuos en los alrededores del apostadero del lobo 

marino. 

Larus dominicanus 

Desde sus primeros registros en LT, ha sido 

señalada como abundante (Coker 1919), incluso, LT fue 

determinada como uno de los centros más grandes de 

reproducción de la especie en el mundo (Murphy 1936). 

En febrero de 1979, Duffy et al. (1984) calcularon un 

máximo de 1000 parejas. En base al registro fotográfico, 

entre el 28 de febrero y 3 de marzo de 2011, la 

población fue estimada entre 500 y 600 individuos. 

Esta disminución estaría relacionada a la constante 

destrucción de sus nidos y huevos, con el objetivo de 

reducir la depredación de las gaviotas sobre los huevos 

y pichones de las aves guaneras (Figueroa 2010). Su 

reproducción ha sido registrada en LT entre noviembre 

y diciembre (Tovar 1968, Figueroa 2010), sin embargo a 

fines de febrero se encontraron pichones recién nacidos 

e incluso huevos recién puestos. 




69 

a 

b 

c 

d 

e 

f 

g 

h 

FIGURA 3. (a) Spheniscus humboldti en la bahía Canevaro. Foto: J. F. (b) Phoebastria irrorata en La Grama. Foto: I. García-Godos. (c) Hembra y 

pichón de Sula granti en el sureste de la isla. Foto: J. F. (d) Sula leucogaster en el muelle de La Grama. Foto: M. Bautista. (e) Hembra de Falco peregrinus 

en El Ñopo. Foto: J. F. (f) Individuo muerto de Calidris alba en el centro de la isla. Foto: J. F. (g) Chroicocephalus cirrocephalus volando sobre la bahía 

Juanchuquita. Foto: J. F. (h) Leucophaeus modestus en El Ñopo. Foto: J. F. 

FIGURE 3. (a) Spheniscus humboldti in the Canevaro bay. Photo: J. F. (b) Phoebastria irrorata in La Grama. Photo: I. García-Godos. (c) Female and 

pigeon of Sula granti in the southeast of the island. Photo: J. F. (d) Sula leucogaster on the dock of La Grama. Photo: M. Bautista. (e) Female of Falco 

peregrinus in El Ñopo. Photo: J. F. (f) Individual died of Calidris alba in the center of the island. Photo: J. F. (g) Chroicocephalus cirrocephalus flying 

over the Juanchuquita bay. Photo: J. F. (h) Leucophaeus modestus in El Ñopo. Photo: J. F. 


70 



Sternula lorata 

Murphy (1936) observó algunos individuos pescando 

en la punta sur de LT en enero de 1920; también los 

observó a 3.5 km de LT, pescando junto con Oceanodroma 

markhami y otras golondrinas de mar (Murphy 1925a). En 

junio de 2006, I. García-Godos (Zavalaga et al. 2009b), 

observó un grupo de aproximadamente 100 individuos 

descansando en una llanura a 1.5 km al noroeste de la 

guardianía de La Grama. A fines de 2010, M. Bautista 

(com. pers.) observó varios individuos descansando en una 

playa de arena al sureste de La Grama. 

Larosterna inca 

Tovar (1968) encontró colonias reproductivas de 

esta especie en LT. M. Bautista y S. García (com. pers.) 

observaron pequeños grupos en los acantilados de la zona 

noroeste de la isla. Se desconoce el estado de la población 

en LT. 

Estado de conservación y problemática 

Del total de especies registradas, trece se encuentran 

consideradas dentro de alguna de las categorías de 

protección del Decreto Supremo 034-2004-AG de 

la legislación peruana (MINAG 2004). Asimismo, 

trece especies están incluidas en las listas de la Unión 

Internacional para la Conservación de la Naturaleza 

(IUCN 2011); tres en la Convención sobre el Comercio 

Internacional de Especies Amenazadas de Fauna y Flora 

Silvestres (CITES 2011); y cinco en la Convención sobre 

la Conservación de las Especies Migratorias de Animales 

Silvestres (CMS 2009) (Tabla 1). Algunas de las especies 

registradas son afectadas directamente por factores como 

disturbios humanos y la presencia de fauna exótica. 

Ingreso de pescadores - Como parte de su riqueza 

marina, la isla LT posee bancos naturales de peces de peña, 

cangrejos, pulpo (Octopus mimus), percebes (Pollicipes 

elegans) y conchas de abanico (Argopecten purpuratus), de 

los cuales la última constituye uno de los más importantes 

del Perú (Carbajal et al. 2004a). Debido a su proximidad 

con la costa, estas especies están sujetas a una fuerte presión 

pesquera (Carbajal et al. 2005a). Desde las primeras 

investigaciones realizadas en la isla relacionadas a las aves, 

se ha señalado a estos pescadores como saqueadores de 

huevos y pichones, principalmente de P. bougainvillii, 

para negociarlos en tierra (Forbes 1914). 

Extracción del guano - Asimismo, desde inicios del 

siglo pasado, Forbes (1914) y Coker (1919) comentaron 

sobre la perturbación de las aves guaneras en las islas del 

norte y su desplazamiento hacia otras islas debido a la 

extracción del guano. En el caso de P. thagus, llegaron a 

abandonar sus huevos y pichones recién nacidos, los cuales 

fueron atacados por L. dominicanus, L. belcheri y C. aura 

(Forbes 1914, Vogt 1942). En la actualidad, la isla LT 

alberga un importante yacimiento de 20,000 t de guano 

fosfatado, del cual a fines de 2010 fueron extraídos solo 

200 t (SERNANP & AGRORURAL 2010). A finales de 

diciembre de 2010, miles de nidos fueron abandonados 

por parte de P. thagus. Es posible que una de las causas 

de este evento sean las perturbaciones producidas por 

el personal que trabajó en la campaña de extracción del 

guano (Figueroa & Stucchi 2012). 

Presencia de gatos cimarrones (Felis catus) - En 

marzo y junio de 1999, se observaron dos gatos en el lado 

sureste de LT. Posteriormente, el 1 de marzo de 2011, se 

encontraron su huellas entre la colonia de P. thagus en 

la bahía Viveros. Al día siguiente, se hallaron en el lado 

centro-norte, heces frescas junto a un nido de S. nebouxii. 

Ese mismo día, a las 09 h 21 min, se observaron dos 

cachorros en el lado centro-oeste, descansando en un área 

sombreada; al vernos uno salió huyendo mientras que 

el otro se quedó observándonos por unos segundos. Al 

parecer, estos gatos cimarrones se distribuyen por toda la 

isla (Figuras 4a y 4b). 

Coker (1919) señaló que los gatos introducidos en 

las islas Lobos, se tornaron cimarrones, alimentándose 

de los huevos y pichones del P. thagus. Asimismo, Vogt 

(1942) y Duffy et al. (1984) observaron que estos 

también atacaron a individuos jóvenes de S. variegata y 

S. nebouxii. Los tres autores recomendaron como acción 

primordial su erradicación. Sin embargo, a pesar que 

el gato cimarrón ha sido identificado como uno de los 

depredadores introducidos por el hombre con mayor 

impacto sobre las poblaciones de aves marinas en las 

islas (Ebenhard 1988, Álvarez-Romero et al. 2008), en 

LT la presencia de este felino data de más de un siglo y, 

al parecer, no ha interferido con las poblaciones de aves 

guaneras y S. nebouxii. Sin embargo, para otras especies 

menos numerosas, podrían representar un peligro. En 

este sentido, será importante desarrollar un estudio 

detallado al respecto, para evaluar si existe un verdadero 

impacto. 

AGRADECIMIENTOS 

A Ignacio García-Godos por sus comentarios 

al manuscrito y el préstamo de la foto de Phoebastria 

irrorata. A Marcelo Stucchi, por su apoyo en el trabajo 

de campo del 2004 y 2011, así como sus comentarios 

al manuscrito y el préstamo de sus fotos. A Gina Mori, 

por compartir sus registros y su apoyo en el trabajo de 

campo en el 2011. A AGRORURAL por permitirnos 

el uso de sus instalaciones, y a sus guarda-islas Mariano 




71 

a 

b 

FIGURA 4. (a) Cachorro de gato en el centro-oeste de la isla. Foto: J. F. (b) Heces de gato en el centro-norte de la isla. Foto: M. Stucchi. 

FIGURE 4. (a) Kitten in the center-west of the island. Photo: J. F. (b) Cat feces in the central-north of the island. Photo: M. Stucchi. 


72 



Bautista, Simón García y Alan Oliva por compartir 

sus observaciones; quienes además conjuntamente con 

Flaviano Zenón nos brindaron las comodidades en la 

isla durante los días de trabajo. A Ismael Ignacio y su 

tripulación por trasladarnos a la isla. A Manuel Plenge 

por el préstamo de bibliografía y sus comentarios sobre 

Thalassarche salvini. A Fernando Angulo por compartir sus 

observaciones de los flamencos en la laguna de Ñapique. 

A Edward Soldaat, por la confirmación de Calidris alba. 

Al revisor anónimo por sus acertados comentarios. 

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March 2013 

article 


Alessandro Pacheco Nunes 1, 2, 11 , Maurício Neves Godoi 1 , Maria Antonietta Castro Pivatto 3 , José Carlos Morante 

Filho 4 , Eduardo Weffort Patrial 5 , Paulo Antonio Silva 6 , Vanessa Katherinne Stavis 7 , Daniel De Granville Manço 3 , 

Marco de Barros Costacurta 8 , Caroline Leuchtenberger 9 e Carlos Rodrigo Lehn 10 

1 

Pós-Graduação em Ecologia e Conservação, Universidade Federal de Mato Grosso do Sul. 

2 

Laboratório de Vida Selvagem, Rua 21 de Setembro, Bairro Nossa Senhora de Fátima, C.P. 109, 79320-900, Corumbá, MS. 

3 

Instituto das Águas da Serra da Bodoquena, Rua 24 de fevereiro, 1507, 2º. Piso, Centro, 79290-000, Bonito, MS. 

4 

Pós-Graduação em Ecologia e Conservação da Biodiversidade. Universidade Estadual de Santa Cruz, Km 16, Rod. Ilhéus-Itabuna, 45662-900, Ilhéus, BA. 

5 

MAE (Meio Ambiente Equilibrado), Rua Piauí, 854, Apto. 401, Centro, 86020-390, Londrina, PR. 

6 

Pós-Graduação em Ecologia e Conservação de Recursos Naturais, Universidade Federal de Uberlândia, Bloco 2D, s/n, 38400-902, Uberlândia, MG. 

7 

Pós-Graduação em Biologia Animal, Universidade Federal de Mato Grosso do Sul, Cidade Universitária s/n, C.P. 549, 79070-900, Campo Grande, MS. 

8 

Pós-Graduação em Meio Ambiente e Desenvolvimento Regional, Universidade para o Desenvolvimento do Estado e Região do Pantanal, Rua 

Ceará, 333, Câmpus I, Bloco II, Bairro Miguel Couto, 79003-010, Campo Grande, MS. 

9 

Pós-Graduação em Ecologia, Instituto Nacional de Pesquisas da Amazônia, C.P. 478, 69011-970, Manaus, AM. 

10 

Instituto Federal de Educação, Ciência e Tecnologia de Mato Grosso do Sul, Rua Pereira Gomes 355, 2º piso, 79400-000, Coxim, MS. 

11 

Autor correspondente: udu@ibest.com.br. 

Recebido em 30 de julho de 2011. Aceito em 19 de janeiro de 2013. 

Abstract: Birds of the Serra de Maracaju, Mato Grosso do Sul, Brazil. The Serra de Maracaju stands out in the orography of 

Mato Grosso do Sul as a watershed between the Upper Paraguay and Upper Parana river basins. In this study, we report on the 

occurrence of 413 species of birds based on historical records and field data collected by us. The records of species such as Aburria 

nattereri, Ictinia mississippiensis, Spizaetus tyrannus, Micrococcyx cinereus, Berlepschia rikeri, Oxyruncus cristatus, Knipolegus lophotes, 

Myiarchus tuberculifer, Tyrannopsis sulphurea, Tityra semifasciata, Cyanerpes cyaneus, and Poospiza cinerea are the first for the state of 

Mato Grosso do Sul. The Serra de Maracaju act as an important dispersal corridor for elements of the Amazon and Atlantic Forest 

in the eastern border of the Pantanal. Fifteen species reported for the area are regarded as globally or nationally endangered, such 

Harpia harpyja, Alectrurus tricolor, and Sporophila maximiliani. Human impacting activities such as agriculture, monoculture of 

exotic trees, and conversion of wood into charcoal, seriously threaten the survival of these and other bird species occurring in the 

region. The creation of a large conservation unit and the proper management of the landscape, so as to maintain the local diversity 

and habitat structure, are crucial to ensure the conservation these species and, therefore, the biodiversity of the surrounding plateaus 

and plains of Pantanal. 

Key-words: bird communities, central Brazil, cerrado, conservation. 

Introdução 

O Cerrado é um dos principais hotspots para a 

conservação da biodiversidade mundial (Myers et al. 

2000). Destaca-se como a mais rica flora dentre as savanas 

tropicais, com mais de 7.000 espécies identificadas, 

das quais 40% são endêmicas (Ratter et al. 1997). No 

entanto, nas últimas três décadas mais da metade dos seus 

2 milhões de km 2 de vegetação original foram suprimidos 

e/ou substituídos por pastagens exóticas (Brachiaria), 

culturas anuais (em especial soja, milho e cana-deaçúcar) 

e monoculturas de Pinus e Eucalyptus (Klink 

& Machado 2005). Originalmente, o Cerrado cobria 

aproximadamente 61% do território de Mato Grosso do 

Sul, entretanto, a ocupação do estado a partir da década 

de 60 reduziu drasticamente a cobertura para apenas 32% 

(Sano et al. 2010). 

No aspecto geomorfológico, o estado de Mato 

Grosso do Sul é caracterizado principalmente pela 

planície do Pantanal e seus planaltos circundantes, tais 

como Urucum, Amolar, Bodoquena, Maracaju - Campo 

Grande e Taquari - Itiquira, sendo estes dois últimos 

formadores da Serra de Maracaju (Boggiani et al. 1998). 

A avifauna da Serra de Maracaju tem sido estudada 

desde 1926, quando o naturalista E. R. Blake, a serviço 

do The Field Museum of Natural History, coletou 

exemplares de várias espécies na Fazenda Capão Bonito 

(Straube 2011). Posteriormente, destacaram-se as 

expedições do Museu de Zoologia da Universidade de São 

Paulo, coordenadas por Olivério Pinto e colaboradores, 

que coletaram vários espécimes em Aquidauana e Coxim 

(Pinto 1932, 1938, 1944, 1964). Os membros do 

Museum of Comparative Zoology também visitaram 

a região em 1931, quando realizaram várias coletas

76 


Alessandro Pacheco Nunes, Maurício Neves Godoi, Maria Antonietta Castro Pivatto, José Carlos Morante Filho, Eduardo Weffort Patrial, Paulo Antonio Silva, 

Vanessa Katherinne Stavis, Daniel De Granville Manço, Marco de Barros Costacurta, Caroline Leuchtenberger e Carlos Rodrigo Lehn 

em Aquidauana (Tubelis & Tomas 2003). Em 1981 o 

naturalista J. Hidasi coletou espécimes em Aquidauana, 

que atualmente encontram-se depositados na coleção da 

Fundação Museu de Zoologia, em Goiânia, Goiás. 

Apesar dessas importantes contribuições, a avifauna 

da Serra de Maracaju ainda é pouco conhecida. Neste 

estudo apresentamos uma lista das espécies de aves 

registradas na região, com base em dados literários e 

registros obtidos em estudos de campo realizados pelos 

autores. 

Métodos 

Área de estudo 

A Serra de Maracaju estende-se no sentido norte-sul 

por todo o estado de Mato Grosso do Sul, atuando como 

um divisor de águas entre as bacias hidrográficas do 

Alto Rio Paraguai, a oeste, e do Alto Rio Paraná, a leste 

(Boggiani et al. 1998). Segundo Damascenso et al. (2000), 

a Serra de Maracaju é coberta principalmente por cerradão 

(savana florestada), florestas estacionais semideciduais, 

vegetação ripária (mata de galeria e mata ciliar) e veredas. 

Entretanto, o desenvolvimento agropecuário reduziu e 

alterou drasticamente a paisagem natural da região, cujos 

remanescentes naturais encontram-se imersos em uma 

matriz de pastagens exóticas e monoculturas (Harris et 

al. 2006). 

O clima na região é tropical subúmido (Aw), 

com estações chuvosa (novembro a abril) e seca (maio 

a outubro) bem definidas. A precipitação anual pode 

atingir até 1.180 mm, com temperatura média mensal 

oscilando entre 21°C e 33°C (Soriano & Alves 2005). 

Amostragem 

A lista de espécies da Serra de Maracaju foi produzida 

com base em revisão de registros históricos obtidos por 

E. R. Blake a serviço do The Field Museum of Natural 

History, bem como dados de bibliografia (Pinto 1940, 

Tubelis & Tomas 2003, Hass 2004, Junqueira 2008, 

Whittaker et al. 2008, Nunes et al. 2012) e estudos de 

campo realizados pelos autores entre 2005 e 2011. No 

entanto, não foi possível resgatar informações referentes 

à natureza dos registros (visual, auditivo ou espécime 

coletado) publicados por Hass (2004). 

Informações sobre os exemplares depositados 

em coleções ornitológicas foram obtidos com base em 

Tubelis & Tomas (2003) e compilado com base nas 

seguintes instituições: Museum of Comparative Zoology 

- MCZ (Cambridge, EUA); The Field Museum of 

Natural History - FMNH (Chicago, EUA); Museu de 

Zoologia da Universidade de São Paulo – MZUSP (São 

Paulo, SP); Fundação Museu de Ornitologia – FMO 

(Goiânia, GO). 

As localidades amostradas, coordenadas, autores, 

período de estudo e esforço amostral estão sintetizados 

na tabela 1. De forma geral, o método utilizado pelos 

autores na coleta de dados foi o censo por observação 

direta (Bibby et al. 1992), que consiste em caminhar por 

diferentes tipos de habitats obtendo registros visuais e 

auditivos do maior número possível de espécies de aves. 

A lista de espécies segue a ordenação taxonômica e a 

nomenclatura científica propostas pelo Comitê Brasileiro 

de Registros Ornitológicos (CBRO 2011), com exceção 

das propostas recentemente indicadas para as famílias 

Caprimulgidae e Thraupidae/Emberizidae e, ainda, de 

situações provisórias (incertae sedis) de alguns grupos 

ou espécies, para os quais utilizamos edição anterior da 

mesma lista (CBRO 2009). Adicionalmente, adotamos 

os limites de espécies para Aburria Reichenbach, 1853 

apresentados por Lopes (2009) contra Grau et al. (2004) 

e CBRO (2011), particularmente na aceitação de A. grayi 

e A. nattereri como espécies plenas. 

Resultados e Discussão 

Neste estudo relacionamos a ocorrência de 413 

espécies distribuídas em 303 gêneros e 66 famílias de aves 

para a Serra de Maracaju (Tabela 2). Aproximadamente 

63% desse total apresentam registros comprobatórios de 

ocorrência na região por meio de coleta de espécime(s), 

fotografia, gravação de áudio e outros tipos de 

documentos que permitam a determinação segura e 

a aferição posterior do táxon. As demais estão inclusas 

na lista secundária, pois ainda aguardam documentação 

comprobatória adequada conforme as normas propostas 

pelo CBRO (2011). 

A região abriga aproximadamente metade das 

espécies de aves ocorrentes no Mato Grosso do Sul 

(Nunes et al. no prelo). Comparada à avifauna de outros 

planaltos de Mato Grosso do Sul, a Serra da Maracaju 

pode ser considerada muito diversificada. No Planalto 

da Bodoquena, Pivatto et al. (2006) relacionaram 

a ocorrência de 353 espécies. Para a borda oeste do 

Pantanal, que inclui a Serra do Amolar e o Maciço do 

Urucum, Tomas et al. (2010) citam a ocorrência de 380 

espécies. 

Entretanto, a riqueza de espécies de aves da Serra 

de Maracaju ainda pode estar sendo subestimada, uma 

vez que o esforço amostral empregado em algumas 

localidades visitadas nesse estudo foi pequeno e pontual. 

Assim como em várias outras regiões do Mato Grosso 

do Sul, ainda existem grandes lacunas de conhecimento 

sobre a avifauna ocorrente nos planaltos e bordas da Serra 

de Maracaju. 





77 

Tabela 1. Localidades com registros de espécies de aves na Serra de Maracaju, Mato Grosso do Sul, Brasil. 

Table 1. Localities with records of bird species in the Serra de Maracaju, state of Mato Grosso do Sul, Brazil. 

N Localidades Coordenadas Município Altitude (m) Fonte Período 

1 PCH Santa Gabriela 17º32’18”S, 

54º26’32”O 

Sonora 540 F, G 2010 (fevereiro, maio, agosto e novembro); 2011 

(fevereiro); 140 horas/observações 

2 Sonora 17º33’45”S, 

54º48’45”O 

Sonora 440 D, F 2008 (março); 2011 (abril, agosto e dezembro); 2012 

(março); 80 horas/observações 

3 Parque Estadual Nascentes 

do rio Taquari 

18º02’25”S, 

53º20’03”O 

Costa Rica e Alcinópolis - C 1998 a 2000; 100 horas/observações 

4 Fazenda Recreio 18º13’27”S, 

54º39’03”O 

Coxim - A 1937 (agosto) 

5 Coxim 18º30’02”S, 

54º48’05”O 

Coxim 260 H 2011 (setembro); 8 horas/observações 

6 Rio Verde 18º56’13”S, 

54º54’07”O 

Rio Verde de Mato Grosso 380 O 2011 (novembro) 

7 Fazenda Trilha do Sol 19º26’40”S, 

54º49’12”O 

Rio Negro 406 I 2011 (março e dezembro); 100 horas/observações 

8 Chapadão 19º28’00”S, 

54º49’20”O 

Rio Negro 630 I 2011 (março e dezembro); 20 horas/observações 

9 Pousada Quinta do Sol 19º46’02”S, 

55º14’35”O 

Corguinho 869 J 2005 (agosto); 2006 (janeiro); 18 horas/observações 

10 Fazenda Rodeio 19º44’59”S, 

55º09’42”O 


11 Fazenda Constantino 19º49’02”S, 

55º15’18”O 


12 RPPN Vale do Bugio 19º56’20”S, 

55º04’26”O 

Corguinho 370 K 2009 (outubro); 25 horas/observações 

13 RPPN Gavião de Penacho 19º57’13”S, 

55º03’48”O 

Corguinho 451 K 2009 (outubro); 31 horas/observações 


78 




N Localidades Coordenadas Município Altitude (m) Fonte Período 

14 Brejão 20º07’05”S, 

55º23’08”O 

Cipolândia 830 J 2005 (setembro); 2006 (fevereiro); 6 horas/observações 

15 Fazenda Bocaina 20º04’00”S, 

55º34’60”O 

Aquidauana 534 J 2005 (setembro); 2006 (fevereiro); 16 horas/observações 

16 Fazenda Taboco 20º04’10”S, 

55º38’46”O 

Aquidauana 524 J 2005 (setembro); 2006 (fevereiro); 20 horas/observações 

17 Fazenda Estância Crioula 20º30’02”S, 

55º31’60”O 

Dois Irmãos do Buriti 680 L 2010 (abril e maio); 20 horas/observações 

18 Aquidauana 20º29’15”S, 

55º48’45”O 

Aquidauana 150 B, E 1930 a 1931; 2003 (outubro) 

19 Fazenda Jatiúca 20º31’57”S, 

55º50’19”O 

Aquidauana 182 K 2008 (fevereiro e setembro); 27 horas/observações 

20 Fazenda Boa Esperança 20º43’32”S, 

56º02’24”O 


21 Fazenda Correntes 20º34’09”S, 

55º24’24”O 


22 Fazenda Vô Fiorindo 20º35’44”S, 

55º25’21”O 


23 Fazenda Taruana 20º30’16”S, 

55º15’51”O 


24 Sidrolândia 20º57’15”S, 

55º03’30”O 

Sidrolândia 350 M 2011 (abril); 4 horas/observações 

25 Fazenda Capão Bonito 21º17’05”S, 

54º49’02”O 

Sidrolândia - N 1926 (julho, agosto e novembro); 1937 (setembro e 

outubro) 

26 Usina Vista Alegre 21º47’30”S, 

55º32’53”O 

Maracaju 560 F 2009 (dezembro); 2010 (janeiro, abril, outubro e 

novembro); 120 horas/observações 

Fonte/Source: A (Pinto 1940); B (Tubelis & Tomas 2003); C (Hass 2004); D (Junqueira 2008); E (Whittaker et al. 2008); F (Maurício Neves Godoi); G (José Carlos Morante Filho); H (Caroline Leuchtenberger 

e Carlos Rodrigo Lehn); I (Eduardo Weffort Patrial); J (Alessandro Pacheco Nunes, Paulo Antonio Silva e Marco de Barros Costacurta); K (Maria Antonieta Castro Pivatto); L (Vanessa Katherinne Stavis); M 

(Daniel De Granville Manço); N (The Field Museum of Natural History, Chicago); O (Nunes et al. 2012). 





79 

Tabela 2. Espécies de aves registradas na Serra de Maracaju, Mato Grosso do Sul. Registros: e (espécime coletado), v (visual), a (auditivo), f (foto), g (zoofonia), i (video). Acrônimos das instituições que abrigam 

espécimes: The Field Museum of Natural History, Chicago (FMNH); Museum of Comparative Zoology, Cambridge (MCZ); Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP); Fundação 

Museu de Ornitologia, Goiânia (FMO). 

Table 2. Bird species recorded in the Serra de Maracaju, state of Mato Grosso do Sul. Records: e (skin), v (visual records), a (vocal records), f (photo), g (taped vocalization), i (vídeo). Institutions with specimens: 

The Field Museum of Natural History, Chicago (FMNH); Museum of Comparative Zoology, Cambridge (MCZ); Museu de Zoologia da Universidade de São Paulo, São Paulo (MZUSP); Fundação Museu de 

Ornitologia, Goiânia (FMO). 

Táxons Localidades Registros Acrônimos 

Struthioniformes 

Rheidae 

Rhea americana (Linnaeus, 1758) 1, 2, 3, 7, 9, 10, 11, 16, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH 

Tinamiformes 

Tinamidae 

Crypturellus undulatus (Temminck, 1815) 1, 2, 3, 4, 7, 11, 12, 13, 15, 16, 19, 20, 21, 22, 23, 26 e, v, a MZUSP 

Crypturellus parvirostris (Wagler, 1827) 1, 2, 3, 7, 9, 10, 11, 12, 13, 18, 19, 20, 21, 22, 23, 25, 26 e, v (f), a (g) FMNH, MZUSP 

Crypturellus tataupa (Temminck, 1815) 1, 2, 3, 10, 15, 19, 21, 26 a 

Rhynchotus rufescens (Temminck, 1815) 1, 2, 3, 7, 8, 9, 10, 13, 20, 21, 22, 23, 25, 26 e, v, a FMNH 

Nothura minor (Spix, 1825) 25 e FMNH 

Nothura maculosa (Temminck, 1815) 2, 3, 8, 9, 10, 14, 23, 25, 26 e, v (f) FMNH 

Anseriformes 

Anhimidae 

Anhima cornuta (Linnaeus, 1766) 2, 3, 14, 23, 26 v (f), a 

Chauna torquata (Oken, 1816) 15, 16 v, a 

Anatidae 

Dendrocygna viduata (Linnaeus, 1766) 2, 3, 10, 16, 20, 21, 25, 26 e, v FMNH 

Dendrocygna autumnalis (Linnaeus, 1758) 2, 15, 16 v 

Cairina moschata (Linnaeus, 1758) 1, 2, 3, 7, 15, 22, 23 v (f) 

Amazonetta brasiliensis (Gmelin, 1789) 2, 3, 11, 13, 14, 15, 18, 20, 22, 25, 26 e, v (f) FMNH, MZUSP 

Anas bahamensis Linnaeus, 1758 2 v (f) 

Nomonyx dominica (Linnaeus, 1766) 18 e MZUSP 

Galliformes 

Cracidae 

Ortalis canicollis (Wagler, 1830) 15, 16, 19, 20, 21 v, a 

Penelope superciliaris Temminck, 1815 1, 3, 13, 20, 21, 22, 26 v (f), a 


80 





Aburria grayi Pelzeln, 1870 7, 11, 15, 16, 20, 22, 23 v (f), a (g) 

Aburria nattereri Reichenbach, 1862 4 e MZUSP 

Crax fasciolata Spix, 1825 1, 3, 9, 12, 16, 20, 25, 26 e, v (f), a (g) FMNH 

Podicipediformes 

Podicipedidae 

Tachybaptus dominicus (Linnaeus, 1766) 2, 3, 18, 20, 25, 26 e, v FMNH, MZUSP, MCZ 

Ciconiiformes 

Ciconiidae 

Ciconia maguari (Gmelin, 1789) 1, 2 v 

Jabiru mycteria (Lichtenstein, 1819) 2, 14, 26 e, v FMNH 

Mycteria americana Linnaeus, 1758 2, 16 v 

Suliformes 

Phalacrocoracidae 

Phalacrocorax brasilianus (Gmelin, 1789) 1, 2, 16, 20 v 

Anhingidae 

Anhinga anhinga (Linnaeus, 1766) 1, 2, 16, 20 v 

Pelecaniformes 

Ardeidae 

Tigrisoma lineatum (Boddaert, 1783) 2, 9, 11, 14, 16, 20, 22, 23, 25, 26 e, v, a FMNH 

Tigrisoma fasciatum (Such, 1825) 6 v (f) 

Cochlearius cochlearius (Linnaeys, 1766) 16 v 

Nycticorax nycticorax (Linnaeus, 1758) 16 v, a 

Butorides striata (Linnaeus, 1758) 1, 2, 3, 7, 9, 10, 11, 16, 20, 22, 25, 26 e, v, a FMNH 

Bubulcus ibis (Linnaeus, 1758) 1, 3, 9, 10, 11, 13, 15, 16, 20, 21, 22 v (f) 

Ardea cocoi Linnaeus, 1766 2, 16, 18 v, a 

Ardea alba Linnaeus, 1758 1, 2, 3, 7, 9, 14, 18, 20, 26 e, v MZUSP 

Syrigma sibilatrix (Temminck, 1824) 1, 2, 3, 7, 9, 10, 11, 16, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH 

Pilherodius pileatus (Boddaert, 1783) 3, 9, 18, 22 e, v MZUSP 

Egretta thula (Molina, 1782) 11, 15, 16, 20, 26 v 

Threskiornithidae 

Plegadis chihi (Vieillot, 1817) 1, 2, 3, 4, 9, 12, 13, 16, 19, 20, 23, 26 e, v, a MZUSP 

Mesembrinibis cayennensis (Gmelin, 1789) 18, 20 v 

Phimosus infuscatus (Lichtenstein, 1823) 2, 16, 18 e, v, a MZUSP 





81 


Theristicus caerulescens (Vieillot, 1817) 1, 3, 7, 9, 10, 12, 13, 16, 18, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Theristicus caudatus (Boddaert, 1783) 1, 2 e, v MZUSP 

Cathartiformes 

Cathartidae 

Cathartes aura (Linnaeus, 1758) 1, 2, 3, 7, 10, 12, 13, 15, 16, 19, 20, 21, 22, 23, 26 v (f) 

Cathartes burrovianus Cassin, 1845 1, 2, 3, 9, 15, 16, 20, 26 v (f) 

Coragyps atratus (Bechstein, 1793) 1, 2, 3, 7, 9, 10, 11, 12, 13, 14, 15, 16, 18, 20, 21, 22, 23, 24 v (f) 

Sarcoramphus papa (Linnaeus, 1758) 2, 3, 7, 8, 10, 12, 15, 26 v (f) 

Accipitriformes 

Pandionidae 

Pandion haliaetus (Linnaeus, 1758) 16 v 

Accipitridae 

Leptodon cayanensis (Latham, 1790) 1, 7 v 

Elanoides forficatus (Linnaeus, 1758) 1 v 

Gampsonyx swainsonii Vigors, 1825 1, 7, 19 v (f) 

Elanus leucurus (Vieillot, 1818) 1, 3 v 

Harpagus diodon (Temminck, 1823) 6 v 

Circus buffoni (Gmelin, 1788) 1, 25, 26 e, v FMNH 

Accipiter striatus Vieillot, 1808 1 v 

Accipiter bicolor (Vieillot, 1817) 4 e MZUSP 

Ictinia mississippiensis (Wilson, 1811) 18 v (i) 

Ictinia plumbea (Gmelin, 1788) 1, 2, 3, 7, 10, 13, 20, 21, 22, 23 v 

Busarellus nigricollis (Latham, 1790) 2, 3, 16, 25 v, a 

Rostrhamus sociabilis (Vieillot, 1817) 16, 20, 25 e, v, a FMNH 

Geranospiza caerulescens (Vieillot, 1817) 1, 7, 9, 18, 22 v 

Heterospizias meridionalis (Latham, 1790) 1, 2, 3, 4, 7, 10, 11, 16, 20, 21, 22, 23, 25 e, v (f), a FMNH, MZUSP 

Urubitinga urubitinga (Gmelin, 1788) 1, 2, 3, 16, 20 v 

Urubitinga coronata (Vieillot, 1817) 3, 5 v (f), a 

Rupornis magnirostris (Gmelin, 1788) 1, 2, 7, 9, 10, 11, 13, 15, 16, 18, 19, 20, 21, 22, 23, 26 e, v (f), a MZUSP 

Geranoaetus albicaudatus Vieillot, 1816 1, 2, 3, 7, 8, 11, 23, 26 v (f) 

Pseudastur albicollis (Latham, 1790) 7 v (f) 

Buteo brachyurus Vieillot, 1816 18, 19 v 

Buteo albonotatus Kaup, 1847 1, 21, 22 v 


82 





Harpia harpyja (Linnaeus, 1758) 17 v (f), a 

Spizaetus melanoleucus (Vieillot, 1816) 7 v 

Spizaetus ornatus (Daudin, 1800) 7, 12, 13 v (f) 

Falconiformes 

Falconidae 

Ibycter americanus (Boddaert, 1783) 3, 4 e, v MZUSP 

Caracara plancus (Miller, 1777) 1, 2, 3, 7, 9, 10, 11, 13, 15, 18, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH 

Milvago chimachima (Vieillot, 1816) 1, 2, 3, 4, 9, 10, 11, 13, 18, 19, 20, 21, 22, 23 e, v (f), a MZUSP, MCZ 

Herpetotheres cachinnans (Linnaeus, 1758) 1, 2, 3, 7, 11, 19, 21, 22, 23, 26 a 

Micrastur ruficollis (Vieillot, 1817) 1, 7 a 

Micrastur semitorquatus (Vieillot, 1817) 7, 15 v, a 

Falco sparverius Linnaeus, 1758 1, 3, 4, 7, 9, 12, 13, 15, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Falco rufigularis Daudin, 1800 7, 11, 15, 19, 21 v 

Falco deiroleucus Temminck 1825 3 

Falco femoralis Temminck, 1822 1, 2, 3, 4, 7, 9, 10, 21, 22, 25 e, v (f) FMNH, MZUSP 

Eurypygiformes 

Eurypygidae 

Eurypyga helias (Pallas, 1781) 3 

Gruiformes 

Aramidae 

Aramus guarauna (Linnaeus, 1766) 16, 18, 25 e, v, a FMNH, MZUSP 

Rallidae 

Aramides cajanea (Statius Muller, 1776) 1, 2, 3, 4, 10, 11, 16, 20, 23, 26 e, v, a MZUSP 

Laterallus viridis (Statius Muller, 1776) 9 v 

Porzana albicollis (Vieillot, 1819) 1, 2, 10, 15, 16, 18, 21, 23, 26 a 

Gallinula galeata (Lichtenstein,1818) 3 

Porphyrio martinica (Linnaeus, 1766) 15, 23, 26 v 

Porphyrio flavirostris (Gmelin, 1789) 16, 26 a 

Heliornithidae 

Heliornis fulica (Boddaert, 1783) 3 

Cariamiformes 

Cariamidae 

Cariama cristata (Linnaeus, 1766) 1, 2, 3, 7, 9, 10, 11, 12, 13, 15, 16, 19, 20, 21, 22, 23, 25, 26 e, v (f), a (g) FMNH 





83 


Charadriiformes 

Charadriidae 

Vanellus chilensis (Molina, 1782) 1, 2, 7, 9, 10, 11, 13, 15, 14, 18, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Recurvirostridae 

Himantopus melanurus Vieillot, 1817 2, 20, 25 e, v FMNH 

Scolopacidae 

Gallinago paraguaiae (Vieillot, 1816) 3 

Bartramia longicauda (Bechstein, 1812) 3, 25 v 

Tringa solitaria Wilson, 1813 3, 7, 20, 22, 23, 25 e, v (f) FMNH 

Tringa melanoleuca (Gmelin, 1789) 20 v 

Tringa flavipes (Gmelin, 1789) 2, 3, 25 e FMNH 

Phalaropus tricolor (Vieillot, 1819) 25 e FMNH 

Jacanidae 

Jacana jacana (Linnaeus, 1766) 2, 3, 11, 15, 16, 18, 20, 23, 25, 26 e, v, a FMNH, MZUSP 

Sternidae 

Phaetusa simplex (Gmelin, 1789) 1, 2 v 

Rynchopidae 

Rynchops niger Linnaeus, 1758 25 v 

Columbiformes 

Columbidae 

Columbina talpacoti (Temminck, 1811) 1, 2, 3, 4, 7, 9, 10, 11, 13, 15, 16, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH, MZUSP 

Columbina squammata (Lesson, 1831) 1, 2, 3, 7, 9, 10, 13, 16, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH 

Columbina picui (Temminck, 1813) 1, 15, 16, 18, 19, 20, 21, 23 v, a 

Claravis pretiosa (Ferrari-Perez, 1886) 2, 4, 7, 12, 19, 21 e, v MZUSP 

Columba livia Gmelin, 1789] 1, 15, 16 v 

Patagioenas speciosa (Gmelin, 1789) 3 

Patagioenas picazuro (Temminck, 1813) 1, 2, 3, 7, 9, 10, 11, 13, 15, 16, 18, 20, 21, 22, 23, 26 v, a 

Patagioenas cayennensis (Bonnaterre, 1792) 1, 2, 3, 7, 9, 10, 11, 12, 13, 16, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH 

Patagioenas plumbea (Vieillot, 1818) 2 v 

Zenaida auriculata (Des Murs, 1847) 1, 2, 3, 4, 7, 10, 15, 18, 21, 22, 23, 26 e, v, a MZUSP 

Leptotila verreauxi Bonaparte, 1855 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 15, 16, 18, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Leptotila rufaxilla (Richard & Bernard, 1792) 1, 2, 3, 10, 11, 12, 25, 26 e, v (f), a FMNH 


84 





Psittaciformes 

Psittacidae 

Anodorhynchus hyacinthinus (Latham, 1790) 3, 7, 9, 12, 13, 16 v (f), a 

Ara ararauna (Linnaeus, 1758) 1, 2, 3, 7, 9, 11, 13, 16, 19, 20, 21, 22, 23 v (f), a 

Ara chloropterus Gray, 1859 1, 2, 3, 7, 11, 12, 13, 16, 19, 20, 21, 23, 26 v (f), a 

Orthopsittaca manilata (Boddaert, 1783) 1, 3, 10, 21, 22, 23 v, a 

Primolius maracana (Vieillot, 1816) 1, 18 v (f), a 

Primolius auricollis (Cassin, 1853) 9, 15, 16 v (f), a 

Diopsittaca nobilis (Linnaeus, 1758) 1, 2, 3, 4, 7, 9, 13, 25 e, v (f), a FMNH, MZUSP 

Aratinga acuticaudata (Vieillot, 1818) 15, 16, 25 e, v FMNH 

Aratinga leucophthalma (Statius Muller, 1776) 1, 3, 7, 9, 10, 12, 15, 16, 19, 20, 21, 22, 23 v (f), a 

Aratinga nenday (Vieillot, 1823) 15, 16 v, a 

Aratinga aurea (Gmelin, 1788) 1, 2, 3, 7, 9, 10, 11, 12, 13, 19, 21, 21, 22, 23, 25, 26 e, v (f), a FMNH 

Pyrrhura devillei (Massena & Souancé, 1854) 7, 11, 12, 13, 15, 16, 19, 20 v (f), a 

Myiopsitta monachus (Boddaert, 1783) 15, 16, 18 v, a 

Forpus xanthopterygius (Spix, 1824) 3 

Brotogeris chiriri (Vieillot, 1818) 1, 3, 4, 7, 9, 10, 11, 13, 15, 16, 18, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH, MZUSP, MCZ 

Alipiopsitta xanthops (Spix, 1824) 1, 2, 3, 9, 11 v (f), a 

Pionus menstruus (Linnaeus, 1766) 3 

Pionus maximiliani (Kuhl, 1820) 3, 11, 15, 16, 19, 20 v, a 

Amazona amazonica (Linnaeus, 1766) 1, 3, 13, 19, 20, 21, 22, 23, 25 e, v (f), a FMNH 

Amazona aestiva (Linnaeus, 1758) 1, 3, 4, 7, 9, 10, 11, 15, 16, 19, 20, 23, 26 e, v, a MZUSP 

Cuculiformes 

Cuculidae 

Micrococcyx cinereus (Vieillot, 1817) 25 e, v FMNH 

Piaya cayana (Linnaeus, 1766) 1, 2, 3, 7, 9, 10, 11, 12, 13, 16, 18, 20, 21, 23, 25, 26 e, v (f) FMNH, MZUSP, FMO 

Coccyzus americanus (Linnaeus, 1758) 2, 12, 21 v (f) 

Crotophaga major Gmelin, 1788 1, 2, 9, 10, 11, 16, 19, 20, 23 v (f), a 

Crotophaga ani Linnaeus, 1758 1, 2, 3, 7, 9, 10, 11, 13, 14, 15, 16, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH 

Guira guira (Gmelin, 1788) 1, 2, 3, 7, 9, 10, 11, 13, 15, 16, 20, 21, 22, 23, 25, 26 e, v, a FMNH 

Tapera naevia (Linnaeus, 1766) 1, 2, 3, 11, 18, 20, 21, 22, 23 e, v (f), a MZUSP 

Dromococcyx phasianellus (Spix, 1824) 12, 13, 15, 20, 23 v (f) 

Dromococcyx pavoninus Pelzeln, 1870 1 v 





85 


Strigiformes 

Tytonidae 

Tyto alba (Scopoli, 1769) 1, 3, 7, 15, 16, 23 a 

Strigidae 

Megascops choliba (Vieillot, 1817) 3, 7, 10, 12, 13, 16, 19, 21, 22, 23 v, a 

Pulsatrix perspicillata (Latham, 1790) 12, 13 v 

Bubo virginianus (Gmelin, 1788) 12, 13, 16 v, a 

Glaucidium brasilianum (Gmelin, 1788) 1, 3, 4, 7, 9, 10, 11, 12, 13, 15, 16, 19, 20, 21, 23, 25, 26 e, v, a (g) FMNH, MZUSP 

Athene cunicularia (Molina, 1782) 1, 2, 3, 7, 9, 10, 13, 16, 20, 21, 22, 23, 25, 26 e, v, a FMNH 

Asio clamator (Vieillot, 1808) 1, 2 a 

Caprimulgiformes 

Nyctibiidae 

Nyctibius grandis (Gmelin, 1789) 10 v 

Nyctibius griseus (Gmelin, 1789) 3, 9, 13, 21, 22 v, a 

Caprimulgidae 

Lurocalis semitorquatus (Gmelin, 1789) 3, 7, 13 v 

Podager nacunda (Vieillot, 1817) 1, 2, 3, 10, 12, 15, 16, 20 v, a 

Nyctidromus albicollis (Gmelin, 1789) 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 16, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Nyctiphrynus ocellatus (Tschudi, 1844) 1 v 

Caprimulgus rufus Boddaert, 1783 12, 22 v 

Caprimulgus parvulus (Gould, 1837) 1, 3, 20, 21, 22, 23, 25, 26 e, v FMNH 

Hydropsalis torquata (Gmelin, 1789) 11 v 

Apodiformes 

Apodidae 

Cypseloides senex (Temminck, 1826) 1 v (f) 

Streptoprocne zonaris (Shaw, 1796) 1, 3, 7, 12, 22 v (f) 

Chaetura meridionalis Hellmayr, 1907 1, 3, 12, 13, 15, 19, 21, 22 v 

Tachornis squamata (Cassin, 1853) 3, 4, 10, 21, 22, 23 e, v MZUSP 

Trochilidae 

Phaethornis pretrei (Lesson & Detattre, 1839) 1, 3, 7, 11, 21, 22, 23, 26 v (f) 

Phaethornis eurynome (Lesson, 1832) 9, 10, 15 v 

Eupetomena macroura (Gmelin, 1788) 1, 2, 3, 4, 7, 9, 11, 13, 16, 19, 22, 23, 26 e, v FMNH, MZUSP 

Florisuga fusca (Vieillot, 1817) 7 v 


86 





Colibri serrirostris (Vieillot, 1816) 1, 3, 4, 7, 8 e, v MZUSP 

Anthracothorax nigricollis (Vieillot, 1817) 1, 2, 3, 12, 15, 18, 21, 23 e, v MZUSP 

Lophornis magnificus (Vieillot, 1817) 4 e MZUSP 

Chlorostilbon lucidus (Shaw, 1812) 1, 2, 3, 7, 9, 10, 11, 12, 13, 15, 16, 19, 21, 22, 23, 25, 26 e, v (f), a FMNH 

Thalurania furcata (Gmelin, 1788) 1, 2, 3, 8, 13, 14, 25, 26 v (f) 

Hylocharis cyanus (Vieillot, 1818) 3 

Hylocharis chrysura (Shaw, 1812) 1, 2, 4, 7, 10, 11, 12, 13, 15, 16, 18, 19, 20, 21, 23 e, v MZUSP, MCZ 

Polytmus guainumbi (Pallas, 1764) 1, 4, 13, 16, 19, 23, 25, 26 e, v FMNH, MZUSP 

Amazilia versicolor (Vieillot, 1818) 4, 7, 9, 10, 12, 19, 21 e, v MZUSP 

Amazilia frimbiata (Gmelin, 1788) 1, 2, 3, 21, 239 v 

Heliactin bilophus (Temminck, 1820) 4 e MZUSP 

Heliomaster furcifer (Shaw, 1812) 1, 4, 9, 19, 21 e, v (f) MZUSP 

Calliphlox amethystina (Boddaert, 1783) 4 e MZUSP 

Trogoniformes 

Trogonidae 

Trogon surrucura Vieillot, 1817 1, 25 v 

Trogon curucui Linnaeus, 1766 2, 3, 4, 7, 9, 10, 12, 13, 15, 16, 19, 10, 21, 22, 23, 26 e, v (f), a MZUSP 

Coraciiformes 

Alcedinidae 

Megaceryle torquata (Linnaeus, 1766) 1, 2, 3, 9, 11, 16, 20, 26 

Chloroceryle amazona (Latham, 1790) 1, 2, 3, 7, 11, 16, 20, 21, 22, 25 e, v (f) FMNH 

Chloroceryle aenea (Pallas, 1764) 1 v 

Chloroceryle americana (Gmelin, 1788) 1, 2, 3, 7, 10, 11, 16, 20, 22 v (f) 

Momotidae 

Momotus momota (Linnaeus, 1766) 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 16, 19, 20, 22, 26 e, v, a MZUSP 

Galbuliformes 

Galbulidae 

Brachygalba lugubris (Swainson, 1838) 4 e MZUSP 

Galbula ruficauda Cuvier, 1816 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 15, 16, 19, 20, 21, 22, 23 e, v (f), a MZUSP 

Bucconidae 

Nystalus chacuru (Vieillot, 1816) 1, 3, 4, 7, 10, 11 e, v (f), a (g) MZUSP 

Nystalus striatipectus (Sclater, 1854) 1, 2, 7, 15, 18, 19, 20, 21, 22, 239 e, v (f) MZUSP, MCZ 

Nonulla rubeculla (Spix, 1824) 3, 12 v (f) 





87 


Monasa nigrifrons (Spix, 1824) 1, 2, 3 v (f), a 

Chelidoptera tenebrosa (Pallas, 1782) 3, 4 e MZUSP 

Piciformes 

Ramphastidae 

Ramphastos toco Statius Muller, 1776 1, 2, 3, 7, 9, 10, 11, 12, 13, 15, 16, 18, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Ramphastos vitellinus Lichtenstein, 1823 3 

Pteroglossus castanotis Gould, 1834 1, 3, 7, 12, 13, 16, 20, 23, 25, 26 v (f), a 

Picidae 

Picumnus cirratus Temminck, 1825 26 v 

Picumnus albosquamatus d’Orbigny, 1840 1, 2, 3, 4, 7, 9, 10, 11, 15, 16, 20, 23 e, v MZUSP 

Melanerpes candidus (Otto, 1796) 1, 3, 4, 7, 10, 13, 15, 16, 18, 19, 20, 21, 22, 23, 24, 26 e, v, a FMNH, MZUSP, MCZ 

Veniliornis passerinus (Linnaeus, 1766) 1, 2, 3, 4, 7, 9, 10, 11, 15, 16, 18, 19, 20, 21, 23 e, v (f), a MZUSP 

Piculus chrysochloros (Vieillot, 1818) 19 v 

Colaptes melanochloros (Gmelin, 1788) 1, 3, 4, 7, 9, 11, 12, 13, 16, 20, 21, 22, 23, 25 e, v (f) FMNH, MZUSP 

Colaptes campestris (Vieillot, 1818) 1, 3, 4, 7, 9, 10, 11, 13, 16, 19, 20, 21, 23, 26 e, v, a FMNH, MZUSP 

Celeus lugubris (Malherbe, 1851) 3, 12, 13, 15, 16, 19, 20, 21, 22, 23, 25 v 

Celeus flavus (Statius Muller, 1776) 3 

Dryocopus lineatus (Linnaeus, 1766) 1, 3, 4, 7, 9, 10, 13, 15, 19, 20, 21, 22, 23 e, v (f), a FMNH, MZUSP 

Campephilus melanoleucos (Gmelin, 1788) 1, 3, 7, 15, 16, 18, 20, 21, 22, 23 e, v (f) FMO 

Passeriformes 

Thamnophilidae 

Formicivora melanogaster Pelzeln, 1868 3 

Formicivora rufa (Wied, 1831) 1, 4, 9, 10, 11, 15, 16, 18, 19, 20, 21, 22, 23 e, v (f), a MZUSP 

Herpsilochmus atricapillus Pelzeln, 1868 3 

Herpsilochmus longirostris Pelzeln, 1868 1, 2, 3, 4, 7, 9, 12, 22, 23, 26 e, v (f), a MZUSP 

Thamnophilus doliatus (Linnaeus, 1764) 1, 2, 3, 4, 7, 9, 10, 11, 13, 15, 16, 18, 19, 20, 21, 22, 23, 25 e, v (f), a FMNH, MZUSP, FMO 

Thamnophilus torquatus Swainson, 1825 3 

Thamnophilus sticturus Pelzeln, 1868 18 e FMO 

Thamnophilus pelzelni Hellmayr, 1924 1, 2, 3, 4, 7, 10, 12, 13, 15, 19, 21, 22, 23 e, v, a MZUSP 

Thamnophilus caerulescens Vieillot, 1816 3, 7, 19, 20, 21 v 

Taraba major (Vieillot, 1816) 2, 3, 10, 11, 15, 16, 18, 19, 20, 21, 23, 26 e, v (f), a MZUSP 

Melanopareiidae 

Melanopareia torquata (Wied, 1831) 3, 4 e MZUSP 


88 





Dendrocolaptidae 

Sittasomus griseicapillus (Vieillot, 1818) 1, 2, 3, 4, 10, 11, 12, 19, 20, 21, 22, 23 e, v MZUSP 

Xiphorhynchus guttatus (Lichtenstein, 1820) 2, 10, 11 v, a 

Campylorhamphus trochilirostris (Lichtenstein, 1820) 1, 4, 7, 9, 11, 15, 16, 19, 20, 23 e, v, a MZUSP 

Dendroplex picus (Gmelin, 1788) 1, 2 v (f), a (g) 

Lepidocolaptes angustirostris (Vieillot, 1818) 1, 2, 3, 7, 9, 10, 13, 16, 18, 19, 20, 21, 22, 23, 26 e, v (f), a MZUSP, MCZ 

Dendrocolaptes picumnus Lichtenstein, 1820 9, 10, 15, 18 e, v FMO 

Dendrocolaptes platyrostris Spix, 1825 3, 4, 12, 13, 19, 20, 21, 22, 23, 26 e, v MZUSP 

Xiphocolaptes major (Vieillot, 1818) 2, 15, 16, 20, 21, 22 v, a 

Furnariidae 

Xenops rutilans Temminck 1821 3, 19 v 

Berlepschia rikeri (Ridgway, 1886) 3 

Furnarius leucopus Swainson, 1838 16, 18 e, v MZUSP 

Furnarius rufus (Gmelin, 1788) 1, 2, 3, 4, 7, 9, 10, 11, 13, 15, 16, 18, 19, 10, 21, 23, 25, 26 e, v (f), a FMNH, MZUSP, MCZ 

Lochmias nematura (Lichtenstein, 1823) 3, 7 v 

Hylocryptus rectirostris (Wied, 1831) 3, 4, 7, 9, 19, 21, 22, 23 e, v (f) MZUSP 

Philydor rufum (Vieillot, 1818) 4, 12, 13 v (f) 

Syndactyla dimidiata (Pelzeln, 1859) 4 e MZUSP 

Pseudoseisura unirufa (d’Orbigny & Lafresnaye, 1838) 16 v, a 

Phacellodomus rufifrons (Wied, 1821) 3, 16, 18, 19, 20, 21, 22, 23 e, v, a MZUSP, MCZ 

Phacellodomus ruber (Vieillot, 1817) 3, 9, 10, 18, 19, 20, 21, 22, 23 v, a (g) 

Schoeniophylax phryganophilus (Vieillot, 1817) 1, 20 v 

Certhiaxis cinnamomeus (Gmelin, 1788) 1, 2, 3, 9, 11, 16, 20, 22, 23, 26 v, a 

Synallaxis frontalis Pelzeln, 1859 1, 2, 3, 4, 7, 9, 10, 19, 21, 22, 23, 25 e, v (f), a FMNH, MZUSP 

Synallaxis albecens Temminck, 1823 1, 19, 22, 26 v, a (g) 

Synallaxis hypospodia Sclater, 1874 20, 21, 22, 23 v, a 

Synallaxis albilora Pelzeln, 1856 15, 16 v 

Synallaxis scutata Sclater, 1859 3 

Pipridae 

Neopelma pallescens (Lafresnaye, 1853) 1, 3, 11, 25 v, a 

Pipra fasciicauda Hellmayr, 1906 3, 12, 22, 26 v (f), a 

Antilophia galeata (Lichtenstein, 1823) 1, 2, 3, 4, 8, 11, 13, 14, 16, 17, 25, 26 e, v (f), a FMNH, MZUSP 





89 


Tityridae 

Oxyruncus cristatus Swainson, 1821 3 

Schiffornis virescens (Lafresnaye, 1838) 10 v 

Tityra inquisitor (Lichtenstein, 1823) 3, 11, 13, 16, 19, 20 v, a 

Tityra cayana (Linnaeus, 1766) 3, 10, 11, 19, 22, 26 v, a 

Tityra semifasciata (Spix, 1825) 1, 3, 7, 26 v (f) 

Pachyramphus viridis (Vieillot, 1816) 2, 10, 11, 18, 20, 21, 23 e, v (f), a MZUSP 

Pachyramphus polychopterus (Vieillot, 1818) 1, 3, 7, 26 v, a 

Pachyramphus validus (Lichtenstein, 1823) 3, 11, 12, 13, 19, 22 v 

Xenopsaris albinucha (Burmeister, 1869) 3 

Rhynchocyclidae 

Leptopogon amaurocephalus Tschudi, 1846 2, 3, 7, 11, 12, 13, 26 v, a 

Corythopis delalandi (Lesson, 1830) 12 v 

Hemitriccus striaticollis (Lafresnaye, 1853) 1, 7, 18 e, v (f) FMO 

Hemitriccus margaritaceiventer (d’Orbigny & Lafresnaye, 1837) 1, 2, 3, 4, 7, 10, 13, 15, 16, 18, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH, MZUSP 

Poecilotriccus latirostris (Pelzeln, 1868) 1, 9, 7, 10, 15, 16, 18 e, a MZUSP 

Tolmomyias sulphurescens (Spix, 1825) 1, 3, 9, 7, 10, 12, 13, 16, 19, 20, 21, 22, 23, 26 a 

Platyrinchus mystaceus Vieillot, 1818 20 v 

Todirostrum cinereum (Linnaeus, 1766) 1, 2, 3, 4, 15, 16, 20, 25, 26 e, v, a FMNH, MZUSP 

Tyrannidae 

Hirundinea ferruginea (Gmelin 1788) 3, 7 v 

Inezia inornata (Salvadori, 1897) 21 v 

Euscarthmus meloryphus Wied, 1831 18, 21 e, v MZUSP 

Camptostoma obsoletum (Temminck, 1824) 1, 2, 3, 7, 9, 10, 11, 13, 16, 20, 21, 22, 23, 26 v (f), a 

Elaenia flavogaster (Thunberg, 1822) 1, 2, 3, 4, 7, 9, 10, 13, 15, 16, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH, MZUSP 

Elaenia spectabilis Pelzeln, 1868 10, 20, 21, 23, 25, 26 e, v, a FMNH 

Elaenia chilensis Hellmayr, 1927 2, 25 e FMNH 

Elaenia chiriquensis Lawrence, 1865 1, 3, 7, 9, 11, 13 v, a (g) 

Suiriri suiriri (Vieillot, 1818) 1, 3, 4, 7, 19, 21, 23 e, v, a (g) MZUSP 

Myiopagis gaimardii (d’Orbigny, 1839) 3 

Myiopagis caniceps (Swainson, 1835) 1, 3, 19, 21, 26 v, a 

Myiopagis viridicata (Vieillot, 1817) 3, 9, 10, 12, 16, 19, 20, 21, 22 v, a 


90 





Capsiempis flaveola (Lichtenstein, 1823) 19, 20, 21 v 

Phaeomyias murina (Spix, 1825) 1, 2, 7, 21, 25 e, v (f), a FMNH 

Phyllomyias fasciatus (Thunberg, 1822) 1, 10 v (f) 

Serpophaga subcristata (Vieillot, 1817) 10, 11, 15, 16, 25, 26 e, v, a FMNH 

Legatus leucophaius (Vieillot, 1818) 1, 2, 3, 13, 26 v (f), a (g) 

Myiarchus tuberculifer (d’Orbigny & Lafresnaye, 1837) 1 v (f) 

Myiarchus swainsoni Cabanis & Heine, 1859 3, 18, 20, 21, 22, 23 e, v, a FMNH 

Myiarchus ferox (Gmelin, 1789) 1, 2, 3, 4, 9, 10, 11, 12, 13, 18, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Myiarchus tyrannulus (Statius Muller, 1776) 1, 2, 3, 4, 7, 9, 10, 13, 14, 16, 19, 20, 21, 25 e, v (f), a FMNH, MZUSP 

Sirystes sibilator (Vieillot, 1818) 3, 12, 19, 21, 23 a 

Casiornis rufus (Vieillot, 1816) 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 15, 16, 19, 20, 21, 22, 23, 26 e, v, a FMNH, MZUSP 

Pitangus sulphuratus (Linnaeus, 1766) 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 15, 14, 18, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Philohydor lictor (Lichtenstein, 1823) 1, 11, 16 v (f), a 

Machetornis rixosa (Vieillot, 1819) 1, 2, 3, 7, 9, 10, 11, 13, 15, 16, 18, 19, 20, 21, 22, 23, 24, 25 e, v FMNH, MZUSP 

Myiodynastes maculatus (Statius Muller, 1776) 1, 2, 3, 9, 10, 12, 13, 16, 19, 18, 21, 22, 23 v (f), a 

Tyrannopsis sulphurea (Spix, 1825) 1 v (f) 

Megarynchus pitangua (Linnaeus, 1766) 1, 2, 4, 7, 9, 10, 11, 12, 13, 15, 16, 19, 20, 21, 22, 23, 26 e, v (f), a MZUSP 

Myiozetetes cayanensis (Linnaeus, 1766) 1, 2, 3, 9, 11, 12, 14, 19, 20, 22, 23, 25 a 

Myiozetetes similis (Spix, 1825) 1, 2, 26 v, a 

Tyrannus albogularis Burmeister, 1856 1, 2, 7, 20, 21, 22, 23 e, v (f) FMNH 

Tyrannus melancholicus Vieillot, 1819 1, 2, 3, 9, 10, 11, 12, 13, 15, 16, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH 

Tyrannus savana Vieillot, 1808 1, 2, 3, 9, 10, 11, 19, 20, 21, 22, 23, 25, 26 e, v FMNH 

Griseotyrannus aurantioatrocristatus (d’Orbigny & Lafresnaye, 1837) 1, 2, 9, 13, 15, 19, 21, 25 e, v (f) FMNH 

Empidonomus varius (Vieillot, 1818) 1, 2, 3, 7, 12, 13, 19, 20, 21, 26 v (f), a 

Colonia colonus (Vieillot, 1818) 3, 4, 11, 21, 23 e, v MZUSP 

Myiophobus fasciatus (Statius Muller, 1776) 1, 2, 4, 19, 23, 26 e, v, g (a) MZUSP 

Sublegatus modestus (Wied, 1831) 10, 18 e, v MZUSP 

Pyrocephalus rubinus (Boddaert, 1783) 1, 2, 3, 4, 9, 11, 18, 19, 20, 21, 22, 23, 25, 26 e, v (f) FMNH, MZUSP 

Fluvicola albiventer (Spix, 1825) 2, 9, 15, 16, 18, 20, 22, 23, 25, 26 e, v (f), a FMNH 

Arundinicola leucocephala (Linnaeus, 1764) 1, 9, 18, 20, 21, 22, 23, 25, 26 e, v FMNH 

Alectrurus tricolor (Vieillot, 1816) 25 e FMNH 

Cnemotriccus fuscatus (Wied, 1831) 1, 7, 9, 10, 11, 13, 15, 16, 19, 20, 21, 22, 23 v, a 





91 


Lathrotriccus euleri (Cabanis, 1868) 3, 7, 9, 10, 16, 21 v, a 

Contopus cinereus (Spix, 1825) 4, 19, 21 e, v (f), a MZUSP 

Knipolegus lophotes Boie, 1828 4 e MZUSP 

Satrapa icterophrys (Vieillot, 1818) 12, 18, 23 v 

Xolmis cinereus (Vieillot, 1816) 1, 2, 3, 7, 10, 13, 18, 19, 20, 21, 22, 23 e, v (f), a MZUSP 

Xolmis velatus (Lichtenstein, 1823) 2, 3, 7, 9, 10, 13, 15, 16, 19, 20, 21, 22, 23, 26 e, v (f) FMNH 

Vireonidae 

Cyclarhis gujanensis (Gmelin, 1789) 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 15, 16, 18, 19, 20, 21, 22, 23, 25, 26 e, v, a (g) FMNH, MZUSP 

Vireo olivaceus (Linnaeus, 1766) 1, 2, 3, 10, 11, 12, 15, 16, 18, 19 e, v (f), a (g) FMO 

Corvidae 

Cyanocorax cyanomelas (Vieillot, 1818) 2, 9, 10, 11, 15, 16, 18, 21, 22, 23 e, v, a MZUSP 

Cyanocorax cristatellus (Temminck, 1823) 1, 3, 4, 7, 9, 21, 22, 25, 26 e, v, a FMNH, MZUSP 

Cyanocorax chrysops (Vieillot, 1818) 1, 2, 4, 7, 9, 10, 11, 12, 13, 15, 16, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Hirundinidae 

Pygochelidon cyanoleuca (Vieillot, 1817) 1 v 

Alopochelidon fucata (Temminck, 1822) 1, 3 v 

Stelgidopteryx ruficollis (Vieillot, 1817) 1, 2, 3, 4, 9, 13, 18, 20, 21, 22, 23 e, v (f) MZUSP 

Progne tapera (Vieillot, 1817) 1, 2, 7, 9, 10, 11, 14, 15, 18, 19, 20, 21, 22, 23, 26 e, v (f), a FMO 

Progne chalybea (Gmelin, 1789) 1, 7, 15, 19, 20, 23, 25, 26 e, v FMNH 

Tachycineta albiventer (Boddaert, 1783) 1, 16, 20, 21, 22 v 

Tachycineta leucorrhoa (Vieillot, 1817) 3, 22, 23 v 

Hirundo rustica Linnaeus, 1758 2, 20 v 

Troglodytidae 

Troglodytes musculus Naumann, 1823 2, 3, 4, 7, 9, 10, 11, 15, 16, 26 e, v, a MZUSP 

Campylorhynchus turdinus (Wied, 1831) 9, 7, 10, 11, 13, 15, 16, 18, 19, 20, 21, 22, 23 e, v, a MZUSP 

Cantorchilus leucotis (Lafresnaye, 1845) 2, 3, 4, 15, 20, 21, 22, 23 e, v, a MZUSP 

Donacobiidae 

Donacobius atricapilla (Linnaeus, 1766) 2, 10, 16, 20, 21, 26 v (f), a 

Polioptilidae 

Polioptila dumicola (Vieillot, 1817) 1, 2, 3, 4, 9, 10, 11, 18, 19, 20, 21, 26 e, v (f), a MZUSP, MCZ 

Turdidae 

Catharus fuscescens 7 v (f) 

Turdus rufiventris Vieillot, 1818 1, 2, 3, 7, 9, 11, 12, 13, 15, 16, 18, 19, 20, 22, 23, 26 e, v, a MZUSP 


92 





Turdus leucomelas Vieillot, 1818 1, 2, 3, 4, 7, 9, 10, 12, 13, 15, 16, 20, 21, 22, 23, 25, 26 e, v, a FMNH, MZUSP 

Turdus amaurochalinus Cabanis, 1850 1, 2, 3, 4, 7, 15, 16, 19, 20, 21, 22, 23 e, v, a MZUSP 

Turdus subalaris (Seebohm, 1887) 3 

Turdus albicollis Vieillot, 1818 10 v 

Mimidae 

Mimus saturninus (Lichtenstein, 1823) 1, 3, 4, 7, 9, 10, 11, 13, 15, 16, 18, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP, FMO 

Mimus triurus (Vieillot, 1818) 25 e FMNH 

Motacilidae 

Anthus lutescens Pucheran, 1855 2, 3, 7, 23, 25, 26 e, v, a FMNH 

Coerebidae 

Coereba flaveola (Linnaeus, 1758) 1, 2, 3, 4, 7, 18 e, v, a MZUSP 

Thraupidae 

Saltator maximus (Statius Muller, 1776) 3 

Saltator coerulescens Vieillot, 1817 16, 20, 23 v, a 

Saltator similis d’Orbigny & Lafresnaye, 1837 1, 2, 3, 4, 7, 9, 12, 13, 15, 19, 20, 21 v, a 

Saltatricula atricollis (Vieillot, 1817) 1, 3, 4, 7, 9, 11, 15, 19, 20, 21, 22, 25 e, v (f), a (g) FMNH, MZUSP 

Schistochlamys melanopis (Latham, 1790) 3, 4 e, v MZUSP 

Schistochlamys ruficapillus (Vieillot, 1817) 1, 21, 23 v (f), a 

Nemosia pileata (Boddaert, 1783) 1, 3, 4, 9, 10, 11, 15, 19, 20, 22 e, v, a MZUSP 

Thlypopsis sordida (d’Orbigny & Lafresnaye, 1837) 4, 21 e, v MZUSP 

Cypsnagra hirundinacea (Lesson, 1831) 15, 26 v (f), a (g) 

Trichothraupis melanops (Vieillot, 1818) 3, 4, 7 e, v MZUSP 

Eucometis penicillata (Spix, 1825) 1, 2, 3, 7, 9, 11, 12, 16, 19, 20, 22, 23, 26 v (f), a 

Tachyphonus rufus (Boddaert, 1783) 1, 2, 3, 4, 11, 12, 13, 19, 20, 21, 22, 23, 25 e, v (f), a FMNH, MZUSP 

Ramphocelus carbo (Pallas, 1764) 1, 2, 3, 4, 7, 10, 11, 16, 18, 19, 20, 22, 23, 25 e, v, a MZUSP, FMO 

Thraupis sayaca (Linnaeus, 1766) 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 15, 16, 17, 19, 20, 21, 22, 23, 25, 26 e, v, a FMNH, MZUSP, FMO 

Thraupis palmarum (Wied, 1823) 1, 2, 3, 4, 9, 10, 11, 13, 15, 16, 19, 20, 22, 23, 25 e, v, a FMNH, MZUSP 

Tangara cayana (Linnaeus, 1766) 1, 2, 3, 4, 7, 9, 10, 12, 13, 20, 21, 22, 23, 26 e, v (f) MZUSP 

Tersina viridis (Illiger, 1811) 1, 3, 4, 7, 9, 12, 13, 21, 22, 23, 26 e, v MZUSP 

Dacnis cayana (Linnaeus, 1766) 1, 2, 3, 7, 9, 11, 12, 13, 19, 20, 21, 22, 23, 26 v (f) 

Cyanerpes cyaneus (Linnaeus, 1766) 1, 2, 3, 7, 23 v 

Hemithraupis guira (Linnaeus, 1766) 1, 2, 3, 4, 7, 9, 10, 11, 19, 20, 21, 23, 26 e, v, a MZUSP 

Conirostrum speciosum (Temminck, 1824) 1, 9, 10, 13, 15, 18, 19, 20 e, v (f) MZUSP 





93 


Emberizidae 

Zonotrichia capensis (Statius Muller, 1776) 3, 4, 10, 20, 21, 22 e, v, a MZUSP 

Ammodramus humeralis (Bosc, 1792) 1, 2, 3, 4, 7, 9, 10, 11, 13, 15, 16, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Porphyrospiza caerulescens (Wied, 1830) 3 

Poospiza cinerea Bonaparte, 1850 4 e MZUSP 

Sicalis citrina Pelzeln, 1870 3, 8 v 

Sicalis flaveola (Linnaeus, 1766) 1, 2, 3, 7, 10, 13, 15, 16, 18, 19, 20, 21, 22, 23 e, v (f), a MZUSP, MCZ 

Emberizoides herbicola (Vieillot, 1817) 1, 3, 4, 8, 9, 11, 20, 21, 22, 23, 26 e, v (f) MZUSP 

Volatinia jacarina (Linnaeus, 1766) 1, 2, 3, 7, 9, 10, 11, 13, 16, 20, 21, 22, 23, 26 v, a 

Sporophila plumbea (Wied, 1830) 1, 2, 3, 4, 20 e, v (f) MZUSP 

Sporophila collaris (Boddaert, 1783) 2, 11, 15, 16, 20, 22, 23, 26 v 

Sporophila lineola (Linnaeus, 1758) 1, 9, 10, 15 v, a 

Sporophila nigricollis (Vieillot, 1823) 1, 2, 3, 21 v (f) 

Sporophila caerulescens (Vieillot, 1823) 1, 2, 3, 19, 21, 23 v (f), a 

Sporophila leucoptera (Vieillot, 1817) 1, 15, 18, 20, 21, 23 e, v (f), a MZUSP, MCZ 

Sporophila bouvreuil (Statius Muller, 1776) 2, 3 v 

Sporophila angolensis (Linnaeus, 1766) 1, 2, 3, 9, 15, 20, 22, 23, 25 e, v (f), a FMNH 

Sporophila maximiliani (Cabanis, 1851) 15 v 

Arremon taciturnus (Hermann, 1783) 3 

Arremon flavirostris Swainson, 1838 4, 12, 13, 20, 21, 23 e, v MZUSP 

Charitospiza eucosma Oberholser, 1905 11 v 

Coryphospingus cucullatus (Statius Muller, 1776) 1, 2, 3, 4, 9, 10, 11, 13, 15, 16, 18, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Paroaria coronata (Miller, 1776)] 16 v 

Paroaria capitata (d’Orbigny & Lafresnaye, 1837) 15, 16 v 

Cardinalidae 

Piranga flava (Vieillot, 1822) 4, 18 e, v MZUSP 

Cyanoloxia brissonii (Lichtenstein, 1823) 3, 18, 19, 21, 23, 25 e, v (f) FMNH, MZUSP 

Parulidae 

Parula pitiayumi (Vieillot, 1817) 1, 7, 9, 10, 11, 13, 16, 18, 19, 20, 21, 23, 26 e, v (f), a MZUSP 

Geothlypis aequinoctialis (Gmelin, 1789) 1, 3, 16, 19, 20, 21, 23, 25, 26 e, v (f), a FMNH 

Basileuterus culicivorus (Deppe, 1830) 26 v 

Basileuterus hypoleucus Bonaparte, 1830 1, 2, 4, 9, 10, 11, 12, 13, 16, 18, 19, 20, 21, 22, 23, 26 e, v MZUSP 


94 





Basileuterus flaveolus (Baird, 1865) 1, 2, 3, 4, 7, 9, 10, 11, 12, 13, 15, 16, 18, 19, 20, 21, 22, 26 e, v (f), a MZUSP 

Basileuterus leucophrys Pelzeln, 1868 1, 3 v 

Icteridae 

Psarocolius decumanus (Pallas, 1769) 3, 7, 11, 12, 15, 16, 20 v, a 

Procacicus solitarius (Vieillot, 1816) 11, 15, 16, 26 v, a 

Cacicus chrysopterus (Vigors, 1825) 15, 16, 19, 20 v 

Cacicus haemorrhous (Linnaeus, 1766) 3, 19, 20, 21, 25, 26 e, v FMNH 

Cacicus cela (Linnaeus, 1758) 3 

Icterus pyrrhopterus (Vieillot, 1819) 2, 4, 9, 10, 11, 15, 16, 18, 19, 21, 22, 23, 25 e, v, a FMNH, MZUSP 

Icterus croconotus (Wagler, 1829) 3, 15, 16, 19, 20, 21, 22, 23, 25 v, a 

Gnorimopsar chopi (Vieillot, 1819) 1, 3, 4, 7, 10, 12, 13, 15, 16, 19, 20, 21, 22, 23, 25 e, v (f), a FMNH, MZUSP 

Agelasticus cyanopus (Vieillot, 1819) 16 v 

Pseudoleistes guirahuro (Vieillot, 1819) 1, 3, 23, 25, 26 e, v (f) FMNH 

Agelaioides badius (Vieillot, 1819)] 7, 15, 16, 23 v 

Molothrus rufoaxillaris Cassin, 1866 1, 10, 13, 15, 16, 23 v (f), a 

Molothrus oryzivorus (Gmelin, 1788) 20, 23 v, a 

Molothrus bonariensis (Gmelin, 1789) 1, 3, 7, 8, 10, 11, 13, 15, 16, 18, 26 e, v (f), a MZUSP 

Sturnella superciliaris (Bonaparte, 1850) 1, 3, 7, 16, 23, 26 v 

Fringillidae 

Sporagra magellanica (Vieillot, 1805) 4, 16, 18 e, v MZUSP 

Euphonia chlorotica (Linnaeus, 1766) 1, 2, 3, 4, 7, 9, 10, 11, 12, 15, 16, 19, 20, 21, 22, 23, 25, 26 e, v (f), a FMNH, MZUSP 

Passeridae 

Passer domesticus (Linnaeus, 1758) 15, 16 v 





95 

Registros duvidosos 

D’Horta et al. (2008) admitem a existência de 

quatro espécies antes agregadas em Icterus cayanensis: I. 

pyrrhopterus, I. valenciobuenoi, I. periporphyrus e I. tibialis, 

sendo as três primeiras ocorrentes no Mato Grosso do Sul 

(Omland et al. 1999, Sturge et al. 2009). No entanto, com 

base no padrão de coloração da dragona dos indivíduos 

observados em campo, optamos por converter todos os 

registros de I. cayanensis em favor de I. pyrrhopterus, táxon 

aceito como espécie plena (CBRO 2011). 

Alguns registros mencionados por Tubelis & Tomas 

(2003) para Aquidauana, bem como aqueles citados por 

Hass (2004) para a vertente do Rio Taquari, no entorno 

do Parque Nacional de Emas, e Nunes et al. (2008) para 

a Fazenda Taboco, foram reavaliados e necessitam de 

revisão e comprobação documentada, por se tratarem de 

equívocos de identificação e incoerências biogeográficas. 

Nesse contexto, enquadram-se os seguintes táxons: 

Phaethornis ruber: menção em lista de espécies 

(Hass 2004), porém, não há registro comprobatório e a 

distribuição é incompatível. 

Thalurania glaucopis: menção em lista de espécies 

(Tubelis & Tomas 2003); sem registro comprobatório. O 

registro não é passível de reavaliação. 

Melanerpes flavifrons: menção em lista de espécies 



Celeus flavescens: menção em lista de espécies 



Formicivora grisea: menção em lista de espécies (Hass 

2004), sem registro comprobatório. As espécies do gênero 

são muito similares, fato que pode ter gerado equívoco de 

identificação. O referido táxon possui distribuição ampla 

distribuição na América do Sul (norte), principalmente na 

Amazônia (Sick 1997). Desta forma, o registro atribuído 

a esse táxon foi convertido em favor de Formicivora 

melanogaster, que conforme a literatura consultada (Sick 

1997), possui distribuição compatível para ocorrência na 

Serra de Maracaju. 

Myiornis auricularis: menção em lista de espécies 

(Nunes et al. 2008), porém, não há circunstanciação e a 


Euphonia violacea: menção em lista de espécies (Hass 

2004), porém, não há circunstanciação e a distribuição é 

incompatível. 

Aspectos biogeográficos 

Do ponto de vista biogeográfico, a Serra de Maracaju 

atua como um importante corredor para dispersão de 

espécies típicas do Cerrado. Neste sentido enquadram-se 

espécies tidas como endêmicas do bioma Cerrado (Silva 

1995, Silva & Santos 2005), tais como Nothura minor, 

Alipiopsitta xanthops, Melanopareia torquata, Antilophia 

galeata, Cyanocorax cristatellus e Saltatricula atricollis. 

A ocorrência de táxons como Aburria nattereri, 

Pseudastur albicollis, Hylocharis cyanus, Celeus flavus, 

Dendroplex picus, Myiarchus tuberculifer, Tyrannopsis 

sulphurea, Tityra semifasciata e Cyanerpes cyaneus é 

notável, uma vez que tais aves possuem a bacia do Rio 

Amazonas como centro de distribuição (Silva 1996). 

Provavelmente as matas de galeria ao longo dos principais 

rios que cortam a Serra de Maracaju e a borda leste da 

planície pantaneira atuam como importantes corredores 

para a dispersão desses elementos amazônicos na região, 

tal como proposto por alguns autores (Brown 1986, 

Nunes & Tomas 2004). 

Segundo Straube et al. (1996) e Pivatto et al. 

(2006), grande parte das espécies endêmicas da Mata 

Atlântica cessam suas distribuições para oeste nas florestas 

estacionais semidecíduas do sul do Mato Grosso do Sul, 

oeste do Paraná, Paraguai oriental e extremo nordeste 

da Argentina. Elementos típicos desse bioma, tais como 

alguns membros da família Thraupidae (como Tangara 

seledon, Pyrrhocoma ruficeps e Orthogonys chloricterus), 

estão ausentes na Serra de Maracaju. No entanto, 

algumas espécies atlânticas podem ser avistadas na região, 

como Florisuga fusca, Lophornis magnificus e Phyllomyias 

fasciatus. 

Espécies como Ortalis canicollis, Aratinga nenday, 

Celeus lugubris, Nystalus striatipectus, Xiphocolaptes 

major, Pseudoseisura unirufa, Mimus triurus e Agelaioides 

badius apresentam distribuição centrada na bacia do alto 

rio Paraguai, na planície do Pantanal e borda oeste do 

território brasileiro, contígua ao leste e norte da Bolívia. 

Tais aves podem ser consideradas elementos de influência 

chaquenha na região (Short 1975, Straube et al. 2006). 

Apesar da proximidade com o Pantanal, algumas 

espécies tidas como abundantes e comuns na planície 

pantaneira, notadamente aves paludícolas, são incomuns 

na Serra de Maracaju. Nesse contexto encontram-se 

Chauna torquata, Jabiru mycteria, Mycteria americana, 

Phalacrocorax brasilianus, Anhinga anhinga, Theristicus 

caerulescens, Rostrhamus sociabilis, Busarellus nigricollis 

e Agelasticus cyanopus. Essas aves foram avistadas com 

mais frequência em localidades próximas à planície de 

inundação, como Aquidauana e Fazenda Taboco. Mesmo 

espécies não relacionadas a ambientes aquáticos como 

Paroaria capitata, Paroaria coronata e Agelaioides badius 

têm suas distribuições restritas às regiões de ecótono entre 

o planalto e a planície do Pantanal. 

Espécies ameaçadas 

Quinze espécies presentes na Serra de Maracaju 

constam em listas de espécies criticamente ameaçadas, 

vulneráveis, ameaçadas ou quase-ameaçadas de extinção 

em âmbito global (BirdLife International 2009) e nacional 


96 




(Silveira & Straube 2008). Nesse sentido encontram-se 

Rhea americana, Nothura minor, Tigrisoma fasciatum, 

Urubitinga coronata, Harpia harpyja, Anodorhynchus 

hyacinthinus, Primolius maracana, Pyrrhura devillei, 

Alipiopsitta xanthops, Culicivora caudacuta, Alectrurus 

tricolor, Porphyrospiza caerulescens, Poospiza cinerea, 

Sporophila maximiliani e Charitospiza eucosma. 

A espécie A. hyacinthinus é tida como ameaçada de 

extinção em âmbito global e vulnerável à extinção em 

âmbito nacional e sua distribuição na Serra de Maracaju 

parece estar restrita às regiões de contato do planalto com 

a planície pantaneira, que abriga suas maiores populações 

no estado de Mato Grosso do Sul (Nunes 2010). 

O desmatamento e a descaracterização da paisagem 

são as principais intervenções humanas apontadas como 

ameaças à conservação de outras espécies de psitacídeos 

ocorrentes na região (Silveira & Straube 2008, BirdLife 

International 2009), dentre as quais podemos destacar P. 

maracana, P. devillei e A. xanthops. 

Apesar da espécie P. caerulescens não estar presente 

em nenhuma categoria de ameaça de extinção em âmbito 

nacional, seu estado de conservação no Mato Grosso do 

Sul deve ser considerado preocupante (Nunes 2009). 

Sua área de ocorrência é disjunta e grande parte das 

populações ocorrentes em território sul mato-grossense 

está restrita aos campos nos topos de morros do Maciço 

do Urucum, na borda oeste do Pantanal (Nunes 2009, 

Lopes 2012). Nunes (2009) relata ainda que tais habitats 

têm sido drasticamente reduzidos e alterados pela extração 

de minério de ferro. 

Outro caso preocupante é o da espécie S. maximiliani 

que se encontra criticamente ameaçado de extinção 

em âmbito nacional devido a sua captura na natureza 

e comércio no tráfico de animais silvestres (Silveira & 

Straube 2008). 

Para espécies raras e ecologicamente exigentes 

quanto ao uso de hábitat, como T. fasciatum, a situação 

é preocupante. Esta espécie ocorre exclusivamente 

em rios límpidos com corredeiras e orlados por densas 

florestas (Faria 2008). Desta forma, o desmatamento 

e a descaracterização da vegetação ripária ao longo dos 

rios que cortam a região, bem como a implantação de 

pequenas centrais hidroelétricas (PCHs) podem ser 

apontados como as principais ameaças à sua conservação 

(Nunes et al. 2012). 

Algumas espécies relacionadas para a região são 

conhecidas apenas por registros históricos, podendo 

ter sido extintas localmente devido às alterações no 

hábitat. Nesse contexto enquadram-se o L. magnificus, 

C. caudacuta e P. cinerea. Por outro lado, a ocorrência de 

grandes rapinantes na região, tais como U. coronata, H. 

harpyja e S. ornatus é um bom indicativo da qualidade 

do habitat, uma vez que tais aves necessitam de grandes 

territórios para sobreviver e se reproduzir (Robinson 

1994). 

Registros notáveis 

Nothura minor: espécie vulnerável à extinção em 

âmbito global e nacional, com registros históricos para 

a Fazenda Capão Bonito (FMNH-110492) e o Parque 

Estadual das Nascentes do rio Taquari (Hass 2004). Os 

demais registros no Mato Grosso do Sul incluem a região 

de Campo Grande (Sick 1997), onde provavelmente a 

espécie já esteja extinta. 

Anas bahamensis: marreca errante tida como rara 

e incomum no Centro Oeste do Brasil (Gwynne et al. 

2010). Há apenas um registro na Serra de Maracaju, no 

município de Sonora (Junqueira 2008). Considerando o 

Mato Grosso do Sul, essa marreca era conhecida apenas 

no Pantanal do Abobral (Nunes et al. 2010). 

Aburria nattereri: espécie típica da bacia amazônica 

que estende sua área de distribuição até as matas de galeria 

dos planaltos da região de Coxim (Fazenda Recreio), bem 

como os pantanais do Paiaguás, Miranda e Aquidauana 

(Nunes 2011), onde possivelmente ocorre em simpatria 

com sua congênere A. grayi. 

Tigrisoma fasciatum: espécie rara, ameaçada de 

extinção e de ocorrência restrita a rios encachoeirados 

e com lajedos rochosos escorregadios em meio às águas 

turbulentas (Gwynne et al. 2010). Há poucos registros 

dessa espécie em território sul-mato-grossense, que 

incluem além da região do Rio Verde, o Rio Sucuriú, o 

Planalto da Bodoquena (Pivatto et al. 2006, Nunes et 

al. 2012) e a Planície de Inundação do Alto Rio Paraná 

(Gimenes et al. 2007). 

Ictinia mississippiensis: migrante setentrional 

registrado em um grande bando sobrevoando a região de 

Aquidauana (Whittaker et al. 2008). Os demais registros 

desse gavião no Mato Grosso do Sul incluem algumas 

áreas da planície pantaneira e o Maciço do Urucum 

(Vasconcelos et al. 2008, Nunes 2011). 

Pseudastur albicollis: espécie típica da bacia 

amazônica, até então registrada apenas na Serra do Amolar, 

na borda oeste da planície do Pantanal. A ocorrência dessa 

espécie na Fazenda Trilhas do Sol pode ser considerada o 

registro mais ao sul da área de sua distribuição no Brasil. 

Harpia harpyja: espécie quase-ameaçada de extinção 

(Birdlife Internacional 2009) que recentemente era 

conhecida apenas no Planalto da Bodoquena (Pivatto 

et al. 2006). No entanto, essa ave tem sido registrada 

constantemente nos arredores da Fazenda Estância 

Crioula, no município de Dois Irmãos do Buriti. A 

ocorrência dessa espécie evidencia o bom estado de 

conservação da região, uma vez que depende de grandes 

áreas para sobreviver e se reproduzir.Spizaetus tyrannus: 

o avistamento desse gavião em Sidrolândia pode ser 

considerado o primeiro registro documentado da espécie 

em território sul-mato-grossense. Até então a espécie só 

era conhecida para a Fazenda Porto Conceição, no Chaco 

brasileiro (Straube et al. 2006). 





97 

Spizaetus ornatus: espécie tida como rara e incomum 

no Brasil (Carlos & Girão 2006). Apesar de não constar 

nas listas de espécies ameaçadas de extinção em âmbito 

global e nacional, seu estado de conservação é preocupante 

em detrimento da perda de hábitat e abate como forma de 

retaliação devido o ataque a animais domésticos (Nunes 

2010). Há poucos registros desse gavião no Mato Grosso 

do Sul, que incluem regiões como o Complexo Aporé- 

Sucuriú (Silva et al. 2006), Porto Murtinho (Tubelis & 

Tomas 2003) e o Maciço do Urucum (Tomas et al. 2010). 

Desta forma, os registros obtidos na Fazenda Trilha do Sol 

e nas Reservas Privadas do Patrimônio Natural (RPPNs) 

Vale do Bugio e Gavião de Penacho são muito importantes 

para a conservação da espécie no estado. 

Falco deiroleucus: falcão raro e incomum no Mato 

Grosso do Sul, com apenas dois registros, Fazenda 

Rio Negro (Donatelli 2005) e Fazenda Barranco Alto 

(Leuzinger 2009), ambas localizadas no Pantanal. 

Embora não esteja presente em nenhuma lista de espécies 

ameaçadas de extinção em âmbito global e nacional, seu 

estado de conservação em território sul-mato-grossense 

requer atenção especial devido a sua raridade. 

Patagioenas speciosa: espécie florestal comum na 

bacia amazônica e tida como rara no Mato Grosso do 

Sul, sendo registrada apenas na região conhecida como 

complexo Aporé-Sucuriú (Silva et al. 2006) e em Três 

Lagoas (J. C. Morante Filho com. pess.), ambas na região 

leste do estado. 

Pionus menstruus: psitacídeo incomum em território 

sul-mato-grossense, com registros na região leste na 

Fazenda Pouso Frio (Silva et al. 2006) e Pantanal, em 

localidades como Salobra (Tubelis & Tomas 2003), 

Fazenda Rio Negro (Donatelli 2005) e Fazenda Barranco 

Alto (Leuzinger 2009). 

Celeus flavus: Antas & Palo Jr. (2009) destacam 

que a RPPN SESC Pantanal é o limite sul da área de 

ocorrência dessa espécie típica da bacia amazônica. No 

entanto, tem sido registrada nos planaltos do entorno do 

Pantanal (Parque Estadual das Nascentes do Rio Taquari) 

e na planície pantaneira (Nunes 2011), onde ocorre em 

simpatria com o congênere chaquenho C. lugubris. 

Formicivora melanogaster: embora Gwynne et al. 

(2010) relacionem a ocorrência dessa espécie em grande 

parte do território sul-mato-grossense, há apenas um 

registro para o estado, no Parque Estadual das Nascentes 

do Rio Taquari. 

Berlepschia rikeri: espécie típica de veredas e 

buritizais do Brasil Central cuja ocorrência no Parque 

Estadual das Nascentes do Rio Taquari é esperada e pode 

ser considerado o primeiro registro em território sulmato-grossense. 

Synallaxis scutata: espécie rara e incomum no Mato 

Grosso do Sul. Além do Parque Nacional das Nascentes 

do Rio Taquari, há registros para o Maciço do Urucum 

(Naumburg 1930), região leste do estado (Piratelli 

1999) e Pantanal da Nhecolândia (Tubelis & Tomas 

2003). 

Oxyruncus cristatus: ave incomum na Bacia do Alto 

Paraguai, sendo registrada apenas na Serra das Araras, 

Mato Grosso (Willis & Oniki 1990). O avistamento no 

Parque Estadual das Nascentes do Rio Taquari é o único 

registro oficial dessa espécie no Mato Grosso do Sul. 

Tityra semifasciata: sua área de ocorrência no Mato 

Grosso do Sul é muito interessante, pois ocorre desde os 

cerrados no norte do estado até regiões de transição entre 

o Cerrado e as florestas estacionais da Bacia do Alto Rio 

Paraná (sul e sudeste do estado), o que amplia em mais 

de 600 km ao sul sua área de distribuição conhecida no 

Brasil (Godoi et al. 2011). 

Tyrannopsis sulphurea: ocorre na Amazônia, 

Maranhão, Piauí, Tocantins, Goiás e oeste de Minas 

Gerais (Pacheco et al. 2010). O registro desse tiranídeo no 

município de Sonora (Pequenas Centrais Hidroelétricas 

- PCH Santa Gabriela) pode ser considerado o primeiro 

para o Mato Grosso do Sul e o limite mais ao sul de sua 

área de ocorrência conhecida no Brasil. 

Myiarchus tuberculifer: trata-se do primeiro registro 

dessa espécie no Mato Grosso do Sul. No Brasil, segundo 

Sick (1997) ocorre em toda a Amazônia e porção oriental 

do país (de Alagoas ao Rio de Janeiro), sendo a região da 

PCH Santa Gabriela possivelmente o limite mais ao sul 

conhecido de sua área de distribuição. 

Catharus fuscescens: migrante setentrional até o 

momento registrado apenas no Maciço do Urucum, em 

Corumbá (Vasconcelos et al. 2008). Seu avistamento na 

Fazenda Trilhas do Sol amplia em mais de 280 km para 

o leste sua área de ocorrência conhecida no Mato Grosso 

do Sul. 

Saltator maximus: táxon típico do cerrado e 

considerado raro no Mato Grosso do Sul. Registrado 

apenas para os cerrados da região leste do estado (Pinto 

1932) e em algumas áreas do Pantanal (Donatelli 2005). 

Porphyrospiza caerulescens: espécie de ocorrência 

disjunta e restrita a campos e cerrados de topos de 

morros no Mato Grosso do Sul, sendo mais comum no 

Maciço do Urucum, borda oeste do Pantanal (Nunes 

2009). Classificada na categoria de quase ameaçada 

(BirdLife International 2009), segundo Nunes (2009), 

a maior ameaça às populações dessa espécie na Bacia 

do Alto Paraguai e Mato Grosso do Sul é a perda e a 

descaracterização do hábitat. 

Poospiza cinerea: espécie rara e de ocorrência localizada 

nos cerrados e campos cerrados do Brasil Central e estados 

de São Paulo (região norte) e Minas Gerais (Sick 1997). No 

Mato Grosso do Sul, existe apenas um registro oriundo de 

um espécime coletado na Fazenda Recreio. Provavelmente 

é substituída geograficamente por sua congênere P. 

melanoleuca, que é típica do Chaco (Straube et al. 2006). 

Consta como vulnerável à extinção em âmbito global de 

acordo com a BirdLife International (2009). 


98 




Sporophila maximiliani: táxon raro e incomum 

no Mato Grosso do Sul, com um único registro obtido 

na Fazenda Bocaina, região de transição para a planície 

do Pantanal (brejos do rio Taboco). É considerada 

criticamente ameaçada de extinção no Brasil (Silveira 

& Straube 2008) devido a sua captura na natureza para 

abastecer o comércio ilegal de animais silvestres, atividade 

comum na região (Nunes 2010). 

Arremon taciturnus: emberezídeo raro e incomum 

no Mato Grosso do Sul, sendo até então registrado apenas 

na Fazenda Lagoinha (Silva et al. 2006). O registro no 

Parque Estadual das Nascentes do Rio Taquari pode ser 

considerado o primeiro para a Bacia do Alto Rio Paraguai 

e amplia a área de distribuição conhecida dessa ave no 

estado. 

Charitospiza eucosma: espécie rara e quaseameaçada 

de extinção em âmbito global (BirdLife 

International 2009) com poucos registros em território 

sul-matogrossense. Há apenas um registro para a área 

de estudo, a Fazenda Constantino, no município de 

Corguinho. Os demais registros estão concentrados na 

região leste do estado, como Três Lagoas (MZUSP 12677, 

Pinto 1932) e Inocência (Bucci 2009). 

Considerações finais 

Comparada a outras regiões de Mato Grosso do 

Sul, notadamente o sul e leste do estado, os planaltos da 

Serra de Maracaju ainda mantém mosaicos de paisagens 

importantes para a conservação de aves no Mato Grosso 

do Sul e Cerrado do Brasil Central. A região abriga várias 

espécies ameaçadas e quase-ameaçadas de extinção e de 

distribuição restrita, atuando como uma área de extrema 

importância para a conservação de aves no Mato Grosso 

do Sul. 

As principais ameaças à biodiversidade local são o 

desmatamento promovido pela expansão da atividade 

agropecuária, ação de carvoarias e silvicultura, como 

plantações de Eucalyptus, que tem causado a perda e 

fragmentação de habitats naturais (Harris et al. 2006). A 

instalação de Pequenas Centrais Hidroelétricas (PCHs), 

além de aumentar as taxas de desmatamento e permitir 

o aumento na caça de animais silvestres, pode alterar o 

regime hidrológico dos rios da região e em longo prazo 

promover um grande desastre ambiental no interior da 

planície pantaneira, tal como o que ocorreu com o rio 

Taquari (Galdino et al. 2005, Tomas et al. 2007). 

Apesar da Serra de Maracaju estar inserida no 

Corredor de Biodiversidade Maracaju-Negro (Machado 

et al. 2009) e ser considerada área prioritária para a 

conservação da biodiversidade do Cerrado e Pantanal 

(MMA 2007), poucas são as unidades de conservação 

existentes. Estas áreas estão representadas principalmente 

por parques estaduais (Parque Estadual da Serra de 

Sonora e Parque Estadual das Nascentes do rio Taquari) 

e Reservas Particulares do Patrimônio Natural (RPPNs 

Gavião de Penacho, Vale do Bugio, Lajedo e Morro da 

Peroba/Fazenda Capão Bonito). Estas RPPNs e parques 

estaduais protegem apenas pouco mais de 10.000 ha de 

áreas naturais. Nesse sentido, faz-se urgente a criação 

de uma grande unidade de conservação que contemple 

os diferentes habitats existentes na paisagem regional, 

de modo a garantir a manutenção e sobrevivência no 

longo prazo da comunidade de aves presente na Serra 

de Maracaju. 

Agradecimentos 

Os autores são gratos a Conservação Internacional 

do Brasil, Universidade Federal de Mato Grosso do 

Sul e Fundação Manoel de Barros pelo apoio logístico 

ao projeto “Inventário Rápido da Biodiversidade no 

Corredor Serra de Maracaju-Negro”. Aos proprietários das 

fazendas Rodeio, Quinta do Sol, Constantino, Taboco e 

Bocaina e das RPPNs Gavião de Penacho e Vale do Bugio 

pela oportunidade de estudos na região. A Fundação 

Neotrópica do Brasil e a MMX Minerais Metálicos 

Ltda pelo apoio logístico durante o “Projeto Reflorestar 

Legal”. A Luis Fábio da Silveira e aos revisores que muito 

contribuiram para a versão final desse manuscrito. 

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Editor Associado: Luís Fábio Silveira 



March 2013 

Short-Communication 

On the nest, eggs and nestlings of the Short-tailed 

Antthrush (Chamaeza campanisona) 

Ismael Franz 

Laboratório de Ornitologia, Museu de Ciências e Tecnologia - MCT, Pontifícia Universidade Católica do Rio Grande do Sul – PUCRS, Avenida 

Ipiranga, 6681, CEP 90619-900, Porto Alegre, Rio Grande do Sul, Brazil; e-mail: ismaelfranz@gmail.com 

Received on 5 July 2012. Accepted on 20 January 2013. 

ABSTRACT: I characterize the nest, eggs and nestlings of the Short-tailed Antthrush (Chamaeza campanisona) based on a nest 

found in an Atlantic semideciduous forest in southern Brazil. The nest was in a natural cavity 1.5 m above the ground and it was 

87 cm deep. Three pure white eggs were recorded on a bed of leaves. The nestlings had pink skin, blackish gray down and they left 

the nest 16-19 days after hatching. Images from inside the nest, the eggs and nestlings are first presented, even for the genus. The 

Chamaeza nest type and egg coloration resemble nest describe for the family Formicariidae, supporting the separation from the 

family Grallariidae. 

KEY-WORDS: breeding, cavity nest, formicarids, Formicariidae, nesting 

The new Neotropical family Formicariidae, after 

the recent separation of the antpittas (Grallariidae), 

comprises 12 species (genera Formicarius and Chamaeza), 

and is closely related to Rhinocryptidae and Furnariidae 

(Irestedt et al. 2002, Krabbe & Schulenberg 2003, 

Chesser 2004, Rice 2005, Moyle et al. 2009). Mainly 

due to the difficulty of finding nests, basic reproductive 

aspects are poorly known in ground antbirds, including 

common species (Krabbe & Schulenberg 2003). This is 

the case to the Chamaeza antthrushes with only some 

scatter and poorly documented nesting records (Cadena 

et al. 2000). I characterize the nest, eggs and nestlings of 

the Short-tailed Antthrush (C. campanisona). Although 

C. campanisona is the most widespread Chamaeza, it 

have a patchy distribution between northern Venezuela 

and central Bolivia, also in the tepuis of southern 

Venezuela to Guyana, and more widely in Brazil, eastern 

Paraguay and northern Argentina, and there are 12 

recognized subspecies (Ridgely & Tudor 1994, Krabbe & 

Schulenberg 2003). 

I found the nest in a forest patch surrounded by a 

rural landscape in Sapiranga city, state of Rio Grande do 

Sul, southern Brazil (29º37’S; 51º02’W; 150m altitude). 

The semi-deciduous forest patch, with 80 ha, form part 

of the Atlantic Forest biome (sensu Teixeira et al. 1986). 

The landscape is a mosaic dominated by agricultural/ 

livestock with sparse forest patches. In the study site, 

the forest seems to have an advanced stage of succession, 

with canopy reaching between 10 and 20 m height. 

The vegetation is characterized by trees such as Trichilia 

claussenii, Cabralea canjerana, Cedrela fissilis (Meliaceae), 

Sorocea bonplandii (Moraceae), Nectandra megapotamica, 

N. oppositifolia (Lauraceae), Inga marginata (Fabaceae), 

Trema micrantha (Cannabaceae) and Allophylus edulis 

(Sapindaceae). Common birds are Red-crowned Anttanager 

(Habia rubica), Rufous-breasted Leaftosser 

(Sclerurus scansor), Golden-crowned Warbler (Basileuterus 

culicivorus), White-browed Warbler (B. leucoblepharus), 

Buff-browed Foliage-gleaner (Syndactyla rufosuperciliata), 

Yellow-olive Flycatcher (Tolmomyias sulphurescens), 

Surucua Trogon (Trogon surrucura), and Olivaceous 

Woodcreeper (Sittasomus griseicapillus), among others. 

I measured the nest with ruler and tape-measure 

to the nearest 0.5 cm. I conducted weekly inspection of 

the nest and took photographs of the cavity interior to 

document its contents. I conducted two short videos (30 

min each) at mid-day with a small video camera located 

6 m from the nest. Because the distance from the cavity 

entrance to bottom was greater than my arm, I could not 

access the eggs or nestling to take measurements, thus all 

my data is descriptive. The nest was classified based on 

Simon & Pacheco (2005). 

The nest was discovered on 12 November 2011 

when I flushed a bird during an avifaunal inventory. It 

was 15 m from the forest edge. The nest cavity was in a 

5 m tall living Allophylus edulis (Sapindaceae) tree, which 

had 17 cm in diameter at breast height and was partially 

tangled by a bamboo (Chusquea tenella). The ground

102 


Ismael Franz 

was fully covered with litter and the forest understory is 

moderately lightened. The cavity entrance was vertical 

with an irregular entrance 9.5 x 12.5 cm (Figure 1a), 

facing north. The bottom margin of the cavity entrance 

was 151 cm above the ground (Figure 1b) and 87 cm 

deep. At the bottom of the cavity (incubation chamber), 

the internal diameter was 11.5 x 9.5 cm. 

When located, three white eggs rested on a “bed” 

of green leaves of 1.5 cm of deep. As it was still green, I 

assumed that the birds led the leaves to cover the bottom 

of the cavity. The nest can be classified as cavity/withtunnel/simple/platform 

(sensu Simon & Pacheco 2005), 

although the “platform” is just a layer of loose leaves rather 

than a well-defined and built platform. The eggs are ovoid 

and pure white opaque without spots (Figure 1c). 

On 19 November 2011 there were three small 

nestlings on the bed of dried leaves, therefore, less than 6 

days old. Their eyes were closed, pink skin, light yellow 

bill and rictus and a blackish gray down covering the 

entire body. After knocking tree, I observed that one 

of the nestlings curved the neck, suggesting that it was 

dead (Figure 1d), remaining in this position for at least 

5 min, when I left the place. I returned after 30 min and 

the three nestlings were in normal position, always quiet 

(like a dark “spot” amid the leaves). No provisioning visit 

was recorded. 

FIGURE 1: Nest cavity (a and b), eggs (c), and nestlings


Ismael Franz 

103 

On 26 November 2011 9-12s day old nestlings 

still have a lot of down but their eyes were open, gray 

pin feathers were appearing in the tail, wings, and chest 

(apparently with some brownish), their bills were dark 

gray with light yellow on the rictus and yellow on the 

commissure and the distal end (Figure 1e). A 27 s feeding 

event was observed where one parent entered quickly and 

quietly the cavity, apparently relying on the cavity walls 

with his long legs. When the bird was leaving the nest it 

remained 4 s in the entrance of the cavity, then it flew 

quietly. On 3 December 2011 the nest was empty and I 

could not determine the nest fate. But it is estimated that 

nestlings left the nest when 16-19 days old, assuming the 

nest was not predated. 

The photos of the nest cavity inside, eggs and 

nestlings of the Short-tailed Antthrush are first presented, 

even considering the genus Chamaeza. Recently, Maders 

& Matuchaka (2011) described the nest, eggs and nestlings 

of C. campanisona tshororo (the same subspecies of this 

study) based on a nest found in September 2002 in the 

province of Misiones, Argentina, without present images. 

Additionally, a photo of an adult of the species in the nest 

cavity entrance was presented in Krabbe & Schulenberg 

(2003), from Alagoas, Brazil (presumably the nominal 

subspecies). Although there is mention that antthrushes 

also nests among plants in the ground or on tree trunks 

(Sick 1997), nests in cavities appears to be a tendency, 

at least for Chamaeza. The nest described here resembles 

that of Argentina, but with shorter distance from the 

ground (1.5 x 3 m) and greater depth of the cavity (87 x 

21 cm). These also resembles a nest of Striated Antthrush 

(C. nobilis) recorded in the Colombian Amazon, in 

a cavity 3 m above the ground with a depth of 30 cm 

(Cadena et al. 2000). Representatives from another genus 

of Formicariidae, Formicarius, also use natural cavities in 

trees for nesting (Cherrie 1916, Skutch 1945, Krabbe & 

Schulenberg 2003). The difficulty of finding nests and 

monitor its content does seem to be the main reason 

why so little information about the reproduction of 

Formicariidae is available. 

Instead of a clutch size of two eggs, as reported in 

the literature as apparently normal for the family (Sick 

1997, Krabbe & Schulenberg 2003), I found three, as well 

as in the nest found in Argentina (Maders & Matuchaka 

2011). As also noted by these authors, the eggs of 

Chamaeza campanisona are clearly unequal poles instead 

of almost spherical (Sick 1997, Krabbe & Schulenberg 

2003). It was not possible to measure eggs to compare 

with existing data, which refer to averages between 27 and 

28 x 22-24 mm (Ihering 1900, Schönwetter 1979, Fraga 

& Narosky 1985) or slightly larger (32 x 23 mm; Maders 

& Matuchaka 2011). All eggs of Formicariidae (Chamaeza 

+ Formicarius) described have white color without spots. 

The only previous description of the Short-tailed 

Antthrush´s nestlings was by Maders & Matuchaka 

(2011): “grayish feathers on the head, wings and tail, the 

rest of the body naked.” Thus, this study has contributed 

with more information and greater detail, presenting the 

first images of the nestlings. If confirmed as such, the 

observed “play possum/dead” behavior can be equivalent 

to the one recorded in the Rusty-breasted Antpitta 

(Grallaricula ferrugineipectus) by Schwartz (1957), but 

see Niklison et al. (2008). It would be interesting to 

manipulate antthrush nestlings to ascertain the occurrence 

of such behavior. I estimated in 16-19 days the time to 

the nestlings leave the nest, which is consistent with the 

value found for Formicarius analis (18 days; Sick 1997). 

The color and general appearance of young Chamaeza 

campanisona when leave the nest remains unknown. 

When compared with the representatives of 

the Grallariidae (revision in Greeney et al. 2008), 

Formicariidae have remarkable differences in relation to 

reproductive attributes. The most evident are the nest 

type (open cup vs. cavity, respectively) and the egg color 

(bluish, greenish, turquoise, pale buff or brown with or 

without spots vs. white without spots). In antthrushes, 

these features seem to be quite consistent and corroborate 

their separation from antpittas (Rice 2005). The 

tendency to hide the eggs (mostly nesting in cavities) 

and whitish eggs are shared by the large group composed 

by Formicariidae + Rhinocryptidae + Furnariidae sensu 

lato + Scleruridae + Dendrocolaptidae, corroborating 

the molecular phylogenies (Irestedt et al. 2002, Chesser 

2004, Moyle et al. 2009). Thamnophilidae, Grallariidae 

and Conopophagidae, more basal Furnariides, have open 

nests and lay colored and/or spotted eggs (Rice 2005). 

Finally, I emphasize that information such as duration 

of incubation, nestling development and parental care 

remain virtually nonexistent for the representatives of the 

Formicariidae and therefore worthy of study efforts. 


I am grateful to Giovanni N. Maurício and Carla 

S. Fontana for comments on the manuscript and 

bibliography provided. José E. Simon for literature 

recommendation. Coordenação de Aperfeiçoamento de 

Pessoal de Nível Superior - CAPES for providing funding. 

Rodney Schmidt for the bamboo identification. And to 

Antonio, Ilani, Bárbara, Valdomiro, Cenira and Fabiano 

for the logistic support in the study area (Sítio Bonanza) 

and friendship. 

REFERENCES 

Cadena, C. D.; Londoño, G. A. & Parra, J. L. 2000. Nesting records 

of five antbird species from the Colombian Amazon. Wilson 

Bulletin, 112: 313-317. 


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Ismael Franz 

Cherrie, G. K. 1916. A contribution to the ornithology of the 

Orinoco region. Museum of the Brooklyn Institute of Arts and 

Sciences, Scientific Bulletin, 2: 133-374. 

Chesser, R. T. 2004. Molecular systematics of New World suboscine 

birds. Molecular Phylogenetics and Evolution, 32: 11-24. 

Fraga, R. & Narosky, S. 1985. Nidificación de las aves argentinas 

(Formicariidae a Cinclidae). Buenos Aires: Asociación Ornitológica 

del Plata. 

Greeney, H. F.; Dobbs, R. C.; Martin, P. R. & Gelis, R. A. 2008. 

The breeding biology of Grallaria and Grallaricula antpittas. 

Journal of Field Ornithology, 79: 113-129. 

Ihering, H. von. 1900. Catálogo crítico-comparativo dos ninhos e 

ovos das aves do Brasil. Revista do Museu Paulista, 4: 191-300. 

Irestedt, M.; Fjeldså, J.; Johansson, U. S. & Ericson, P. G. P. 2002. 

Systematic relationships and biogeography of the tracheophone 

suboscines (Aves: Passeriformes). Molecular Phylogenetics and 

Evolution, 23: 499-512. 

Krabbe, N. K. & Schulenberg, T. S. 2003. Family Formicariidae 

(ground-antbirds), p. 748-788. In: del Hoyo, J.; Elliott, A. 

& Christe, D. A. (eds.). Handbook of the birds of the world, 

Broadbills to Tapaculos, v. 8. Barcelona: Lynx Edicions. 

Maders, C. J. & Matuchaka, V. 2011. Descripción de nido, huevos y 

pichones de la Tovaca Común (Chamaeza campanisona tshororo). 

Nuestras Aves, 56: 10-11. 

Moyle, R. G.; Chesser, R. T.; Brumfield, R. T.; Tello, J. G.; Marchese, 

D. J. & Cracraft, J. 2009. Phylogeny and phylogenetic classification 

of the antbirds, ovenbirds, woodcreepers, and allies (Aves: 

Passeriformes: infraorder Furnariides). Cladistics, 25: 386-405. 

Niklison, A. M.; Areta, J. I.; Ruggera, R. A.; Decker, K. L.; Bosque, 

C. & Martin, T. E. 2008. Natural history and breeding biology 

of the Rusty-breasted Antpitta (Grallaricula ferrugineipectus). The 

Wilson Journal of Ornithology, 120: 345-352. 

Rice, N. H. 2005. Further evidence for paraphyly of the Formicariidae 

(Passeriformes). The Condor, 107: 910-915. 

Ridgely, R. S. & Tudor, G. 1994. The birds of South America, The 

suboscine passerines, v. 2,. Austin: University of Texas Press. 

Schönwetter, M. 1979. Handbuch der oologie (Passeriformes 1), v. 2. 

Berlin: Akademie-Verlag. 

Schwartz, P. 1957. Observaciones sobre Grallaricula ferrugineipectus. 

Boletín de la Sociedad Venezolana de Ciencias Naturales, 18: 42-62. 

Sick, H. 1997. Ornitologia brasileira. Rio de Janeiro: Nova Fronteira. 

Simon, J. E. & Pacheco, S. 2005. On the standardization of nest 

descriptions of neotropical birds. Revista Brasileira de Ornitologia, 

13: 143-154. 

Skutch, A. F. 1945. On the habits and nest of the Ant-thrush 

Formicarius analis. Wilson Bulletin, 57: 122-128. 

Teixeira, M. B.; Coura-Neto, A. B.; Pastore, U. & Rangel-Filho, A. 

L. R. 1986. Vegetação, p. 541-620. In: IBGE. Levantamento de 

recursos naturais, v. 33. Rio de Janeiro: IBGE. 

Associate Editor: Caio Graco Machado 



March 2013 

Short-Communication 

Registros documentados da gaivota-de-franklin 

Leucophaeus pipixcan (Wagler, 1831) no Nordeste do Brasil 

e da gaivota-de-cabeça-cinza Chroicocephalus cirrocephalus 

(Vieillot, 1817) nos Estados de Alagoas e Pernambuco 

Sergio Leal 1 , Lúcio Cláudio H. Serapião 2 e Glauco Alves Pereira 3,4,5 

1 

Organização de Preservação Ambiental – OPA. 

2 

Graduando em Medicina Veterinária pela Fundação Universidade do Vale do São Francisco - UNIVASF. 

3 

Observadores de Aves de Pernambuco (OAP). 

4 

Programa de Pós-Graduação em Etnobiologia e Conservação da Natureza – UFRPE. 

5 

Autor correspondente: glaucoapereira@yahoo.com.br 

Recebido em 11 de outubro de 2011. Aceito em 24 de agosto de 2012. 

ABSTRACT: First documented records of Franklin’s Gull Leucophaeus pipixcan (Wagler, 1831) from north east Brazil and 

Grey-headed Gull Chroicocephalus cirrocephalus (Vieillot, 1817) in the states of Alagoas and Pernambuco. We report on the 

first documented record of Franklin’s Gull in north-eastern Brazil, when one individual was found dead in Maceió, state of Alagoas, 

on 27 July 2010, and other was found and photographed very weak on the same beach on 8 May 2011. Since most records of this 

species in South America come from the western part of the continent, this individual probably reached northeastern Brazil through 

the Caribbean. About 25 Grey-headed gulls were photographed at Barra de São Miguel, state of Alagoas, on 5 March 2010, and two 

individuals were observed at the same place on 23 April 2010. In the state of Pernambuco, two birds were photographed at Itapissuma 

on 1 October 2010. These are the first records of the Grey-headed Gull for Pernambuco and Alagoas states in northeastern Brazil. 

KEY-WORDS: gulls, migration, photographed records. 

No Brasil há nove espécies de gaivotas (CBRO 2011), 

dos quais seis ocorrem na região Nordeste: Chroicocephalus 

maculipennis, C. cirrocephalus, Leucophaeus atricilla, 

L. pipixcan, Larus dominicanus e L. fuscus (Sick 1997, 

Azevedo-Junior et al. 2004, Girão et al. 2006, Albano et 

al. 2007, Farias et al. 2008, Souza & Borges 2008, Lepage 

& Pereira 2010, Lima et al. 2010). Alguns desses registros, 

como o de L. pipixcan em Fernando de Noronha (Antas 

et al. 1988), não foram acompanhados de nenhum tipo 

de documentação científica (fotografia, gravação da voz 

ou coleta de espécimes), gerando questionamentos sobre 

suas legitimidades, conforme exposto em Nacinovic & 

Teixeira (1989). 

Nesse trabalho, divulgamos o primeiro registro 

documentado de L. pipixcan no Nordeste do Brasil e os 

primeiros registros de C. cirrhocephalus para os estados de 

Alagoas e Pernambuco. Os indivíduos foram identificados 

através das pranchas, fotos e informações morfológicas 

disponíveis em Harrison (1983, 1987), Burger & 

Gochfeld (1996), Enticott & Tipling (1997), Erize et al. 

(2006) e Howell & Dunn (2007). 

Leucophaeus pipixcan 

A gaivota-de-franklin Leucophaeus pipixcan ocorre no 

interior da América do Norte, onde se reproduz, migrando 

durante o inverno para o continente sulamericano, 

permanecendo na costa do Peru, Chile, Patagônia, Tristão 

da Cunha, Terra do Fogo e Argentina. Vagantes já foram 

encontrados na Europa, África, Cazaquistão, Emirados 

Árabes Unidos, Israel, China, Japão e Austrália (Harrison 

1983, 1987, Burger & Gochfeld 1996, Enticott & 

Tipling 1997, Olsen & Larsson 2003, Smith 2004, Holt 

2005, Howell & Dunn 2007, Dias et al. 2010, Wassink et 

al. 2011). No Brasil, há registros da espécie em Fernando 

de Noronha, em maio de 1988 (Antas et al. 1988), no 

Amazonas, em março de 1994 (Pacheco 1995), em 

São Paulo, em setembro de 2002 (Almeida 2003) e no 

Rio Grande do Sul, em abril de 1999, março de 2007, 

janeiro e abril de 2009, e em maio de 2010 (Dias et al. 

2010). O registro em Fernando de Noronha, como não 

foi documentado, foi considerado duvidoso, havendo a 

possibilidade de confusão com L. atricilla, mais comum 

na região (Nacinovic & Teixeira 1989).

106 

Registros documentados da gaivota-de-franklin Leucophaeus pipixcan (Wagler, 1831) no Nordeste do Brasil e da gaivota-de-cabeça-cinza 


Sergio Leal, Lúcio Cláudio H. Serapião e Glauco Alves Pereira 

Um indivíduo de Leucophaeus pipixcan foi 

encontrado morto na praia de Jacarecica, Maceió, Alagoas 

(9°36’S; 35°41’W), em 27 de julho de 2010. Um segundo 

indivíduo foi fotografado enquanto estava bastante 

debilitado na mesma praia em 8 de maio de 2011 (Figura 

1). O bico curto e avermelhado, as primárias negras com 

pontas brancas e as medidas de comprimento (38 cm) e 

envergadura (90 cm) indicam ser um indivíduo adulto 

com a plumagem reprodutiva (Harrison 1983, Howell 

& Dunn 2007), não correspondendo a L. atricilla, que 

ocorre em áreas do Nordeste do Brasil (Albano et al. 2007). 

Esses e outros indivíduos vagantes na costa brasileiras 

podem ser oriundos do leste dos Andes, podendo cruzar 

a cordilheira e entrar no continente pelo Caribe durante 

sua migração para o sul ou como vagantes que cruzam o 

Oceano Atlântico em direção à África (Dias et al. 2010). 

Figura 1: Leucophaeus pipixcan, Praia de Jacarecica, Maceió, Alagoas, Brasil (Foto de Sergio Leal). 

Figure 1: Franklin’s Gull, Jacarecica Beach, Maceió, Alagoas, Brazil (Photo by Sergio Leal). 

Chroicocephalus cirrocephalus 

A gaivota-de-cabeça-cinza Chroicocephalus cirrocephalus, 

ao contrário da anterior, é uma espécie localmente 

residente nos continentes sulamericano e africano, 

havendo registros de indivíduos vagantes no Panamá, 

Norte do Mar Vermelho, América do Norte e Europa 

(Harrison 1983, 1987, Enticott & Tipling 1997, Olsen 

& Larsson 2003, Howell & Dunn 2007). No Brasil, há 

registros para os Estados do Rio de Janeiro, São Paulo, 

Maranhão, Mato Grosso do Sul, Santa Catarina, Rio 

Grande do Sul, Piauí, Ceará e Rio Grande do Norte (Sick 

1997, Azevedo-Junior et al. 2004, Albano et al. 2007, 

Barbieri et al. 2010). 

Um bando com 25 indivíduos adultos, com 

plumagem reprodutiva e em descanso reprodutivo, foi 

fotografado na Praia das Conchas, Barra de São Miguel, 

Alagoas (9°50’S; 35°53’W), em 5 de março de 2010 

(Figura 2). Dois indivíduos foram registrados no mesmo 

local, em 23 de abril do mesmo ano. Em Pernambuco, 

dois indivíduos jovens foram fotografados em Itapissuma 

(07°46’S; 34°56’W), em 1 de outubro de 2011 (Figura 

3). Os dois não possuíam as pontas das coberteiras das 

primárias brancas, no entanto, de acordo com Howell & 

Dunn (2007), alguns indivíduos podem não apresentar 

essa característica. Esses são os primeiros registros dessa 

espécie para os estados de Alagoas e Pernambuco. C. 

cirrocephalus se dispersa para o norte do continente 

durante o inverno austral, principalmente entre os meses 

de maio a outubro (Howell & Dunn 2007), coincidindo 

justamente com o período em que os registros desse 

trabalho foram realizados. 





107 

Figura 2: Chroicocephalus cirrocephalus, Praia das Conchas, Barra de São Miguel, Alagoas, Brasil (Foto de Sergio Leal). 

Figure 2: Grey-headed Gull, Conchas Beach, Barra de São Miguel, Alagoas, Brazil (Photo by Sergio Leal). 

Figura 3: Chroicocephalus cirrocephalus, Itapissuma, Pernambuco, Brasil (Foto de Lúcio Cláudio Serapião). 

Figure 3: Grey-headed Gull, Itapissuma, Brazil (Photo by Lúcio Cláudio Serapião). 


108 




AGRADECIMENTOS 

Aos revisores anônimos da Revista Brasileira de 

Ornitologia pelas correções e sugestões. Ao amigo John 

Medcraft pela ajuda no abstract. Aos demais amigos que 

direta ou indiretamente ajudaram na obtenção dos dados 

e publicação deste manuscrito. 

REFERÊNCIAS 

Albano, C.; Girão, W. & Campos, A. 2007. Aves Costeiras de Icapuí. 

Icapuí: Editora Fundação Brasil Cidadão. 

Almeida, A. N. F. 2003. First documented record of Franklin’s Gull 

(Larus pipixcan) in Brazil. Ararajuba, 11:116-117. 

Antas, P. T. Z.; Filippini, A. & Azevedo-Junior, S. M. 1988. 

Anilhamento de aves oceânicas e/ou migratórias no arquipélago de 

Fernando de Noronha em 1987 e 1988. Em: Anais do 4° Encontro 

Nacional de Anilhadores de Aves, p.13-17. Recife: Universidade 

Federal Rural de Pernambuco. 

Azevedo-Júnior, S. M.; Larrazábal & Pena, O. 2004. Aves aquáticas 

de ambientes antrópicos (salinas) do Rio Grande do Norte, Brazil, 

p. 255-266. Em: Branco, J. O. (Org.). Aves marinhas e insulares 

brasileiras: biologia e conservação. Itajaí: Editora UNIVALI. 

Barbieri, E.; Gonçalves, C. A.; Silveira, L. F. & Cortez-kiyohara, A. 

L. L. 2010. Registros de duas aves marinhas inéditas no estado de 

São Paulo, Brasil: Chroicocephalus cirrocephalus e Anous minutus 

(Charadriiformes). Revista Brasileira de Ornitologia, 18: 242-244. 

Burger, J. & Gochfeld, M. 1996. Family Laridae (Gulls), p.572-623. 

In: Del Hoyo, J.; Elliott, A. & Sargatal, J. (eds.). Handbook of 

the birds of the world. Vol. 3. Hoatzin to Auks. Barcelona: Lynx 

Ediciones. 

CBRO. Comitê Brasileiro de Registros Ornitológicos. 2011. Listas 

das aves do Brasil. 10ª Edição. www.cbro.org.br (Acesso em 31 de 

novembro de 2011). 

Dias, R. A.; Agne, C. A.; Gianuca, D.; Gianuca, A.; Barcellos- 

Silveira, A. & Bugoni, L. 2010. New records, distribution and 

status of six seabirds species in Brazil. Iheringia, Série. Zoológica., 

100: 379-390. 

Enticott, J. & Tripling, D. 1997. Seabirds of the world: the complete 

reference. New Holland: Stackpole Books. 

Farias, G. B.; Pereira, G. A. & Silva, W. A. G. 2008. Lista das Aves de 

Pernambuco. Recife: Observadores de Aves de Pernambuco. 

Girão, W.; Albano, C.; Pinto, T.; Campos, A.; Meirellis, A. C. 

& Silva, C. P. N. 2006. First record of the Lesser Black-backed 

Gull Larus fuscus Linnaeus, 1758 for Brazil. Revista Brasileira de 


Harrison, P. 1983. Seabirds: an identification guide. Boston: Houghton 

Mifflin Co. 

Harrison, P. 1987. Seabirds of the World: A photographic guide. 

London: Christopher Helm. 

Holt, P. 2005. Franklin’s Gull Larus pipixcan at Tanggu, Tianjin: first 

record for China. Forktail, 21: 171-173. 

Howell, S. N. G. & Dunn, J. 2007. Gulls of the Americas. Boston: 

Houghton Mifflin. 

Lepage, R. & Pereira, G. A. 2010. Registros documentados do 

gaivotão Larus dominicanus (Charadriiformes: Laridae) para o 

Estado de Pernambuco, Brasil. Revista Brasileira de Ornitologia, 

18: 256-257. 

Lima, L. M.; Schunck, F.; Siciliano, S.; Carlos, C. J.; Rennó, B.; 

Fonseca-Neto, F. P.; Fedrizzi, C. E; Albano, C. & Moura, 

J. F. 2010. Distribuição, abundância e sazonalidade de 

Leucophaeus atricilla (Charadriiformes: Laridae) no Brasil. Revista 

Brasileira de Ornitologia, 18: 199-206. 

Nacinovic, J. B. & Teixeira, D. M. 1989. As aves de Fernando de 

Noronha: uma lista sistemática anotada. Revista Brasileira de 

Biologia, 49: 709-729. 

Olsen, K. M. & Larsson, H. 2003. Gulls of North America, Europe 

and Asia. New Jersey: Princeton University Press. 

Pacheco, J. F. 1995. Ocorrência acidental da gaivota-de-franklin, 

Larus pipixcan no médio Solimões, Amazonas. Atualidades 

Ornitológicas, 66: 4. 

Sick, H. 1997. Ornitologia Brasileira. Rio de Janeiro: Nova Fronteira. 

Smith, J. P. 2004. The first Franklin’s Gull Larus pipixcan in Israel and 

the Middle East. Sandgrouse, 26: 65-67. 

Souza, D. G. S. & Borges, O. B. 2008. Lista das Aves do Estado da 

Bahia, Brasil – Versão novembro 2008. www.anor.org.br (Acesso 

em 3 de novembro de 2011). 

Wassink, A.; Ahmed, R.; Busuttil, S.; & Salemgareev, A. 2011. 

Oriental Plover, Franklin’s Gull, Syrian Woodpecker and Masked 

Shrike new to Kazakhstan. Dutch Birding, 33: 239-244. 

Editor Associado: Alexander C. Lees 



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Articles 

Fargione, J.; Hill, J.; Tilman, D.; Polasky, S. & Hawthornez, P. 2008. Land clearing and the biofuel carbon debt. Science, 319: 1235-1238. 

Santos, M. P. D. & Vasconcelos, M. F. 2007. Range extension for Kaempfer’s Woodpecker Celeus obrieni in Brazil, with the first male specimen. 

Bulletin of the British Ornithologists’ Club, 127: 249-252. 

Worthington, A. H. 1989. Adaptations for avian frugivory: assimilation efficiency and gut transit time of Manacus vitellinus and Pipra mentalis. 

Oecologia, 80: 381-389. 

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Sick, H. 1985. Ornitologia brasileira, uma introdução, v. 1. Brasília: Editora Universidade de Brasília. 

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(Studies in Avian Biology 13). 

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Novaes, F. C. 1970. Estudo ecológico das aves em uma área de vegetação secundária no Baixo Amazonas, Estado do Pará. Ph.D. dissertation. Rio Claro: 

Faculdade de Filosofia, Ciências e Letras de Rio Claro. 

Web-Based References 

Dornas, T. 2009a. [XC95575, Celeus obrieni]. www.xeno-canto.org/95575 (access on 25 February 2012). 

Dornas, T. 2009b. [XC95576, Celeus obrieni]. www.xeno-canto.org/95576 (access on 25 February 2012). 

IUCN. 1987. A posição da IUCN sobre a migração de organismos vivos: introduções, reintroduções e reforços. http://iucn.org/themes/ssc/pubs/ 

policy/index.htm (access on 25 August 2005). 

Pinheiro, R. T. 2009. [WA589090, Celeus obrieni Short, 1973]. www.wikiaves.com/589090 (access on 05 March 2012). 

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Revista Brasileira de de Ornitologia 

Volume Volume 20 21 – 20 Número –– Número 2 – 1 Junho 2 –– Março Junho 2012 2013 2012 / Issue // Issue 20 – 21 20 Number – Number 2 – 1 June 2 – March June 2012 2012 2013 

SUMÁRIO / CONTENTS / artigos / papers 

Plumage variation in the Planalto Woodcreeper (Dendrocolaptes platyrostris) and the melanocortin-1 receptor gene 

(MC1R) 

Josmael Corso, Gustavo Sebastián Cabanne, Fernando Mendonça d’Horta, Élgion Lucio da Silva Loreto and Cristina Yumi 

Miyaki............................................................................................................................................................................. 


André Magnani Xavier de Lima and James Joseph Roper...................................................................................................... 


Paulo Rubim..................................................................................................................................................................... 

One hundred and thirty-five years of avifaunal surveys around Santarém, central Brazilian Amazon 

Alexander C. Lees, Nárgila G. de Moura, Christian Borges Andretti, Bradley J. W. Davis, Edson V. Lopes, L. Magalli Pinto 

Henriques, Alexandre Aleixo, Jos Barlow, Joice Ferreira and Toby A. Gardner........................................................................ 


Judith Figueroa.................................................................................................................................................................. 


Alessandro Pacheco Nunes, Maurício Neves Godoi, Maria Antonietta Castro Pivatto, José Carlos Morante Filho, Eduardo 

Weffort Patrial, Paulo Antonio Silva, Vanessa Katherinne Stavis, Daniel De Granville Manço, Marco de Barros Costacurta, 

Caroline Leuchtenberger e Carlos Rodrigo Lehn.................................................................................................................. 

1 

5 

10 

16 

58 

75 

NotaS / Short-CommunicationS 


Ismael Franz..................................................................................................................................................................... 

Registros documentados da gaivota-de-franklin Leucophaeus pipixcan (Wagler, 1831) no Nordeste do Brasil e da gaivota-decabeça-cinza 


Sergio Leal, Lúcio Cláudio H. Serapião e Glauco Alves Pereira............................................................................................. 

101 

105 

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