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Accepted by T. Chesser: 12 Nov. 2010; published: 17 Jan. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2011 · Magnolia Press
Zootaxa 2742: 1–33 (2011)
www.mapress.com/zootaxa/Article
1
Taxonomy and geographic variation of Formicivora serrana (Hellmayr, 1929)
and Formicivora littoralis Gonzaga and Pacheco, 1990
(Aves: Passeriformes: Thamnophilidae)
DANIEL H. FIRME1 & MARCOS A. RAPOSO1
Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Setor de Ornitologia, Quinta da Boa Vista,
20940-040, Rio de Janeiro, RJ, Brazil. E-mail: 1danielfirme@hotmail.com
Abstract
Formicivora serrana was described by Hellmayr in 1929 from specimens collected in Minas Gerais state, and subspecies
F. s . i n t e r p o s i t a and F. s . l i t t o r a li s by Gonzaga and Pacheco (1990) from specimens from Minas Gerais and Rio de Janeiro
states and Rio de Janeiro state, respectively. According to the latter authors, interposita is intermediate both geographical-
ly and morphologically in relation to F. s . s e r r a n a and F. s . l i t t o r a l i s . Two years after its description, F. s . l i t t o r a l i s was
elevated to species status (Collar et al. 1992) and it has subsequently been considered a species of conservation concern
because of its restricted geographical range and threatened habitat. Formicivora s. interposita, however, was not elevated
and is generally considered a subspecies of serrana. In this study we review the taxonomy of F. s e r r an a , testing the va-
lidity of F. l i t t o r a l i s and proposing the best taxonomic treatment for F. s . i n t e r p o s i t a . The results of morphometric, vocal
and plumage analyses show that F. s . i n t e r p o s i t a and F. l i t t o r a l i s lack diagnostic characters, and are therefore not valid
phylogenetic species. Further, our results indicate that F. l i t t o r a l i s is not a valid biological species, but that it and inter-
posita could be considered subspecies of F. serrana under this species concept. These results stress the need for detailed
systematic studies in support of taxonomic decisions.
Key words: taxonomy, geographic variation, Formicivora serrana, Formicivora littoralis, Thamnophilidae
Resumo
Formicivora serrana foi descrita por Hellmayr em 1929 a partir espécimes coletados no estado de Minas Gerais, enquanto
as subespécies F. s . i n t e r p o s i t a e F. s . l i t t o r a l i s foram descritas por Gonzaga e Pacheco (1990) para espécimes coletados
nos estados de Minas Gerais e Rio de Janeiro e Rio de Janeiro, respectivamente. Segundo esses autores, F. s . i n t e r p o s i t a
foi descrita como subespécie de F. s e r r a n a devido à sua condição intermediária tanto geográfica quanto morfológica em
relação a F. s . s e r r a n a e F. s . l i t t o r a l i s . Dois anos após sua descrição como subespécie de F. s e r r a n a , F. s . l i t t o r a l i s passou
a ser tratada como uma espécie válida (Collar et al. 1992) e subseqüentemente considerada ameaçada devido à sua dis-
tribuição restrita pela perda de hábitat. Formicivora s. interposita, porém, não foi elevada a espécie, sendo considerada
uma subespécie de F. s e r r a n a . Neste estudo, nós revisamos a taxonomia de F. s e r r a n a , testando a validade de F. l i t t o r a l i s
e verificando o melhor tratamento a ser dado a F. s . i n t e r p o s i t a . Os resultados das análises morfométricas, de plumagem
e vocais mostraram que F. s . i n t e r p o s i t a e F. l i t t o r a l i s não são espécies filogenéticas válidas devido à falta de caracteres
diagnósticos. Além disso, nossos resultados indicam que F. l i t t o r a l i s , sob o ponto de vista do conceito biológico de es-
pécie, poderia ser considerada, juntamente com interposita, subespécie de F. s e r r a n a . Esses resultados enfatizam a ne-
cessidade da realização de estudos sistemáticos detalhados para que decisões taxonômicas bem suportadas sejam tomadas.
Palavras-chave: Taxonomia, variação geográfica, Formicivora serrana, Formicivora littoralis, Thamnophilidae
Introduction
The genus Formicivora Swainson comprises eight species (Gonzaga et al. 2007), four of which are considered
polytypic (Zimmer & Isler 2003). These species inhabit a variety of non-forest habitats throughout South America
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and part of Central America (Gonzaga 2001). Formicivora species are sexually dimorphic in plumage (Ridgely &
Tudor 1994; Sick 1997) and juvenile males are similar to females in color pattern (Gonzaga 2001).
Neorhopias (Formicivora) serrana was described by Hellmayr in 1929 based on a specimen collected by Rein-
hardt in Sete Lagoas, Minas Gerais state. This species was distinguished from F. melanogaster by its mars brown
upperparts, the cinnamon brown or russet fringes to the outer webs of its tertials, and its posteriorly widened super-
ciliaries (Hellmayr 1929) (Figure 1).
Gonzaga and Pacheco (1990) described two new subspecies of F. serrana, F. s . i n t e r p o s i t a and F. s . l i t t o r a l i s .
Under this taxonomy, nominate serrana was considered to inhabit the mountains of Minas Gerais and Espírito
Santo states, at altitudes above 1000 m; interposita the Vale do Paraíba, in Minas Gerais and Rio de Janeiro states,
at altitudes above 700 m; and littoralis the restingas of the municipalities of Saquarema through to Cabo Frio and
Comprida and Cabo Frio islands, ca. sea level, in Rio de Janeiro state (Gonzaga & Pacheco 1990) (Figure 2).
According to these authors, the extremes of variation in these forms are readily diagnosable. Males of F. s. serrana
have brown upperparts and a well marked white supercilium (Figure 1), whereas males of F. s . l i t t o r a l i s , at the
other extreme, are characterized by dark upperparts, which vary from Vandyke Brown to Sepia (Figure 3A), and
the lack of a supercilium (Figure 3B). Additionally, the white spots of the median and lesser wing coverts are
diminutive or absent in littoralis. The geographically intermediate taxon, F. s . i n t e r p o s i t a , is also morphologically
intermediate, as demonstrated by its upperparts, which vary from Vandyke Brown to Sepia (Gonzaga & Pacheco
1990) (Figure 3A), the diminutive white spots on the middle and lesser wing coverts, and a thin and interrupted
supercilium (Figure 3B) (Gonzaga & Pacheco 1990).
Apart from these intermediate specimens, other factors, including similarity in vocalizations and similarity of
female plumage in these three taxa (Gonzaga & Pacheco 1990), persuaded the authors to consider littoralis and
interposita as subspecies of F. serrana, although they noted that morphological and ecological characters might
argue for species status for littoralis.
Two years after i ts d escription, F. s . l i t t o r a l i s was elevated to species status by Collar et al. (1992), based
largely on the argument of Gonzaga and Pacheco (1990). Subsequently, F. l i t t o r a l i s has been considered both
threatened (Tobias & Williams 1996; BirdLife International 2004; Rocha et al. 2005; Marini & Garcia 2005;
Vechhi & Alves 2008) and a valid species (e. g. Sick 1997; Zimmer & Isler in del Hoyo et al. 2003), as well as the
only endemic bird species of the restingas (Sick 1997; Reis & Gonzaga 2000; Gonzaga 2001; Rocha et al. 2005;
Vecchi & Alves 2008), while F. s . i n t e r p o s i t a continued to be considered a subspecies of F. s e r r a n a .
In this study, the taxonomy of the Formicivora serrana complex is reviewed. We test the validity of F. l i t t o r a -
lis and consider the best taxonomic treatment for F. s . i n t e r p o s i t a .
Material and methods
Morphological analysis. A total of 146 specimens (99 males and 47 females) was examined (Appendix 1). Col-
lections consulted were the Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ), Museu de Zoologia
da Universidade de São Paulo (MZUSP), Laboratório de Bioacústica, Universidade Federal do Rio de Janeiro
(UFRJ), Departamento de Zoologia da Universidade Federal de Minas Gerais (DZUFMG), Museu de Biologia
Professor Mello Leitão (MBML) and Museu de Zoologia João Moojen de Oliveira (MZUFV). We examined the
holotypes of F. serrana (housed at the Zoological Museum, Copenhagen) and F. d e l u z a e Ménétriés, 1835 (housed
at the Russian Academy of Science) and consulted the original descriptions of all relevant names (F. n i g r i c o l l is
Ménétriés, 1835; F. m e l a n o g a s t r a Reinhardt, 1870; F. r u f a t r a Cabanis 1874 and F. g r i s ea Ihering & Ihering 1907).
The analyses of skins focused on plumage color (using Smithe 1975 as a standard for color descriptions),
plumage patterns, and morphometric characters (length of wing and tail, exposed culmen, bill width at the nares,
and bill height at the base). The software STATISTICA (Statsoft 2001) was used for the descriptive analyses, tests
of normality (Kolmogorov-Smirmov tests), analyses of variance (one-way ANOVAs), and discriminant-function
analyses. Statistical significance was accepted at p< 0.05.
Adults were identified based largely on skull ossification. Specimens with completely ossified skulls were con-
sidered adults and those with partially or non-ossified skulls were considered juveniles.
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VARIATION OF FORMICIVORA SERRANA & FORMICIVORA LITTORALIS
FIGURE 1. Holotype of F. s e r r a n a (Hellmayr, 1929), housed in the Copenhagen Zoology Museum, showing some of its diag-
nostic features. A. Wide white supercilium. B. Smoke grayish flanks. C. Mars Brown upperparts. (Photo: Marcos Raposo).
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FIGURE 2. Distribution and some diagnostic features (supercilium, wing coverts and rectrices) of the subspecies of F. s e r r a n a
(Gonzaga and Pacheco 1990). The open triangle, square and circle represent the type locality of F. s . s e r r an a , F. s . i n t e r p o s i t a
and F. s . l i t t o r a l i s , respectively. (Original plate from Gonzaga and Pacheco (1990).
Three analyses were conducted for each plumage character, two analyses for the adults and one for juveniles.
The first analysis of adults focused on all specimens with completely ossified skulls; the second excluded all spec-
imens with completely ossified skulls whose upperparts were brown where the upperparts of other specimens were
predominantly black or grey. These specimens were treated as “possible juveniles” because F. s . l i t t o r a l i s juveniles
have brown upperparts (Soneghet 1991) in contrast to the Vandyke Brown to Sepia upperparts of adults (Gonzaga
& Pacheco 1990). The analyses of the juveniles focused only on specimens with partially or non-ossified skulls.
Vo ca l a na ly si s. Vocalizations were obtained from the researcher’s personal files (Carlos Rodrigo Meireles de
Abreu (CRMA), José Eduardo Simon (JES), Faunativa (FN), Jeremy Minns (JM), Arquivo Sonoro Elias Coelho
(ASEC/UFRJ), and from fieldwork (listed as DHF on Appendix 2). These latter vocalizations were digitized using
a sampling rate of 44.1 kHz and 16 bits per sample in the mono pattern. The sound analyses were made using the
programs Syrinx 2.6hs and Soundruler 0.9.6.0. Statistical analysis of vocalizations were conducted using the soft-
ware STATISTICA (Statsoft 2001) for the descriptive analyses, tests of normality (Kolmogorov-Smirmov tests),
and analyses of variance (one-way ANOVAs). Statistical significance was accepted at p< 0.05.
The vocalizations were divided into song and calls, the latter divided into call 1 (= “alarm call” of Gonzaga
2001) and call 2 (= “contact call” of Gonzaga 2001). The characters analyzed for song were number of syllables
(defined as a series of units which occur together in a particular pattern; Catchpole & Slater 1995); duration of
phrase (measured in seconds from the beginning of the first syllable to the end of the last syllable); pace of the
phrase (calculated as the number of syllables of the phrase divided by its duration); duration and frequency of the
fundamental harmonic (kHz) of the second, central and last syllables; duration of the interval between these sylla-
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VARIATION OF FORMICIVORA SERRANA & FORMICIVORA LITTORALIS
bles and the previous ones; structure of the syllables and elements; and presence of brief pauses in song phrases
(Tobias and Williams 1996). The analysis of call 1 focused on the number of syllables, duration and pace of the
phrases, duration and frequency of the fundamental harmonic, and structure of the central syllable. The analysis of
call 2 focused on the duration and frequency of the fundamental harmonic of the central syllable. Decisions regard-
ing diagnosability of characters followed the methodology of Isler et al. (1998); that is, continuous variables, such
as duration of the phrases, could not overlap, and discrete variables, such as syllable structure, must be used
unequivocally to identify every spectogram to species.
Results
Names related to F. serrana. The examination of the type specimens and the original descriptions of F. d e l u z a e
Ménétriés, 1835; F. nigricollis Ménétriés, 1835; F. melanogastra Reinhardt, 1870; F. r u f a t r a Cabanis, 1874; F. g r i -
sea Ihering and Ihering, 1907 and other relevant names did not reveal any senior synonym available for F. serrana.
Plumage (males). The character states for plumage of the males are described in Table 1. Given the lack of
significant differences among analyses, only the results of the first analysis (of adults) are discussed here.
White supercilium. Six character states were observed (Figure 4). Adult specimens of F. s. serrana (Figure 5)
presented only the “large and uninterrupted” white supercilium. This character state was also found in some speci-
mens of F. s . i n t e r p o si t a , which presented five states of supercilium varying from “large and uninterrupted” to
“post ocular white macula” (Figure 6). Formicivora s. littoralis presented four patterns of supercilium: “thin and
interrupted”; “thin and interrupted with only a few white feathers”; “post ocular white macula”, which was also
found in specimens of F. s . i n t e r p o s i t a ; and “post ocular thin and uninterrupted” white supercilium, the only state
restricted to specimens of F. s . l i t t o r a l i s . The original description of F. s . l i t t o r a l i s (Gonzaga and Pacheco, 1990) did
not mention the presence of white feathers at the superciliary area.
Upperparts. Seven patterns of upperparts coloration were observed (Figure 7). Upperparts of adult specimens
of F. s. serrana varied only from rufous brown to cinnamon brown (Figure 8), in contrast to those of F. s . i n t e r p o s -
ita and F. s . l i t t o r a l i s , which showed greater variation. This variation is present even in the type series of F. s . i n t e r -
posita and F. s . l i t t o r a l i s (Figure 9), in which the same upperparts colors can be seen in specimens of F. s .
interposita (MPEG 46315) and F. s . l i t t o r a l i s (MPEG 46319) and different colors in specimens of F. s . l i t t o r a l i s
(compare specimens MPEG 46319 with MPEG 46317 and 46318). All upperparts colors were present in F. s . i n t e r -
posita, except for gray, which was found exclusively in specimens of F. s . l i t t o r a l i s . Specimens of F. s . l i t t o r a l i s
also had the cinnamon brown upperparts found in specimens of F. s. serrana and F. s . i n t e r p o s i t a . In addition,
upperparts of some specimens of F. s . i n t e r p o s i t a were brown olive and others were black shaded with gray.
Rectrices. For the description of each observed pattern of rectrices see Table 1 and Figure 10. The adult males
of F. s. serrana showed all patterns (Figure 11), whereas F. s . i n t e r p o s i t a presented only patterns 1 and 2 and F. s .
littoralis presented only patterns 2 and 3.
Wing coverts. Four patterns of wing coverts were observed (Figure 12). Adult males of F. s. serrana and F. s .
interposita presented the same patterns (Figure 13): “large white spots”, “intermediate white spots” and “small
white spots” on the tips of the lesser and median wing coverts. F. s . l i t t o r a l i s showed the same three patterns as well
as the pattern “without white spots”, which was restricted to this taxon.
Flanks. The color of the flanks varied from white shaded with brown or gray, to dark gray (Figure 14). Adult
males of F. s. serrana presented white flanks, white flanks shaded with brown, and white flanks shaded with gray
(Figure 15). Formicivora s. interposita showed the same colors as well as dark gray flanks, only found in a few
specimens. The majority of specimens of this taxon had the white flanks shaded with gray. All four character states
were found in specimens of F. s . l i t t o r a l i s .
Tibia feathers. The three character states of tibia feathers (Figure 16) were found in adult specimens of all
taxa (Figure 17). The majority of specimens of F. s . l i t t o r a l i s had entirely black tibia feathers.
None of the plumage characters examined were useful for diagnosis of taxa, given the overlap of character
states between taxa in analyses of both adults and juveniles.
Plumage (females). Table 2 describes the characters and their respective states for female specimens. There
were only three juvenile specimens, according to skull ossification data on the labels; therefore, only adults were
analyzed.
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FIGURE 3. Holotypes of F. s . i n t e r p o s i t a (MNRJ 36750, above) and of F. s. littoralis (MNRJ 36751, below) illustrating some
of their diagnostic features. A. Upperparts. B. White supercilium, wing coverts and flanks.
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VARIATION OF FORMICIVORA SERRANA & FORMICIVORA LITTORALIS
FIGURE 4. Observed patterns of the character white supercilium. A. large and uninterrupted (F. s . s e r r a n a , MNRJ 27247). B.
thin and uninterrupted (F. s . i n t e r p o s i t a , MNRJ 44422). C. post ocular thin and uninterrupted (F. s. littoralis, MNRJ 39171). D.
thin and interrupted (MNRJ 36750—holotype of F. s . i n t e r p o s i t a ). E. thin and interrupted with only a few feathers (F. s . l i t t or a -
lis, MNRJ 36800). F. post ocular white macula (F. s . l i t t o r a l i s , MNRJ 43737).
Upperparts. The upperparts varied from dark brown to grayish horn (Figure 18). Females of F. s. serrana and
F. s . l i t t o r a l i s showed all upperparts colors, whereas those of F. s . i n t e r p o s i t a presented only dark brown and light
brown (Figure 19).
Abdomen. The observed colors for belly were faint yellow, cream yellow, and buff yellow (Figure 20). The
specimens of all three taxa presented all three belly colors (Figure 21).
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VARIATION OF FORMICIVORA SERRANA & FORMICIVORA LITTORALIS
TAB LE 2. Descriptions of characters analyzed for female specimens.
Rectrices. Female specimens presented three rectrix patterns (patterns 1, 2 and 3; see Figure 10). Specimens of
F. s. serrana had only pattern 1, F. s . i n t e r p o s i t a patterns 1 and 2, and F. s . l i t t o r a l i s all three patterns (Figure 22).
Wing coverts. None of the female specimens presented the pattern “without white spots” on lesser and
median wing coverts (Figure 23). Specimens of F. s . s e r r a n a had “large white spots” and “intermediate white
spots”, whereas F. s . i n t e r p o s i t a and F. s . l i t t o r a l i s showed these patterns as well as “small white spots” (Figure 24).
As with the males, it was not possible to diagnose these taxa based on analysis of specimens of females (see
also Gonzaga and Pacheco 1990, Ridgely and Tudor (1994).
Vo ca l an al ys is . For descriptions of the vocal characters, see Appendix 2. Both normality and homoscedasticity
tests allowed the use of parametric tests for song and call analyses.
Song. No structural differences in either syllables or elements were found among the three taxa (Figures 25–
27).
Formicivora s. serrana had the longest phrase duration (mean 3.53 seconds), followed by F. s . i n t e r p o s it a
(mean 3.20 seconds) and F. s . l i t t o r a l i s (mean 2.73 seconds), but no significant difference between these taxa was
observed (F2,31=1.7022, p=0.198).
The number of syllables ranged from 2 to 51 and there were no significant differences among the taxa
(F2,31=0.2518, p=0.778). The mean value for the number of syllables was somewhat lower than that found by Gon-
zaga (2001) for F. s . l i t t o r a l i s (15.9 vs. 16.6 syllables). No comparison was made between this study and Gonzaga
(2001) for F. s. serrana and F. s . i n t e r p o s i t a because they were analyzed together as F. serrana by Gonzaga (2001).
According to Gonzaga (2001), the pace of F. s . l i t t o r a l i s phrases is faster than that of F. s. serrana phrases, a
statement corroborated by our results. The pace (F2,29=14.677, p<0.000) of phrases of F. s . l i t t o r a l i s was signifi-
cantly faster than that of phrases of F. s. serrana (p<0.000) and F. s . i n t e r p o s i t a (p=0.001), but no significant differ-
ence was observed between F. s . s e r r a n a and F. s . i n t e r p o s i t a (p=0.481). The faster pace of F. s . l i t t or a l i s relative to
F. s . s e r r a n a and F. s . i n t e r p o s i t a is confirmed by the duration and number of syllables in F. s . l i t t o r a l i s (see Table
3).
Character Character state
F. s . s e r r a n a F. s . i n t e r p o s i t a F. s . l i t t o r a l i s
Upperparts Dark brown (color 121 A);
light brown (color 37); gray-
ish horn (color 91)
Dark brown (color 121 A); light
brown (color 37)
Dark brown (color 121 A); light
brown (color 37); grayish horn
(color 91)
Belly Faint yellow (color 92);
cream yellow (color 54); buff
yellow (color 124)
Faint yellow (color 92); cream yellow
(color 54); buff yellow (color 124)
Faint yellow (color 92); cream yel-
low (color 54); buff yellow (color
124)
Rectrices Tip and outer vane of 6th and
5th pairs in white and 4th pair
only with tip in white (pattern
1)
Tip and outer vane of 6th and 5th pairs
in white and 4th pair only with tip in
white (pattern 1); tip and outer vane of
6th pair in white, 5th with edge and tip
with white and 4th pair only with tip in
white (pattern 2);
Tip and outer vane of 6th and 5 th pairs
in white and 4th pair only with tip in
white (pattern 1); tip and outer vane
of 6th pair in white, 5th with edge and
tip with white and 4th pair only with
tip in white (pattern 2); tip and outer
vane of 6th pair in white, 5th and 4th
pair only with tip in white (pattern 3)
Wing coverts Tips of lesser and median
wing coverts with great white
spots (large white spots); tips
of lesser and median wing
coverts with intermediate
white spots (intermediate
white spots)
Tips of lesser and median wing
coverts with great white spots (large
white spots); tips of lesser and median
wing coverts with intermediate white
spots (intermediate white spots); tips
of lesser and median wing coverts
with small white spots (small white
spots)
Tips of lesser and median wing
coverts with great white spots (large
white spots); tips of lesser and
median wing coverts with intermedi-
ate white spots (intermediate white
spots); tips of lesser and median
wing coverts with small white spots
(small white spots)
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FIGURE 5. Map showing the distribution of the patterns of the white supercilium character in adult males. ES, MG and RJ
mean, respectively, Espírito Santo, Minas Gerais and Rio de Janeiro states.
The duration of the interval between the first two syllables (F2,31=6.2426, p=0.005) was significantly slower in
F. s. littoralis when compared with F. s . s e r r a n a (p=0.013) and F. s . i n t e r p o s i t a (p=0.020) but there was no signifi-
cant difference between F. s. serrana and F. s . i n t e r p os i t a (p=0.993). The results of the interval duration between
the central syllables (F2,31=6.8440, p=0.003) was similar to that of the first two syllables, where the duration of this
interval of F. s . l i t t or a l i s was significantly slower relative to F. s . s e r r a na (p=0.005) and F. s . i n t e r p o s it a (p=0.029);
however, no significant difference was found between the latter two taxa (p=0.825). No significant difference was
found among the three taxa in the interval duration of the last two syllables (F2,31=2.4794, p=0.100).
There was no significant difference among the taxa in the duration of the second syllable (F2,29=2.7808,
p=0.078). A significant difference was observed only between F. s. serrana and F. s . l i t t o r a l i s in duration of the
central syllable (p=0.014) and last syllable (p=0.023). The duration of the syllables (second, central and last) was
longest in F. s . l i t t o r a l i s and shortest in F. s. serrana (see Table 3).
The fundamental harmonic frequency (independent of the syllable) ranged from 1.67 to 2.71 kHz. No signifi-
cant difference was observed among the taxa in the fundamental harmonic frequency of the second element of the
second syllable (F2,30=2.2515, p=0.122), the central syllable (F2,31=0.1088, p=0.897), or the last syllable
(F2,31=0.6315, p=0.538).
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VARIATION OF FORMICIVORA SERRANA & FORMICIVORA LITTORALIS
TABLE 3. Descriptive statistics for song characters analyzed: mean, standard deviation (SD), minimum value (Min), maximum value (Max), and number of observations (n).
F. s. serrana F. s. interposita F. s. littoralis
Characters Mean SD Min Max n Mean SD Min Max n Mean SD Min Max n
Duration of the phrase (sec) 3.538 1.954 0.270 8.101 67 3.208 2.067 0.553 9.874 51 2.734 1.909 0.355 8.794 105
Nº of syllables of the phrase 17.373 9.382 2 39 67 17.098 10.870 4 50 51 15.952 10.497 3 51 105
Pace of the phrase (sec) 5.019 0.633 3.857 7.407 67 5.400 0.519 4.550 7.226 51 6.097 0.776 4.705 8.792 105
Interval duration between
the 1st two syllables (sec)
0.171 0.028 0.132 0.275 64 0.164 0.022 0.130 0.237 48 0.143 0.017 0.097 0.219 93
Duration of the 2nd syllable
(sec)
0.063 0.062 0.045 0.055 64 0.051 0.004 0.042 0.060 48 0.048 0.006 0.024 0.060 93
Fundamental harmonic
frequency of the 2nd
element of the 2nd syllable
(kHz)
2.100 0.189 1.679 2.454 62 2.173 0.190 1.765 2.627 46 2.099 0.156 1.765 2.713 89
Interval duration between
the central syllables (sec)
0.157 0.028 0.114 0.275 64 0.146 0.021 0.114 0.210 50 0.126 0.017 0.099 0.201 93
Duration of the central
syllable (sec)
0.066 0.062 0.048 0.055 64 0.054 0.004 0.041 0.063 50 0.051 0.005 0.028 0.060 93
Fundamental harmonic
frequency of the 2nd
element of the central
syllable (kHz)
2.174 0.169 1.765 2.454 64 2.165 0.126 1.851 2.454 49 2.166 0.169 1.765 2.713 91
Interval duration between
the last two syllables (sec)
0.171 0.044 0.115 0.333 64 0.150 0.022 0.106 0.243 50 0.139 0.032 0.096 0.316 93
Duration of the last syllable
(sec)
0.058 0.004 0.049 0.066 64 0.054 0.004 0.040 0.064 50 0.052 0.006 0.031 0.063 92
Fundamental harmonic
frequency of the 2nd
element of the last syllable
(kHz)
2.160 0.188 1.679 2.454 63 2.185 0.171 1.765 2.627 49 2.177 0.165 1.679 2.713 92
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FIGURE 6. Patterns of the white supercilium character found on specimens of F. s . i n t e r p o s i t a . From above to below: large
and uninterrupted (MNRJ 16979), thin and uninterrupted (MNRJ 44422), thin and interrupted (MNRJ 36750 - holotype) and
post ocular white macula (MNRJ 44657).
FIGURE 7. Observed patterns of the upperparts character. From the left to the right: rufous brown (F. s. serrana, MNRJ
38595), cinnamon brown (F. s. serrana, MNRJ 44073), olive brown (F. s . i n t e r p o s i t a , MNRJ 44438), black shaded with olive
brown (F. s . i nt e r p o s i t a, MNRJ 44610), black (F. s . i n t e r p o s i t a , MNRJ 44424), black shaded with gray (F. s . i n t e r p o s i t a , MNRJ
44439) and gray (F. s . l i t t o r a l i s , MNRJ 39172).
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FIGURE 8. Map showing the distribution of the patterns of the upperparts character in adult males.
FIGURE 9. Paratypes of F. s . i n t e r p o s i t a and F. s . l i t t o r a l i s showing the color variation of the upperparts between these taxa.
From above to below: F. s . l i t t o r a l i s (MPEG 46319), F. s . i n t e r p o s i t a (MPEG 46315), F. s. littoralis (MPEG 46317) and F. s.lit-
toralis (MPEG 46318). Note the same upperparts color for F. s. littoralis (MPEG 46319) and F. s . i n t e r p o s i t a (MPEG 46315)
and different upperparts colors for the two specimens of F. s. littoralis (MPEG 46319) and (MPEG 46317).
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FIGURE 10. Observed patterns of the rectrices character. From left to right: pattern 1 (F. s . s e r r a n a , MNRJ 38592), pattern 2
(F. s . i n t e r p o s i t a , MNRJ 44387), pattern 3 (F. s . s e r r a n a , MNRJ 44413), pattern 4 (F. s . l it t o r a l i s , MNRJ 37438) and pattern 5
(F. s . l i t t o r a l i s , MNRJ 39172). (Plate by Jorge Nacinovic).
FIGURE 11. Map showing the distribution of the patterns of the rectrices character in adult males.
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FIGURE 12. Observed patterns of the wing coverts character. From above to below: large white spots (F. s . s e r r a n a , MNRJ
44417), intermediate white spots (F. s . i n t e r p o s i t a , MNRJ 44657), small white spots (F. s . s e r r a n a , MNRJ 44613) and without
white spots (F. s. littoralis, MNRJ 37538).
FIGURE 13. Map showing the distribution of the patterns of the wing coverts character in adult males.
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FIGURE 14. Observed patterns of the flanks character. From left to right: white flanks (F. s. serrana, MNRJ 27249), flanks
white shaded with brown (F. s . s e r r a n a , MNRJ 44073), flanks white shaded with gray (F. s . i n t e r p o s i t a , MNRJ 44423) and dark
gray flanks (F. s . l i t t o r a l i s , MNRJ 37166).
FIGURE 15. Map showing the distribution of the patterns of the flanks character in adult males.
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FIGURE 16. Observed patterns of the tibia feathers character. From left to right: black feathers with white tips (F. s. serrana,
MNRJ 27247), black feathers with some sparse white feathers (F. s. serrana, MNRJ 38595) and entirely black feathers (F. s . l i t -
toralis, MNRJ 36801).
FIGURE 17. Map showing the distribution of the patterns of the tibia feathers character in adult males.
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FIGURE 18. Observed patterns of the upperparts character. From left to right: dark brown (F. s. serrana, MNRJ 38594), light
brown (F. s. serrana, MNRJ 27278) and grayish horn (F. s . l i t t o r a l i s , MNRJ 39173).
FIGURE 19. Map showing the distribution of the patterns of the upperparts character in adult females.
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FIGURE 20. Observed patterns of the abdomen character. From left to right: faint yellow (F. s . li t t o r a l i s, MNRJ 39173), cream
yellow (F. s. littoralis, MNRJ 37054) and buff yellow (F. s. littoralis, MNRJ 37408).
FIGURE 21. Map showing the distribution of the patterns of the abdomen character in adult females.
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FIGURE 22. Map showing the distribution of the patterns of the rectrices character in adult females.
FIGURE 23. Observed patterns of the wing coverts character in adult females. From above to below: large white spots (F. s .
littoralis, MNRJ 37474), intermediate white spots (F. s. serrana, MNRJ 27278) and small white spots (F. s . i n t e r p o s i t a , MNRJ
16977).
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FIGURE 24. Map showing the distribution of the patterns of the wing coverts character in adult females.
FIGURE 25. Song phrase of F. s . s e r r a n a (LPG 102 08). A. Oscillogram of a song phrase. B. Detail of the structure of a song
syllable composed of two distinct elements. C. Sonogram of a song phrase. D. Sonogram of a song syllable.
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FIGURE 26. Song phrase of F. s . i n t e r p o s i t a (LPG D 15ab GDAC). A. Oscillogram of a song phrase. B. Detail of the structure
of a song syllable composed of two distinct elements. C. Sonogram of a song phrase. D. Sonogram of a song syllable.
FIGURE 27. Song phrase of F. s. littoralis (LPG 096 04). A. Oscillogram of a song phrase. B. Detail of the structure of a song
syllable composed of two distinct elements. C. Sonogram of a song phrase. D. Sonogram of a song syllable.
The pauses in song phrases (Figure 28), described by Tobias and Williams (1996) as “brief pauses as if a single
note [syllable] is missing”, were found in 19.4%, 5.8%, and 34.2% of F. s. serrana, F. s . i n t e r p o s i t a and F. s . l i t t o -
ralis phrases, respectively. This result corroborates Tobias and Williams’ (1996) observation that these pauses are
more frequent in F. s . l i t t o r a l i s than in F. s. serrana.
Despite several significant differences, the overlap between the minimum and maximum values in all continu-
ous variables, as well as the absence of a structural difference in syllables among the three taxa, did not allow a
vocal diagnosis of these taxa.
The intermediate position occupied by F. s . i n t e r p o s i t a in all vocal characters related to duration is remarkable
(see mean values in Table 3). This could be interpreted as clinal variation in song similar to that noted for the plum-
age characters.
Call 1. No significant difference was observed in phrase duration among the taxa (F2,21=0.0093, p=0.990).
The number of the syllables ranged from 2 to 15 (Table 4). The mean value of F. s . l i t t o r a l i s was the highest
(6.24 syllables), followed by F. s . i n t e r p o s i t a (4.66 syllables) and F. s. serrana (4.15 syllables). As indicated by
Gonzaga (2001), in longer phrases, the syllables could present a variation characterized by “an abrupt initial attack
of rapidly descendant modulations (Figures 30D and 31D) but not ascendant (Figure 29D) with a frequency and
tone a little more high-pitched”. This variation was found in all three taxa, corroborating Gonzaga (2001), but it
was not useful for distinguishing one from another.
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FIGURE 28. Sonograms showing the pauses in song phrases. A. Sonogram of a phrase of F. s . s e r r a n a (LPG 102 08). The
pause is between 0.5 and 1 second. B. Sonogram of a phrase of F. s . i n t e r p o s i t a (LPG 021 08) illustrating the pause between 0
and 1 second. C. Sonogram of a phrase of F. s. littoralis (LPG c 015 24). Notice the pause between 0.5 and 1 second.
TAB LE 4. Descriptive statistics for charcters analyzed for Call 1: mean, standard deviation (SD), minimum value (Min), max-
imum value (Max), number of observations (n).
F. s. serrana F. s. interposita
Characters Mean SD Min Max n Mean SD Min Max n
Duration of the phrase (sec) 1.294 0.695 0.300 3.178 20 1.310 0.410 0.397 1.870 24
Nº of syllables of the phrase 4.150 1.182 2 6 20 4.666 1.493 2 8 24
Pace of the phrase (sec) 3.452 0.788 1.887 4.787 19 3.634 0.592 2.886 5.031 24
Duration of the central syllable (sec) 0.121 0.020 0.090 0.159 16 0.116 0.015 0.087 0.156 23
Fundamental harmonic frequency of
the central syllable (kHz)
2.375 0.384 1.851 3.402 16 2.025 0.270 1.593 2.497 16
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24 · Zootaxa 2742 © 2011 Magnolia Press
continued.
The pace of the phrases was significantly different (F2,21=4.7385, p=0.020) only between F. s . l i t t o r a li s and F. s .
interposita (p=0.044). The pace of the phrases of F. s . s e r r a n a was the slowest (3.45 seconds), followed by F. s .
interposita (3.63 seconds) and F. s . l i t t o r a l i s (5.33 seconds) (Table 4).
The duration of the central syllable varied from 0.07 to 0.15 seconds (Table 4). The central syllables of F. s .
serrana were the longest ones whereas those of F. s . l i t t o ra l i s were shortest (see Table 4). A significant difference
(F2,19=5.5650, p=0.012) was observed in this character between F. s. serrana and F. s . l i t t o r a l i s (p=0.020).
The fundamental harmonic frequency of the central syllable ranged from 1.85 to 3.40 kHz (Table 4) and a sig-
nificant difference (F2,18=7.0462, p=0.005) was evident only between F. s. serrana and F. s . l i t t o r a l i s (p=0.005).
As in the analyses of song, overlaps between the minimum and maximum values in all continuous variables
(Table 4) and a lack of structural differences among the taxa, did not allow a diagnosis of the characters of Call 1.
Call 2. The duration of the central syllable ranged from 0.085 to 0.181 seconds (Table 5) and did not differ sig-
nificantly among the taxa (F2,22=0.0307, p=0.969). Some degree of variation in the structure of this syllable was
observed (Figures 32–34) as indicated by Gonzaga (2001). This variation is a consequence of the change in modu-
lation of the syllables and it was not related to any specific taxon.
TAB LE 5. Descriptive statistics for characters analyzed for Call 2: mean, standard deviation (SD), minimum value (Min), max-
imum value (Max) and number of observations (n).
continued.
The fundamental harmonic frequency of the central syllable ranged from 3.057 to 5.383 kHz (Table 5). The
mean value of this character was 4.379 kHz in F. s. serrana, 4.306 kHz in F. s . i n t e r p o s i t a , and 4.183 kHz in F. s .
littoralis. No significant difference was observed between the taxa for this character (F2.21=2.9724, p=0.073).
The characters of Call 2 were not diagnosable among taxa due the lack of structural difference and the overlap of
values in continuous variables.
Morphometrics. Results of the descriptive analyses are available in Table 6. The normality test and the homo-
scedasticity test allowed the use of parametric tests in our analyses for both sexes.
Formicivora s. littoralis was significantly larger than F. s . i n t e r p o s i ta and F. s. serrana in all characters except
tail length (Tables 6, 7). F. s . i n t e r p o s i t a was intermediate in bill length and bill width, and F. s . s e r r a n a in bill
F. s . l i t t o r a l i s
Characters Mean SD Min Max n
Duration of the phrase (sec) 1.230 0.645 0.456 2.692 33
Nº of syllables of the phrase 6.242 2.872 3 15 33
Pace of the phrase (sec) 5.335 1.021 3.319 7.554 33
Duration of the central syllable (sec) 0.097 0.017 0.072 0.133 33
Fundamental harmonic frequency of the central
syllable (kHz)
2.432 0.492 1.593 3.402 31
F. s. serrana F. s. interposita
Characters Mean SD Min Max n Mean SD Min Max n
Duration of the central syllable (sec) 0.124 0.018 0.100 0.181 59 0.124 0.007 0.111 0.142 60
Fundamental harmonic frequency of
the central syllable (kHz)
4.379 0.327 3.919 5.383 44 4.306 0.238 3.746 4.866 59
F. s. littoralis
Characters Mean SD Min Max n
Duration of the central syllable (sec) 0.131 0.013 0.085 0.148 98
Fundamental harmonic frequency of
the central syllable (kHz)
4.183 0.501 3.057 5.211 97
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height, wing length and tail length. These results corroborate the morphometric data described by Gonzaga and
Pacheco (1990). No significant difference was found between F. s. serrana and F. s . i n t e r p o s i t a , except in bill
length, which was significantly longer in F. s . i n t e r p o s i t a (p=0.044). Despite the significant differences between
taxa, overlaps in values make it impossible to use these as diagnostic characters.
TAB LE 6. Mensural data of males and females of Formicivora s. serrana, F. s . i n t e r p o s i t a and F. s . l i t t o r a l i s : male specimens
(M), female specimens (F), mean, standard deviation (SD), minimum value (Min), maximum value (Max) and number of spec-
imens examined (n).
continued.
TAB LE 7. Results of ANOVA for morphometric comparison between F. s e r r a n a , F. s . i n te r p o s i t a and F. s. littoralis. Values in
bold indicate significant differences (p<0.05). Male specimens (M), female specimens (F).
Bill length (mm) Bill height (mm) Bill width (mm)
Mean SD Min Max n Mean SD Min Max n Mean SD Min Max n
F. s . s e r r a n a
M15.070.9012.1316.39383.930.283.374.70374.000.303.084.6138
F14.020.8213.7016.17153.750.253.194.21154.040.273.664.5915
F. s . i n t e r p o s i t a
M15.690.6214.3116.62213.820.253.504.36214.060.293.394.5121
F14.730.8214.0516.7693.650.163.423.8873.840.353.394.529
F. s . l i t t o r a l i s
M16.390.7814.5617.72294.120.303.615.09304.340.313.734.8430
F16.440.5615.2317.07144.030.183.724.37154.240.393.214.7115
Wing length (m m) Tail length ( mm)
Mean SD Min Max n Mean SD Min Max n
F. s . s e r r a n a
M52.911.9050.2058.704155.411.9352.0059.0033
F51.921.4349.9355.401654.582.7451.0059.0013
F. s . i n t e r p o s i t a
M51.771.1449.8653.941254.001.9152.0057.0012
F49.251.8146.1252.10851.292.3148.0054.007
F. s . l i t t o r a l i s
M56.331.5453.2259.062655.792.2951.5061.0026
F53.671.3051.0956.751454.572.0451.0058.0014
F. s. serrana F. s. interposita
MFM F
F. s . i n t e r p o s i t a Bill length p=0.044 p=0.920
Bill height p=0.419 p=0.608
Bill width p=0.758 p=0.357
Wing length p=0.128 p=0.001
Tai l le ngth p= 0.1 36 p =0.0 36
F. s . l i t t o r a l i s Bill length p=0.000 p<0.001 p=0.013 p<0.001
Bill height p=0.016 p=0.003 p=0.001 p=0.001
Bill width p<0.001 p=0.277 p=0.005 p=0.026
Wing length p=0.000 p=0.005 p=0.000 p=0.000
Tai l le ngth p= 0.7 65 p =0. 963 p =0. 050 p=0. 022
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FIGURE 29. Call 1 phrase of F. s . s e r r a n a (LPG 102 12e). A. Oscillogram of a call 1 phrase. B. Detail of the structure of a call
1 syllable. C. Sonogram of a call 1 phrase. D. Detail of a call 1 syllable. Notice the rapid ascendant modulation in the beginning
of the syllable followed by a long descendant modulation.
FIGURE 30. Call 1 phrase of F. s . i n t e r p o s i t a (LPG 021 08). A. Oscillogram of a call 1 phrase. B. Detail of the structure of a
call 1 syllable. C. Sonogram of a call 1 phrase. D. Detail of a call 1 syllable. Notice the rapid abrupt descendant modulation in
the beginning of the syllable followed by a smoother descendant modulation.
The Principal Components Analysis (PCA) for males indicated that Factor 1 and Factor 2 were responsible for
68.0% of the variation. The greatest contributing characters for Factor 1 (46.0%) and Factor 2 (22.0%) were,
respectively, bill length and tail length. The measurements of F. s . i n t e r p o s i t a fell within the variation of F. s . s e r -
rana, as did those of some specimens of F. s . l i t t o r a l i s (Figure 35). The discriminant function analysis did not sig-
nificantly identify F. s. serrana (93.8%), F. s . i n t e r p o s i t a (22.2%), or F. s . l i t t o r a l i s (89.4%).
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FIGURE 31. Call 1 phrase of F. s. littoralis (LPG 021 07c). A. Oscillogram of a call 1 phrase. B. Detail of the structure of a
call 1 syllable. C. Sonogram of a call 1 phrase. D. Detail of a call 1 syllable. Notice the rapid abrupt descendant modulation in
the beginning of the syllable followed by a smoother descendant modulation.
FIGURE 32. Contact call syllables of F. s . s e r r a n a (DHF 07) showing the variation in its structure. A. Sonogram of a syllable.
B. Sonogram illustrating the variation in syllable structure.
Females showed significant differences among taxa in most morphometric characters (Table 7). Females of F.
s. littoralis were largest, F. s . i n t e r p o s i t a intermediate, and F. s. serrana smallest (Table 6). Again, however, there
was overlap in values among taxa.
The PCA for female specimens indicated that Factor 1 and Factor 2 were responsible for 70.9% of the varia-
tion. The greatest contributing characters for Factor 1 (47.7%) and Factor 2 (23.2%) were, respectively, wing
length and bill length. Overlap in measurements of F. s . i n t er p o s i t a , F. s . l i t t o r a l i s , and F. s. serrana make it impos-
sible to distinguish these taxa (Figure 36).
The discriminant function analysis did not significantly identify F. s. serrana (92.3%), F. s . i n t e r p o s i t a (80%)
and F. s. l i t t o r a li s (83.3%).
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FIGURE 33. Contact call syllables of F. s . i n t e r p o s i t a (Inter 07) showing the variation in its structure. A. Sonogram of a sylla-
ble. B. Sonogram illustrating the variation in syllable structure.
FIGURE 34. Contact call syllables of F. s . l i t t o r a l i s (LPG 023 04) showing the variation in its structure. A. Sonogram of a syl-
lable. B. Sonogram illustrating the variation in syllable structure.
Discussion
Taxo no my. Neither F. s . i n t e r p o s i t a nor F. s . l i t t o r a l i s proved to be a valid taxon under the phylogenetic species
concept (PSC) (sensu Nelson and Platnick 1981), due to the lack of diagnostic characters in relation to F. serrana,
which has nomenclatural priority over these two taxa. Under the PSC, which we advocate, interposita and littoralis
must be treated as synonyms of F. serrana (Hellmayr, 1929).
Under the biological species concept (BSC), these populations would likewise be considered a single species.
They lack diagnostic features and are probably not reproductively isolated. Thus, the treatment of the three popula-
tions would likely be the same as that proposed by Gonzaga and Pacheco (1990): that is, subspecies of F. serrana.
Formicivora s. serrana, F. s . i n t e r p os i t a and F. s . l i t t o r a l i s would correspond to the mountain population, Vale do
Paraiba population, and littoral population, respectively, of F. serrana. Under the evolutionary species concept
(Wiley 1981), each population could be considered a valid species, depending on the author’s interpretation of the
intermediate characters. The statistical differences between the populations could be considered an indication of
unique evolutionary trajectories.
Morphometrics. The morphometric data demonstrated geographic variation in both males and females, but
not all characters exhibited the same pattern of variation. In females, all characters presented mosaic variation
(sensu Futuyma 1992). Lowest values were found in the central, or Vale do Paraiba population, with higher values
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FIGURE 35. Principal Component Analysis (PCA) for male specimens. The blue, red and green ellipses represent, respec-
tively, the groups of F. s . s e r r a n a , F. s . i n t e r p o s i t a and F. s . l i t t o r a l i s .
FIGURE 36. Principal Component Analysis (PCA) for female specimens.
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in the littoral population (with the highest mean values except for tail length) and in the mountain population
(which was morphometrically intermediate in all characters except tail length; see Table 6). Among males, bill
height, wing length and tail length presented mosaic variation similar to that of females, increasing from the center,
represented by the Vale do Paraíba population, to the extremes represented by the littoral population, with the high-
est mean values, and mountain population, the morphometric intermediate. The remaining characters (bill length
and bill width) showed clinal variation (sensu Endler 1977) in males, characterized by an increase of size from the
mountains to the littoral population.
Although there were significant differences in morphometric characters among populations, they were not
diagnostic (see Table 6).
Plumage. In plumage coloration, F. serrana is a polymorphic species, varying geographically as a north-south
cline. This species shows both individual and inter-populational variation. Examples of this variation include the
reduction of the white coloration in the supercilium (Figure 5), the change in upperparts coloration from brown to
black to grey (Figure 8), and a reduction of white in the rectrices (Figure 11). The remaining characters (wing
coverts, flanks and tibia feathers) varied in the same direction but in a smoother way than the previous characters,
which presented a more abrupt and well marked morphological break. This variation could be interpreted as a step
cline pattern.
The population of Vale do Paraíba was the most polymorphic of the three populations, which is best seen in the
variation in the supercilium (Figures 5 and 6) and upperparts (see Figure 8). A similar polymorphism in an interme-
diate population was observed by Miller and McCabe (1935) for Passerella lincolnii, which included character
states found in specimens of both extreme populations.
These morphological differences, more noteworthy in the littoral population of F. serrana, can also be related
to the probable lack of gene flow between this population and the mountain and Vale do Paraíba populations, due to
its relative geographic isolation. Recently, Vecchi and Alves (2008) extended the range of the littoral population to
the north bank of the Araruama lake in the municipality of Iguaba Grande, representing the most “interior” record
for this population. These authors also pointed out the need to confirm the existence of the geographic gap between
the populations of the littoral and Vale do Paraíba, as noted by Gonzaga and Pacheco (1990). In spite of this range
extension, the existence of this gap was supported by our fieldwork in the municipalities between these two popu-
lations (Duas Barras, Bom Jardim, Nova Friburgo, Teresópolis, Macaé, Casimiro de Abreu, Silva Jardim, Cachoe-
ira de Macacu and Guapimirim), where no individuals were recorded.
Vo ca li za ti on s. The results of our vocal analyses demonstrated the similarity of the vocalizations of the three
populations, as noted by Gonzaga and Pacheco (1990). This was reinforced by playback, in which vocalizations
(song and calls) of each population elicited response from the other populations of F. serrana (Firme and Raposo,
unpubl. data). This similarity is reflected in the lack of diagnostic characters in F. serrana. In spite of the lack of
diagnostic vocal characters, the results of the analyses (Tables 3–5) suggest a cline similar to that found for the
plumage.
Conservation. Taxonomic st udies ar e cr ucial for c onse rvational efforts (Z ink & Mc Ki tri ck 1 995; Zi nk et al.
2000; Raposo 2001; Valdecasas & Camacho 2003; Agapow et al. 2004; Mace 2004; Winker et al. 2007). Our
results have demonstrated that the littoral population of F. serrana, formerly considered a threatened species (Col-
lar et al. 1992, Tobias & Williams 1996; Sick 1997; IBAMA 2003; Zimmer & Isler 2003; BirdLife International
2004; Rocha et al. 2005; Marini & Garcia 2005; Vechhi & Alves 2008), is not a valid species. Future conservation
and behavioral studies focusing on the littoral populations of F. serrana must be mindful of its relationship to the
other populations of this species.
Acknowledgements
We are grateful to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for thei r st ude nt-
ship support of Daniel H. Firme (Bolsa de Mestrado 134848/2006-3) and The Instituto Brasileiro de Meio Ambi-
ente e Recursos Naturais Renováveis (IBAMA) who provided collection permits for the authors. To Neotropical
Ornithological Society (NOS) for François Vuilleumier fund for research on neotropical Birds. Marcos A. Raposo
research is also supported by CNPq (pq 301350/2008-6). Dante Teixeira and Renato Gaban for their comments on
this work. To Aline Abreu for collecting permission in RPPN do Caraça, Minas Gerais. Jorge Nacinovic kindly
Zootaxa 2742 © 2011 Magnolia Press · 31
VARIATION OF FORMICIVORA SERRANA & FORMICIVORA LITTORALIS
provided the plate of the rectrices and reviewed the abstract. To Luís. F. Silveira (MZUSP), Luíz P. Gonzaga
(UFRJ), Marcos Rodrigues (DZUFMG), Leonardo Lopes (DZUFMG), Marcelo F. Vasconcelos (DZUFMG),
Rômulo Ribon (UFV) and Gustavo M. Prado (MBML) for their assistance in the ornithological collection under
their care. To Jeremy Minns, Ricardo Parrini, José E. Simon, Ana C. Venturini, Pedro R. da Paz and Luíz P. Gon-
zaga and Arquivo Sonoro Elias Coelho (ASEC) for providing vocalizations. We also thank Renata Stopiglia,
Claydson P. Assis, Gustavo Silveira, Piero Ruschi, Carlos R. M. de Abreu, Lorian C. Straker, David Straker for the
English review, Teresa Straker, Maurício Vecchi, Juliana Mattos, Flávia Chaves, Guilherme R. R. Brito, Marcelo
Brito, Bentinho, Gustavo Pedersoli and Guy Kirwan for helping, somehow, to carrying out this work. We are also
indebted to Terry Chesser, Morton L. Isler and Joe Tobias for their valuable comments and suggestions.
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APPENDIX 1. Specimens examined.
F. d e l u z a e : Brazil: Rio de Janeiro: Teresópolis: F, ZISP 145082.
Formicivora s. serrana: Brazil: Espírito Santo: Baixo Guandu: M, MNRJ 16983, 16978; F, MNRJ 16990, 16984. Santa
Ter esa: M, MNR J 44 043, 263 16, 4407 3, MBML 6 980 , s/ n; F, M NRJ 385 93, 262 66, MBM L 72 68. Jat iboc a: M, M NRJ
27247, 27249; F, MNRJ 27278. Minas Gerais: Sete Lagoas: M, ZMUC 9/1056 nº 156 (holotype). São Gonçalo do Rio
Preto: M, DZUFMG 4267. Timóteo: M, MZUSP 25243, 25244, 25245, 25247. Marliéria: M, MNRJ 38595; F, MNRJ
38594, MZUSP 10385. Nova Era: M, MZUSP 25246. São Gonçalo do Rio Abaixo: M, DZUFMG 3679, 3680. Córrego
Novo: F, MZUSP 10384. Vargem Alegre: M, MZUSP 1563. Santa Bárbara: M, MNRJ 38590, DZUFMG 3624, 3955,
1047, 1048; F, DZUFMG 1049. Catas Altas: M, MNRJ 44413, 44414, 44416, 44417, 44418; F, MNRJ 44415. Rio Acima:
M, MNRJ 38592. São Pedro dos Ferros: M, MZUFV s/n; F, MZUFV s/n. Mariana: M, MNRJ 16980, 16988, 16989,
DZUFMG 4275, 4358; F, MNRJ 16985. Araponga: M, MZUFV 1412. Viçosa: M, MNRJ 38591, MZUFV 839; F,
MZUFV 823, 840.
F. s . i n t e r p o s i t a : Brazil: Minas Gerais: Muriaé: M, MNRJ 16979, 16982, 16986 (paratypes); F, MNRJ 16976, 16977, 16981,
16987 (paratypes). Volta Grande: M, MNRJ 23989 (paratype); F, MNRJ 23978, 23979 (paratypes). Rio de Janeiro: Cam-
buci: M, MNRJ 44657, 44658; F, MNRJ 44659. Carmo: M, MNRJ 44424, 44611, 44612. Sapucaia: M, MNRJ 44438,
44439, 44610, 44656. Três Rios: M, MNRJ 44440. Santa Maria Madalena: M, MNRJ 44403; F, MNRJ 44402. São
Sebastião do Alto: M, MNRJ 44422, 44613. Macuco: M, MNRJ 44387, 44423. Trajano de Morais: M, MNRJ 36750
(holotype), 44404, UFRJ 0813; F, UFRJ 0814.
F. s . l i t t o r a l i s : Brazil: Rio de Janeiro: Cabo Frio: M, MNRJ 26540 (paratype), 43737, 44394, UFRJ 0797; F, UFRJ 0811.
Arraial do Cabo: M, MNRJ 36800, 37166, 37187, 38362, 36799, 37440, 36801, 37538, 37438, 37055, 37472, 37473,
43497, 37470, 37404, 36751 (holotype), 38365, 37165, 38364, MZUSP 73506 (paratype), UFRJ 0812, 0669; F, MNRJ
37407, 37405, 37054, 37474, 39410, 38363, 37406, 37439, 37539, 37409, 37408, 37471, 43736, MZUSP 73507 (para-
type). Araruama: M, MNRJ 39172, 39171, 39170, UFRJ 0354, 0352, 0356, 0355, 0358; F, MNRJ 39173, UFRJ 0353,
0360, 0359, 0357. Saquarema: M, UFRJ 0361.
APPENDIX 2. Recordings examined. ASEC (Arquivo Sonoro Elias Coelho), CRMA (Carlos Rodrigo Meireles
de Abreu), DHF (Daniel Honorato Firme), FN (Faunativa), JES (José Eduardo Simon), JM (Jeremy Minns).
F. s. serrana: Brazil: Espírito Santo: Fundão: FN: F. serrana 01. Santa Teresa: JES: JES 01, 02, 03. Minas Gerais: Aimorés:
FN: F. serrana 02. Santa Bárbara: ASEC: LPG D 034, LPG 102 04, 102 06, 102 08, 102 012e, 102 13, 102 14, 102 15, 102
16, LPG D - 002 36, 002 37a, 002 39ac, 006 14, 062 14. Rio Doce: ASEC: 097 26. Viçosa: ASEC: 109 09. Rio de
Janeiro: Var r e - S ai : A S E C: C B E 0 0 2 1 8, 0 0 4 02 .
F. s . i n t e r p o s i t a : Brazil: Rio de Janeiro: Cambuci: ASEC: LPG D 076 15 ab GDAC. Carmo: DHF: DHF 04, 022. Sapucaia:
DHF: DHF 01. Santa Maria Madalena: DHF: DHF 01, 07, 08, 18. São Sebastião do Alto: DHF: DHF 03. Macuco: DHF:
DHF 08, 09, 10a, 10b, 12, 18. Trajano de Morais: ASEC: LPG 021 08, 021 10a; DHF: DHF 02, 15.
F. s. littoralis: Brazil: Rio de Janeiro: Cabo Frio: ASEC: LPG 021 04, 021 07c, 023 04, 021 03, 021 06, 021 05; DHF: DHF
01, 05; CRMA: CRMA 05. Arraial do Cabo: ASEC: LPG 023 05. Araruama: ASEC: LPG 095 01, 095 06, 095 10, 096 01,
096 04, 096 03, 095 12, 096 02, 096 13 ac, 096 09; JCM: JCM 042 29. Saquarema: ASEC: LPG c 015 24.
