ZOOLOGIA 35: e12993

ISSN 1984-4689 (online)

zoologia.pensoft.net

RESEARCH ARTICLE

Morphological evidence for the taxonomic status of the Bridge’s Guan,

Penelope bridgesi, with comments on the validity of P. obscura bronzina
(Aves: Cracidae)
Oscar Diego Evangelista-Vargas1, Luís Fábio Silveira1

1
Seção de Aves, Museu de Zoologia da Universidade de São Paulo, São Paulo, Brasil. Avenida Nazaré 481, Ipiranga,
04263-000 São Paulo, SP, Brazil.
Corresponding author: Diego Evangelista-Vargas (diegoevangelistav@gmail.com)
http://zoobank.org/C4A2862F-20E9-47CE-AA81-AE74C6AFEC84

ABSTRACT. Penelope obscura Temminck, 1815 is a forest guan found in Brazil, Paraguay, Uruguay, Argentina and Bolivia.
Three subspecies are currently recognized: Penelope o. obscura Temminck, 1815, P. o. bridgesi Gray, 1860, and P. o. bronzina
Hellmayr, 1914. The limits between Penelope taxa are poorly understood since few studies have evaluated their differences
in plumage, distribution and taxonomy. Based on 104 specimens deposited in ornithological collections we studied the
variations in the plumage of P. obscura, including all characters that have been used to describe the included subspecies.
Our results show that the plumage of these birds is extremely variable in southern and southeastern Brazil. Without any
morphological and morphometric characters to support P. o. bronzina as a valid taxon, we synonymized it with P. obscura.
Conversely, P. o. bridgesi, which occurs in the Yungas and the Chaco, is a distinct taxon and should be treated as a separate
species from P. obscura.
KEY WORDS. Distribution, morphology, Penelope, taxonomy.

INTRODUCTION Based on the “Yacuhú” of Azara (1805), Temminck (1815)

described P. obscura. Paraguay is considered as its type locality.
Penelope obscura Temminck, 1815 is distributed from Gray (1860) described Penelope bridgesi, type locality in Bolivia
eastern Minas Gerais and northern Espírito Santo in Brazil, (restricted to Villa Montes, Tarija by Vaurie 1966). Ogilvie-Grant
to southeastern Paraguay, northeastern Argentina (extending (1893) suggested that P. bridgesi should be considered a synonym
to central Bolivia) and Uruguay (Del Hoyo and Kirwan 2016). of P. obscura; however, Hartert and Venturi (1909) considered P.
As it is often the case among members of the genus, P. obscura bridgesi as a subspecies of P. obscura, a placement that is currently
is syntopic with other species throughout its distribution: on accepted. Finally, Hellmayr (1914) described P. obscura bronzina
the eastern slopes of the Andes of Bolivia and northwestern based on a specimen collected in Santa Catarina (Colonia Hansa,
Argentina, this species shares its habitat with P. montagnii and currently Corupá). The author also delimited the geographic dis-
P. dabbenei, and in southeastern Brazil, eastern Paraguay and tribution of the three subspecies. Hellmayr’s hypotheses (1914)
northeastern Argentina, it is usually found with P. superciliaris have been adopted up to now, and no revisions of Penelope have
(Vaurie 1968). been published since the taxonomic study of Vaurie (1968).
The body plumage of P. obscura is usually dark brown; Penelope obscura bronzina is an endemic subspecies of the
neck, mantle, upper wing coverts and breast are edged with dull Brazilian Atlantic Forest and has a wide distribution, occurring
white, as the forehead and eyebrow. The tarsus is blackish brown from east of Minas Gerais to Santa Catarina through Espírito
or black in life (Vaurie 1968). It presents sexual dimorphism in Santo, Rio de Janeiro, São Paulo and Paraná (Sick and Teixeira
the color of the iris, red to orange in males and brown or dark 1979). The nominate form is distributed in southern Brazil, in
brown in females, which is already well differentiated in the the states of Santa Catarina and Rio Grande do Sul, also including
first year of life (Sick 1997, Chalukian 1997, Nacinovic 2012). the forested regions in Paraguay, Uruguay and northeastern

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 D. Evangelista-Vargas & L.F. Silveira

Argentina (in the provinces of Misiones, Chaco, Corrientes, following institutions: Museum für Naturkunde (MFN), Berlin,
Entre Ríos and Santa Fe); the exact range of this subspecies in Germany; Field Museum of Natural History (FMNH), Chicago,
Brazil is unclear, but it is most likely restricted to the forests of USA; Rijksmuseum van Natuurlijke Historie (RNHL), Leiden,
the Paraná and Uruguay Rivers. Lastly, P. o. bridgesi occurs in Netherlands; Zoologische Staatssammlung München (ZSM),
western South America, being found in the Bolivian Yungas Munich, Germany; and the Natural History Museum (BMNH),
in the provinces of Cochabamba, Santa Cruz, Chuquisaca Tring, UK (Fig. 1).
and Tarija, and towards the south in Argentina in the forests Localities and geographic coordinates were obtained from the
of eastern slopes of the Andes in the provinces of Jujuy, Salta, labels attached to the specimens. When the coordinates were not
Tucumán and Catamarca. The gap between the ranges of this and available they were obtained by consulting ornithological gazetteers
the two other subspecies may have been caused by an increase (Paynter and Traylor 1991, Paynter 1992, 1995, Vanzolini 1992).
of the aridity in the Chaco, which has isolated this taxon in the The maps were designed using the software QGis 2.6.0, showing
Andean foothills (Vaurie 1968). the divisions of countries, Brazilian states and the biogeographic
Vaurie (1966) recognized that the nominate subspecies provinces proposed by Morrone (2009) to differentiate the areas
differs from P. o. bronzina by having the feathers of the head where the taxa occur. In order to perform detailed analysis of
entirely blackish with only a suggestion of light edgings on geographic variations in body size and plumage coloration we
the forehead and brows. Penelope o. bronzina, according to this defined nine geographic groups considering morphological
author, has the head with long and bordered feathers, especially homogeneity, geographic proximity and ecological similarity.
on the anterior half of the crown, with silvery-white. Also, the Specimens identified as juveniles by distinct plumage and smaller
race obscura would be smaller in size and lighter, with plumage sizes were excluded from the morphometric and plumage analyses,
browner and throughout duller. Other authors such as Delacour and were considered only to compile the distribution of the taxa.
and Amadon (2004) documented that P. o. bridgesi is similar to We measured the length of the exposed culmen, bill from
P. o. bronzina, being larger and having the crest feathers less nostril, tarsus, middle toe, wing and tail using calipers with
edged with white and little or no trace of eyebrow stripe, with precision of 0.01 mm following Baldwin et al. (1931). Morpho-
the plumage duller and browner. metric data were analyzed to test the sexual dimorphism and
Del Hoyo and Kirwan (2016) mentioned that plumage intra- and inter-geographic group variations. Shapiro-Wilk and
differences among races suggest that more than one species Lilliefors (Kolmogorov-Smirnov) tests were performed to test
may be involved in the P. obscura complex. They also pointed the normality of data. Sexual dimorphism in each group was
out that the possibility of contact and intergradation of the two evaluated through the Student’s t test, testing the differences be-
eastern races, P. o. bronzina and P. o. obscura, required detailed tween sexes for each selected character. For multivariate analyses,
study. Vaurie (1968) suggested that these races do not intergrade, the data requires log-transformed variables as input so that all
and Del Hoyo and Motis (2004) noted a gap between the two of them have the same relevance for the analysis. Multivariate
forms in the state of Santa Catarina, Brazil, and also recognized analyses are based on multiple variables that are measured in
the need of studies to evaluate the taxonomy of this complex. the same individual and are not independent from each other,
and we tested the general morphology of P. obscura with all the
MATERIAL AND METHODS morphometric variables taken together. To corroborate the sex-
ual dimorphism we performed the Hotelling’s T square test, the
We analyzed and measured 80 museum specimens of all multivariate counterpart of Student’s t, testing the differences
taxa currently considered as belonging to the P. obscura complex between the multivariate means. A Lineal Discriminant Analysis
(36 males, 29 females and 15 of unknown sex), all of which are (LDA) was also used; this test identifies all the elements that
distributed in Brazil, Argentina, Paraguay and Bolivia. These belong to each geographic group (including each measure for
specimens are housed in the following institutions: Museu de each variable) and generates a linear combination of variables
Zoologia da Universidade de São Paulo (MZUSP, including the that can find the pattern within the data and effectively classi-
topotype of P. o. bronzina), São Paulo, Brazil; Museu Nacional da fy it. It represents in percentages the possibility of individuals
Universidade Federal do Rio de Janeiro (MNRJ), Rio de Janeiro, belonging to the pre-determined group or meeting the pattern
Brazil; Museu de História Natural do Capão da Imbuia (MHNCI), of another group. The Canonical Variates Analysis (CVA) was
Curitiba, Brazil; Museu de Ciências Naturais da Fundação Zoobo- performed to check whether the groups form significantly dis-
tânica do Rio Grande do Sul (MCNRS), Porto Alegre, Brazil; and tinct clusters morphometrically. The univariate part of the tests
Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” was conducted in R software statistical package v. 3.2.1 and the
(MACN), Buenos Aires, Argentina. Photographs of the specimens multivariate analysis was carried out in Past v. 2.15.
were taken against a white background using 10 watt LED spot- The plumage was evaluated according to the Munsell
lights, always in the same place to standardize the lightening. Color Guide (1994), based on traits usually considered as
We also examined photographs of 24 specimens, including the diagnostic for the three subspecies: supercilium, crown, throat,
holotypes of P. o. bronzina and P. o. bridgesi, deposited in the breast, abdomen, back, tail and wing coverts. We also analyzed

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 Morphological evidence for the taxonomic status of P. bridgesi

Figure 1. Distribution of Penelope obscura. Black squares: specimens analyzed personally; Black triangles: specimens analyzed by photo-
graphs; White circles: living individuals (photos).

photographs of living individuals available in the online data- 2). Table 2 presents the descriptive statistical analysis including
bases Wiki Aves (http://www.wikiaves.com.br) and Ecoregistros mean, standard deviation and minimum and maximum values.
(http://www.ecoregistros.org). From WikiAves were reviewed Univariate analysis of normality suggests that data of the
4.342 images from the states of Espírito Santo (122), Minas Gerais six variables meet the criteria of the parametric tests. Student’s
(976), Rio de Janeiro (711), São Paulo (1.582), Paraná (333), Santa t-test was not performed for the groups with small sample
Catarina (274) and Rio Grande do Sul (344) in Brazil; and 100 number; it was used to test the null hypothesis that the mean
images from Ecoregistros, for the records in Uruguay (18) and value of each variable does not differ between the two sexes for
northeast (31) and northwest (51) of Argentina (Fig. 1). Some each group. Based on the test, the vast majority of the means
of the examined photographs were excluded from our analysis of the two sexes do not differ significantly; only in five specific
because they had been taken from a long distance, were very cases was there a significant difference between the sexes, with
dark, or portrayed partially or wrongly identified taxa. males being larger than females (Table 3). The T2 Hotelling test
has as null hypothesis that the vectors of the means of the sexes
RESULTS are equal, and shows that there is no sexual dimorphism in the
same geographic groups tested for Student’s t-test (p > 0.05).
Statistical analyzes: nine geographic groups were defined: Table 4 presents the results of Lineal Discriminant Analysis
three from the Atlantic forest, two from the Araucaria forest, and shows that everything in the off-diagonal are the frequencies
and one from the Parana forest, Pampa, Chaco and Yungas (Ta- of individuals that also meet the pattern of other groups. Only
ble 1). For statistical analyses, we examined 77 adult specimens 33 (44 %) of the individuals were correctly classified within
distributed in the aforementioned biogeographic provinces (Fig. their pre-determined group. The MANOVA/CVA explained

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 D. Evangelista-Vargas & L.F. Silveira

Figure 2. Geographic groups of Penelope obscura separated by biogeographic provinces: Parana forest (1), Atlantic Forest (2,3,4), Araucaria
Forest (5,6), Pampa (7), Chaco (8) and Yungas (9).

Table 1. Geographic groups and respective number of skins used the variations between the geographic groups in the first two
for statistical analysis. axes, and most of them could not be differentiated by any of
the morphometric variables. Groups 6 and 7 from southern
Geographic groups Number and gender Localities Brazil and northeastern Argentina (Araucaria forest and Pampa,
1. Parana Forest 8 (4 males, 4 females) Southeast of Minas Gerais (Brazil) respectively) differed slightly from the others in the length of
2. Atlantic Forest 1
24 (14 males, 8
Northeast of São Paulo (Brazil) the tail (Fig. 3).
females, 2 unknown)
Specimens collected in eastern South America showed
11 (5 males, 6
3. Atlantic Forest 2
females)
East of São Paulo (Brazil) continuous distribution with no gap in the state of Santa Ca-
East of Paraná and Northeast of
tarina in Brazil. Their plumage showed no sexual dimorphism
4. Atlantic Forest 3 5 (1 male, 4 unknown)
Santa Catarina (Brazil) in any geographic group. The characters used to diagnose the
5. Araucaria Forest 1
5 (1 male, 2 females, 2 Central Paraná and North of Santa subspecies P. o. obscura were found in specimens from various
unknown) Catarina (Brazil)
parts of its wide range. For instance, the whitish edges of head
Center Santa Catarina and Northeast
6. Araucaria Forest 2 4 (1 male, 3 females)
of Rio Grande do Sul (Brazil)
feathers are wide (MZUSP 31577, from Varjão de Guaratuba, São
Paulo, BR), thin (MZUSP 34130, Maromba, Rio de Janeiro, BR) or
7 (4 males, 2 females, Southwest of Rio Grande do Sul
7. Pampa
1 unknown) (Brazil) and Misiones (Argentina) almost imperceptible (MACN 44036, Chantecler, Misiones, ARG),
11 (6 males, 4 without any apparent geographic trend. The overall color of the
8. Chaco Tucumán and Catamarca (Argentina)
females, 1 unknown)
crown feathers affects the appearance of the head, which can be
9. Yungas 1 (unknown) Salta (Argentina) black, gray, or brown. In addition, the prominence of the white

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 Morphological evidence for the taxonomic status of P. bridgesi

Table 2. Mean, standard deviation, minimum and maximum values of body measurements (mm) from the geographic groups of Penelope
obscura.

Geographic
Exposed culmen Nostril Tarsus Middle Toe Wing Tail
groups
1 (n = 8) 29.04 + 2.62 (26.2–33.1) 25.41 + 1.83 (23.3–27.6) 75.41 + 5.83 (70.2–88.6) 62.38 + 2.39 (59–67) 314.75 + 14.51 (292–338) 357.38 + 10.25 (342–372)
2 (n = 24) 30.24 + 2.70 (24.9–35.1) 24.52 + 1.66 (21.6–27.3) 76.35 + 4.10 (68.4–83.1) 66.75 + 3.35 (62–73) 324.3 + 15.89 (295–360) 356.4 + 19.23 (321–410)
3 (n = 11) 32.16 + 1.49 (30–34.7) 25.21 + 1.22 (23.7–27.8) 80.73 + 2.07 (76.2–82.8) 69.54 + 1.86 (67–72) 324.7 + 25.68 (293–362) 346.2 + 23.62 (322–385)
4 (n = 5) 32.1 + 2.41 (29–35) 25.68 + 1.70 (23.1–27.2) 84.32 + 3.54 (78.7–87.2) 71 + 4.74 (63–75) 326.8 + 14.20 (306–343) 346 + 16.98 (321–368)
5 (n = 5) 27.42 + 2.74 (23.2–30.4) 22.66 + 1.98 (20.5–24.8) 79.96 + 3.90 (77–86.7) 66.8 + 4.15 (61–72) 320.2 + 20.90 (299–347) 356.6 + 19.77 (330–384)
6 (n = 4) 25.95 + 0.87 (25.2–27.2) 22.3 + 0.22 (22–22.5) 77.65 + 3.13 (74–81.2) 64.75 + 2.5 (62–68) 311.25 + 16.68 (287–324) 311.5 + 16.98 (288–327)
7 (n = 7) 25.81 + 1.23 (23.9–27.7) 22.7 + 0.81 (21.4–23.7) 79.01 + 2.33 (77.1–83.3) 65.86 + 2.91 (61–70) 322.4 + 17.35 (297–345) 323 + 15.06 (299–339)
8 (n = 11) 27.16 + 1.03 (25.9–28.5) 22.66 + 1.18 (21.1–24.5) 76.86 + 3.32 (71.2–81.4) 63.4 + 3.96 (58.2–70) 329.5 + 18.13 (295–346) 355.3 + 15.03 (328–376)
9 (n = 1) 26.1 23.3 81.8 68 346 368

Table 3. Student t-test comparing males and females of each geo- Table 4. Frequency of percentages of predictive performance from
graphic group with more than four individuals for each sex. Asterisks the Lineal Discriminant Analysis (LDA) for all the geographic groups.
indicates significant differences between both sexes at p < 0.05.
Original
Predicted
Geographic Exposed Middle 1 2 3 4 5 6 7 8
Nostril Tarsus Wing Tail
group culmen Toe
1 50.0 12.5 – – – – – 9.1
t = -1.730 t = -2.006 t = -0.118 t = 0.137 t = -0.971 t = 0.921
1 (♂: 4, ♀: 4) 2 37.5 62.5 27.3 – 20 – 28.6 36.4
p = 0.134 p = 0.091 p = 0.91 p = 0.895 p = 0.369 p = 0.392
3 – 8.3 54.5 80 40 – – –
t = -1.448 t = -0.354 t = -1.132 t = -3.248 t = -2.388 t = -1.047
2 (♂: 14, ♀: 8)
p = 0.163 p = 0.727 p = 0.27 p = 0.004* p = 0.027* p = 0.307 4 12.5 – 9.1 – – – – –
t = -2.342 t = 0.116 t = -0.409 t = -0.084 t = -1.160 t = -1.430 5 – 4.2 – – – – – 9.1
3 (♂: 5, ♀: 6)
p = 0.044* p = 0.910 p = 0.692 p = 0.935 p = 0.276 p = 0.186
6 – – – – – 25 28.6 –
t = -0.468 t = -1.756 t = -0.533 t = -0.620 t = -3.367 t = -2.968 7 – 8.3 9.1 20 – 75 28.6 –
8 (♂: 6, ♀: 4)
p = 0.652 p = 0.117 p = 0.608 p = 0.552 p = 0.010* p = 0.018*
8 – 4.2 – – 40 – 14.2 45.4

Figure 3. CVA Plot of geographic groups according to morphometric measurements. Black circles: Parana Forest (group 1); Plus: Atlantic
Forest 1 (group 2); White squares: Atlantic Forest 2 (group 3); Black squares: Atlantic Forest 3 (group 4); Exes: Araucaria Forest 1 (group 5);
White circles: Araucaria Forest 2 (group 6); White diamonds: Pampa (group 7); Inverted black triangles: Chaco (group 8); White triangle:
Yungas (group 9).

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 D. Evangelista-Vargas & L.F. Silveira

Figure 4. From top to bottom, dorsal view of P. o. bronzina (MZUSP 49384, São Paulo, southeastern BR), P. o. obscura (MACN 48384,
Corrientes, northeastern ARG), and P. o. bridgesi (MACN 8148a, Tucumán, northern ARG). Scale bars: 10.0 cm.

hue in the edge of the feathers appears to vary considerably. blackish (MZUSP 49384, Rocha, São Paulo, BR). Apparently,
Specimens from Potreirinhos in Rio Grande do Sul (MCNRS 682, specimens from the northern part of the range have the white
BR) to the south often have less noticeable whitish edges and streaks on the back more noticeable when compared to the
tend to have a blackish general coloration, but this pattern is also southern specimens. However, we also found specimens in the
recorded in birds from the northern portion of the range, which south with wide white streaks (MCNRS 1765, Bagé, Rio Grande
present general blackish coloration and feathers with white do Sul), and in the north the white streaks in some specimens
edges reduced or absent (WA700594, WA823706). On the other are almost absent (MN 45542, Simão Pereira, Minas Gerais, BR)
hand, specimens in the south of the range may present feath- (Fig. 4).
ers with noticeable white edges and general grayish coloration Some specimens showed a contrasting dark chest with a
(WA914588, WA1763524, WA1782826; ER10072, ER10141). paler belly (MN 47672, Matias Barbosa, Minas Gerais; MHNCI
The plumage of the back also showed considerable vari- 204, Tibagi, Paraná, BR), whereas other specimens had a uni-
ation, without any apparent geographical pattern. Across the formly colored chest with either a dark belly (MN 49619, Levi
entire range of the species we found individuals with the back Gasparian, Rio de Janeiro; MZUSP 49382, Morretinho, São Paulo;
greenish-brown (MZUSP 32906, Teresópolis, Rio de Janeiro, BR), MCNRS 1765, Bagé, Rio Grande do Sul, BR) or a paler, uniform
brownish (MZUSP 7021, Fazenda Monte Alegre, Paraná, BR) or belly (MHNCI 399, Vale do Ivaí, Paraná, BR). Both phenotypes

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 Morphological evidence for the taxonomic status of P. bridgesi

Figure 5. From top to bottom: lateral view of P. o. bronzina (MZUSP 49384, São Paulo, southeastern BR), P. o. obscura (MACN 48384,
Corrientes, northeastern ARG), and P. o. bridgesi (MACN 8148a, Tucumán, northern ARG). Scale bars: 10.0 cm.

were found in photographs taken at the same locality (e.g. tently uniform and slightly paler than the back, whereas in P. o.
WA1380991 and WA1423978, from Afonso Cláudio, Espírito obscura and P. o. bronzina the coloration of the upperparts was
Santo, BR). The color of the belly was also variable, ranging highly variable. The wing coverts of P. o. bridgesi show broad
from brown (MZUSP 1911, Colonia Hansa, Santa Catarina, BR, white margins, a character state not found in P. o. obscura and
topotypical) to orangish (MZUSP 78284, Bananal, São Paulo, BR). P. o. bronzina, which have indistinct white margins to the wing
It is common to find individuals with different overall coverts (Fig. 5). Overall, P. o. bridgesi shows less pronounced
coloration in the same place, demonstrating the existence of a individual variation in plumage coloration than P. o. obscura
large individual variation (e. g. WA1681568, Pouso Alegre, Minas and P. o. bronzina.
Gerais; WA450514, Nova Friburgo, Rio de Janeiro; WA1518511,
Itatiba, São Paulo; WA1084879 and WA1090972, Tibagi, Paraná, DISCUSSION
BR). As mentioned before we located specimens with plumage
characters supposedly diagnostic for the nominate subspecies Misidentifications of Penelope obscura are still common,
in specimens found in the range of P. o. bronzina (WA1765142, especially where it is sympatric with P. superciliaris. The range
Bocaina de Minas, Minas Gerais, BR) and specimens with char- of P. obscura in eastern South America is continuous, without
acters considered diagnostic for P. o. bronzina in the geographic any gap between the distribution of the nominate race and P. o.
range of nominate subspecies (ER10141; Ribera del Norte, Buenos bronzina in southern Brazil (Fig. 1). We suspect that the absence
Aires, Argentina). of specimens collected between the states of Santa Catarina (from
On the other hand, all specimens attributed to P. o. north, called P. o. bronzina) and Rio Grande do Sul (to south, P. o.
bridgesi lacked olive brown coloration with a green tinge to obscura), deposited in North American and European museums
the upperparts (unlike P. o. obscura and P. o. bronzina) (Fig. 4). may have led to the assumption that there was a large geographic
Instead, specimens of P. o. bridgesi showed chestnut upperparts gap between the ranges of P. o. obscura and P. o. bronzina. As
contrasting with the black head. The underparts were consis- no detailed specimen-based study has been conducted to date,

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 D. Evangelista-Vargas & L.F. Silveira

including analyses of individual and geographic variation, the guay, and the holotype from Bolivia), excluding the characters of
validity of P. obscura bronzina remained uncontested for many bridgesi cited by Gray (1860) that are diagnosable to considered
decades in the ornithological literature. The same applies to the it as a species; the same situation was also observed in Hartert
taxonomic rank of P. o. bridgesi, an isolated taxon occurring in and Venturi (1909), which considered it as a subspecies of P.
the biogeographic provinces of Yungas and Chaco on the eastern obscura. Our results clearly show that Penelope bridgesi is a taxon
slopes of the Andes in Bolivia and Argentina, which also was that can be differentiated from all others without overlapping
never studied in detail. with any other Penelope. It is geographically isolated and should
The eastern population (P. o. obscura and P. o. bronzina) be treated as a valid species (see below). The populations in east-
showed considerable variation in plumage coloration, with ern and southern Brazil are continuously distributed and show
many individuals presenting characters previously considered considerable individual variation in plumage, without any clear
as diagnostic for one of the subspecies in the range of the other. geographic trend. There are any morphological or morphometric
The plumage in P. o. bridgesi was much less variable throughout characters to support the continued existence of more than one
its range than that of P. o. obscura/bronzina. Statistical analyses taxon in the region. As P. o. bronzina cannot be distinguished
showed no morphological differences between males and fe- from the nominate subspecies, we here treat Penelope obscura
males within the groups, and the few differences among some bronzina Hellmayr, 1914 as a synonym of Penelope obscura Tem-
geographic groups do not represent a pattern of biogeographic minck, 1815. The treatment proposed here is consistent with
variation in the eastern or western populations. We cannot be the current proposals found for the other species of Penelope.
sure that there is a marked difference in size between the afore-
mentioned subspecies of P. obscura, as was previously mentioned TAXONOMY
by Vaurie (1968). Total size measurements attached to some
specimens’ labels suggest that males are larger than females, Penelope obscura Temminck, 1815
but the color of the iris is the only character that effectively
differentiates the two sexes. Holotype. The description of the species is based on the
The occurrence of P. obscura is still questionable in some ‘Yacuhú’, made by Azara (1805). Type locality: Paraguay.
regions. Sousa (1999) mentioned the presence of this species Common names. Portuguese: Jacuaçu. Spanish: Pava
in Elísio Medrado (13°S, 39°45’W) in the state of Bahia (Brazil), oscura. English: Dusky-legged Guan.
being the north limit of its distribution. However, as this record Diagnosis. Distinguished from P. bridgesi by olive brown
was based on a personal communication from a third part, and coloration with green tinge in upperparts, and generally darker
there are no records from naturalists in the 19th and 20th centu- overall coloration. Underparts very variable, unlike P. bridgesi.
ries, when the habitat was undisturbed, we consider unlikely that White margins of wing coverts markedly reduced. Head, hind-
P. obscura occurs in the state of Bahia since there is no reliable neck, and back coloration more uniform than in P. bridgesi.
evidence for it. In Paraguay, Hayes (1995) mentioned Penelope Re-description. No sexual dimorphism found in plumage
obscura as a hypothetical species. It was visually reported in the coloration. Whitish edges extension of head feathers varies from
Chaco (President Hayes, Boquerón) and Orient (Central, Alto wide to almost imperceptible. Some individuals show a moder-
Paraná and Ñeembucú), but unfortunately those reports had no ately distinct superciliary stripe formed by short feathers with
other details. The only specimens mentioned by the author are white edges. Overall color of crown feathers variable, ranging
deposited at the National Museum of Natural History, Smithso- from black, gray to brown. White hue of the edge of the feathers
nian Institution (USNM 256875, 58994), and they lack precise varies individually. Brown back may show greenish, brownish
information on collecting dates and localities, and were obtained or blackish tinge. Neck, breast, upper back, and shoulders
at a time when Paraguay included parts of what is now Brazil conspicuously edged with white. Birds in the northern parts of
and Argentina. Smith and Derna (2015) recently reviewed the the species’ range tend to have more noticeable white streaks
records in the country and showed that museum skins attributed on the back, but streaks also may be lacking. The coloration of
to P. obscura actually represent members of P. superciliaris. Recent the underparts is variable and may show a contrastingly darker
records from Paraguay are photographs taken in the Refugio chest than belly, but underparts may also be uniform dark or
de Vida Silvestre Atinguy, in Misiones department (27°20’S, lighter than the breast. The belly feathers have contrasting
56°41’W), claimed by the authors to be wild-origin birds. horizontal scaling, which may range from brown to orangish,
After the review of a large series of specimens of P. o. and may vary in intensity. Tail feathers are dark brown with
bridgesi, and by comparing it with P. o. obscura and P. o. bron­ bronze or purplish sheen. Bare skin of face slaty, legs dusky gray
zina, we find that what is currently defined in the literature as to blackish, and wattle red.
Penelope obscura does not include the characters observed in the Distribution. Southeastern and southern Brazil (from the
individuals of P. o. bridgesi. When Ogilvie-Grant (1893) proposed east of Minas Gerais and north of Espírito Santo to Rio Grande
bridgesi as a synonym of Penelope obscura, he only reviewed seven do Sul), northeastern Argentina (Misiones, Chaco, Corrientes,
specimens (four from Brazil, one from Uruguay, one from Para- Entre Ríos, Santa Fe and Buenos Aires), and the forested regions

8 / 10 ZOOLOGIA 35: e12993 | DOI: 10.3897/zoologia.35.e12993 | March 29, 2018

 Morphological evidence for the taxonomic status of P. bridgesi

of Uruguay, with recent photographic records for Paraguay. In- Ecoregistros.org (n = 2). ARGENTINA: 10072 (Reserva
habits the biogeographic provinces of Atlantic Forest, Cerrado, Ecológica de Ribera del Norte – San Isidro, Buenos Aires); 10141
Paraná Forest, Araucaria Forest and Pampa. (Reserva Ecológica de Ribera del Norte – San Isidro, Buenos Aires.
Examined material (n = 84). BRAZIL. Minas Gerais: Muriaé
(1 ♀- MNRJ 36791); Juiz de Fora Km.118 (1 ♂- MNRJ 45036); Juiz
de Fora Km.788 (1 ♂- MNRJ 48346); Matias Barbosa Km.795 (1 Penelope bridgesi Gray, 1860
♂- MNRJ 47672); Matias Barbosa Km.801 (1 ♀- MNRJ 48384);
Simão Pereira Km.807 (1 ♀- MNRJ 49558); Simão Pereira Km.817 Holotype. BMNH 1846.9.9.42 (sex unknown, Bolivia;
(1 ♀- MNRJ 45542, 1 ♂- MNRJ 48997); Simão Pereira Km.825 (1 examined).
♀- MNRJ 48383); Estação do Túnel (2 ♀- BMNH 1901.3.28.65, Common name. Spanish: Pava de Bridge. English:
BMNH 1901.3.28.64); Rio de Janeiro: 2-BMNH 1889.6.1.215, Bridge’s Guan.
RNHL no number, 1 ♂- 1895.4.1.3, 1 ♀- 1895.4.1.4; Rio de Janeiro Diagnosis. Distinguished from all members of Penelope by
(1 – ZSM no number); Levi Gasparian Km.792 (1 ♂- MNRJ 49619); having chestnut-olive brown coloration without green tinge,
Levi Gasparian (1 ♀- MNRJ 48338); Tres Rios Km.27 (1 ♀- MNRJ and overall coloration browner than P. obscura. Underparts
49080); Areal Km.44 (1 ♂- MNRJ 49081); Petropolis Km.52 (1 ♂- are uniform and slightly paler than the back, unlike P. obscura.
MNRJ 49574); Petropolis Km.64 (1 ♂- MNRJ 44389); Teresopolis Wing coverts with contrasting large white margins. Black head
Km.52 (1 ♀- MNRJ 47743); Teresopolis Km.82 (1 ♀- MNRJ 48354); and hindneck contrast with brownish back, being distinct of
Teresopolis (2 – MNRJ 9321, MNRJ 9322, 3 ♂- MZUSP 32905, P. obscura.
MZUSP 32906, MZUSP 32907, 2 ♀- MZUSP 32904, MZUSP 32908; Re-description. No sexual dimorphism found in plumage
Guapimirim Km.96 (1 ♂- MNRJ 48337); Parnaso (1 ♂- MNRJ coloration. General coloration chestnut, with limited indi-
48336); Inga (1 ♂- MNRJ 47552); Morro Redondo (1 ♀- MNRJ vidual variation. Crest feathers dark with whitish margins,
9405); Maromba (1 ♀- MZUSP 34129, 1 ♂- MZUSP 34130); Paraty most strongly marked on the forehead; little or no trace of an
(1 ♂- MNRJ 36927); São Paulo: 1 – BMNH 1889.6.1.203; Bananal superciliary stripe. Upperparts chestnut, contrasting with the
(1 ♂- MZUSP 78284, 1 ♀- MZUSP 78285); Varjão de Guaratuba black head and hindneck. Lower parts uniform dull reddish
(3 ♀- MNRJ 31062, MZUSP 31576, MZUSP 43763, 1 ♂- MZUSP brown and slightly paler than the back. The belly feathers
31577); Cabreúva (1 ♂- MZUSP 93098); Morretinho e Rocha (2 have contrasting horizontal scaling, which may range from
♂- MZUSP 49382, MZUSP 49383, 1 ♀- MZUSP 49384); Rio Ribeira brown to orangish and may vary in intensity. Feathers of the
(1 ♀- MZUSP 66890, 1 ♂- MZUSP 66891); Icapara (1 ♀- MZUSP neck, mantle and chest are margined with white, wider on
73215); Paraná: Vale do Ivai (1 – MHNCI 399); Santa Barbara the mantle and wing coverts. Tail feathers dark brown with
(1 ♀- MHNCI 251); Fazenda Monte Alegre (1 ♂- MZUSP 7021); brownish sheen. Bare skin of face slaty, legs dusky gray to
Tibagi (1 ♀- MHNCI 2041); Guaratuba-Caiobá (1 ♂- MHNCI blackish, and wattle red.
1189, 2 – MHNCI 256, MHNCI 2519); Santa Catarina: Rio Negro Distribution. Forests of the eastern slopes of the Andes, from
(1 – MHNCI 5776); Joinville (1 ♂- RNHL 1292-3, 1 – MZUSP central Bolivia (Cochabamba, Chuquisaca and Tarija) to northwest
4863); Colonia Hansa (1 ♂- ZSM 12415 Penelope obscura bronzina Argentina (Jujuy, Salta, Tucumán and Catamarca). Found in the
HOLOTYPE, 2 – MZUSP 1911, MFN 2759); Catanduvas (1 ♂- biogeographic provinces of the Yungas and the Chaco.
MNRJ 36928); Anita Garibaldi (1 ♀- MCNRS 2768); Jaguariaiva, Examined material (n = 20). ARGENTINA, Jujuy: Cerro
Fazenda Morungava (1 – FMNH no number, 1 ♂- 110740); Rio Calilegua (1 ♂- ZSM 21118); Valle Grande, Ledesma (1 ♀- ZSM
Grande do Sul: Potreirinhos (1 ♀- MCNRS 682); Bom Jesus (1 59536); Salta: Río Santa María (1 – MACN 30575); Salta (1 ♀-
♀- MCNRS 2544); São Francisco de Assis (1 – MCNRS 1975); BMNH 1899.1.27.214); Tucumán: Vipos (1 ♀- MACN 8148a);
Bagé (1 ♂- MCNRS 1765). URUGUAY. 1 ♂- BMNH 1878.1.25.1; Tafí Viejo (1 ♂- MACN 8884); Sierra del Aconquija (1 ♀- MACN
Payssandu: 1 – BMNH 1901.2.10.24. PARAGUAY. 1 – BMNH no 42153, 4 ♂- MACN 42156, MACN 42157, MACN 42158, MACN
number. ARGENTINA. Misiones: Chantecler (2 ♂- MACN 44034, 42159, 1 – MACN 42160); Aconquija (1 ♀- MACN 32840);
MACN 44036, 1 ♀- MACN 44035); Corrientes: Rincón Mercedes Catamarca: El Suncho (1 ♂- MACN 42154, 1 ♀- MACN 42155).
(1 ♂- MACN 48384, 1 ♀- MACN 48385). BOLIVIA: Penelope bridgesi HOLOTYPE (BMNH 1846.9.9.42);
Mentioned photographs. Wikiaves.com.br (n = 13): BRA- Cochabamba: Tin-Tin (1 ♂- FMNH 413837); Tarija: Villa Montes
ZIL. 700594 (Guarapuava, Paraná); 823706 (Caputira, Minas (2 – ZSM 27977, ZSM 27978, 1 ♀- MFN 313633).
Gerais); 914588 (Taquara, Rio Grande do Sul); 1763524 (Aceguá,
Rio Grande do Sul); 1782826 (São Martinho da Serra, Rio Grande ACKNOWLEDGMENTS
do Sul); 1380991 (Afonso Cláudio, Espirito Santo); 1423978
(Afonso Cláudio, Espirito Santo); 1681568 (Pouso Alegre, Mi- We thank Fundação de Amparo à Pesquisa do Estado
nas Gerais); 450514 (Nova Friburgo, Rio de Janeiro); 1518511 de São Paulo (FAPESP), Conselho Nacional de Desenvolvi-
(Itatiba, São Paulo); 1084879 (Tibagi, Paraná); 1090972 (Tibagi, mento Científico e Tecnológico (CNPq) and Coordenação de
Paraná); 1765142 (Bocaina de Minas, Minas Gerais). Aperfeiçoamento do Pessoal de Nível Superior (CAPES) for the

ZOOLOGIA 35: e12993 | DOI: 10.3897/zoologia.35.e12993 | March 29, 2018 9 / 10

 D. Evangelista-Vargas & L.F. Silveira

grants received and for the financial support; Idea Wild for the Munsell AH (1994) Soil Color Charts. Macbeth Division of Kollmorgan
equipment donated to DEV; the photographers who posted the Instruments Corporation, New York, 10 pp.
photos in EcoRegistros and WikiAves websites. The curators Nacinovic JB (2012) Uma contribuição iconográfica para a iden-
and staff of the following institutions kindly welcomed us and tificação dos Jacus (Aves, Cracidae, Penelope) que ocorrem no
allowed us to study the specimens under their care: MFN and Brasil. Agência 2A Comunicação, Rio de Janeiro, 80 pp.
ZSM (Germany); FMNH (USA); RNHL (Netherlands); BMNH Ogilvie-Grant WR (1893) Catalogue of the Birds in the British Mu-
(UK); MACN (Argentina); MNRJ, MHNCI and MCNRS (Brazil). seum 22. Order of the Trustees, London, 585 pp.
We also thank Instituto Chico Mendes de Conservação da Bio- Paynter RA (1992) Ornithological Gazetteer of Bolivia. Museum of
diversidade (ICMBio/SISBIO) for issuing collecting permits. LFS Comparative Zoology of Harvard University, Cambridge, 185 pp.
is supported by a CNPq research productivity fellowship. George https://doi.org/10.5962/bhl.title.14591
Sangster kindly reviewed the manuscript and offered important Paynter RA (1995) Ornithological Gazetteer of Argentina. Muse-
contributions to this text. um of Comparative Zoology of Harvard University, Cambridge,
1043 pp. https://doi.org/10.5962/bhl.title.14627
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Morrone JJ (2009) Evolutionary Biogeography: An Integrative the ms and gave final approval for publication.
Approach with Case Studies. Columbia University Press, New Competing Interests: The authors have declared that no competing
York, 304 pp. interests exist.

10 / 10 ZOOLOGIA 35: e12993 | DOI: 10.3897/zoologia.35.e12993 | March 29, 2018