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Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil

Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Revista Brasileira de Ornitologia, 22(2), 95-101 ARTICLE June 2014 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt Department of Ornithology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024, USA. E-mail: sclaramunt@amnh.org Received on 8 January 2014. Accepted on 10 May 2014. ABSTRACT: Mazar Barnett & Buzzetti (2014) described a new species, Cichlocolaptes mazarbarnetti, from the Atlantic forests of northeastern Brazil. The holotype of the new species is a female that shows remarkable similarities with the sympatric Philydor novaesi. Here I analyze eight morphometric variables to assess phenotypic similarities among specimens of Philydor novaesi, Cichlocolaptes mazarbarnetti and other species in these two genera. The holotype of C. mazarbarnetti differs from Philydor novaesi and falls in a region of the morphometric space occupied by specimens of Cichlocolaptes leucophrus. Therefore, morphometric data is consistent with other morphological, behavioral, and vocal data that suggest the existence of a new species Cichlocolaptes in the northern Atlantic forest. KEYWORDS: Cichlocolaptes mazarbarnetti, Philydor novaesi, Furnariidae, measurements, taxonomy. INTRODUCTION originally identified as P. novaesi is an exemplar of the new foliage-gleaner; diagnostic characters include a larger The avifauna of the northern stretch of the Atlantic body size, a longer beak, absence of buffy orbital feathers, Forest is one of the least known, yet most endangered on and rounded tail feather tips. Although Mazar Barnett & Buzzetti (2014) measured and discussed some biometric Earth. Habitat destruction has outpaced ornithological research and conducting research on some species today data, comparisons were limited to single variables is difficult because of their scarcity and vulnerability. The and few species. As part of a large-scale morphometric inventory of the taxonomic composition of this avifauna analysis of the Furnariidae (Claramunt 2010; Claramunt is incomplete, as demonstrated by continuous discoveries et al. 2013), I examined and measured all specimens identified as P. novaesi at the Museu Nacional, including of new species in the last decades (e.g. Teixeira 1987; Teixeira & Gonzaga 1983, Silva et al. 2002). One such the holotypes of P. novaesi and C. mazarbarnetti. Here I species is the Alagoas foliage-gleaner Philydor novaesi, present a morphometric analysis of those specimens and discovered at Pedra Branca, Muricí, a relict of foothill discuss its implications regarding their taxonomic status. forest in Alagoas State (Teixeira & Gonzaga 1983b), and found later in a small reserve in Pernambuco State (Mazar METHODS Barnett et al. 2005). The species is considered critically endangered (BirdLife International 2013). In a surprising turn of events, Mazar Barnett & Buzzetti (2014) Measurements were taken with a Mitutoyo Digimatic described yet another new foliage-gleaner from Murici, Point Caliper (resolution: 0.01 mm) with an output interface. Variables were the same as in previous Cichlocolaptes mazarbarnetti, based on field observations morphometric studies of the Furnariidae (Claramunt and examination of museum specimens. The new species is extremely similar to P. novaesi in plumage pattern and et al. 2010; Claramunt et al. 2013). Here, I analyzed color but its behavior and vocalizations resemble those of only two variables from each body region because some Cichlocolaptes leucophrus. In particular, like C. leucophrus, measurements were highly correlated in foliage-gleaners: 1) wing length; 2) wing length to the first secondary the new species shows a preference for foraging on feather; 3) tail length to central rectrices; 4) tail length to bromeliads. Upon examination of specimens identified as P. novaesi at the Museu Nacional in Rio de Janeiro, Mazar the most external rectrix; 5) bill length from the anterior Barnett & Buzzetti (2014) concluded that one specimen border of the nostril to the tip of the bill; 6) bill width at Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt the level of the anterior border of the nostrils; 7) tarsus Cichlocolaptes examined. Given a ratio of 1.32, rectrix 1 length; and 8) hallux length with claw (Baldwin et al. should be around 82.6 mm when fully-grown, not much 1931; Claramunt et al. 2010). longer than the actual measurement (Table 1). All known specimens of Philydor novaesi and the Two principal components explained 86% of holotype of Cichlocolaptes mazarbarnetti were examined the variance among all specimens measured (Table 2). and measured (Appendix). For comparison, I included Principal component 1 is positively correlated with all in the analysis other Philydor and Cichlocolaptes taxa. variables; thus, it is associated with overall size. Principal Although DNA sequences of P. novaesi are not available, component 2 is a contrast between variables of the bill its overall morphology and plumage suggest a close and variables of the flight apparatus (wings and tail). The relationship with P. atricapillus (Teixeira & Gonzaga three genera, Philydor, Cichlocolaptes, and Pseudocolaptes, 1983b) from southeastern Brazil, which belongs to a occupy different parts of the morphospace defined clade including P. pyrrhodes, Cichlocolaptes leucophrus, by components 1 and 2, with limited overlap (Figure and Heliobletus contaminatus (Derryberry et al. 2011; 1). Within each genus, females and males are almost Appendix). I did not include H. contaminatus in the completely segregated. Females tend to have lower scores analysis because of its small size and very distinctive than males in both components, indicating smaller morphology compared to the other species examined. overall size but relatively larger bills. P. novaesi is in an In addition, I analyzed specimens of P. fuscipenne and intermediate position among the three genera. The three P. erythrocercum (Appendix), which are more distantly males of P. novaesi are tightly clustered, with intermediate related but share several phenotypic similarities with scores on component 1 (intermediate size) and high scores P. novaesi, and cannot be discarded as potential close on component 2 (relatively long wings and tail but short relatives. Finally, I also analyzed specimens of the genus bill). Females, in contrast, are heterogeneous. Female Pseudocolaptes (Appendix), another genus of Furnariidae 33873 has lower scores in both components, compared that specialized in bromeliad foraging (Sillet et al. 1997, to males, and its position matches the relative position Martínez 2003), to investigate patterns of morphological of females due to sexual dimorphism. Female 34531, variation related to bromeliad specialization. I examined in contrast, is positioned near males P. novaesi. Female scatter plots of all variables obtained and conducted a 34530, the type of C. mazarbarnetti, has a slightly larger Principal Components Analysis on the covariance matrix component 1 score but a considerably lower component of log-transformed values for all measured specimens. 2 score compared to male P. novaesi; has a result, it is positioned in a region of the morphospace occupied by Cichlocolaptes leucophrus, and closer to females of RESULTS Pseudocolaptes than to males of P. novaesi. The use of the estimated fully-grown tail length of 34530 instead of its All species analyzed showed sexual dimorphism in actual length had a negligible effect on the position of this which males are larger than females (Table 1). The size specimen in the multivariate space. dimorphism is particularly pronounced in wing and A simple plot of wing and bill length shows a similar tail length, for which males of some species average 10 pattern (Figure 2). P. novaesi specimens are closer to other mm longer than females. Exceptions to this pattern are species of Philydor. Note, again, the nearly complete Cichlocolaptes and Pseudocolaptes, which show reversed segregation of males and females along the wing-length axis, sexual dimorphism in bill length, with females having and, for Cichlocolaptes and Pseudocolaptes, along the bill- longer bills (Table 1). P. novaesi is larger than other species length axis. Again, female 34530 is closer to Cichlocolaptes of Philydor. Whereas males are relatively uniform (small and Pseudocolaptes rather than to other specimens of P. standard deviations), females previously assigned to P. novaesi and specimens of Philydor in general. novaesi (including the holotype of C. mazarbarnetti) are unusually heterogeneous, particularly in wing, bill, and tarsus length (Table 1). Female 33873 is about DISCUSSION 90% smaller than males’ averages in wing, tail, and feet variables. Female 34531 is about the same size as males in The morphometric analysis presented here provides a all variables. Female 34530, the type of C. mazarbarnetti, quantitative assessment of all specimens of P. novaesi and has long wings and bill, surpassing all specimens of P. C. mazarbarnetti (Mazar Barnett & Buzzetti 2014). The novaesi, including males. The tail of 34530 is not longer analysis indicates that P. novaesi is larged compared to the than that of P. novaesi, however, rectrices 1 and 2 may not other species of Philydor, evident in both univariate and be fully grown, as their rachises still show basal sheaths. It multivariate analyses (Table 1, Fig. 1 and 2). The three is possible to estimate the expected tail length of 34530 male specimens of P. novaesi are fairly homogeneous, from the length of rectrix 6, using the average ratio between occupying a small sector of the morphospace, but the these two measurement among specimens of Philydor and three females are unusually heterogeneous. Female 33873 Revista Brasileira de Ornitologia, 22(2), 2014 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt FIGURE 1. Principal component analysis of eight morphometric variables for male (plain symbols) and female (cross) specimens of Philydor, Cichlocolaptes, and Pseudocolaptes, including Philydor novaesi (black squares), the holotype of Cichlocolaptes mazarbarnetti (black star) and a specimen of uncertain affinities (black circle). Numbers below symbols are Museu Nacional specimen numbers. An approximate interpretation is given for each component but see Table 2 for variable loadings. FIGURE 2. Position of male (plain) and female (cross) specimens of Philydor, Cichlocolaptes, and Pseudocolaptes in the morphometric space defined by wing length and bill length, includine specimens of Philydor novaesi (black squares),dthe holotype of Cichlocolaptes mazarbarnetti (black star) and a specimen of uncertain affinities (black symbol). Numbers below symbols are Museu Nacional(specimen number). Revista Brasileira de Ornitologia, 22(2), 2014 Santiago Claramunt fj Revista Brasileira de Ornitologia, 22(2), 2014 TABLE 1. Morphometric data for species of Philydor, Cichlocolaptes, and Pseudocolaptes, including individual data for all specimens of Philydor novaesi and Cichlocolaptes mazarbarneti (holotypes marked with *), and average and standard deviations for a combination of female specimens of P. novaesi and C. mazarbarnetti, which were originally identified as the former species only. See discussion for the placement of MN 34531 in P. novaesi or C. mazarbarnetti. wing secondary 1 rectrix 1 rectrix 6 bill length bill width tarsus hallux _ _ _ _ _ _ _ _ Taxon sex X, SD X, SD X, SD X, SD X, SD X, SD X, SD X, SD Cichlocolaptes mazarbarnetti 34530* f 96.5 77.7 81.8 62.5 15.3 4.9 22.7 16.7 Cichlocolaptes mazarbarnetti? 34531 90.1 76.0 84.0 64.4 12.8 4.6 22.7 17.7 Philydor novaesi 33873 f 83.5 68.5 76.3 58.8 12.8 4.8 20.8 16.7 32029* m 94.2 76.6 84.2 65.5 13.1 4.1 22.7 18.7 32028 m 94.9 77.6 84.8 67.8 12.3 4.3 22.6 18.5 33872 m 91.4 75.0 79.9 62.7 12.9 4.0 22.2 17.9 m 93.5 1.8 76.4 1.3 82.9 2.7 65.3 2.5 12.8 0.4 4.2 0.2 22.5 0.3 18.4 0.4 P. novaesi + C. mazarbarnetti f 90.0 6.5 74.1 4.9 80.7 4.0 61.9 2.8 13.6 1.5 4.8 0.2 22.1 1.1 17.0 0.6 Philydor atricapillus f 77.9 2.7 63.8 3.0 72.1 2.3 50.7 3.1 11.4 0.4 4.0 0.1 19.7 1.0 16.4 0.7 m 84.2 0.7 69.0 1.5 76.6 2.9 55.1 1.8 12.1 0.4 3.8 0.2 20.1 0.4 16.7 0.5 Philydor pyrrhodes f 77.5 3.9 63.9 2.6 59.1 1.5 46.3 1.2 12.3 0.8 4.4 0.3 19.5 0.6 17.7 0.5 m 87.8 3.9 72.6 2.9 64.9 3.5 53.1 4.3 12.2 0.6 4.1 0.1 21.1 0.3 18.5 0.3 Philydor erythrocercum f 83.3 3.5 66.2 2.6 65.1 4.6 58.3 4.4 11.2 0.2 4.2 0.3 18.9 0.5 14.0 0.4 m 88.7 2.3 71.8 1.9 69.6 3.6 62.2 3.3 10.9 0.7 4.0 0.2 19.8 0.8 15.1 0.7 Philydor fuscipenne f 81.3 0.9 65.3 0.5 63.9 1.6 54.6 2.7 11.2 0.2 4.3 0.2 18.3 0.6 14.5 0.7 m 90.2 1.6 72.9 1.9 69.9 1.2 63.7 1.8 12.4 0.2 4.1 0.1 19.3 0.3 15.6 0.1 Cichlocolaptes l. leucophrus f 98.3 3.0 77.5 2.7 87.0 3.3 64.8 3.0 19.8 0.4 5.4 0.3 23.1 0.5 18.6 0.6 m 103.3 2.8 83.3 2.0 86.5 2.2 68.9 1.6 16.8 0.7 5.1 0.4 23.7 1.1 19.2 1.2 Cichlocolaptes l. holti f 82.9 3.1 67.9 1.8 74.9 2.0 55.7 1.5 16.6 0.4 4.8 0.4 21.5 0.5 17.2 0.3 m 89.0 3.1 73.6 1.8 79.5 2.1 59.1 3.0 15.3 0.5 4.5 0.3 22.5 0.9 17.9 0.6 Pseudocolaptes boissonneautii f 100.9 2.7 83.4 2.4 90.6 4.2 65.0 3.0 17.1 2.1 4.5 0.1 24.4 0.3 18.3 0.5 m 110.4 5.5 89.6 3.0 93.8 6.3 69.2 7.0 13.7 0.8 4.2 0.2 26.1 0.9 20.4 1.9 Pseudocolaptes lawrencii f 103.0 3.4 83.3 3.7 92.1 3.6 66.5 4.3 15.9 0.3 4.6 0.3 25.3 0.4 20.8 0.5 m 3.0 88.4 2.1 99.9 4.9 71.8 4.7 13.6 0.9 4.2 0.3 25.7 0.2 21.1 0.8 110.4 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt TABLE 2. Principal components analysis loadings and variance That would explain behavioral similarities between explained. P. novaesi and Cichlocolaptes noted in the field, and a strong reversed bill size dimorphism that would explain PC 1 PC2 the longer bill of female 34530. Although P. novaesi has always been considered related to P. atricapillus based on Wing length 0.37 0.23 plumage similarity (Teixeira & Gonzaga 1983b; Remsen Secondary 1 0.36 0.22 2003), it has never been the subject of phylogenetic Tail length 0.48 0.15 analysis. P. atricapillus and Cichlocolaptes leucophrus Rectrix 6 0.36 0.27 are part of a clade that also includes P. pyrrhodes and Bill length 0.38 -0.78 Heliobletus contaminatus (Derryberry et al. 2011). P. novaesi may belong to this clade but, at least from a Bill width 0.12 -0.44 morphometric perspective, it is not clear whether it is Tarsus length 0.35 0.04 closer to P. atricapillus or C. leucophrus since it occupies Hallux length 0.35 -0.01 an intermediate position in the morphometric space % variance 67 19 (Figure 1). However, this hypothesis does not explain the morphometric disparity between female specimens is considerably smaller than males, particularly in wing 33873 and 34530 (both adult), neither explain the long and tail lengths; however, the magnitude and direction of wings of female 34530, since even among bromeliad the difference in the morphometric space is as expected specialists, Cichlocolaptes and Pseudocolaptes, females given the strong sexual dimorphism in Philydor. In have considerable shorter wings than males. Although plumage coloration, this specimen is indistinguishable bromeliad specialists Cichlocolaptes and Pseudocolaptes tend from males of P. novaesi (Mazar Barnett & Buzzetti 2014; to be morphometrically heterogeneous (see morphospace pers. obs.). Therefore, phenotypic data suggest that 33873 occupation in Figure 1 compared to Philydor), except for is a true P. novaesi female. the sexual dimorphism in the bill, this heterogeneity most Measurements of female 34531 do not differ much likely represents geographically structured differentiation. from those of males of P. novaesi. Although Mazar Barnett For example, Cichlocolaptes leucophrus is composed of & Buzzetti (2014) concluded that this specimen is larger two very distinct subspecies, C. l. leucophrus in northern than males of P. novaesi, differences in weight (2 grams Atlantic forests and C. l. holti in the south (specimens heavier than the heaviest male) and total length (2 mm with a PC 1 score lower than 0.2; Figure 1). Within longer than the longest male) are relatively minor and each subspecies, levels of variation among females are involved two variables with high measurement error. not greater than those of species of Philydor (Table 1). Female 34531 had a ca. 2 mm smooth ovary according to Pseudocolaptes is also composed of multiple lineages that its label data, indicating immaturity. In addition to its size, vary geographically (Remsen 2003). In contrast, females the plumage of 34531 is closer to that of 34530, the type 33873 and 34530 are from the same geographic locality. of C. mazarbarnetti, than other specimens of P. novaesi, A third possibility is that specimen 34530 is some which led Mazar Barnett & Buzzetti (2014) to speculate sort of aberrant individual of P. novaesi, and similarities that it may represent a juvenile C. mazarbarnetti. Another with Cichlocolaptes are just a coincidence. This hypothesis possibility is that 34531 represents a young male P. novaesi is difficult to test since, in principle, it does not predict and its small testicle was mistaken for a small, smooth a particular pattern of variation or position in the ovary. In any case, because of its young age, it is safer to morphospace in relation to other individuals or species. set this specimen aside for taxonomic considerations. However, it is not a simple case of gigantism in which Female 34530, the holotype of C. mazarbarnetti, an individual is isometrically larger than others in all is different from all specimens of P. novaesi examined. variables. Specimen 34530 is larger than P. novaesi in It is similar to males of P. novaesi in most measurement bill and wing lengths but not in tarsus or hallux lengths. except for its much longer bill. However, given the strong This pattern is unusual for intraspecific variation in birds sexual dimorphism in Philydor, females are not expected since genetic changes that affect forelimb length will to be similar to males. The multivariate analysis placed more likely affect hindlimb length rather than bill length this specimen apart from P. novaesi and in a region of the (Nemeschkal 1999; Magwene 2001). Other aspects of morphospace occupied by Cichlocolaptes. Therefore, the this hypothesis can be tested with additional data. For morphometric data is at least consistent with two aspects example, comparison of levels of asymmetry can be used of Mazar Barnett & Buzzetti’s (2014) hypothesis: that to evaluate whether specimen 34530 experienced an female 34530 is not a specimen of P. novaesi, and that it anomalous development (Palmer & Strobeck 1986). belongs to Cichlocolaptes. I conclude that the existence of a new Cichlocolaptes One alternative to this hypothesis is that P. novaesi species, C. mazarbarnetti, is a plausible explanation for itself is a Cichlocolaptes, or at least, a bromeliad specialist. the morphometric data analyzed. The reduced number Revista Brasileira de Ornitologia, 22(2), 2014 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt Derryberry, E. P.; Claramunt, S.; Chesser, R. T.; Remsen Jr., J. of specimens available limits any definite conclusion and V.; Cracraft, J.; Aleixo, A. & Brumfield, R. T. 2011. Lineage any taxonomic recommendation should consider also diversification and morphological evolution in a large-scale the morphological and behavioral evidence presented by continental radiation: the Neotropical ovenbirds and woodcreepers Mazar Barnett & Buzzetti (2014). A source of information (Aves: Furnariidae). Evolution, 65: 2973-2986. that may be powerful in this case is genetic material Magwene, P. M. 2001. New tools for studying integration and modularity. Evolution, 55: 1734-1745. extracted from study skins (Mundy et al. 1997) with which Martínez, O. 2003. Composición por especies y uso de sustratos the phylogenetic affinities of P. novaesi and specimen por las bandadas mixtas de aves en un bosque nublado andino de 34530 can be determined with more confidence. Given Bolivia. Ecología en Bolivia, 38: 99-119. the high levels of genetic divergence between members of Mazar Barnett, J.; Carlos, C. J. & Roda, S. A. 2005. Renewed hope for the threatened avian endemics of northeastern Brazil. the Cichlocolaptes-Philydor clade (Derryberry et al. 2011), Biodiversity and Conservation, 14: 2265-2274. even a short fragment of mitochondrial DNA may prove Mazar Barnett, J. & Buzzetti, D. 2014. The ‘Gritador-do-Nordeste’, useful in determining relationships. an undescribed trace of the fading life in northeastern Brazil. Revista Brasileira de Ornotologia. Mundy, N. I.; Unitt, P. & Woodruff, D. S. 1997.Skin from feet of museum specimens as a non-destructive source of DNA for avian ACKNOWLEDGEMENTS genotyping. Auk, 114: 126-129. Nemeschkal, H. L. 1999. Morphometric correlation patterns of adult I am grateful to curators, researchers, and personnel that birds (Fringillidae: Passeriformes and Columbiformes) mirror the facilitated the examination of specimens for this study, expression of developmental control genes. Evolution, 53: 899-918. Palmer, A. R. & Strobeck, C. 1986. Fluctuating asymmetry: in particular to Marcos Raposo (Museu Nacional, Rio measurements, analysis and patterns. Annual Review of Ecology, de Janeiro), and Luis Fábio Silveira and Vítor de Q. Evolution, and Systematics, 17: 391-421. Piacentini (Universidade de São Paulo). I would also like Remsen, J. V., Jr. 2003. Family Furnariidae (ovenbirds), p. 162-357. to thank Cathy Bechtoldt and two anonymous reviewers In: del Hoyo, J.; Elliott, A.; & Christie, D. A. (eds.) Handbook of for comments and suggestions. My work at the American the birds of the world, Vol. 8, broadbills to tapaculos. Barcelona: Lynx Edicions. Museum of Natural History was supported by a Chapman Sillett, T. S.; James, A. & Sillett, K. B. 1997. Bromeliad foraging Postdoctoral Fellowship. specialization and diet selection of Pseudocolaptes lawrencii (Furnariidae). Ornithological Monographs, 48: 733-742. Silva, J. M. C.; Coelho, G. & Gonzaga, L. P. 2002. Discovered on the brink of extinction: a new species of Pygmy-Owl (Strigidae: REFERENCES Glaucidium) from Atlantic Forest of northeastern Brazil. Ararajuba, 10: 123-130. Baldwin, S. P.; Oberholser, H. C. & Worley, L. G. 1931. Teixeira, D. M. 1987. A new tyrannulet (Phylloscartes) from Measurements of birds. Scientific Publications of the Cleveland northeastern Brazil. Bulletin of the British Ornithologists’ Club, Museum of Natural History, 2:1–165. 107: 37-41. BirdLife International. 2013. Species factsheet: Philydor novaesi. Teixeira, D. M. & Gonzaga, L. P. 1983a. A new antwren from http://www.birdlife.org (access on 27 November 2013). northeastern Brazil. Bulletin of the British Ornithologists’ Club, Claramunt, S. 2010. Discovering exceptional diversifications at 103: 133-135. continental scales: the case of the endemic families of Neotropical Teixeira, D. M. & Gonzaga, L. P. 1983b. Um novo Furnariidae do suboscine passerines. Evolution, 64:2004-2019. nordeste do Brasil: Philydor novaesi sp. nov. Boletim do Museu Claramunt, S.; Derryberry, E. P.; Cadena, C. D.; Cuervo, A. M.; Paraense Emílio Goeldi, 124: 1-22. Sanín, C. & Brumfield, R. T. 2013. Phylogeny and classification of Automolus foliage-gleaners and allies (Furnariidae). Condor, 115: 375-385. Associate Editor: Luciano N. Naka Revista Brasileira de Ornitologia, 22(2), 2014 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt APPENDIX: Specimens examined and measured in the collections of the American Museum of Natural History (AMNH), Louisiana State University Museum of Natural Science (LSUMZ), Museu Nacional, Rio de Janeiro (MN), and Museu de Zoologia, Universidade de São Paulo (MZUSP): Cichlocolaptes mazarbarnetti. MN 34530 (holotype) adult female collected on 16 January 1986, 48 g, globulous 12 mm ovary with one ovum > 2 mm, ossified skull, 221 mm total length; MN 34531 (tentative assignation), young female collected on 20 January 1986, 36 g, smooth 2 mm ovary, according to a drawing in the label, 207 mm total length. Cichlocolaptes leucophrus leucophrus. Males: AMNH 243297, 317611, MZUSP 33324, 34530. Females: AMNH 317612, 316817, MZUSP 33322, 28506. Cichlocolaptes leucophrus holti. Males: AMNH 314749, 314748, 524356, LSUMZ 68015, 31666, MZUSP 49762. Females: AMNH 314750, 314751, 314752, 524357, LSUMZ 53001, 63359, MZUSP 32147, 54949. Philydor novaesi. MN 32029 (holotype) adult male collected on 7 November 1979, 32 g., enlarged testis, 205 mm total length; MN 32028 (paratype) adult male, 34 g, enlarged testis, 195 mm total length; MN 33872, male, 32 g, small testis; 33873: adult female collected on 21 November 1983, 30 g, granulated 9 mm ovary, ossified skull, 195 mm total length. Philydor atricapillus. Males: LSUMZ 63354, 70433, AMNH 314701, 243301. Females: LSUMZ 62951, AMNH 243302, 243303, 317614, 524190. Philydor pyrrhodes. Males: LSUMZ 109768, 115010, 156423, AMNH 286828. Females:LSUMZ 137071, AMNH 430943, 824593, 142492, 274140. Philydor erythrocercum. Males: LSUMZ 132520, 87888, 175383, 96027. Females: AMNH: 429578, 256099, 283973, 819937. Philydor fuscipenne. Males: LSUMZ 108297, AMNH 246775, 136619. Females: AMNH 135827, 136620, 135826. Pseudocolaptes boissonneautii. Males: AMNH 124519, 167340, 124520, 820955, 820474, LSUMZ 45349, 178990, 169854. Females: AMNH 820956, 820779, 820420, LSUMZ 81936. Pseudocolaptes lawrencii. Males: LSUMZ 63643, 154029, AMNH 524041, 102195, 102196. Females: AMNH 811839, LSUMZ 154031. Revista Brasileira de Ornitologia, 22(2), 2014 http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Ornithology Research Springer Journals

Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil

Ornithology Research , Volume 22 (2) – Jun 1, 2014

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Revista Brasileira de Ornitologia, 22(2), 95-101 ARTICLE June 2014 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt Department of Ornithology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024, USA. E-mail: sclaramunt@amnh.org Received on 8 January 2014. Accepted on 10 May 2014. ABSTRACT: Mazar Barnett & Buzzetti (2014) described a new species, Cichlocolaptes mazarbarnetti, from the Atlantic forests of northeastern Brazil. The holotype of the new species is a female that shows remarkable similarities with the sympatric Philydor novaesi. Here I analyze eight morphometric variables to assess phenotypic similarities among specimens of Philydor novaesi, Cichlocolaptes mazarbarnetti and other species in these two genera. The holotype of C. mazarbarnetti differs from Philydor novaesi and falls in a region of the morphometric space occupied by specimens of Cichlocolaptes leucophrus. Therefore, morphometric data is consistent with other morphological, behavioral, and vocal data that suggest the existence of a new species Cichlocolaptes in the northern Atlantic forest. KEYWORDS: Cichlocolaptes mazarbarnetti, Philydor novaesi, Furnariidae, measurements, taxonomy. INTRODUCTION originally identified as P. novaesi is an exemplar of the new foliage-gleaner; diagnostic characters include a larger The avifauna of the northern stretch of the Atlantic body size, a longer beak, absence of buffy orbital feathers, Forest is one of the least known, yet most endangered on and rounded tail feather tips. Although Mazar Barnett & Buzzetti (2014) measured and discussed some biometric Earth. Habitat destruction has outpaced ornithological research and conducting research on some species today data, comparisons were limited to single variables is difficult because of their scarcity and vulnerability. The and few species. As part of a large-scale morphometric inventory of the taxonomic composition of this avifauna analysis of the Furnariidae (Claramunt 2010; Claramunt is incomplete, as demonstrated by continuous discoveries et al. 2013), I examined and measured all specimens identified as P. novaesi at the Museu Nacional, including of new species in the last decades (e.g. Teixeira 1987; Teixeira & Gonzaga 1983, Silva et al. 2002). One such the holotypes of P. novaesi and C. mazarbarnetti. Here I species is the Alagoas foliage-gleaner Philydor novaesi, present a morphometric analysis of those specimens and discovered at Pedra Branca, Muricí, a relict of foothill discuss its implications regarding their taxonomic status. forest in Alagoas State (Teixeira & Gonzaga 1983b), and found later in a small reserve in Pernambuco State (Mazar METHODS Barnett et al. 2005). The species is considered critically endangered (BirdLife International 2013). In a surprising turn of events, Mazar Barnett & Buzzetti (2014) Measurements were taken with a Mitutoyo Digimatic described yet another new foliage-gleaner from Murici, Point Caliper (resolution: 0.01 mm) with an output interface. Variables were the same as in previous Cichlocolaptes mazarbarnetti, based on field observations morphometric studies of the Furnariidae (Claramunt and examination of museum specimens. The new species is extremely similar to P. novaesi in plumage pattern and et al. 2010; Claramunt et al. 2013). Here, I analyzed color but its behavior and vocalizations resemble those of only two variables from each body region because some Cichlocolaptes leucophrus. In particular, like C. leucophrus, measurements were highly correlated in foliage-gleaners: 1) wing length; 2) wing length to the first secondary the new species shows a preference for foraging on feather; 3) tail length to central rectrices; 4) tail length to bromeliads. Upon examination of specimens identified as P. novaesi at the Museu Nacional in Rio de Janeiro, Mazar the most external rectrix; 5) bill length from the anterior Barnett & Buzzetti (2014) concluded that one specimen border of the nostril to the tip of the bill; 6) bill width at Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt the level of the anterior border of the nostrils; 7) tarsus Cichlocolaptes examined. Given a ratio of 1.32, rectrix 1 length; and 8) hallux length with claw (Baldwin et al. should be around 82.6 mm when fully-grown, not much 1931; Claramunt et al. 2010). longer than the actual measurement (Table 1). All known specimens of Philydor novaesi and the Two principal components explained 86% of holotype of Cichlocolaptes mazarbarnetti were examined the variance among all specimens measured (Table 2). and measured (Appendix). For comparison, I included Principal component 1 is positively correlated with all in the analysis other Philydor and Cichlocolaptes taxa. variables; thus, it is associated with overall size. Principal Although DNA sequences of P. novaesi are not available, component 2 is a contrast between variables of the bill its overall morphology and plumage suggest a close and variables of the flight apparatus (wings and tail). The relationship with P. atricapillus (Teixeira & Gonzaga three genera, Philydor, Cichlocolaptes, and Pseudocolaptes, 1983b) from southeastern Brazil, which belongs to a occupy different parts of the morphospace defined clade including P. pyrrhodes, Cichlocolaptes leucophrus, by components 1 and 2, with limited overlap (Figure and Heliobletus contaminatus (Derryberry et al. 2011; 1). Within each genus, females and males are almost Appendix). I did not include H. contaminatus in the completely segregated. Females tend to have lower scores analysis because of its small size and very distinctive than males in both components, indicating smaller morphology compared to the other species examined. overall size but relatively larger bills. P. novaesi is in an In addition, I analyzed specimens of P. fuscipenne and intermediate position among the three genera. The three P. erythrocercum (Appendix), which are more distantly males of P. novaesi are tightly clustered, with intermediate related but share several phenotypic similarities with scores on component 1 (intermediate size) and high scores P. novaesi, and cannot be discarded as potential close on component 2 (relatively long wings and tail but short relatives. Finally, I also analyzed specimens of the genus bill). Females, in contrast, are heterogeneous. Female Pseudocolaptes (Appendix), another genus of Furnariidae 33873 has lower scores in both components, compared that specialized in bromeliad foraging (Sillet et al. 1997, to males, and its position matches the relative position Martínez 2003), to investigate patterns of morphological of females due to sexual dimorphism. Female 34531, variation related to bromeliad specialization. I examined in contrast, is positioned near males P. novaesi. Female scatter plots of all variables obtained and conducted a 34530, the type of C. mazarbarnetti, has a slightly larger Principal Components Analysis on the covariance matrix component 1 score but a considerably lower component of log-transformed values for all measured specimens. 2 score compared to male P. novaesi; has a result, it is positioned in a region of the morphospace occupied by Cichlocolaptes leucophrus, and closer to females of RESULTS Pseudocolaptes than to males of P. novaesi. The use of the estimated fully-grown tail length of 34530 instead of its All species analyzed showed sexual dimorphism in actual length had a negligible effect on the position of this which males are larger than females (Table 1). The size specimen in the multivariate space. dimorphism is particularly pronounced in wing and A simple plot of wing and bill length shows a similar tail length, for which males of some species average 10 pattern (Figure 2). P. novaesi specimens are closer to other mm longer than females. Exceptions to this pattern are species of Philydor. Note, again, the nearly complete Cichlocolaptes and Pseudocolaptes, which show reversed segregation of males and females along the wing-length axis, sexual dimorphism in bill length, with females having and, for Cichlocolaptes and Pseudocolaptes, along the bill- longer bills (Table 1). P. novaesi is larger than other species length axis. Again, female 34530 is closer to Cichlocolaptes of Philydor. Whereas males are relatively uniform (small and Pseudocolaptes rather than to other specimens of P. standard deviations), females previously assigned to P. novaesi and specimens of Philydor in general. novaesi (including the holotype of C. mazarbarnetti) are unusually heterogeneous, particularly in wing, bill, and tarsus length (Table 1). Female 33873 is about DISCUSSION 90% smaller than males’ averages in wing, tail, and feet variables. Female 34531 is about the same size as males in The morphometric analysis presented here provides a all variables. Female 34530, the type of C. mazarbarnetti, quantitative assessment of all specimens of P. novaesi and has long wings and bill, surpassing all specimens of P. C. mazarbarnetti (Mazar Barnett & Buzzetti 2014). The novaesi, including males. The tail of 34530 is not longer analysis indicates that P. novaesi is larged compared to the than that of P. novaesi, however, rectrices 1 and 2 may not other species of Philydor, evident in both univariate and be fully grown, as their rachises still show basal sheaths. It multivariate analyses (Table 1, Fig. 1 and 2). The three is possible to estimate the expected tail length of 34530 male specimens of P. novaesi are fairly homogeneous, from the length of rectrix 6, using the average ratio between occupying a small sector of the morphospace, but the these two measurement among specimens of Philydor and three females are unusually heterogeneous. Female 33873 Revista Brasileira de Ornitologia, 22(2), 2014 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt FIGURE 1. Principal component analysis of eight morphometric variables for male (plain symbols) and female (cross) specimens of Philydor, Cichlocolaptes, and Pseudocolaptes, including Philydor novaesi (black squares), the holotype of Cichlocolaptes mazarbarnetti (black star) and a specimen of uncertain affinities (black circle). Numbers below symbols are Museu Nacional specimen numbers. An approximate interpretation is given for each component but see Table 2 for variable loadings. FIGURE 2. Position of male (plain) and female (cross) specimens of Philydor, Cichlocolaptes, and Pseudocolaptes in the morphometric space defined by wing length and bill length, includine specimens of Philydor novaesi (black squares),dthe holotype of Cichlocolaptes mazarbarnetti (black star) and a specimen of uncertain affinities (black symbol). Numbers below symbols are Museu Nacional(specimen number). Revista Brasileira de Ornitologia, 22(2), 2014 Santiago Claramunt fj Revista Brasileira de Ornitologia, 22(2), 2014 TABLE 1. Morphometric data for species of Philydor, Cichlocolaptes, and Pseudocolaptes, including individual data for all specimens of Philydor novaesi and Cichlocolaptes mazarbarneti (holotypes marked with *), and average and standard deviations for a combination of female specimens of P. novaesi and C. mazarbarnetti, which were originally identified as the former species only. See discussion for the placement of MN 34531 in P. novaesi or C. mazarbarnetti. wing secondary 1 rectrix 1 rectrix 6 bill length bill width tarsus hallux _ _ _ _ _ _ _ _ Taxon sex X, SD X, SD X, SD X, SD X, SD X, SD X, SD X, SD Cichlocolaptes mazarbarnetti 34530* f 96.5 77.7 81.8 62.5 15.3 4.9 22.7 16.7 Cichlocolaptes mazarbarnetti? 34531 90.1 76.0 84.0 64.4 12.8 4.6 22.7 17.7 Philydor novaesi 33873 f 83.5 68.5 76.3 58.8 12.8 4.8 20.8 16.7 32029* m 94.2 76.6 84.2 65.5 13.1 4.1 22.7 18.7 32028 m 94.9 77.6 84.8 67.8 12.3 4.3 22.6 18.5 33872 m 91.4 75.0 79.9 62.7 12.9 4.0 22.2 17.9 m 93.5 1.8 76.4 1.3 82.9 2.7 65.3 2.5 12.8 0.4 4.2 0.2 22.5 0.3 18.4 0.4 P. novaesi + C. mazarbarnetti f 90.0 6.5 74.1 4.9 80.7 4.0 61.9 2.8 13.6 1.5 4.8 0.2 22.1 1.1 17.0 0.6 Philydor atricapillus f 77.9 2.7 63.8 3.0 72.1 2.3 50.7 3.1 11.4 0.4 4.0 0.1 19.7 1.0 16.4 0.7 m 84.2 0.7 69.0 1.5 76.6 2.9 55.1 1.8 12.1 0.4 3.8 0.2 20.1 0.4 16.7 0.5 Philydor pyrrhodes f 77.5 3.9 63.9 2.6 59.1 1.5 46.3 1.2 12.3 0.8 4.4 0.3 19.5 0.6 17.7 0.5 m 87.8 3.9 72.6 2.9 64.9 3.5 53.1 4.3 12.2 0.6 4.1 0.1 21.1 0.3 18.5 0.3 Philydor erythrocercum f 83.3 3.5 66.2 2.6 65.1 4.6 58.3 4.4 11.2 0.2 4.2 0.3 18.9 0.5 14.0 0.4 m 88.7 2.3 71.8 1.9 69.6 3.6 62.2 3.3 10.9 0.7 4.0 0.2 19.8 0.8 15.1 0.7 Philydor fuscipenne f 81.3 0.9 65.3 0.5 63.9 1.6 54.6 2.7 11.2 0.2 4.3 0.2 18.3 0.6 14.5 0.7 m 90.2 1.6 72.9 1.9 69.9 1.2 63.7 1.8 12.4 0.2 4.1 0.1 19.3 0.3 15.6 0.1 Cichlocolaptes l. leucophrus f 98.3 3.0 77.5 2.7 87.0 3.3 64.8 3.0 19.8 0.4 5.4 0.3 23.1 0.5 18.6 0.6 m 103.3 2.8 83.3 2.0 86.5 2.2 68.9 1.6 16.8 0.7 5.1 0.4 23.7 1.1 19.2 1.2 Cichlocolaptes l. holti f 82.9 3.1 67.9 1.8 74.9 2.0 55.7 1.5 16.6 0.4 4.8 0.4 21.5 0.5 17.2 0.3 m 89.0 3.1 73.6 1.8 79.5 2.1 59.1 3.0 15.3 0.5 4.5 0.3 22.5 0.9 17.9 0.6 Pseudocolaptes boissonneautii f 100.9 2.7 83.4 2.4 90.6 4.2 65.0 3.0 17.1 2.1 4.5 0.1 24.4 0.3 18.3 0.5 m 110.4 5.5 89.6 3.0 93.8 6.3 69.2 7.0 13.7 0.8 4.2 0.2 26.1 0.9 20.4 1.9 Pseudocolaptes lawrencii f 103.0 3.4 83.3 3.7 92.1 3.6 66.5 4.3 15.9 0.3 4.6 0.3 25.3 0.4 20.8 0.5 m 3.0 88.4 2.1 99.9 4.9 71.8 4.7 13.6 0.9 4.2 0.3 25.7 0.2 21.1 0.8 110.4 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt TABLE 2. Principal components analysis loadings and variance That would explain behavioral similarities between explained. P. novaesi and Cichlocolaptes noted in the field, and a strong reversed bill size dimorphism that would explain PC 1 PC2 the longer bill of female 34530. Although P. novaesi has always been considered related to P. atricapillus based on Wing length 0.37 0.23 plumage similarity (Teixeira & Gonzaga 1983b; Remsen Secondary 1 0.36 0.22 2003), it has never been the subject of phylogenetic Tail length 0.48 0.15 analysis. P. atricapillus and Cichlocolaptes leucophrus Rectrix 6 0.36 0.27 are part of a clade that also includes P. pyrrhodes and Bill length 0.38 -0.78 Heliobletus contaminatus (Derryberry et al. 2011). P. novaesi may belong to this clade but, at least from a Bill width 0.12 -0.44 morphometric perspective, it is not clear whether it is Tarsus length 0.35 0.04 closer to P. atricapillus or C. leucophrus since it occupies Hallux length 0.35 -0.01 an intermediate position in the morphometric space % variance 67 19 (Figure 1). However, this hypothesis does not explain the morphometric disparity between female specimens is considerably smaller than males, particularly in wing 33873 and 34530 (both adult), neither explain the long and tail lengths; however, the magnitude and direction of wings of female 34530, since even among bromeliad the difference in the morphometric space is as expected specialists, Cichlocolaptes and Pseudocolaptes, females given the strong sexual dimorphism in Philydor. In have considerable shorter wings than males. Although plumage coloration, this specimen is indistinguishable bromeliad specialists Cichlocolaptes and Pseudocolaptes tend from males of P. novaesi (Mazar Barnett & Buzzetti 2014; to be morphometrically heterogeneous (see morphospace pers. obs.). Therefore, phenotypic data suggest that 33873 occupation in Figure 1 compared to Philydor), except for is a true P. novaesi female. the sexual dimorphism in the bill, this heterogeneity most Measurements of female 34531 do not differ much likely represents geographically structured differentiation. from those of males of P. novaesi. Although Mazar Barnett For example, Cichlocolaptes leucophrus is composed of & Buzzetti (2014) concluded that this specimen is larger two very distinct subspecies, C. l. leucophrus in northern than males of P. novaesi, differences in weight (2 grams Atlantic forests and C. l. holti in the south (specimens heavier than the heaviest male) and total length (2 mm with a PC 1 score lower than 0.2; Figure 1). Within longer than the longest male) are relatively minor and each subspecies, levels of variation among females are involved two variables with high measurement error. not greater than those of species of Philydor (Table 1). Female 34531 had a ca. 2 mm smooth ovary according to Pseudocolaptes is also composed of multiple lineages that its label data, indicating immaturity. In addition to its size, vary geographically (Remsen 2003). In contrast, females the plumage of 34531 is closer to that of 34530, the type 33873 and 34530 are from the same geographic locality. of C. mazarbarnetti, than other specimens of P. novaesi, A third possibility is that specimen 34530 is some which led Mazar Barnett & Buzzetti (2014) to speculate sort of aberrant individual of P. novaesi, and similarities that it may represent a juvenile C. mazarbarnetti. Another with Cichlocolaptes are just a coincidence. This hypothesis possibility is that 34531 represents a young male P. novaesi is difficult to test since, in principle, it does not predict and its small testicle was mistaken for a small, smooth a particular pattern of variation or position in the ovary. In any case, because of its young age, it is safer to morphospace in relation to other individuals or species. set this specimen aside for taxonomic considerations. However, it is not a simple case of gigantism in which Female 34530, the holotype of C. mazarbarnetti, an individual is isometrically larger than others in all is different from all specimens of P. novaesi examined. variables. Specimen 34530 is larger than P. novaesi in It is similar to males of P. novaesi in most measurement bill and wing lengths but not in tarsus or hallux lengths. except for its much longer bill. However, given the strong This pattern is unusual for intraspecific variation in birds sexual dimorphism in Philydor, females are not expected since genetic changes that affect forelimb length will to be similar to males. The multivariate analysis placed more likely affect hindlimb length rather than bill length this specimen apart from P. novaesi and in a region of the (Nemeschkal 1999; Magwene 2001). Other aspects of morphospace occupied by Cichlocolaptes. Therefore, the this hypothesis can be tested with additional data. For morphometric data is at least consistent with two aspects example, comparison of levels of asymmetry can be used of Mazar Barnett & Buzzetti’s (2014) hypothesis: that to evaluate whether specimen 34530 experienced an female 34530 is not a specimen of P. novaesi, and that it anomalous development (Palmer & Strobeck 1986). belongs to Cichlocolaptes. I conclude that the existence of a new Cichlocolaptes One alternative to this hypothesis is that P. novaesi species, C. mazarbarnetti, is a plausible explanation for itself is a Cichlocolaptes, or at least, a bromeliad specialist. the morphometric data analyzed. The reduced number Revista Brasileira de Ornitologia, 22(2), 2014 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt Derryberry, E. P.; Claramunt, S.; Chesser, R. T.; Remsen Jr., J. of specimens available limits any definite conclusion and V.; Cracraft, J.; Aleixo, A. & Brumfield, R. T. 2011. Lineage any taxonomic recommendation should consider also diversification and morphological evolution in a large-scale the morphological and behavioral evidence presented by continental radiation: the Neotropical ovenbirds and woodcreepers Mazar Barnett & Buzzetti (2014). A source of information (Aves: Furnariidae). Evolution, 65: 2973-2986. that may be powerful in this case is genetic material Magwene, P. M. 2001. New tools for studying integration and modularity. Evolution, 55: 1734-1745. extracted from study skins (Mundy et al. 1997) with which Martínez, O. 2003. Composición por especies y uso de sustratos the phylogenetic affinities of P. novaesi and specimen por las bandadas mixtas de aves en un bosque nublado andino de 34530 can be determined with more confidence. Given Bolivia. Ecología en Bolivia, 38: 99-119. the high levels of genetic divergence between members of Mazar Barnett, J.; Carlos, C. J. & Roda, S. A. 2005. Renewed hope for the threatened avian endemics of northeastern Brazil. the Cichlocolaptes-Philydor clade (Derryberry et al. 2011), Biodiversity and Conservation, 14: 2265-2274. even a short fragment of mitochondrial DNA may prove Mazar Barnett, J. & Buzzetti, D. 2014. The ‘Gritador-do-Nordeste’, useful in determining relationships. an undescribed trace of the fading life in northeastern Brazil. Revista Brasileira de Ornotologia. Mundy, N. I.; Unitt, P. & Woodruff, D. S. 1997.Skin from feet of museum specimens as a non-destructive source of DNA for avian ACKNOWLEDGEMENTS genotyping. Auk, 114: 126-129. Nemeschkal, H. L. 1999. Morphometric correlation patterns of adult I am grateful to curators, researchers, and personnel that birds (Fringillidae: Passeriformes and Columbiformes) mirror the facilitated the examination of specimens for this study, expression of developmental control genes. Evolution, 53: 899-918. Palmer, A. R. & Strobeck, C. 1986. Fluctuating asymmetry: in particular to Marcos Raposo (Museu Nacional, Rio measurements, analysis and patterns. Annual Review of Ecology, de Janeiro), and Luis Fábio Silveira and Vítor de Q. Evolution, and Systematics, 17: 391-421. Piacentini (Universidade de São Paulo). I would also like Remsen, J. V., Jr. 2003. Family Furnariidae (ovenbirds), p. 162-357. to thank Cathy Bechtoldt and two anonymous reviewers In: del Hoyo, J.; Elliott, A.; & Christie, D. A. (eds.) Handbook of for comments and suggestions. My work at the American the birds of the world, Vol. 8, broadbills to tapaculos. Barcelona: Lynx Edicions. Museum of Natural History was supported by a Chapman Sillett, T. S.; James, A. & Sillett, K. B. 1997. Bromeliad foraging Postdoctoral Fellowship. specialization and diet selection of Pseudocolaptes lawrencii (Furnariidae). Ornithological Monographs, 48: 733-742. Silva, J. M. C.; Coelho, G. & Gonzaga, L. P. 2002. Discovered on the brink of extinction: a new species of Pygmy-Owl (Strigidae: REFERENCES Glaucidium) from Atlantic Forest of northeastern Brazil. Ararajuba, 10: 123-130. Baldwin, S. P.; Oberholser, H. C. & Worley, L. G. 1931. Teixeira, D. M. 1987. A new tyrannulet (Phylloscartes) from Measurements of birds. Scientific Publications of the Cleveland northeastern Brazil. Bulletin of the British Ornithologists’ Club, Museum of Natural History, 2:1–165. 107: 37-41. BirdLife International. 2013. Species factsheet: Philydor novaesi. Teixeira, D. M. & Gonzaga, L. P. 1983a. A new antwren from http://www.birdlife.org (access on 27 November 2013). northeastern Brazil. Bulletin of the British Ornithologists’ Club, Claramunt, S. 2010. Discovering exceptional diversifications at 103: 133-135. continental scales: the case of the endemic families of Neotropical Teixeira, D. M. & Gonzaga, L. P. 1983b. Um novo Furnariidae do suboscine passerines. Evolution, 64:2004-2019. nordeste do Brasil: Philydor novaesi sp. nov. Boletim do Museu Claramunt, S.; Derryberry, E. P.; Cadena, C. D.; Cuervo, A. M.; Paraense Emílio Goeldi, 124: 1-22. Sanín, C. & Brumfield, R. T. 2013. Phylogeny and classification of Automolus foliage-gleaners and allies (Furnariidae). Condor, 115: 375-385. Associate Editor: Luciano N. Naka Revista Brasileira de Ornitologia, 22(2), 2014 Morphometric insights into the existence of a new species of Cichlocolaptes in northeastern Brazil Santiago Claramunt APPENDIX: Specimens examined and measured in the collections of the American Museum of Natural History (AMNH), Louisiana State University Museum of Natural Science (LSUMZ), Museu Nacional, Rio de Janeiro (MN), and Museu de Zoologia, Universidade de São Paulo (MZUSP): Cichlocolaptes mazarbarnetti. MN 34530 (holotype) adult female collected on 16 January 1986, 48 g, globulous 12 mm ovary with one ovum > 2 mm, ossified skull, 221 mm total length; MN 34531 (tentative assignation), young female collected on 20 January 1986, 36 g, smooth 2 mm ovary, according to a drawing in the label, 207 mm total length. Cichlocolaptes leucophrus leucophrus. Males: AMNH 243297, 317611, MZUSP 33324, 34530. Females: AMNH 317612, 316817, MZUSP 33322, 28506. Cichlocolaptes leucophrus holti. Males: AMNH 314749, 314748, 524356, LSUMZ 68015, 31666, MZUSP 49762. Females: AMNH 314750, 314751, 314752, 524357, LSUMZ 53001, 63359, MZUSP 32147, 54949. Philydor novaesi. MN 32029 (holotype) adult male collected on 7 November 1979, 32 g., enlarged testis, 205 mm total length; MN 32028 (paratype) adult male, 34 g, enlarged testis, 195 mm total length; MN 33872, male, 32 g, small testis; 33873: adult female collected on 21 November 1983, 30 g, granulated 9 mm ovary, ossified skull, 195 mm total length. Philydor atricapillus. Males: LSUMZ 63354, 70433, AMNH 314701, 243301. Females: LSUMZ 62951, AMNH 243302, 243303, 317614, 524190. Philydor pyrrhodes. Males: LSUMZ 109768, 115010, 156423, AMNH 286828. Females:LSUMZ 137071, AMNH 430943, 824593, 142492, 274140. Philydor erythrocercum. Males: LSUMZ 132520, 87888, 175383, 96027. Females: AMNH: 429578, 256099, 283973, 819937. Philydor fuscipenne. Males: LSUMZ 108297, AMNH 246775, 136619. Females: AMNH 135827, 136620, 135826. Pseudocolaptes boissonneautii. Males: AMNH 124519, 167340, 124520, 820955, 820474, LSUMZ 45349, 178990, 169854. Females: AMNH 820956, 820779, 820420, LSUMZ 81936. Pseudocolaptes lawrencii. Males: LSUMZ 63643, 154029, AMNH 524041, 102195, 102196. Females: AMNH 811839, LSUMZ 154031. Revista Brasileira de Ornitologia, 22(2), 2014

Journal

Ornithology ResearchSpringer Journals

Published: Jun 1, 2014

Keywords: Cichlocolaptes mazarbarnetti; Philydor novaesi; Furnariidae; measurements; taxonomy

References