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Circumscription of a Monophyletic Family for the Tapaculos (Aves: Rhinocryptidae): Psiloramphus in and Melanopareia Out

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Circumscription of a Monophyletic Family for the Tapaculos (Aves: Rhinocryptidae): Psiloramphus in and Melanopareia Out J Ornithol (2010) 151:337–345 DOI 10.1007/s10336-009-0460-9 ORIGINAL ARTICLE Circumscription of a monophyletic family for the tapaculos (Aves: Rhinocryptidae): Psiloramphus in and Melanopareia out Per G. P. Ericson • Storrs L. Olson • Martin Irestedt • Herculano Alvarenga • Jon Fjeldsa˚ Received: 22 February 2009 / Revised: 3 August 2009 / Accepted: 21 September 2009 / Published online: 14 October 2009 Ó Dt. Ornithologen-Gesellschaft e.V. 2009 Abstract The tapaculos (Rhinocryptidae) are tracheo- within the Rhinocryptidae, Melanopareia falls far outside phone, suboscine birds restricted to South and Central that clade. A new family is erected for Melanopareia. America. Most tapaculos share a number of internal and external characteristics that have been used to define the Keywords Melanopareia Á Psiloramphus Á family taxonomically. The genera Melanopareia and Rhinocryptidae Á Tapaculos Á Molecular systematics Á Psiloramphus do not fully fit this pattern and have caused Taxonomy Á South America considerable dispute among taxonomists since they were first described. In this paper we delimit the systematic boundaries of the tapaculos and assess their generic rela- Introduction tionships by analysis of molecular sequence data. The results show that whereas Psiloramphus is nested well The impudently named tapaculos (Rhinocryptidae) are a small group of tracheophone, suboscine passerines whose greatest generic diversity is in southern South America. Most are large-footed, strong-legged ground birds remi- Communicated by M. Wink. niscent of some of the ground-dwelling antthrushes (For- micariidae), with which they were often associated. The & P. G. P. Ericson ( ) family is generally well defined by the presence of oper- Department of Vertebrate Zoology, Swedish Museum of Natural History, culate nostrils, a tracheophone syrinx, a somewhat curved P.O. Box 50007, 10405 Stockholm, Sweden humerus, and a four-notched sternum (Ames 1971; e-mail: [email protected] Feduccia and Olson 1982; Maurı´cio et al. 2008). Thus, Krabbe and Schulenberg (2003), p. 748, considered that the S. L. Olson Division of Birds, Department of Vertebrate Zoology, ‘‘tapaculos constitute a well-knit group the members of Smithsonian Institution, National Museum of Natural History, which are united by several derived characters. Only the P.O. Box 37012, Washington, DC 20013-7012, USA genera Melanopareia and Psiloramphus differ to such a degree that their systematic position as tapaculos could be M. Irestedt Molecular Systematics Laboratorium, disputed.’’ Furthermore, the phylogenetic relationships Swedish Museum of Natural History, within the family Rhinocryptidae are poorly known, par- P.O. Box 50007, 10405 Stockholm, Sweden ticularly regarding the placement of Liosceles and Acropternis, and the large austral species of Pteroptochos H. Alvarenga Museu de Histo´ria Natural de Taubate´, and Scelorchilus. Rua Juvenal Dias de Carvalho, 111, Taubate´, The early taxonomic history of the group was ably SP 12070-640, Brazil summarized by Sclater (1874). d’Orbigny (1837) first erected a family ‘‘Rhinomyidaeae’’ (sic., p. 192) for Pter- J. Fjeldsa˚ Zoological Museum, University of Copenhagen, optochos and his Rhinomya (= Rhinocrypta) using the Universitetsparken 15, 2100 Copenhagen, Denmark operculate nostril to separate them from the Formicariidae. 123 338 J Ornithol (2010) 151:337–345 However, there was no coherent understanding of the su- boscine groups until their distinction from the oscines was established by the pioneering work of Mu¨ller (1847) on the syrinx. Mu¨ller showed that Scytalopus was a tracheophone suboscine and not a wren (Troglodytidae) and also that Scytalopus and Pteroptochus differed from other known passerines in having a four-notched sternum. Sclater’s (1874) ‘‘Pteroptochidae’’ comprised Scytal- opus (including the type species of what later became Myornis Chapman 1915), Merulaxis, Liosceles, Pteropto- chos (including the type species of what later became Teledromas Wetmore and Peters 1922, and Scelorchilus Oberholser 1923), Rhinocrypta, Hylactes (now included in Pteroptochos), Acropternis, and Triptorhinus (= Eugralla). Except for the problem genera Psiloramphus and Mela- nopareia, this composition of the group was essentially maintained until Peters (1951). The three or four species of Melanopareia differ from other tapaculos by their rather slender build and boldly and attractively patterned plumage, and by sharing a semi- concealed white dorsal patch with various true antbirds (Thamnophilidae). They were originally described in the genus Synallaxis (Furnariidae), in which Sclater (1890) later submerged the genus. Salvin (1876) described a new species from Ecuador as Formicivora speciosa, duly rec- ognized in that combination by Sclater (1890), p. 251, and Fig. 1 The bamboo-wren Psilorhamphus guttatus bears little external resemblance to typical members of the tapaculo family. With its grey others until Hellmayr (1906), p. 334, showed that this was a iris, facial expression, bill shape, and wing-coverts with white dots synonym of Synallaxis elegans Lesson, in which genus Psilorhamphus instead resembles some antbirds (Dysithamnus, Myr- Hellmayr continued to place it while regarding Salvin’s motherula) with which early ornithologists consequently placed it. allocation of it to Formicivora with incredulity. Ridgway Unlike other tapaculos Psilorhamphus spends most of the time above the ground. Photo: Edson Endrigo (1909) seems to have overlooked this when he created a new genus Rhoporchilis for Formicivora speciosa. It was Hellmayr (1921) who eventually established the modern comment that these genera ‘‘might perhaps be more natu- concept of the genus in showing that Synallaxis elegans, S. rally placed as a distinct subfamily of Pteroptochidae torquata, and S. maximiliani were congeneric and would [= Rhinocryptidae]’’ despite the fact that ‘‘there is little all fall under Reichenbach’s earlier generic name Mela- external difference between the appearance of these birds nopareia and ‘‘find their natural place in the Formicarii- and the true Wrens [Troglodytidae]’’. We are not aware, dae’’, where they stayed for only a few years (Cory and however, of any instance in which Psilorhamphus was Hellmayr 1924). Next came the observation of W.D.W. placed in either the Troglodytidae or in a family with only Miller that the sternum of Melanopareia was four-not- sylviid-like genera, as might be inferred from Krabbe and ched—information that was conveyed to and presented by Schulenberg (2003). Psilorhamphus continued to be asso- Wetmore (1926), p. 292. On this basis, Peters (1951) ciated with Ramphocaenus in the Formicariidae—e.g. included Melanopareia in the Rhinocryptidae, where it has Sclater (1890) and Cory and Hellmayr (1924), p. 205— resided since. although in the latter reference it was noted that W.D.W. The other problem species, the Bamboo-wren Psilor- Miller would show Psilorhamphus and Ramphocaenus to hamphus guttatus, is a small bamboo specialist with a ‘‘constitute a separate family’’ in ‘‘a paper shortly to be rather long, slender bill, a long tail, and relatively weak published.’’ Peters (1951), p. 213, later explained that feet, so it bears little resemblance to large-footed terrestrial Miller’s death prevented publication of his results but that tapaculos (Fig. 1). From the beginning (Me´ne´trie´s 1835)it Wetmore (1943), p. 306, had shown Ramphocaenus to was placed with the antbirds in the Myiotherinae (= For- have an oscine syrinx, and had told Peters that Microbates micariidae). Sclater (1858), p. 243, associated Psilorham- likewise was oscine and that he believed, on the basis of phus with Ramphocaenus (which is now in the oscine external morphology, that Psilorhamphus was also proba- family Polioptilidae) in the Formicariidae with the bly oscine. Therefore, Peters postponed his treatment of 123 J Ornithol (2010) 151:337–345 339 those genera for a future volume treating Sylviidae. Sick 1994; Krabbe and Schulenberg 2003), including represen- (1954) placed Ramphocaenus in the Sylviidae while pro- tatives of the large genus Scytalopus, including one rep- visionally referring Psilorhamphus to the Formicariidae. resentative (indigoticus) of the ‘‘blue’’ species, which were Then, Plo´tnick (1958) revealed that Psilorhamphus had a recently placed in a separate genus Eleoscytalopus four-notched sternum, a tracheophone syrinx, and had other (Maurı´cio et al. 2008). Three of the authors have significant characters, including an operculate nostril, indicating that it field experience of the biology and vocalizations of should be placed in the Rhinocryptidae. Thus, in Peters’ tapaculos, and this was supplemented with comments and Checklist Psilorhamphus appears as an addendum to the analyses of sound archives by Niels Krabbe (personal Rhinocryptidae that appeared in the volume on Sylviidae communication). Representatives of the main lineages (Paynter 1964). within the tracheophone radiation serve as outgroups Heimerdinger and Ames (1967) confirmed that the rhi- (Ridgely and Tudor 1994; Irestedt et al. 2002; Krabbe and nocryptids they examined all had a four-notched sternum Schulenberg 2003; Chesser 2004). but also showed that this condition obtained in at least two Three nuclear gene regions, myoglobin intron 2, orni- genera of grallarine Formicariidae, which was confirmed thine decarboxylase (ODC) introns 6 to 7, and glycer- by Feduccia and Olson (1982). aldehyde-3-phosphodehydrogenase (G3PDH) intron 11, Ames (1971) made a thorough study of the anatomy of were
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