http://asociacioncolombianadeornitologia.org/revista-ornitologia-colombiana/ Ornitología Colombiana Ornitología Colombiana Colombiana Ornitología ety eadda te ot relcal protec- irreplaceable most the as regarded cently re- was Park National SNSM the endemism, king Hernández Cleef 1922, Carriker & (Todd groups and plant multiple for endemism of center to up level sea from rises that Colombia Northern in massif mountain a Nevada MartaisolateddeSanta (SNSM), Sierra The Palabras clave: estarecomendación. hacemosmicaAquíformal. tal recomendación describimosy en detallelas diferencias enplumajemediciones y apoyar para unataxonó-recomendaciónpresentamosno medidas morfométricasyplumaje en diferencias las detalle enpresentamos embargo,no Sin taxon. delotro vs.lossimpatría,propios mostraronellosrespuestassusastrechadecantos diferentes na la Además,suszo- encontramoscantos.enuna fenotípicasde enqueyy genéticasdiferencas base H. bangsi,con bajas, más elevaciones de táxon del distinta especie una de rango el ameritaanachoreta, H. Marta, Santa de Nevada Sierra la de res previo,elelevacionesestudioHenichorhinacucarachohabitaevidenciaunpresentamos delgénero que Enquelas superio- Resumen words:Key thisofrecommendation. morphometrics support andinplumage differencesin the descriptiondetailedofmore present andhere, so do We status. taxonomic their regarding recommendationtaxonomic formal a make not did and detail, in morphometrics and plumage in differencesthe present not did weHowever, form. other the vs. own their of songs the to responsesdifferential showed sympatrynarrowthatzoneMoreover,wetheyfoundof welldifferences and as a geneticphenotypicsong.evidencein in as Marta, Santa de da wepreviouspresentedstudy, evidence a thethatIn Abstract 5 4 3 TheNetherlands 2 1 Slabbekoorn Hans Cadena Daniel Carlos Behavioral Biology,Institute Biology,of Leiden University, Leiden,The Netherlands Museum Vertebrateof Zoology andDepartment Integrativeof Biology,University California,of Berkeley,USA DepartmentEcologyof & Evolutionary Biology,Tulane University, NewOrleans, USA BiodiversityDynamics,EcosystemEvolutionEcologyAmsterdam, and Universiteit Program, and Amsterdam,Institute vanof DepartamentodeCiencias Biológicas, Universidad delos Andes,Bogotá, Colombia. anachoreta anachoreta Henicorhina [email protected] for the Colombia Sierra speciesfor Nevada Marta, Santa de

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merits status as a species distinct from the lower élevation taxon, élevation lower the from distinct species a as status merits especiación,Sierra Nevada de Santa Marta,taxonomía. 1 et al. al. et , Paula C. Paula , Caycedo (Troglodytidae), otra de(Troglodytidae), especie endémicaave (Troglodytidae),endemicanother Henicorhina Henicorhina 1984, 82 ( phenotypic of patterns on based recognized been have range mountain this to endemic subspecies 50 than more , In work. revisionary insufficient and treatments taxonomic historical to to adherence due underestimated considerably be still may SNSM the of endemism species true the markably, Saout (Le World the in area ted

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Cadena et al. vocalizations and patterns of genetic differentia- bed as a distinct species, H. anachoreta (Bangs tion, it has become increasingly apparent that a 1899), whereas the form occurring at lower eleva- number of endemic taxa remain undescribed tions (bangsi) was described as a subspecies of H. (Rheindt et al. 2013), and that several populations hilaris (Ridgway 1903), a taxon later considered endemic to the SNSM and traditionally regarded conspecific with H. leucophrys (e.g., Hellmayr as subspecies of widespread species are suffi- 1934). Although bangsi was listed as a separate ciently differentiated to be accorded full species species by Brabourne & Chubb (1912), most status (Krabbe & Schulenberg 1997, Krabbe 2008, authors initially treated both anachoreta and Cadena & Cuervo 2010, Isler et al. 2012, Collar & bangsi as subspecies of H. hilaris and then more Salaman 2013, Lozano-Jaramillo et al. 2014). Here, broadly as part of H. leucophrys (e.g., Ridgway we propose the restitution of species status for yet 1904, Todd & Carriker 1922, Hellmayr 1934, Mayr another taxon endemic to the SNSM. & Greenway 1960).

Wrens in the Henicorhina (Troglodytidae) Caro et al. (2013) conducted a morphological, are widely distributed in the Neotropical region. genetic, and behavioral study of wood- po- Although their taxonomic diversity at the subspe- pulations along an elevational gradient in the cies level is high (33 taxa), only four species are northwestern SNSM, and found that anachoreta currently recognized (Kroodsma & Brewer 2005, and bangsi are not only phenotypically and gene- Remsen et al. 2015). For many years, the genus tically distinct, but also that they are sympatric was thought to include only two species, the over a narrow elevational range. In addition, based White-breasted Wood-Wren (H. leucosticta) occu- on response patterns to local and foreign songs rring in the lowlands and the Grey-breasted assessed using playback experiments, Caro et al. Wood-Wren (H. leucophrys) in montane areas. (2013) hypothesized that vocal differences likely The other two species were discovered in recent served as a behavioral barrier to gene flow bet- decades: the Bar-winged Wood-Wren (H. leucop- ween forms. Taken together, these results de- tera) in southern Ecuador and northern Peru monstrated marked divergence and evidence of (Fitzpatrick et al. 1977), and the Munchique Wood reproductive isolation in sympatry (see also Burbi- -Wren (H. negreti) in the Western Cordillera of dge et al. 2015 for confirmation of vocal recogni- Colombia (Salaman et al. 2003). The existence of tion patterns), which indicated that the two wood- marked geographic variation in plumage, vocali- wren populations from the SNSM represent sepa- zations and mitochondrial DNA sequences within rate species. However, formal taxonomic recom- both H. leucosticta and H. leucophrys suggests mendations have not been proposed in light of that each of these widespread taxa may comprise these findings. more than one species (Winker et al. 1996; Dingle et al. 2006, 2008). Phylogeographic analyses based on mitochondrial DNA sequences revealed evidence that bangsi In the SNSM, two forms of H. leucophrys descri- and anachoreta are relatively distant relatives bed from museum specimens on the basis of sub- within H. leucophrys, implying they did not derive tle but consistent differences in morphometrics from a single ancestor colonizing the SNSM but and coloration replace each other along elevatio- rather from separate colonization events (Caro et nal gradients (Bangs 1899, Ridgway 1903, Todd & al. 2013). Furthermore, bangsi, the form of the Carriker 1922, Hilty & Brown 1986). The form oc- foothills and lower montane forests up to ca. 2270 curring at higher elevations was originally descri- m in the SNSM, is relatively weakly differentiated in

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Figure 1. Henicorhina anachoreta (Hermit Wood-Wren), an overlooked species endemic to upper montane forests of the Sierra Nevada de Santa Marta, Colombia. Photograph by N. Athanas taken at El Dorado Nature Reserve, San Lorenzo Ridge. mitochondrial DNA from populations of H. leu- já, and the other to anachoreta (Caro et al. 2013). cophrys occurring on the nearby western slopes of the Serranía de Perijá (Caro et al. 2013). The We recognize that considerable additional wood-wren populations from Perijá are likely as- research centered on genetic divergence and ge- signable to H. l. manastarae (López-O. et al. 2014), ne flow and differentiation in vocal and behavioral a subspecies described from the eastern signals (no such analyses outside of the SNSM (Venezuelan) slope of that mountain range were conducted in the earlier study, but see Din- (Aveledo Hostos & Ginés 1952). In turn, the form gle et al. 2008, Burbidge et al. 2015) is still requi- occurring in upper montane forest above ca. 2270 red to acquire a complete understanding of spe- m in the SNSM, anachoreta, is more divergent ciation in Henicorhina wood-wrens and species from all other sampled populations of H. leucop- limits in H. leucophrys. Nonetheless, the present hrys and appears to have been an earlier colonist evidence firmly shows that the two populations to Santa Marta likely derived from Andean stock from the SNSM referred to H. leucophrys belong (Caro et al. 2013, J. L. Pérez-Emán et al., unpublis- to different species. In addition, available data in- hed data). In addition, variation at six microsatellite dicate that the population from higher elevations loci revealed the existence of two genetic clusters is more divergent from populations outside the in the SNSM and Perijá, one of which correspon- SNSM referable to H. leucophrys than is the popu- ded to bangsi and manastarae from western Peri- lation from lower elevations. Based on the above,

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Figure 2. Box plots summarizing measurements of individuals captured in the field in the Sierra Nevada de Santa Marta and assigned to Henicorhina anachoreta (gray) or H. leucophrys bangsi (white) based on multivariate analyses of morphometric and molecular data (data from Caro et al. 2013). For each trait and taxon, plots show median, lower and upper quartiles, 5% and 95% percentiles, and outliers. Numbers on each plot indicate sample sizes per taxon. Note the smaller overall size of anachoreta in body and bill dimensions, with the exception of tarsus length. we recommend treating the highland population The holotype of H. anachoreta (housed at the Mu- occurring in the SNSM as a distinct species, Heni- seum of Comparative Zoology, MCZ 106494) is corhina anachoreta Bangs, 1899 (Figure 1). Based from Páramo de Chiruqua, 12,000 ft (ca. 3940 m; on the meaning of the latin name anachoreta, we Bangs 1899), whereas the type of H. l. bangsi propose Hermit Wood-Wren as an English name. (housed at the National Museum of Natural Histo-

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Figure 3. Representative specimens of wood-wrens from the Sierra Nevada de Santa Marta illustrating overall differences in plumage coloration between H. l. bangsi and H. anachoreta. All specimens were prepared by M. A. Carriker, Jr. and are housed at the USNM. H. l. bangsi (from Depto. Magdalena, Hacienda Cincinnati): 387788 (female, ca. 1520 m) and 374501 (male, 1060-2020 m); H. anachoreta (from Depto. Cesar, upper Río Guatapurí): 387761 (female, 3050-3300 m) and 387771 (male, 2450-2900 m). Note the overall darker coloration of H. anachoreta ventrally; dorsally, it tends to be more lightly colored than H. l. bangsi (see text for details). Photographs by David Ocampo. ry, USNM 163791) was collected above Pueblo themselves because there is some overlap bet- Viejo, Depto. Magdalena, in the foothills of the ween forms, so we suggest the species are best SNSM. Hence, there is no doubt that the name diagnosed based on a combination of measure- anachoreta applies to the upper-montane species. ments. In terms of plumage coloration (Figure 3), H. anachoreta differs from H. l. bangsi in its lighter, Todd & Carriker (1922) noted that "[H. anachore- more russet tone of the flanks and upperparts ta] is so similar to H. hilaris bangsi as to be practi- (including tail, rump, wings and mantle), tending cally indistinguishable in life". The similarity of the to sooty or ferruginous brown instead of deep two species, compounded with the possibility of chestnut. In addition, the nape and pileum in H. finding individuals with intermediate phenotypes anachoreta are olive brown and may blend into due to limited hybridization and introgression gray towards the forehead, instead of being uni- (Todd & Carriker 1922, Hellmayr 1934, Caro et al. formly dark brown as in H. l. bangsi. Ventrally, the 2013), implies that reliable identification in the field throat of H. anachoreta is whitish gray with indis- is challenging. Nonetheless, noticeable differences tinct streaks instead of the uniform grayish white between the two taxa exist in morphology and co- throat of H. l. bangsi, and the breast is decidedly loration. As noted by Caro et al. (2013), H. ana- darker and less suffused with buffy tones relative choreta is smaller overall (contra Bangs 1899), with to H. l. bangsi (cf. Bangs 1899, Ridgway 1903, lower body mass and shorter wings, and a smaller Todd & Carriker 1922, Hellmayr 1934). The songs bill (cf. Ridgway 1903, Hellmayr 1934; Figure 2). of H. anachoreta and H. l. bangsi differ signifi- However, none of these traits are diagnostic by cantly (i.e., the former sings at a higher frequency

http://asociacioncolombianadeornitologia.org/ 86 2016 | Número 15 Cadena et al. and over a greater frequency range), but because an important task for taxonomists working on vocalizations are variable within H. l. bangsi and Neotropical montane birds is to examine the sta- because songs vary more gradually than morpho- tus of populations currently treated as subspecies logy along elevational gradients (Caro et al. 2013), differing in elevational distributions. We believe care must be exercised when identifying indivi- that elevation is likely an important geographic duals based only on vocal cues. dimension along which cryptic species diversity has been overlooked. We note that our proposed recognition of H. ana- choreta as a species would imply recognizing a Finally, because H. anachoreta is widespread and paraphyletic species H. leucophrys (Caro et al. common in the SNSM, tolerates some habitat dis- 2013). Species-level paraphyly is expected when turbance, and occurs in several protected areas in speciation occurs in the periphery of the geo- the region including the SNSM National Park, we graphic ranges of widespread lineages (Funk & do not consider it to be threatened from a conser- Omland 2003) and the existing taxonomy of wrens vation standpoint. However, we encourage moni- (Lara et al. 2012, Remsen et al. 2015) already ac- toring of populations especially given looming cepts paraphyletic species. In particular, H. leucos- threats faced by montane birds from the SNSM as ticta is paraphyletic with respect to H. leucoptera a consequence of climatic change (Jankowski et al. (Dingle et al. 2006). In addition, ongoing work in- 2010, Velásquez-Tibatá et al. 2013, Ramírez- dicates that H. negreti is nested within H. leucop- Villegas et al. 2014) hrys (J. L. Pérez-Emán et al. unpubl. data). We be- lieve that paraphyly in Neotropical wren species Acknowledgements partly reflects speciation in the periphery of ran- ges, but also faulty taxonomy (Ross 2014). We sus- We thank V. Piacentini and J. V. Remsen for com- pect that future analyses will likely conclude that H. ments on the manuscript, R. T. Chesser for allo- leucophrys comprises multiple species-level taxa, wing us to examine specimens at the National Mu- and we may arrive at a stable classification in the seum of Natural History - Smithsonian Institution future in which species are accurately delimited (USNM), and N. Athanas and D. Ocampo for the and monophyletic. For the time being, however, photos accompanying this note. we consider it best to begin by recognizing the well-differentiated H. anachoreta as distinct in a References first step towards a revised classification of the group. AVELEDO HOSTOS, R., & H. GINÉS. 1952. Cuatro aves nuevas y dos extensiones de distribución para Venezuela, de

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Recibido: 07 de octubre de 2014 Aceptado: 25 de agosto de 2015

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