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PHYLOGENETIC RELATIONSHIPS of the WRENTIT BASED on MITOCHONDRIAL CYTOCHROME B SEQUENCES

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PHYLOGENETIC RELATIONSHIPS of the WRENTIT BASED on MITOCHONDRIAL CYTOCHROME B SEQUENCES The Condor 104:740±749 q The Cooper Ornithological Society 2002 PHYLOGENETIC RELATIONSHIPS OF THE WRENTIT BASED ON MITOCHONDRIAL CYTOCHROME b SEQUENCES DINO N. BARHOUM AND KEVIN J. BURNS1 Department of Biology, San Diego State University, San Diego, CA 92182-4614 Abstract. The phylogenetic relationship of the Wrentit (Chamaea fasciata) to other pas- serine birds is understood poorly. A variety of taxa have been proposed as closely related to the Wrentit, but in general this species is placed in a monotypic taxon to emphasize its morphological distinctiveness and lack of clear relationship to other groups of birds. We used the mitochondrial cytochrome b gene to infer the relationship of this monotypic genus to other avian groups including representatives of Sylvioidea, Muscicapoidea, and Passe- roidea. Results of this study corroborate in part earlier proposals based on DNA-DNA hybridization. Maximum parsimony, maximum likelihood, and Bayesian phylogenetic anal- yses support the Wrentit, Sylvia, and babblers (Garrulax, Illadopsis, Leiothrix, Pomatorhin- us, Stachyris, and Yuhina) as a clade, with the Wrentit sharing a more recent common ancestor with Sylvia than with babblers. A Sylvia and Wrentit association is further supported by similarities in morphology and ecology. In addition to ®ndings on Wrentit relationships, our study in general agrees with the major groups of oscine passerines identi®ed by earlier DNA hybridization studies. Key words: babbler, Chamaea fasciata, mitochondrial DNA, phylogeny, Sylvia, Wrentit. Relaciones FilogeneÂticas de Chamaea fasciata Basadas en Secuencias del Citocromo b Mitocondrial Resumen. La relacioÂn ®logeneÂtica de Chamaea fasciata con otros paserinos es vaga- mente entendida. Una variedad de taxa ha sido propuesta como relacionada cercanamente a C. fasciata, pero en general esta especie es situada en un taxoÂn monotõÂpico para enfatizar su morfologõÂa distintiva y la falta de claridad en la relacioÂn con otros grupos de aves. Utilizamos el gen del citocromo b mitocondrial para inferir la relacioÂn de este geÂnero mo- notõÂpico con otros grupos de aves incluyendo representantes de Sylviodea, Muscicapoidea y Passeroidea. Resultados de este estudio corroboran parcialmente propuestas anteriores basadas en hibridacioÂn de ADN-ADN. AnaÂlisis ®logeneÂticos de maÂxima parsimonia, de maÂxima probabilidad y bayesianos respaldan a C. fasciata, Sylvia, Garrulax, Illadopsis, Leiothrix, Pomatorhinus, Stachyris y Yuhina como un clado, con C. fasciata compartiendo un ancestro comuÂn maÂs reciente con Sylvia. Una asociacioÂn entre Sylvia y C. fasciata es respaldada por similitudes en morfologõÂa y ecologõÂa. En adicioÂn a los descubrimientos en las relaciones de C. fasciata, nuestro estudio coincide en general con los grupos mayores de paserinos oscinos identi®cados en estudios previos mediante hibridacioÂn de ADN. INTRODUCTION blers (Timaliidae), and Old World warblers The Wrentit (Chamaea fasciata) is a small song- (Sylviidae) have all been considered the Wren- bird that frequents brushy habitats from northern tit's closest relatives (Grinnell 1913, Sibley Baja California to Oregon. This species is un- 1970, Sibley and Ahlquist 1982, 1990). Reasons usual among North American birds in that it ap- for this uncertainty stem from the lack of de®n- parently is not closely related to any other New able similarities in behavior, ecology, and mor- World species. The evolutionary relationship of phology to other North American species. Be- the Wrentit to other birds has been debated since cause of its lack of clear relationship to other Gambel (1845) ®rst described the species. Tit- birds, the Wrentit has been assigned to a mono- mice (Paridae), wrens (Troglodytidae), bushtits typic taxon at various taxonomic levels (genus, (Psaltriparus), mockingbirds (Mimidae), bab- tribe, family). Most recent classi®cations consider the Wren- tit to be related to Old World warblers and bab- Manuscript received 5 February 2002; accepted 23 blers (traditionally de®ned as the families Sylvii- July 2002. 1 Corresponding author. E-mail: kburns@sunstroke. dae and Timaliidae, respectively; Wetmore sdsu.edu 1960, Howard and Moore 1991). For example, [740] WRENTIT PHYLOGENETIC RELATIONSHIPS 741 Sibley (1970) compared electrophoretic mobility World warblers are found in three families of the of egg-white proteins and was unable to un- Sylvioidea, including Sylviidae. Sibley and equivocally de®ne the sister taxon, but narrowed Monroe (1990) also placed most traditional bab- the possibilities to either Timaliidae or Sylvii- blers into two of the four subfamilies of Sylvii- dae. However, this result is not very enlighten- dae (Garrulacinae and Sylviinae). One of these ing because the limits and monophyly of these subfamilies (Sylviinae) includes the majority of two families are uncertain (Sibley and Ahlquist the babblers (tribe Timaliini), the Old World 1982, 1990, Cibois et al. 1999). Babblers are a warbler genus Sylvia (tribe Sylviini) and the diverse group that primarily inhabit tropical ar- Wrentit (tribe Chamaeini). These results were eas of Asia and Africa. They do not have well- consistent with Sibley and Ahlquist's (1982) ear- recognized synapomorphies, and genera without lier DNA hybridization study in which they con- other obvious taxonomic af®nities have been in- cluded that the Wrentit is approximately equi- cluded among the babblers (Sibley and Monroe distant from babblers and from Sylvia. Thus, 1990, Howard and Moore 1991). Although bab- DNA hybridization evidence indicates that the bler monophyly has been questioned, Cibois et Wrentit is closely related to the genus Sylvia and al. (1999, 2001) demonstrated the monophyly of to a clade containing most traditional babblers several babblers including species in the genera (Timaliini). Garrulax, Illadopsis, Leiothrix, Pomatorhinus, Although the classi®cation of Sibley and Stachyris, and Yuhina. Old World warblers Monroe (1990) is becoming more widely used, (Sylviidae) are another extensive radiation of several authors (e.g., Harshman 1994) have crit- birds that occupy a variety of niches and are icized some of the methods of analysis used by abundant in Europe, Asia, and Africa. Although Sibley and Ahlquist (1990) to derive this tax- the group is not monophyletic (Sibley and Ahl- onomy. Several recent DNA sequencing studies quist 1990), recent systematic studies have iden- (Groth 1998, Ericson et al. 2000, Lovette and ti®ed several monophyletic assemblages within Bermingham 2000, Barker et al. 2001) have di- Old World warblers. These include the reed war- rectly tested predictions made from these DNA blers (Acrocephalus, Chloropeta, and Hippolais) hybridization studies. However, none of these and some leaf warblers (Phylloscopus), based on studies have addressed the phylogenetic position studies by Helbig and Seibold (1999) and Price of the Wrentit. In this study, we use mitochon- et al. (1997, 1998), respectively. drial DNA sequence data to identify the closest The only recent, comprehensive systematic living relative of the Wrentit. In addition, we study of both Timaliidae and Sylviidae that in- compare our phylogenies of oscine passerine re- cluded the Wrentit was the DNA-DNA hybrid- lationships based on DNA sequence data to the ization study of Sibley and Ahlquist (1982, DNA-DNA hybridization phylogenies produced 1990). The taxonomic classi®cation based on by Sibley and Ahlquist (1982, 1990) and more Sibley and Ahlquist (1990) was outlined in Sib- recent DNA sequencing studies. ley and Monroe (1990). Sibley and Ahlquist (1990) found the families Sylviidae and Tima- METHODS liidae as traditionally de®ned (Wetmore 1960, TAXON SAMPLING Howard and Moore 1991) to be polyphyletic. Thus, Sibley and Monroe (1990) altered the Because of the taxonomic uncertainty surround- classi®cation of these groups. Polyphyly of the ing higher-level oscine passerine relationships, a traditionally recognized Sylviidae and Timalii- broad sample of taxa was necessary to investi- dae was indicated by the inclusion of the genus gate phylogenetic relationships of the Wrentit. Pomatostomus, traditionally a babbler, in the Forty-four representatives of parvorder Passeri- parvorder Corvida and close association of the da (sensu Sibley and Monroe 1990) were sam- remaining babblers (tribe Timaliini) to Sylvia (an pled, including representatives of all three Pas- Old World warbler genus). Both Old World war- serida superfamilies (Muscicapoidea, Passeroi- blers and babblers are included in Sibley and dea, and Sylvioidea). Fourteen genera of Old Monroe's (1990) superfamily Sylvioidea (199 World warblers (Acrocephalus, Bradypterus, genera, 1195 species). Sylvioidea consists of 11 Cettia, Chloropeta, Cisticola, Hippolais, Hyper- families, one of which Sibley and Monroe gerus, Locustella, Orthotomus, Phylloscopus, (1990) designated as Sylviidae. However, Old Regulus, Scotocerca, Sylvia, and Thamnornis) 742 DINO N. BARHOUM AND KEVIN J. BURNS and seven genera of babblers (Illadopsis, Garru- isolated using Chelext (5% solution, incubated lax, Leiothrix, Neomixis, Pomatorhinus, Stachy- for 20 min at 958C, Walsh et al. 1991) from fresh ris and Yuhina) were chosen to represent the liver, heart, or breast muscle stored at 2808C. Sylvioidea. Seven genera of Muscicapoidea Cytochrome b was ampli®ed using three primer (Catharus, Copsychus, Erithacus, Ficedula, pairs (H15298/L14830, H1570/L15184, and Muscicapa, Toxostoma, and Turdus) and six H16108/L15635; Groth 1998) for three overlap- genera of Passeroidea (Anthus,
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