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- W2056979431 abstract "The Hawaiian “honeyeaters,” five endemic species of recently extinct, nectar-feeding songbirds in the genera Moho and Chaetoptila, looked and acted like Australasian honeyeaters (Meliphagidae), and no taxonomist since their discovery on James Cook's third voyage has classified them as anything else [1Merrem, B. (1786). Avium rariorum et minus cognitarum, Icones et Descriptiones. Lipsiae.Google Scholar, 2Lesson R.P. Traité d'Ornithologie. F.G. Levraulet, Paris1831Google Scholar, 3Peale T.R. United States Exploring Expedition, Vol. 8, Mammalia and Ornithology. C. Sherman, Philadelphia1848Google Scholar, 4Rothschild W. The Avifauna of Laysan and the Neighbouring Islands with a Complete History to Date of the Birds of the Hawaiian Possessions. R.H. Porter, London1893–1900Google Scholar, 5Wilson S.B. Evans A.H. Aves Hawaiienses: The Birds of the Sandwich Islands. R.H. Porter, London1890–1899Crossref Google Scholar, 6Munro C. Birds of Hawaii. C.E. Tuttle, Co., Rutland, VT1944Google Scholar, 7Perkins R.C.L. Fauna Hawaiiensis: Vertebrata. Cambridge University Press, UK1903Google Scholar, 8Amadon D. The Hawaiian honeycreepers (Aves:Drepaniidae).Bull. Am. Mus. Nat. Hist. 1950; 95: 157-262Google Scholar]. We obtained DNA sequences from museum specimens of Moho and Chaetoptila collected in Hawaii 115–158 years ago. Phylogenetic analysis of these sequences supports monophyly of the two Hawaiian genera but, surprisingly, reveals that neither taxon is a meliphagid honeyeater, nor even in the same part of the songbird radiation as meliphagids. Instead, the Hawaiian species are divergent members of a passeridan group that includes deceptively dissimilar families of songbirds (Holarctic waxwings, neotropical silky flycatchers, and palm chats). Here we designate them as a new family, the Mohoidae. A nuclear-DNA rate calibration [9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar] suggests that mohoids diverged from their closest living ancestor 14–17 mya, coincident with the estimated earliest arrival in Hawaii of a bird-pollinated plant lineage [10Givnish T.J. Millam K.C. Mast A.R. Paterson T.B. Theim T.J. Hipp A.L. Henss J.M. Smith J.F. Wood K.R. Sytsma K.J. Origin, adaptive radiation and diversification of the Hawaiian lobeliads (Asterales: Campanulaceae).Proc. Biol. Sci. 2008; (Published online October 14, 2008)https://doi.org/10.1098/rspb.2008.1204Crossref Scopus (255) Google Scholar]. Convergent evolution, the evolution of similar traits in distantly related taxa because of common selective pressures, is illustrated well by nectar-feeding birds [11Wallace A.R. Who are the humming bird's relations?.Zoologist. 1863; 21: 8486-8491Google Scholar], but the morphological, behavioral, and ecological similarity of the mohoids to the Australasian honeyeaters makes them a particularly striking example of the phenomenon. The Hawaiian “honeyeaters,” five endemic species of recently extinct, nectar-feeding songbirds in the genera Moho and Chaetoptila, looked and acted like Australasian honeyeaters (Meliphagidae), and no taxonomist since their discovery on James Cook's third voyage has classified them as anything else [1Merrem, B. (1786). Avium rariorum et minus cognitarum, Icones et Descriptiones. Lipsiae.Google Scholar, 2Lesson R.P. Traité d'Ornithologie. F.G. Levraulet, Paris1831Google Scholar, 3Peale T.R. United States Exploring Expedition, Vol. 8, Mammalia and Ornithology. C. Sherman, Philadelphia1848Google Scholar, 4Rothschild W. The Avifauna of Laysan and the Neighbouring Islands with a Complete History to Date of the Birds of the Hawaiian Possessions. R.H. Porter, London1893–1900Google Scholar, 5Wilson S.B. Evans A.H. Aves Hawaiienses: The Birds of the Sandwich Islands. R.H. Porter, London1890–1899Crossref Google Scholar, 6Munro C. Birds of Hawaii. C.E. Tuttle, Co., Rutland, VT1944Google Scholar, 7Perkins R.C.L. Fauna Hawaiiensis: Vertebrata. Cambridge University Press, UK1903Google Scholar, 8Amadon D. The Hawaiian honeycreepers (Aves:Drepaniidae).Bull. Am. Mus. Nat. Hist. 1950; 95: 157-262Google Scholar]. We obtained DNA sequences from museum specimens of Moho and Chaetoptila collected in Hawaii 115–158 years ago. Phylogenetic analysis of these sequences supports monophyly of the two Hawaiian genera but, surprisingly, reveals that neither taxon is a meliphagid honeyeater, nor even in the same part of the songbird radiation as meliphagids. Instead, the Hawaiian species are divergent members of a passeridan group that includes deceptively dissimilar families of songbirds (Holarctic waxwings, neotropical silky flycatchers, and palm chats). Here we designate them as a new family, the Mohoidae. A nuclear-DNA rate calibration [9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar] suggests that mohoids diverged from their closest living ancestor 14–17 mya, coincident with the estimated earliest arrival in Hawaii of a bird-pollinated plant lineage [10Givnish T.J. Millam K.C. Mast A.R. Paterson T.B. Theim T.J. Hipp A.L. Henss J.M. Smith J.F. Wood K.R. Sytsma K.J. Origin, adaptive radiation and diversification of the Hawaiian lobeliads (Asterales: Campanulaceae).Proc. Biol. Sci. 2008; (Published online October 14, 2008)https://doi.org/10.1098/rspb.2008.1204Crossref Scopus (255) Google Scholar]. Convergent evolution, the evolution of similar traits in distantly related taxa because of common selective pressures, is illustrated well by nectar-feeding birds [11Wallace A.R. Who are the humming bird's relations?.Zoologist. 1863; 21: 8486-8491Google Scholar], but the morphological, behavioral, and ecological similarity of the mohoids to the Australasian honeyeaters makes them a particularly striking example of the phenomenon. The Australasian honeyeaters (Meliphagidae) are a group of songbirds that branch off within the passeriform (perching bird) phylogeny basal to both the “core Corvoidea” and the Passerida [9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar]. They have classical adaptations for nectarivory, including scroll-edged, forked, brush-tipped tongues (Figure 1) and long, often decurved, bills (Figure 2). The 182 species of Meliphagidae occur south of Wallace's line in New Guinea and Australia, with a few genera such as Myzomela, Foulehaio, and Gymnomyza spilling out onto the islands of Micronesia and Polynesia. Also traditionally included in the Meliphagidae were the Hawaiian Moho (four species of `o`o, each found on a different island; Figures 2A and 2E) and the rather differently appearing Chaetoptila angustipluma (the kioea; Figure 2C). All five species were nectarivores with meliphagid-like tongues (Figure 1). Taxonomists have never doubted that Moho and Chaetoptila were meliphagids, and have only expressed uncertainty about whether they arose from a single colonization of the Hawaiian Islands (i.e., are monophyletic) and which particular meliphagid taxa might be their closest relatives ([4Rothschild W. The Avifauna of Laysan and the Neighbouring Islands with a Complete History to Date of the Birds of the Hawaiian Possessions. R.H. Porter, London1893–1900Google Scholar, 5Wilson S.B. Evans A.H. Aves Hawaiienses: The Birds of the Sandwich Islands. R.H. Porter, London1890–1899Crossref Google Scholar, 6Munro C. Birds of Hawaii. C.E. Tuttle, Co., Rutland, VT1944Google Scholar, 7Perkins R.C.L. Fauna Hawaiiensis: Vertebrata. Cambridge University Press, UK1903Google Scholar, 8Amadon D. The Hawaiian honeycreepers (Aves:Drepaniidae).Bull. Am. Mus. Nat. Hist. 1950; 95: 157-262Google Scholar]; summarized in Supplemental Data available online).Figure 2Illustrations of Three of the Five Species of Hawaiian “Honeyeaters” and Three Representative Meliphagid HoneyeatersShow full captionThe three Hawaiian taxa represent the three primary morphological types found in Hawaiian “honeyeaters” (Mohoidae: [A], Moho nobilis; [C], Chaetoptila angustipluma; and [E], Moho braccatus). The three meliphagids include one from New Zealand ([B], Prosthemadera novaeseelandiae), one from Australia ([D], Anthochaera carunculata), and one from Samoa ([F], Gymnomyza samoensis). Paintings are by John Anderton and are used here with permission.View Large Image Figure ViewerDownload Hi-res image Download (PPT) The three Hawaiian taxa represent the three primary morphological types found in Hawaiian “honeyeaters” (Mohoidae: [A], Moho nobilis; [C], Chaetoptila angustipluma; and [E], Moho braccatus). The three meliphagids include one from New Zealand ([B], Prosthemadera novaeseelandiae), one from Australia ([D], Anthochaera carunculata), and one from Samoa ([F], Gymnomyza samoensis). Paintings are by John Anderton and are used here with permission. The five historically known Hawaiian “honeyeaters” unfortunately all became extinct between the 1850s and the 1980s, and molecular analysis is limited to DNA from relatively old museum specimens. Here we evaluate the phylogenetic position of Moho and Chaetoptila within the order of perching birds, by using up to 1923 bp of nuclear and 717 bp of mitochondrial DNA sequences obtained from multiple specimens of Moho and Chaetoptila collected during the 1800s (Table S1). Although our phylogenetic analyses of mtDNA sequences provide strong support for the monophyly of all of the Hawaiian taxa (Figure 3A), we were surprised to find no support for the placement of this group within the family Meliphagidae on the basis of mtDNA (Figure 3A), nuclear RAG-1 (Figure 3B), or nuclear intron sequences (Figure 3C). Nor was there support for including them within the basal oscine songbird clade [9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar] that contains meliphagids along with related families of fairy wrens, chats, and pardalotes. Instead, there was strong support (Figure 3) for including Moho and Chaetoptila in another great and secondary radiation of songbirds, the Passerida, and more specifically, within an unusual passeridan clade containing three avian families: waxwings (Bombycillidae), New World silky flycatchers (Ptiligonatidae), and the monotypic palm chat of Hispaniola (Dulidae). All of these species are frugivores or insectivores, and are not nectarivores like the two Hawaiian genera. In addition to the high bootstrap and Bayesian support for the relationship (Figure 3), we found that RAG-1 trees constrained to include the Hawaiian taxa within the Meliphagidae were significantly less likely by Shimodaira-Hasegawa test (p < 0.0001; Supplemental Data) than the unconstrained maximum likelihood (ML) tree as shown in Figure 3B. These DNA results prompted us to re-evaluate the morphological characteristics of Moho and Chaetoptila in relation to Australasian honeyeaters and other songbirds. Many of the traits that prompted systematists to place them in the Meliphagidae are adaptive trophic structures: long tarsi and strong perching feet for reaching flowers, long decurved bills and extendable tongues to probe floral nectaries, tubular or semitubular brush-tipped tongues that use capillary attraction to move nectar up into the throat (Figure 1), and an operculum over the nares to protect the nasal cavity from pollen. The Hawaiian and Australasian nectarivores also display parallels in plumage (Figure 2), behavior, and song [4Rothschild W. The Avifauna of Laysan and the Neighbouring Islands with a Complete History to Date of the Birds of the Hawaiian Possessions. R.H. Porter, London1893–1900Google Scholar, 5Wilson S.B. Evans A.H. Aves Hawaiienses: The Birds of the Sandwich Islands. R.H. Porter, London1890–1899Crossref Google Scholar, 6Munro C. Birds of Hawaii. C.E. Tuttle, Co., Rutland, VT1944Google Scholar, 7Perkins R.C.L. Fauna Hawaiiensis: Vertebrata. Cambridge University Press, UK1903Google Scholar, 8Amadon D. The Hawaiian honeycreepers (Aves:Drepaniidae).Bull. Am. Mus. Nat. Hist. 1950; 95: 157-262Google Scholar] that indicate an even broader convergence in their life histories as part of defending ephemeral or widely spaced nectar sources. This convergence is so pervasive that, without the molecular sequence data, it would probably never have been possible to recognize the closest relatives of the Hawaiian lineage as being the waxwings and allies. Our results indicate that the Hawaiian birds were derived from Holarctic or Neotropical, and not South Pacific, ancestors. This further strengthens Mayr's contention that the Hawaiian avifauna is more American than otherwise [12Mayr E. The zoogeographic position of the Hawaiian Islands.Condor. 1943; 45: 45-48Crossref Google Scholar, 13Fleischer R.C. McIntosh C.E. Molecular systematics and biogeography of the Hawaiian avifauna.Studies Avian Biol. 2001; 22: 51-60Google Scholar]. Our molecular analyses also show that Moho and Chaetoptila are unique taxonomically and relatively divergent from any of their closest relatives (Table 1), necessitating the recognition of a new family-level rank.Table 1Date Estimates Based on Independent Rate Calibrations with the Program r8sA. Comparison-RAG-1NPRS (SE)PL (SE)Moho versus Phainoptila14.35 (0.58)16.01 (0.50)Moho versus Phainopepla16.16 (0.55)17.27 (0.42)Moho versus Dulus16.17 (0.75)16.89 (0.57)Moho versus Bombycilla18.11 (0.84)19.95 (0.63)M. nobilis versus M. bishopi0.88 (0.03)0.56 (0.02)B. Comparison-mtDNANPRS (SE)PL (SE)Moho versus Phainoptila19.91 (3.07)12.28 (3.20)Moho versus Phainopepla19.67 (3.07)12.25 (3.20)Moho versus Dulus18.13 (3.30)10.62 (2.35)Moho versus Bombycilla21.28 (3.22)13.72 (4.35)M. nobilis versus M. bishopi2.44 (0.25)2.13 (0.19)(A) Date estimates of nodes from the RAG-1 tree for comparisons of different close relatives of Moho (Ptilogonatidae, Dulidae, Bombycillidae), estimated by NPRS and PL approaches in r8s 17Sanderson M.J. Estimating absolute rates of molecular evolution and divergence times: A penalized likelihood approach.Mol. Biol. Evol. 2002; 19: 101-109Crossref PubMed Scopus (1465) Google Scholar. The RAG1 calibration, as in 9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar, is based on an 82 million year split between New Zealand's Acanthisitta and other passeriforms. Standard error (SE) was calculated from mean of dates at nodes of trees derived from 50 bootstrap repetitions.(B) Dates of nodes from mtDNA sequences for comparisons of available close relatives of Moho, estimated as above, but with an internal rate calibration based on the estimated subaerial age of Oahu (Supplemental Data). Open table in a new tab (A) Date estimates of nodes from the RAG-1 tree for comparisons of different close relatives of Moho (Ptilogonatidae, Dulidae, Bombycillidae), estimated by NPRS and PL approaches in r8s 17Sanderson M.J. Estimating absolute rates of molecular evolution and divergence times: A penalized likelihood approach.Mol. Biol. Evol. 2002; 19: 101-109Crossref PubMed Scopus (1465) Google Scholar. The RAG1 calibration, as in 9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar, is based on an 82 million year split between New Zealand's Acanthisitta and other passeriforms. Standard error (SE) was calculated from mean of dates at nodes of trees derived from 50 bootstrap repetitions. (B) Dates of nodes from mtDNA sequences for comparisons of available close relatives of Moho, estimated as above, but with an internal rate calibration based on the estimated subaerial age of Oahu (Supplemental Data). Type genus: Moho Lesson, 1831.Included genera: Moho, Chaetoptila Gray, 1869.Diagnosis: Passerida with the nectar-feeding adaptations mentioned above, and a single pneumotricipital fossa of the humerus with a large pneumatic opening. The Mohoidae present one of the most deceptive cases of convergent evolution in birds. Their closest relatives, and presumably their common ancestor, look nothing like meliphagids, yet Chaetoptila and Moho have such typical meliphagid characteristics (e.g., Figure 1) that they fooled generations of taxonomists into placing them in the Meliphagidae without equivocation [1Merrem, B. (1786). Avium rariorum et minus cognitarum, Icones et Descriptiones. Lipsiae.Google Scholar, 2Lesson R.P. Traité d'Ornithologie. F.G. Levraulet, Paris1831Google Scholar, 3Peale T.R. United States Exploring Expedition, Vol. 8, Mammalia and Ornithology. C. Sherman, Philadelphia1848Google Scholar, 4Rothschild W. The Avifauna of Laysan and the Neighbouring Islands with a Complete History to Date of the Birds of the Hawaiian Possessions. R.H. Porter, London1893–1900Google Scholar, 5Wilson S.B. Evans A.H. Aves Hawaiienses: The Birds of the Sandwich Islands. R.H. Porter, London1890–1899Crossref Google Scholar, 6Munro C. Birds of Hawaii. C.E. Tuttle, Co., Rutland, VT1944Google Scholar, 7Perkins R.C.L. Fauna Hawaiiensis: Vertebrata. Cambridge University Press, UK1903Google Scholar, 8Amadon D. The Hawaiian honeycreepers (Aves:Drepaniidae).Bull. Am. Mus. Nat. Hist. 1950; 95: 157-262Google Scholar]. New Zealand's endemic stitchbird (Notiomystis cincta) is another “honeyeater” that does not fall within the Meliphagidae on the basis of nuclear and mtDNA sequence analysis [14Driskell A. Christidis L. Gill B.J. Boles W.E. Barker F.K. Longmore N.W. A new endemic family of New Zealand passerine birds: Adding heat to a biodiversity hotspot.Aust. J. Zool. 2007; 55: 73-78Crossref Scopus (47) Google Scholar, 15Ewen J.G. Flux I. Ericson P.G.P. Systematic affinities of two enigmatic New Zealand passerines of high conservation priority, the hihi or stitchbird Notiomystis cincta and the kokako Callaeas cinerea.Mol. Phylogenet. Evol. 2006; 40: 281-284Crossref PubMed Scopus (27) Google Scholar]. It represents another deceptive case of convergent evolution; but this species is placed among the basal songbird lineages, along with the meliphagids, as opposed to the Hawaiian taxa, which are placed deep within the Passerida. In addition, whereas the stitchbird does have meliphagid characteristics, other aspects of its morphology and biology had led taxonomists to question its placement in Meliphagidae prior to the molecular analyses [16Higgins P.J. Peter J.M. Steele W.K. Handbook of Australian, New Zealand and Antarctic Birds Volume 5: Tyrant-Flycatchers to Chats. Oxford University Press, Melbourne2001Google Scholar]. Also, the convergence we report is not limited to a single mohoid and a single meliphagid morphotype; instead, at least three distinct morphotypes in the Mohoidae are also represented in the Meliphagidae, suggesting parallel adaptations across two independent radiations (Figure 2). Although the degree of convergence between the Mohoidae and the Meliphagidae may seem remarkable, one must take into account the amount of time since the Hawaiian lineage diverged from a mainland ancestor. On the basis of a RAG-1 external rate calibration [9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar] with nonparametric rate smoothing (NPRS) and penalized likelihood (PL) approaches [17Sanderson M.J. Estimating absolute rates of molecular evolution and divergence times: A penalized likelihood approach.Mol. Biol. Evol. 2002; 19: 101-109Crossref PubMed Scopus (1465) Google Scholar], we estimated the divergence time between Moho and its closest mainland relatives, the silky flycatchers, to range from about 14 to 17 million years (Table 1). A divergence time based on mtDNA divergences and island age is less precise, estimating a split from silky flycatchers or the palm chat at 10–20 million years (Table 1). Either of these estimated timeframes would presumably provide ample opportunity to evolve the adaptations for nectarivory that make Moho and Chaetoptila appear so similar in gestalt to the distantly related Australasian honeyeaters. In addition, if either one of these estimated time periods corresponds to the presence of the Mohoidae in the Hawaiian Islands, they would be the oldest avian lineage in the Hawaiian Islands [13Fleischer R.C. McIntosh C.E. Molecular systematics and biogeography of the Hawaiian avifauna.Studies Avian Biol. 2001; 22: 51-60Google Scholar, 18Price J.P. Clague D.A. How old is the Hawaiian biota? Geology and phylogeny suggest recent divergence.Proc. Biol. Sci. 2002; 269: 2429-2435Crossref PubMed Scopus (236) Google Scholar], and the more recent estimates would coincide well with the earliest postulated arrival of bird-pollinated plant lineages [10Givnish T.J. Millam K.C. Mast A.R. Paterson T.B. Theim T.J. Hipp A.L. Henss J.M. Smith J.F. Wood K.R. Sytsma K.J. Origin, adaptive radiation and diversification of the Hawaiian lobeliads (Asterales: Campanulaceae).Proc. Biol. Sci. 2008; (Published online October 14, 2008)https://doi.org/10.1098/rspb.2008.1204Crossref Scopus (255) Google Scholar, 18Price J.P. Clague D.A. How old is the Hawaiian biota? Geology and phylogeny suggest recent divergence.Proc. Biol. Sci. 2002; 269: 2429-2435Crossref PubMed Scopus (236) Google Scholar]. Unfortunately, the Mohoidae are the only family of songbirds to suffer complete extinction during the past few hundred years, and their extinction resulted in greater loss of avian phylogenetic diversity than if had they been merely a far-flung lineage of the Meliphagidae [15Ewen J.G. Flux I. Ericson P.G.P. Systematic affinities of two enigmatic New Zealand passerines of high conservation priority, the hihi or stitchbird Notiomystis cincta and the kokako Callaeas cinerea.Mol. Phylogenet. Evol. 2006; 40: 281-284Crossref PubMed Scopus (27) Google Scholar]. Detailed experimental procedures are provided in Supplemental Data, but summarized here. We sampled museum specimens of at least one individual of each species of Moho, a Chaetoptila angustipluma, a Samoan meliphagid (Gymnomyza samoenisis), and a crow (Corvus nasicus) (Table S1). DNA was isolated from the samples in isolated ancient-DNA laboratories (in the UK and USA) via standard phenol-chloroform and centrifugal-dialysis protocols with extreme care and controls to avoid or detect contamination. Primers were designed to amplify small segments from three nuclear genes (Table S2), and existing primers were used to amplify from mtDNA 12 s RNA, cytochrome b, and ATPase6 and ATPase8 genes. These products were sequenced, providing up to 1502 bp of RAG-1, 717 bp of mtDNA, 250 bp of β-fibrinogen intron 5, and 171 bp of β-fibrinogen intron 7. Comparative sequences were obtained from GenBank and relied mostly on two large RAG-1 datasets [9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar, 19Cibois A. Cracraft J. Assessing the passerine “tapestry”: Phylogenetic relationships of the Muscicapoidea inferred from nuclear DNA sequences.Mol. Phylogenet. Evol. 2004; 32: 264-273Crossref PubMed Scopus (48) Google Scholar]. Phylogenies were estimated from the data sets via maximum-parsimony, maximum-likelihood, and Bayesian approaches. Support for nodes was assessed by bootstrapping for the MP and ML trees, and by posterior probabilities for the Bayesian trees. In addition, we used Shimodaira-Hasegawa tests to test whether trees obtained through heuristic searches differed from ones constraining the position of Moho within the Meliphagidae. We did not combine the different sequence partitions (except for the β-fib sequences) because we had mostly different comparative taxa or individuals. Dates of particular nodes were estimated from the RAG-1 and mtDNA data sets (Table 1 and Supplemental Data) via NPRS and PL methods [17Sanderson M.J. Estimating absolute rates of molecular evolution and divergence times: A penalized likelihood approach.Mol. Biol. Evol. 2002; 19: 101-109Crossref PubMed Scopus (1465) Google Scholar] with a calibration date from Barker et al. of 82 million years for RAG1 [9Barker F.K. Cibois A. Schikler P. Feinstein J. Cracraft J. Phylogeny and diversification of the largest avian radiation.Proc. Natl. Acad. Sci. USA. 2004; 101: 11040-11045Crossref PubMed Scopus (576) Google Scholar]. This is the estimated date of the separation of Acanthisitta from the other Passeriformes, which was based on estimates of the timing of isolation of New Zealand from Antarctica. A calibration point internal to the genus Moho was used for the mtDNA data set and was based on the age of the island of Oahu [20Fleischer R.C. Tarr C.L. McIntosh C.E. Evolution on a volcanic conveyor belt: Using phylogeographic reconstructions and K-Ar based ages of the Hawaiian Islands to estimate molecular evolutionary rates.Mol. Ecol. 1998; 7: 533-545Crossref PubMed Scopus (381) Google Scholar]. We thank C. McIntosh, A.R. Hoelzel, and M. Haynie for logistical assistance; A. Driskell for discussion; J. Anderton for the beautiful bird illustrations; B. Schmidt for help with Figure 2; and curators of the American (J. Cracraft, G. Barrowclough), British (R. Prys-Jones), Cambridge University (M. de L. Brooke), Harvard University (D. Siegel-Causey), Liverpool (C. Fisher), and Smithsonian (G. Graves) museums for permission to sample rare specimens. The Smithsonian Institution Genetics Program and National Science Foundation (NSF DEB-0643291) provided funding. Download .pdf (.05 MB) Help with pdf files Document S1. A Taxonomic Summary, Detailed Supplemental Experimental Procedures, Supplemental Results, and Two Tables" @default.
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- W2056979431 title "Convergent Evolution of Hawaiian and Australo-Pacific Honeyeaters from Distant Songbird Ancestors" @default.
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