Advertisement
Browse Subject Areas
?

Click through the PLOS taxonomy to find articles in your field.

For more information about PLOS Subject Areas, click here.

  • Loading metrics

The New Zealand Thrush: An Extinct Oriole

The New Zealand Thrush: An Extinct Oriole

  • Ulf S. Johansson, 
  • Eric Pasquet, 
  • Martin Irestedt
PLOS
x

Abstract

The New Zealand Thrush, or Piopio, is an extinct passerine that was endemic to New Zealand. It has often been placed in its own family (Turnagridae), unresolved relative to other passerines, but affinities with thrushes, Australaian magpies, manucodes, whistlers, birds-of-paradise and bowerbirds has been suggested based on morphological data. An affinity with the bowerbirds was also indicated in an early molecular study, but low statistical support make this association uncertain. In this study we use sequence data from three nuclear introns to examine the phylogenetic relationships of the piopios. All three genes independently indicate an oriole (Oriolidae) affinity of the piopios, and the monophyly of the typical orioles (Oriolus), figbirds (Sphecotheres), and the piopios is strongly supported in the Bayesian analysis of the concatenated data set (posterior probability = 1.0). The exact placement of the piopios within Oriolidae is, however, more uncertain but in the combined analysis and in two of the gene trees the piopios are placed basal to the typical orioles while the third gene suggest a sister relationship with the figbirds. This is the first time an oriole affinity has been proposed for the piopios. Divergence time estimates for the orioles suggest that the clade originated ca 20 million years ago, and based on these estimates it is evident that the piopios must have arrived on New Zealand by dispersal across the Tasman Sea and not as a result of vicariance when New Zealand separated from Gondwana in the late Cretaceous.

Introduction

The New Zealand thrush, or Piopio, was an endemic, but now extinct, passerine on New Zealand. It was widely distributed on both the North and the South Island, from north of Auckland to Steven Island [1]. The birds on the two main islands were morphologically quite similar and have historically often been considered conspecific. They did, however, show some differences in plumage patterns as well as in size and other morphological characters [2] and are today often treated as two different species; the South Island Piopio (Turnagra capensis) and the North Island Piopio (T. tanagra) [3], [4]. The piopios apparently favoured forest undergrowth and fed on a wide range of food items, including berries, seeds, various invertebrates, eggs and other birds [1]. Of the two species the South Island Piopio was the first to be described in 1787 by Anders Sparrman but in the hundred years that followed it declined from being “common” to virtually extinct by the late 1800s. The North Island Piopio went through a similar decline, and for both species the last confirmed sightings were made at the turn of the century. The primary cause for the decline and final extirpation was apparently caused by predation from introduced predators such as cats, dogs, ferrets, stoats and rats [3].

Historically, the Piopio has often been referred to as the “New Zealand thrush” and was in some 19th century classifications placed in Turdidae [5], [6] (see [2] for a detailed review). However, Buller [7] questioned this association and placed the piopios in their own family, Turnagridae. Later studies have also shown that the thrush-like appearance does not reflect its phylogenetic affinity, but so far no consensus about its actual position within the passerine tree has been reached [4]. For instance, Oliver [1], [8] noted that the palate of the piopios indicated affinity with the Australian magpies (Gymnorhina tibicen) or manucodes (Manucodia). Mayr and Amandon [9], on the other hand, placed the piopios in their Pachycephalini together with e.g., Pachycephala, Colluricincla, and Falcunculus. However, Mayr and Amandon [9] also placed Pitohui, Oreoica, Hylocitrea, and Pachycare in this group, all of which are now considered part of other passerine radiations [10], [11], [12]. Olson et al. [2] examined the osteology, myology and pterolysis of the piopios and concluded that, albeit with considerable conflict among the characters, the piopios were related to the “birds-of-paradise and bowerbird assemblage”. Recent molecular studies, however, have shown that this assemblage is not monophyletic, but rather consists of three separate, unrelated lineages; bowerbirds (Ptilonorhynchidae), birds-of-paradise (Paradisaeidae) and satinbirds (Cnemophilidae) [13].

Nevertheless, a possible relationship with the bowerbirds was also indicated in a molecular study based on 900 base pair (bp) of cytochrome b by Chrisitidis et al. [14]. Unfortunately, low bootstrap support for the indicated relationships as well as a limited taxon sampling makes it difficult to draw any firm conclusions from these results. Furthermore, Gibb [15] has recently questioned the accuracy of the sequence used by Chrisitidis et al. [14] and noted that this sequence differs in 45 out of 307 positions (14.7%) compared to a new cytochrome b sequence from another individual. In a re-analysis based on 1783 bp of mitochondrial DNA, Gibb [15] instead concluded that the piopios likely belong in the “core Corvoidea” radiation, but were unable to confidently place it in any particular clade within that group.

The majority of the land bird species native to New Zealand, excluding those intentionally or unintentionally introduced by humans, appear to be the result of oversea colonization from the Australian continent [16], [17]. These colonization events range from a few decades (e.g. the Pacific Swallow Hirundo tahitica and White-faced Heron Egretta novaehollandiae) to many million years ago. But it has also been suggested that some groups, e.g. moas (Dinornithiformes) and New Zealand Parrots (Strigopidae), became isolated when New Zealand broke off from the Gondwanan continent, 80-60 million years ago (mya) [18], [19]. Among passerines, the New Zealand wrens (Acanthisittidae), which represent the first split in the passerine tree, may be another of these groups [13], [20], [21], [22], and it has also been suggested that the New Zealand Wattlebirds (Callaeatidae) became isolated at this event [23] (but see [24]).

The fact that it has been difficult to place the Piopios relative to other groups of passerines could possibly indicate that they are another of these “ancient lineages” that became isolated when New Zealand separated from Gondwana. [23]. The purpose of this study is to resolve the phylogenetic position of the piopios and discern whether this lineage represents one of these potential early divergences or a more recent dispersal.

Methods

Taxon sampling, amplification and sequencing

We examined the phylogenetic affinity of the South Island Piopio, Turnagra capensis, by analyzing DNA sequences from three nuclear introns; myoglobin intron 2, ornithine decarboxylase introns 6 to 7 (ODC), and glyceraldehyde-3-phosphodehydrogenase intron 11 (GAPDH). DNA was extracted from a foot pad sample obtained from one specimen of Turnagra capensis (MNHN 1999-1258), housed in the collections of Muséum National d'Histoire Naturelle, Paris. We used the Qiagen DNeasy Tissue Micro Kit for the extraction following the manufacturer's recommendation, except for that 20 µl of 1 molar DTT (dithiothreitol) was added during the lysis stage and the sample was heated to 72°C for 10 minutes after the buffer AL had been added. To minimize the risk of contamination, extraction where done in a special facility dedicated to the preparation of DNA samples from museum specimen and prior to extraction all equipment and buffers were sterilized with UV light.

Amplification and sequencing of fragmented DNA from old study skins require careful primer design, as target regions usually need to be divided into short, overlapping fragments. We amplified fragments of ca 200–250 bp, generally by using primer combinations that have previously successfully amplified a broad selection of passerine birds, e.g. [25], [26], but for some fragments new primers were also designed. The glyceraldehyde-3-phosphodehydrogenase intron 11 GAPDH was amplified in two fragments using the primer combinations G3Pcora1R [26]/G3PintL1 [27] and G3Pcora1F [26]/G3P14b [27], myoglobin intron 2 was amplified in four fragments using the primer combinations Myo2 [28]/Myo-cora182H [26], Myo-cora159L [26]/Myo344H [22], Myo-TurnF343 (AGT GAC TGG ACA CAA GGG ACA)/Myo-TurnR515 (GCA GAA GCA CTG GGC TCT AT), and Myo-cora491L [26]/Myo3F [29], and ornithine decarboxylase introns 6 to 7 (ODC) was amplified in three fragments using the primer combinations ODintF2 [25]/OD-TurnR1 (CAT GGA AAC TAC AAA AAG ATA CAA AC), OD-TurnF3 (TGT GTG TTT GAT ATG GGA GTA AGT)/OD-TurnR3 (GTA ATA GTC ATT TGA GTT TGA GCT G), and OD-TurnF4 (CTC ATC TAC AGA TGC ACT AAA ATT G)/ODintR4 [25]. We used hot-start touchdown PCR, with annealing temperatures for the first cycles generally just 1–2°C below the melting temperature of the primer with the lowest melting temperature. A representative thermocycling program for a given primer combination started with an initial denaturation at 95°C for 5 min, followed by two cycles of 95°C for 30 s, 59°C for 30 s, 72°C for 60 s, and two sets of cycles, each repeated two times, were the annealing temperature was lowered to 57°C and 55°C, respectively (all other temperatures and intervals identical). The thermocycling program was completed with 34 cycles with the annealing temperature set to 53°C and a final 72°C for 5 min. The extractions, amplifications, and sequencing procedures otherwise followed the procedures described in Irestedt et al. [25].

Our taxon sampling includes a broad selection of oscine birds, including representatives of the bowerbirds, satinbirds, birds-of-paradise, thrushes and whistlers. As preliminary assessments of our first sequences from Turnagra capensis indicted an oriole (Oriolidae) affinity, Oriolidae have been particularly densely sampled in the final data set. Menura novaehollandiae was used as outgroup as Menura novaehollandiae has been found to form the sister clade to all other oscine birds [13]. Voucher and GenBank accession numbers are given in Table 1.

thumbnail
Table 1. List of samples, with specimen numbers and GenBank accession numbers.

https://doi.org/10.1371/journal.pone.0024317.t001

Phylogenetic analyses

We used Bayesian inference to estimate phylogenetic relationships. The models for nucleotide substitutions used in the analyses were selected for each gene individually by the Akaike Information Criterion using the program MRMODELTEST 2.2 [30] in conjunction with PAUP* [31]. The number of indels was low and the sequences could easily be aligned by eye. The final alignment of the three gene segments included 1744 bp and all gaps were treated as missing data in the analyses.

Posterior probabilities of trees and parameters in the substitution models were approximated with MCMC and Metropolis coupling using the program MRBAYES 3.1.1 [32]. Analyses were performed for each of the individual genes (10 million generations) as well as on the concatenated data set (50 million generations), with trees sampled every 1000 generations. The program AWTY [33] was used to estimate when the chains had reached their apparent target distributions, and trees sampled during the burn-in phase were discarded.

Results

In total we obtained 1504 bp of nuclear DNA sequences from the Turnagra capensis sample (707 bp from myoglobin intron 2, 501 bp from ornithine decarboxylase introns 6 to 7 (ODC), excluding a region of about 100 bp that we were unable to sequence, and 296 bp from glyceraldehyde-3-phosphodehydrogenase intron 11 GAPDH). No mismatches between overlapping fragments were found in any of the target sequences, no heterozygotic sites were found, and none of the sequence fragments turned out to be identical to any other corresponding fragment in any other species checked.

The analysis of the concatenated, three-gene data set strongly supports an oriole affinity of the piopios (Fig. 1). The Turnagra forms a strongly supported clade (posterior probability [PP] = 1.0) together with the two oriole clades Oriolus and Sphecotheres. Within this clade the Turnagra is placed with weak support (PP = 0.69) as the sister of Oriolus. Pitohui dichrous, another species with a proposed oriole affinity [34], [35] is placed as the sister group of these three lineages. In this tree, the whistlers (Pachycephalidae), a group to which the piopios sometimes have been assigned, are placed as the sister group of the orioles.

thumbnail
Figure 1. Bayesian consensus tree of the concatenated, mixed model analysis of three nuclear introns (myoglobin, ODC and GAPDH).

Posterior probabilities are indicated at nodes. An asterisk * indicates a posterior probability of 1.0. The South Island Piopio (Turnagra capensis) is indicated in bold.

https://doi.org/10.1371/journal.pone.0024317.g001

All of the individual gene trees (not shown) indicate an oriole affinity of the piopios, although each gene indicates a slightly different topology. Both myoglobin and G3P place Turnagra as the sister of the three included Oriolus species, but the two gene trees differ in the placement of Sphecotheres and Pitohui relative to this group. In the myoglobin gene tree Sphecotheres and Pitohui are placed as sister taxa, and this clade is in turn placed as the sister group of the Oriolus/Turnagra clade, but in this gene tree there is also weak support for placing the Yellow-green Vireo Vireo flavoviridis with the former two taxa (PP = 0.60). In G3P, neither Sphecotheres nor Pitohui are placed with the Oriolus/Turnagra clade, but are placed with different taxa in different parts of the tree. The support values in the G3P gene tree are generally low (PP<0.95), and this tree is, in those parts of importance for this study, basically unresolved due to low posterior probabilities for the indicated relationships. The ODC gene tree is similar to the combined tree in that Pitohui is placed basal relative to an Oriolus/Turnagra/Sphecotheres clade, but in this tree Turnagra is placed as the sister group of Sphecotheres rather than Oriolus.

Discussion

Our data strongly point at an oriole affinity for the piopios and that they are nested within the Oriolidae. This clade consists of the typical orioles Oriolus (27 species) and figbirds Sphecotheres (3 species) [36], and in our study the piopios are placed with weak support as the sister group of the Oriolus, basal relative to other species of that clade (cf. [35]). Our study also confirms that the Hooded Pithoui Pitoui dichrous is part of this clade, but rather placed basal relative to the other taxa in this group instead of sisters to the figbirds as indicated by Jønsson et al. [35].

Orioles are distributed in the Eurasian, Afrotropical, Indomalayan and Australasian zoogeographical regions [36]. Most species of orioles in the former regions are bright yellow or red, whereas the figbirds and the other orioles of the Australasian region are mostly drab brown or olive green. The piopios, being olive-grey to olive-brown, were in this respect most similar to the Australasian orioles, and the South Island Piopio had brown streaking on the breast similar to e.g. the Australian Olive-backed Oriole (Oriolus sagittatus) as well as females and juveniles of many other oriole species. Very little is known about the biology of the piopios but they appear to have been omnivorous and fed on a wide range of food items, including insects, worms, fruits and berries, much like the orioles. In contrast to the orioles, which rarely descend to the ground for feeding [36], piopios appear to a large extent have foraged on the ground “grubbing with its bill among the dry leaves and other forest debris” [7]. This change in behavior to a more ground-living lifestyle has been rather common among New Zealand birds, and several species of e.g. rails, ducks, parrots and passerines have evolved flightlessness on New Zealand. However, this behavior made them more vulnerable to the introduced predators that arrived in the 19th century and ultimately caused the extinction of many species, including the piopios.

Biogeographical analyses [35] have shown that the Oriolidae likely originated in Australasia, and from there dispersed to other regions. Within the Oriolidae, figbirds, pithouis and the basalmost lineage of Oriolus are confined to the Australia, New Guinea and Wallacea; and the placement of the piopios among these groups indicate that the ancestor of the piopios also lived in this region and at some point dispersed to New Zealand. The dating analysis by Jønsson et al [35] indicate that the split between Sphecotheres/Pithoui and Oriolus took place around 20 Mya, and the earliest split within Oriolus, i.e. between the Australasian clade and all other Oriolus species, took place around 13 Mya. These estimates provide a rough time frame for the dispersal of the ancestor of the piopios and suggest that the dispersal to New Zealand took place no earlier than approximately 20 Mya.

New Zealand is part of the largely submerged continent Zealandia. This continent, which extends from Caledonia to the subantarctic islands off the cost of New Zealand, was in the Cretaceous above sea level and attached to the large southern hemisphere continent Gondwana. By the end of the Cretaceous (ca 82 Mya) the two continents had begun to separate but may have remained connected in what is now northern Australia until the Early Paleocene (65–61 Mya) or the Early Eocene (ca 52 Mya) [37], [38], [39], [40], [41]. Shortly after Zealandia had separated from Gondwana crustal thinning and stretching resulted in marine transgressions in the Eocene and Oligocene, and by the Late Oligocene most of this region was deep under water [39], [42]. The extent of the Oligocene transgression is unknown but it is clear that much of this region was under water and it has even been suggested that New Zealand was completely submerged around 25 Mya [42], [43].

The phylogenetic position of the piopios within the Oriolidae makes it unlikely that they became isolated on New Zealand when this continent broke off from Gondwana in the Cretaceous. Instead, the divergence time estimate for the Oriolidae [35] suggests that the dispersal took place long after the isolation of New Zealand. Based on these estimates it is likely that the piopios arrived after the Oligocene transgressions, which occurred around 25 Mya, but even though this is a reasonable assumption, these estimates are too crude to establish this with certainty. It is, nevertheless, evident that the piopios add to the list of species that colonized New Zealand once the Tasman Sea had opened rather than being Gondwana relicts.

Acknowledgments

We thank Per Ericson for invaluable input and support for the study. We also thank Pia Eldenäs and three anonymous for valuable comments on the manuscript.

Author Contributions

Conceived and designed the experiments: UJ EP MI. Performed the experiments: MI. Analyzed the data: UJ MI. Contributed reagents/materials/analysis tools: EP. Wrote the paper: UJ MI.

References

  1. 1. Oliver WRB (1955) New Zealand birds. Wellington: A.H. & A.W. Reed. WRB Oliver1955New Zealand birdsWellingtonA.H. & A.W. Reed
  2. 2. Olson SL, Parkes KC, Clench MH, Borecky SR (1983) The affinities of the New Zealand passerine genus Turnagra. Notornis 30: 319–336.SL OlsonKC ParkesMH ClenchSR Borecky1983The affinities of the New Zealand passerine genus Turnagra.Notornis30319336
  3. 3. Tennyson AJD, Martinson P (2006) Extinct birds of New Zealand. Wellington: Te Papa Press. AJD TennysonP. Martinson2006Extinct birds of New ZealandWellingtonTe Papa Press
  4. 4. Gill BJ, Bell BD, Chambers GK, Medway DG, Palma RL, et al. (2010) Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, Antarctica. Wellington: Ornithological Society of New Zealand & Te Papa Press. BJ GillBD BellGK ChambersDG MedwayRL Palma2010Checklist of the Birds of New Zealand, Norfolk and Macquarie Islands, and the Ross Dependency, AntarcticaWellingtonOrnithological Society of New Zealand & Te Papa Press
  5. 5. Buller WL (1869) On some new species of New-Zealand birds. Ibis 5: 37–43.WL Buller1869On some new species of New-Zealand birds.Ibis53743
  6. 6. Buller WL (1882) Manual of the birds of New Zealand. Wellington: George Didsbury, Government Printer. WL Buller1882Manual of the birds of New ZealandWellingtonGeorge Didsbury, Government Printer
  7. 7. Buller WL (1888) A History of the Birds of New Zealand (2nd edition). London: Published by the author. WL Buller1888A History of the Birds of New Zealand (2nd edition)LondonPublished by the author
  8. 8. Oliver WRB (1945) Avian evolution in New Zealand and Australia. Part II. Emu 45: 119–152.WRB Oliver1945Avian evolution in New Zealand and Australia. Part II.Emu45119152
  9. 9. Mayr E, Amadon D (1951) A classification of recent birds. American Museum Novitates 1496: 1–42.E. MayrD. Amadon1951A classification of recent birds.American Museum Novitates1496142
  10. 10. Spellman GM, Cibois A, Moyle RG, Winker K, Barker FK (2008) Clarifying the systematics of an enigmatic avian lineage: What is a bombycillid? Molecular Phylogenetics and Evolution 49: 1036–1040.GM SpellmanA. CiboisRG MoyleK. WinkerFK Barker2008Clarifying the systematics of an enigmatic avian lineage: What is a bombycillid?Molecular Phylogenetics and Evolution4910361040
  11. 11. Norman JA, Boles WE, Christidis L (2009) Relationships of the New Guinean songbird genera Amalocichla and Pachycare based on mitochondrial and nuclear DNA sequences. Journal of Avian Biology 40: 640–645.JA NormanWE BolesL. Christidis2009Relationships of the New Guinean songbird genera Amalocichla and Pachycare based on mitochondrial and nuclear DNA sequences.Journal of Avian Biology40640645
  12. 12. Norman JA, Ericson PGP, Jønsson KA, Fjeldså J, Christidis L (2009) A multi-gene phylogeny reveals novel relationships for aberrant genera of Australo-Papuan core Corvoidea and polyphyly of the Pachycephalidae and Psophodidae (Aves: Passeriformes). Molecular Phylogenetics and Evolution 52: 488–497.JA NormanPGP EricsonKA JønssonJ. FjeldsåL. Christidis2009A multi-gene phylogeny reveals novel relationships for aberrant genera of Australo-Papuan core Corvoidea and polyphyly of the Pachycephalidae and Psophodidae (Aves: Passeriformes).Molecular Phylogenetics and Evolution52488497
  13. 13. Barker FK, Cibois A, Schikler P, Feinstein J, Cracraft J (2004) Phylogeny and diversification of the largest avian radiation. Proceedings of National Academy of Sciences of the United States of America 101: 11040–11045.FK BarkerA. CiboisP. SchiklerJ. FeinsteinJ. Cracraft2004Phylogeny and diversification of the largest avian radiation.Proceedings of National Academy of Sciences of the United States of America1011104011045
  14. 14. Christidis L, Leeton PR, Westerman M (1996) Were bowerbirds part of the New Zealand fauna? Proceedings of National Academy of Sciences of the United States of America 93: 3898–3901.L. ChristidisPR LeetonM. Westerman1996Were bowerbirds part of the New Zealand fauna?Proceedings of National Academy of Sciences of the United States of America9338983901
  15. 15. Gibb GC (2010) Birds in a tree: A journey through avian phylogeny, with particular emphasis on the birds of New Zealand. Palmerston North, New Zealand: Massey University. GC Gibb2010Birds in a tree: A journey through avian phylogeny, with particular emphasis on the birds of New ZealandPalmerston North, New ZealandMassey University
  16. 16. Tennyson AJD (2010) The origin and history of the New Zealand's terrestrial verebrates. New Zealand Journal of Ecology 34: 6–27.AJD Tennyson2010The origin and history of the New Zealand's terrestrial verebrates.New Zealand Journal of Ecology34627
  17. 17. Trewick SA, Gibb GC (2010) Vicars, tramps and assembly of the New Zealand avifauna: a review of molecular phylogenetic evidence. Ibis 152: 226–253.SA TrewickGC Gibb2010Vicars, tramps and assembly of the New Zealand avifauna: a review of molecular phylogenetic evidence.Ibis152226253
  18. 18. Wright TF, Schirtzinger EE, Matsumoto T, Eberhard JR, Graves GR, et al. (2008) A multilocus molecular phylogeny of the parrots (Psittaciformes): support for a Gondwanan origin during the Cretaceous. Molecular Biology and Evolution 25: 2141–2156.TF WrightEE SchirtzingerT. MatsumotoJR EberhardGR Graves2008A multilocus molecular phylogeny of the parrots (Psittaciformes): support for a Gondwanan origin during the Cretaceous.Molecular Biology and Evolution2521412156
  19. 19. Cooper A, Lalueza-Fox C, Anderson S, Rambaut A, Austin JJ, et al. (2001) Complete mitochondrial genome sequences of two extinct moas clarify ratite evolution. Nature 409: 704–707.A. CooperC. Lalueza-FoxS. AndersonA. RambautJJ Austin2001Complete mitochondrial genome sequences of two extinct moas clarify ratite evolution.Nature409704707
  20. 20. Edwards SV, Boles WE (2002) Out of Gondwana: the origin of passerine birds. Trends in Ecology & Evolution 17: 347–349.SV EdwardsWE Boles2002Out of Gondwana: the origin of passerine birds.Trends in Ecology & Evolution17347349
  21. 21. Ericson PGP, Christidis L, Cooper A, Irestedt M, Jackson J, et al. (2002) A Gondwanan origin of passerine birds supported by DNA sequences of the endemic New Zealand wrens. Proceedings of the Royal Society of London B Biological Sciences 269: 435–441.PGP EricsonL. ChristidisA. CooperM. IrestedtJ. Jackson2002A Gondwanan origin of passerine birds supported by DNA sequences of the endemic New Zealand wrens.Proceedings of the Royal Society of London B Biological Sciences269435441
  22. 22. Ericson PGP, Irestedt M, Johansson US (2003) Evolution, biogeography, and patterns of diversification in passerine birds. Journal of Avian Biology 34: 3–15.PGP EricsonM. IrestedtUS Johansson2003Evolution, biogeography, and patterns of diversification in passerine birds.Journal of Avian Biology34315
  23. 23. Newton I (2003) The speciation and biogeography of birds. London: Academic Press. I. Newton2003The speciation and biogeography of birdsLondonAcademic Press
  24. 24. Shepherd LD, Lambert DM (2007) The relationships and origins of the New Zealand wattlebirds (Passeriformes, Callaeatidae) from DNA sequences analyses. Molecular Phylogenetics and Evolution 43: 480–492.LD ShepherdDM Lambert2007The relationships and origins of the New Zealand wattlebirds (Passeriformes, Callaeatidae) from DNA sequences analyses.Molecular Phylogenetics and Evolution43480492
  25. 25. Irestedt M, Ohlson JI, Zuccon D, Källersjö M, Ericson PGP (2006) Nuclear DNA from old collections of avian study skins reveals the evolutionary history of the Old World suboscines (Aves, Passeriformes). Zoologica Scripta 35: 567–580.M. IrestedtJI OhlsonD. ZucconM. KällersjöPGP Ericson2006Nuclear DNA from old collections of avian study skins reveals the evolutionary history of the Old World suboscines (Aves, Passeriformes).Zoologica Scripta35567580
  26. 26. Jønsson KA, Irestedt M, Fuchs J, Ericson PGP, Christidis L, et al. (2008) Explosive avian radiations and multi-directional dispersal across Wallacea: evidence from the Campephagidae and other Crown Corvida (Aves). Molecular Phylogenetics and Evolution 47: 221–236.KA JønssonM. IrestedtJ. FuchsPGP EricsonL. Christidis2008Explosive avian radiations and multi-directional dispersal across Wallacea: evidence from the Campephagidae and other Crown Corvida (Aves).Molecular Phylogenetics and Evolution47221236
  27. 27. Fjeldså J, Zuccon D, Irestedt M, Johansson US, Ericson PGP (2003) Sapayoa aenigma: a New World representative of “Old World suboscines”. Proceedings of the Royal Society of London B Biological Sciences (Supplement) 270: S238–S241.J. FjeldsåD. ZucconM. IrestedtUS JohanssonPGP Ericson2003Sapayoa aenigma: a New World representative of “Old World suboscines”.Proceedings of the Royal Society of London B Biological Sciences (Supplement)270S238S241
  28. 28. Slade RW, Moritz C, Heideman A, Hale PT (1993) Rapid assessment of single-copy nuclear DNA variation in diverse species. Molecular Ecology 2: 359–373.RW SladeC. MoritzA. HeidemanPT Hale1993Rapid assessment of single-copy nuclear DNA variation in diverse species.Molecular Ecology2359373
  29. 29. Heslewood MM, Elphinstone MS, Tidemann SC, Baverstock PR (1998) Myoglobin intron variation in the Gouldian Finch Erythrura gouldiae assessed by temperature gradient gel electrophoresis. Electrophoresis 19: 142–151.MM HeslewoodMS ElphinstoneSC TidemannPR Baverstock1998Myoglobin intron variation in the Gouldian Finch Erythrura gouldiae assessed by temperature gradient gel electrophoresis.Electrophoresis19142151
  30. 30. Nylander JAA (2004) MrModeltest. v2 ed: Program distributed by the author. Evolutionary Biology Centre, Uppsala University. JAA Nylander2004MrModeltest. v2 ed: Program distributed by the authorEvolutionary Biology Centre, Uppsala University
  31. 31. Swofford DL (2001) PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Version 4. Sunderland, Massachusetts: Sinauer Associates. DL Swofford2001PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Version 4Sunderland, MassachusettsSinauer Associates
  32. 32. Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19: 1572–1574.F. RonquistJP Huelsenbeck2003MrBayes 3: Bayesian phylogenetic inference under mixed models.Bioinformatics1915721574
  33. 33. Nylander JAA, Wilgenbusch JC, Warren DL, Swofford DL (2008) AWTY (are we there yet?): a system for graphical exploration of MCMC convergence in Bayesian phylogenetics. Bioinformatics 24: 581–583.JAA NylanderJC WilgenbuschDL WarrenDL Swofford2008AWTY (are we there yet?): a system for graphical exploration of MCMC convergence in Bayesian phylogenetics.Bioinformatics24581583
  34. 34. Jønsson KA, Bowie RCK, Norman JA, Christidis L, Fjeldså J (2008) Polyphyletic origin of toxic Pitohui birds suggest widespread occurence of toxity in corvoid birds. Biology Letters 4: 71–74.KA JønssonRCK BowieJA NormanL. ChristidisJ. Fjeldså2008Polyphyletic origin of toxic Pitohui birds suggest widespread occurence of toxity in corvoid birds.Biology Letters47174
  35. 35. Jønsson KA, Bowie RCK, Moyle RG, Irestedt M, Christidis L, et al. (2010) Phylogeny and biogeography of Oriolidae (Aves: Passeriformes). Ecography 33: 232–241.KA JønssonRCK BowieRG MoyleM. IrestedtL. Christidis2010Phylogeny and biogeography of Oriolidae (Aves: Passeriformes).Ecography33232241
  36. 36. Walther B, Jones P (2008) Family Oriolidae (Orioles). In: Del Hoyo J, Elliott A, Christie DA, editors. Handbook of the birds of the world. Barcelona: Lynx Edicions. B. WaltherP. Jones2008Family Oriolidae (Orioles).J. Del HoyoA. ElliottDA ChristieHandbook of the birds of the worldBarcelonaLynx Edicions
  37. 37. Gaina C, Müller RD, Royer J-Y, Stock J, Hardebeck J, et al. (1998) The tectonic history of the Tasman Sea: A puzzle with thirteen pieces. Journal of Geophysical Research 103: 12413–12433.C. GainaRD MüllerJ-Y RoyerJ. StockJ. Hardebeck1998The tectonic history of the Tasman Sea: A puzzle with thirteen pieces.Journal of Geophysical Research1031241312433
  38. 38. Gaina C, Müller RD, Royer J-Y, Symonds P (1999) The evolution of the Louisiade Triple junction. Journal of Geophysical Research 104: 12927–12939.C. GainaRD MüllerJ-Y RoyerP. Symonds1999The evolution of the Louisiade Triple junction.Journal of Geophysical Research1041292712939
  39. 39. Ladiges PY, Cantrill D (2007) New Caledonia-Australian connections: biogeographic patterns and geology. Australian Systematic Botany 20: 383–389.PY LadigesD. Cantrill2007New Caledonia-Australian connections: biogeographic patterns and geology.Australian Systematic Botany20383389
  40. 40. Neall VE, Trewick SA (2008) The age and origin of the Pacific islands: a geological overview. Philosophical Transactions of the Royal Society of London B Biological Sciences 363: 3293–3308.VE NeallSA Trewick2008The age and origin of the Pacific islands: a geological overview.Philosophical Transactions of the Royal Society of London B Biological Sciences36332933308
  41. 41. Schellart WP, Lister GS, Toy VG (2006) A late Cretaceous and Cenozoic reconstruction of the Southwest Pacific region: Tectonics contolled by subduction and slab rollback processes. Earth-Science Reviews 191–233: 191–233.WP SchellartGS ListerVG Toy2006A late Cretaceous and Cenozoic reconstruction of the Southwest Pacific region: Tectonics contolled by subduction and slab rollback processes.Earth-Science Reviews191–233191233
  42. 42. Trewick SA, Paterson AM, Campbell HJ (2007) Hello New Zealand. Journal of Biogeography 34: 1–6.SA TrewickAM PatersonHJ Campbell2007Hello New Zealand.Journal of Biogeography3416
  43. 43. Landis CA, Campbell HJ, Begg JG, Paterson AM, Trewick SA (2006) The drowning of Zealandia: evidence and implications. In: Trewick SA, Phillips MJ, editors. Geological Society of New Zealand Miscellaneous Publication Wellington, New Zealand. 21 p.CA LandisHJ CampbellJG BeggAM PatersonSA Trewick2006The drowning of Zealandia: evidence and implications.SA TrewickMJ PhillipsGeological Society of New Zealand Miscellaneous Publication Wellington, New Zealand21
  44. 44. Fuchs J, Cruaud C, Couloux A, Pasquet E (2007) Complex biogeographic history of the cuckoo-shrikes and allies (Passeriformes: Campephagidae) revealed by mitochondrial and nuclear sequence data. Molecular Phylogenetics and Evolution 44: 138–153.J. FuchsC. CruaudA. CoulouxE. Pasquet2007Complex biogeographic history of the cuckoo-shrikes and allies (Passeriformes: Campephagidae) revealed by mitochondrial and nuclear sequence data.Molecular Phylogenetics and Evolution44138153
  45. 45. Fuchs J, Fjeldså J, Pasquet E (2006) An ancient African radiation of corvoid birds (Aves: Passeriformes) detected by mitochondrial and nuclear sequence data. Zoologica Scripta 35: 375–385.J. FuchsJ. FjeldsåE. Pasquet2006An ancient African radiation of corvoid birds (Aves: Passeriformes) detected by mitochondrial and nuclear sequence data.Zoologica Scripta35375385
  46. 46. Irestedt M, Ohlson JI (2008) The division of the major songbird radiation into Passerida and ‘core Corvoidea’ (Aves: Passeriformes) - the species tree vs gene trees. Zoologica Scripta 37: 305–313.M. IrestedtJI Ohlson2008The division of the major songbird radiation into Passerida and ‘core Corvoidea’ (Aves: Passeriformes) - the species tree vs gene trees.Zoologica Scripta37305313
  47. 47. Ericson PGP, Christidis L, Irestedt M, Norman JA (2002) Systematic affinities of the lyrebirds (Passeriformes: Menura), with a novel classification of the major groups of passerine bird. Molecular Phylogenetics and Evolution 25: 53–62.PGP EricsonL. ChristidisM. IrestedtJA Norman2002Systematic affinities of the lyrebirds (Passeriformes: Menura), with a novel classification of the major groups of passerine bird.Molecular Phylogenetics and Evolution255362
  48. 48. Jønsson KA, Fjeldså J, Ericson PGP, Irestedt M (2007) Systematic placement of an enigmatic Southeast Asian taxon Eupetes macrocercus and implications for the biogeography of a main songbird radiation, the Passerida. Biology Letters 3: 323–326.KA JønssonJ. FjeldsåPGP EricsonM. Irestedt2007Systematic placement of an enigmatic Southeast Asian taxon Eupetes macrocercus and implications for the biogeography of a main songbird radiation, the Passerida.Biology Letters3323326
  49. 49. Irestedt M, Fuchs J, Jønsson KA, Ohlson JI, Pasquet E, et al. (2008) The systematic affinity of the enigmatic Lamprolia victoriae (Aves: Passeriformes)—An example of avian dispersal between New Guinea and Fiji over Miocene intermittent land bridges? Molecular Phylogenetics and Evolution 48: 1218–1222.M. IrestedtJ. FuchsKA JønssonJI OhlsonE. Pasquet2008The systematic affinity of the enigmatic Lamprolia victoriae (Aves: Passeriformes)—An example of avian dispersal between New Guinea and Fiji over Miocene intermittent land bridges?Molecular Phylogenetics and Evolution4812181222
  50. 50. Filardi CE, Moyle RG (2005) Single origin of a pan-Pacific bird group and upstream colonization of Australasia. Nature 438: 216–219.CE FilardiRG Moyle2005Single origin of a pan-Pacific bird group and upstream colonization of Australasia.Nature438216219
  51. 51. Fuchs J, Bowie RCK, Fjeldså J, Pasquet E (2004) Phylogenetic relationships of the African bush-shrikes and helmet-shrikes (Passeriformes: Malaconotidae). Molecular Phylogenetics and Evolution 33: 428–439.J. FuchsRCK BowieJ. FjeldsåE. Pasquet2004Phylogenetic relationships of the African bush-shrikes and helmet-shrikes (Passeriformes: Malaconotidae).Molecular Phylogenetics and Evolution33428439
  52. 52. Ericson PGP, Johansson US (2003) Phylogeny of Passerida (Aves: Passeriformes) based on nuclear and mitochondrial sequence data. Molecular Phylogenetics and Evolution 29: 126–138.PGP EricsonUS Johansson2003Phylogeny of Passerida (Aves: Passeriformes) based on nuclear and mitochondrial sequence data.Molecular Phylogenetics and Evolution29126138
  53. 53. Zuccon D, Cibois A, Pasquet E, Ericson PGP (2006) Nuclear and mitochondrial sequence data reveal the major lineages of starlings, mynas and related taxa. Molecular Phylogenetics and Evolution 41: 333–344.D. ZucconA. CiboisE. PasquetPGP Ericson2006Nuclear and mitochondrial sequence data reveal the major lineages of starlings, mynas and related taxa.Molecular Phylogenetics and Evolution41333344
  54. 54. Zuccon D, Ericson PGP (2010) A multi-gene phylogeny disentangles the chat-flycatcher complex (Aves: Muscicapidae). Zoologica Scripta 39: 213–224.D. ZucconPGP Ericson2010A multi-gene phylogeny disentangles the chat-flycatcher complex (Aves: Muscicapidae).Zoologica Scripta39213224