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John Harshman and Joseph W. Brown
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taxon links [up-->]Rheidae [up-->]Tinamiformes [up-->]Dinornithidae [up-->]Apterygidae [up-->]Casuariidae [up-->]Aepyornithidae [up-->]Struthionidae [up-->]Dromaiidae extinct icon extinct icon Phylogenetic position of group is uncertain[down<--]Neornithes Interpreting the tree
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This tree diagram shows the relationships between several groups of organisms.

The root of the current tree connects the organisms featured in this tree to their containing group and the rest of the Tree of Life. The basal branching point in the tree represents the ancestor of the other groups in the tree. This ancestor diversified over time into several descendent subgroups, which are represented as internal nodes and terminal taxa to the right.

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You can click on the root to travel down the Tree of Life all the way to the root of all Life, and you can click on the names of descendent subgroups to travel up the Tree of Life all the way to individual species.

For more information on ToL tree formatting, please see Interpreting the Tree or Classification. To learn more about phylogenetic trees, please visit our Phylogenetic Biology pages.

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Tree based on Harshman et al. (2008). Position of extinct families is unclear.
Containing group: Neornithes

Discussion of Phylogenetic Relationships

Relationships among paleognaths have been controversial and are likely to remain so for some time, but the tree shown above is the current best estimate, based on analysis of 20 unlinked nuclear genes (Harshman et al. 2008).  This tree shows the ratites (flightless paleognaths, including all extant paleognaths except the tinamous)  to be polyphyletic, a major departure from most recent analyses of molecular (Cooper et al. 2001, Haddrath and Baker 2001, Sibley and Ahlquist 1990) and morphological (Lee et al. 1997, Livezey and Zusi 2007) data.

All possible relationships among the paleognaths, including complete polyphyly, have been proposed at various times (Olson 1985, Houde 1988), but in the past few decades, a few points have been settled. Paleognath monophyly has become clear, and it is universally agreed that Dromaiidae (emus) and Casuaridae (cassowaries) are close relatives. And, until recently, monophyly of the ratites had been generally accepted. Beyond this, however, there has been no agreement. Most morphological analyses have found Apterygidae (kiwis) to be the sister group of all other ratites. Most molecular analyses have found kiwis to be the sister group of emus and cassowaries.

There have been a few holdouts against ratite monophyly; both Elzanowski (1995) and Bock and Buhler (1990), on the basis of morphological characters, found Struthionidae (ostriches) to be the sister group of all other paleognaths, and this is also what the nuclear DNA data show. Nuclear data also find, in accord with previous molecular analyses, that kiwis are the sister group of the emus and cassowaries, the three groups together consituting a clade of Australasian ratites. However, the data are unable to resolve relationships among the Australasian ratites, Rheidae (rheas), and tinamous.

The importance of evolutionary models in phylogenetic analyses is borne out by the most recent analyses of mitochondrial data (Phillips et al., 2010), in which the same data that previously resulted in ratite monophyly have been reanalyzed using a more complex model, with results consistent with the nuclear DNA tree above; that is, with ostrich as the sister group of all other paleognaths, though support is poor.

This tree implies that the various flightless paleognaths lost their ability to fly (and thus their sternal keels) independently of each other. The alternative possibility, that tinamous regained the ability to fly (and their sternal keels) after inheriting flightlessness from a common ratite ancestor, is considered less likely.

Placement of the extinct groups is unclear.  Morphological and mitochondrial DNA data are in disgreement on relationships of Dinornithidae (moas), and there are so far no nuclear DNA sequences for extinct birds. The recent reanalysis of mitochondrial data (Phillips et al., 2010) found moas to be the sister group of tinamous. This is also in agreement with a combined analysis of nuclear and mitochondrial data (Harshman, unpublished), in which moas were represented by mitochondrial data only.

For more information about alternative hypotheses see Paleognath Relationships.

Other Names for Palaeognathae


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Ansari, H., N. Kakagi, and M. Sasaki. 1988. Morphological differentiation of sex chromosomes in three species of ratite birds. Cytogenetics and Cell Genetics 47: 185-188.

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Davies, S. J. J. F. 2002. Ratites and Tinamous. Bird Families of the World. Oxford University Press, Oxford, New York.

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Harrison, G. L., McLenachan, M. J. Phillips, K. E. Slack, A. Cooper, and D. Penny. 2004. Four new avian mitochondrial genomes help get to basic evolutionary questions in the late Cretaceous. Molecular Biology and Evolution 21: 974-983.

Harshman, J., E. L. Braun, M. J. Braun, C. J. Huddleston, R. C. K. Bowie, J. L. Chojnowski, S. J. Hackett, K.-L. Han, R. T. Kimball, B. D. Marks, K. J. Miglia, W. S. Moore, S. Reddy, F. H. Sheldon, D. W. Steadman, S. J. Steppan, C. C. Witt, and T. Yuri. 2008. Phylogenomic evidence for multiple losses of flight in ratite birds. Proc. Natl. Acad. Sci. USA 105:13462-12467.

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Information on the Internet

Title Illustrations
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Scientific Name Struthio camelus
Location Simon's Town, Western Cape, South Africa
Specimen Condition Live Specimen
Source Wild Ostrich
Source Collection Flickr
Image Use creative commons This media file is licensed under the Creative Commons Attribution-NonCommercial-ShareAlike License - Version 2.0.
Copyright © 2005 Josh M
Scientific Name Dromaius novaehollandiae
Location Captive at Colin Mackenzie Sanctuary, Healesville, South Australia
Comments Emus
Creator Dr. G Dallas and Margaret Hanna
Specimen Condition Live Specimen
Source Collection CalPhotos
Copyright © 2004 California Academy of Sciences
Scientific Name Eudromia elegans
Location Biedma, Chubut, Argentina
Specimen Condition Live Specimen
Source Elegant Crested Tinamou
Source Collection Flickr
Image Use creative commons This media file is licensed under the Creative Commons Attribution-NonCommercial-ShareAlike License - Version 2.0.
Copyright ©
About This Page

Joseph W. Brown
University of Michigan Museum of Zoology, Ann Arbor, Michigan, USA

Correspondence regarding this page should be directed to John Harshman at

Page: Tree of Life Palaeognathae. Authored by John Harshman and Joseph W. Brown. The TEXT of this page is licensed under the Creative Commons Attribution-NonCommercial License - Version 3.0. Note that images and other media featured on this page are each governed by their own license, and they may or may not be available for reuse. Click on an image or a media link to access the media data window, which provides the relevant licensing information. For the general terms and conditions of ToL material reuse and redistribution, please see the Tree of Life Copyright Policies.

Citing this page:

Harshman, John and Joseph W. Brown. 2010. Palaeognathae. Version 13 May 2010 (under construction). http://tolweb.org/Palaeognathae/15837/2010.05.13 in The Tree of Life Web Project, http://tolweb.org/

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