Phylogenies of Ratites
Dyke 2003
Neornithes
|--Paleognathae
| |--Tinamidae
| `--+--Lithornis
| `--Ratites
| |--+--Apteryx
| | `--Dinornis
| `--+--Palaeotis
| |--+--Casuarius
| | `--Dromaius
| `--+--Rhea
| `--Struthio
`--Neognathae
|--Galloanserae
| |--Anseriforms
| `--Galliforms`
`--Neoaves “Although controversy still exists regarding precise relationships within Paleognaths, especially within the ratites, most workers agree that tinamous are the basal sister taxon with respect to the remaining ratite birds.” Dyke 2003.
Tuinen, Sibly and Hedges 1998: DNA Study
.
|-----Tinamou
`---+------------Struthio
`-----+--------Rhea
`---+-----Apteryx
|-----Dromaius (Emu)
`-----Casuarius
Study is online at: http://evo.bio.psu.edu/hedgeslab/Publications/PDF-files/98.pdf
Lee, Feinstein and Cracraft: Molecular and Morphological Data Sets
|--------------Galliforms/Anseriformes
`--+-----------Tinamidae
`--+-----+--Dinornithids
| `--Apteryx
`--+--+--Casuarius
| `--Dromaius
`--+--Struthio
`--Rhea
“The debate on where over the placement of Apteryx on the tree seems dependent on where the outgroup joins
the ingroup”
Cooper et al 1992: DNA Study
.
|-----Rhea
`---+--Moa
`-----+----Ostrich
`---+------Apteryx
|-----Emu
`-----Casuarius
“The most surprising result of the phylogenetic analysis is that the two groups of New Zealand ratites have different
origins, the moas representing an earlier divergence among ratites, whereas the kiwi more recently shared an ancestor
with the ostrich and the Australian ratites”
Cooper et al 1991: DNA Study
.
|- Tinamous, chicken
`--+------------Rhea
`-+-------+--Emeus
| `--Dinorinis
`-+------------Ostrich
`-+?????-Elephant-bird
`-+------Kiwi
`--+---Emu
`---Casuarius
“Although these data do not allow a presice placement or an estimate of divergence date, the elephant-bird is
clearly not the result of a recent divergence from the ostrich, or any other ratite lineage.”
“The tight clustering of dates and lack of phylogenetic resolution, despite the long sequences, also suggests
that the derived ostrich, kiwi, and emu/cassowary taxa speciated rapidly, possibly in the remnant Gondwanan
landmass of Antarctica/Australia.”
Cracraft, 1974: Morphological Study
Paleognathiforms
|--Tinami
| `--tinamidae
`Ratiti
|--Apteryges
| |--Apterigidae
| `Diornithidae
`--Struthiones
|--Aepyornithidae
`--Struthionoidae
|--+--Cassuariidae
| `--Dromiceinae
`--Struthionidae
|--Struthioninae
`--Rheinae
Robinson, 2004
Neornithes
|--Paleognathae
| `--Prototinamou(Undiscovered, Parks and Clark 1966)(Def: the common ancestor of ratites and tinamous)
| |--Tinamidae (only earliest fossils are listed)
| | |--Eudromia (Miocene Argentina)
| | |--Eudromia olsoni (Pliocene, Argentina)
| | |--Querandiornis romani (Pliocene, Argentina)
| | `--Nothura parvulus (Pliocene, Argentina)
| |?-Dinornithiformes (Order of Moas)
| | |--Diornithidae (Dinornithis Moas)
| | | |--Dinornis struthoids (Holocene New Zealand)
| | | |--Dinornis novazealandiae (Holocene New Zealand)
| | | `--Dinornis Maximus (Holocene New Zealand)
| | `--Anomalopterygidae(Emeidae)
| | |--Anomalopteryginae
| | | |--Anomalopteryx didiformus (New Zealand)
| | | |--Megalapteryx didinus (Holocene New Zealand)
| | | |--Pachyornis elephantopus (Holocene Stewart Island off South Island, New Zealand)
| | | |--Pachyornis australis (Holocene New Zealand)
| | | `--Pachyornis mappini (Holocene North Island New Zealand)
| | `--Emeinae
| | |--Emeus crassus (Holocene South Island New Zealand)
| | |--Euryapteryx gravis?
| | |--Euryapteryx curtis (Late Pliocene-Holocene North Island, Great Barrier Island)
| | |--Euryapteryx geranoids (Holocene New Zealand)
| | `--Zelornis?
| |?-Remiornis minor (Paleocene France)
| `?-+--Green River Paleognath (Houde 1986)
| `--+--Lithornidae (genus)
| | |--Lithornis celetius (Paleocene North America)(Houde 1988)
| | |--Lithornis plebius (Eocene North America & London Clay)(Houde 1988)
| | |--Lithornis promiscuus(Eocene North America)(Houde 1988)
| | |--Lithornis nasi (Eocene North America & London Clay)(Houde 1988)
| | |--Lithornis hookeri (Eocene London Clay)(Houde 1988)
| | |--Lithornis vulturinus (Eocene London Clay)(Houde 1988)
| |?-Pseudocrypturus cercanaxius (Eocene North America & London Clay)(Houde 1988)
| `--+--Paracathartes howardae (Eocene North America)(Houde 1988)
| ??-Apterygiforms (Order of kiwis)
| | `--Apterygidae (Kiwi family)
| | |--Apteryx australis (Brown Kiwi)
| | |--Apteryx haastii (Great spotted Kiwi)
| | `--Apteryx owenii (Little Spotted Kiwi)
| ??-Casuariiforms (Order of casuaries and emus)
| | `--+--Dromaius gidju (Patterson and Rich, 1987)
| | |--Casuariidae (Cassowary Family)
| | | |--Casuarius casuarius
| | | |--Casuarius unappendiculatus
| | | `--Casuarius bennetti
| | `--Dromiceidae (Emu family)
| | |--Dromaius ater (Holocene of King Island off South Australia)
| | |--Dromaius baudinianus (Holocene of Kangeroo Island off South Australia)
| | |--Dromaius diemenisis (Holocene of Tasmania off South Australia)
| | |--Dromaius ocypus (A.H. Miller, 1963)
| | `--Dromaius novaehollandiae
| ??-Aepyornithidae
| | |--Mullerornis betsilei (Pliestocene Madagascar)
| | |--Mullerornis agilis (Pliestocene Madagascar)
| | |--Mullerornis rudis (Pliestocene Madagascar)
| | |--Aepyornis medius
| | |--Aepyornis hildebrandti
| | |--Aepyornis gracilis
| | `--Aepyornis maximus
| `?-+--Palaeotididae
| | `--Palaeotis weigelti (Eocene Germany)(Houde and Haubold 1987)
| |?-Rheidae (Rhea family)
| | |?-Opisthodactylidae
| | | |--Diogenornis fragilis (Paleocene, Brazil)(Alvarenga 1983)
| | | `--Opisthodactylus patagonicus (Miocene, Argentina)
| | |--Heterorhea dabbenei (Pliocene Argentina)(Tambussi, 1995)
| | |--Hinasuri nehuensis (Pliocene Argentina)(Tambussi, 1995)
| | |--Pterocnemia pennata (Darwin's or Lesser Rhea)
| | `--Rhea americana (Greater Rhea)
| `?-Struthionidae (Ostrich family)
| |--Struthio coppensi (Miocene S. Africa) (Murer Chauvier et al 1996)
| |--Strthio orvoli (Miocene Moldavia)(Kurochkin and Lungu 1970)
| |--Struthio asiaticus (India)
| |--Struthio cheronensis (Greece Ukraine, Kazakhstan)
| |--Struthio wimani (China, Mongolia)
| |--Struthio bradydactylus (Odesa)
| `--Stuthio cameleus
`--Neognathae
|--Galloanserae
| |--+--Anseriforms
| `--Galliforms
`--Neoaves