Information about Paleognathae

Paleognaths Fossil range: Middle Eocene? - Recent
A kiwi A kiwi
Scientific classification
Kingdom: Animalia Phylum: Chordata Class: Aves Subclass: Carinatae Infraclass: Neornithes Superorder: Paleognathae Pycraft, 1900
Orders
Primary Classification Lithornithiformes Dinornithiformes Aepyornithiformes Struthioniformes Rheiformes Casuariiformes Apterygiformes Tinamiformes Alternate Classification Lithornithiformes Struthioniformes Tinamiformes see text for more info
Synonyms
Palaeognathia Huxley, 1867

The Paleognathae or paleognaths ("old jaws") are one of the two living superorders of birds. The other living superorder is Neognathae.

The paleognaths contain several living orders of birds, the Tinamiformes (tinamous), the Apterygiformes (kiwis), Casuariiformes (cassowaries and emus), Rheiformes (rheas), and the Struthioniformes (ostriches). All but the tinamous are flightless. There are also many extinct orders: the Lithornithiformes, the Ambiornithiformes, the Gansuiformes, the Paleocursornithiformes, the Dinornithiformes (moas), and the Aepyornithiformes (elephant birds)

Most paleognaths have long necks and long legs, and are specialized for running rather than flight; indeed, the ratites are all completely flightless (with the partial exception of the tinamous). The group is not separated because of this though, it is rather based on the form of the jaw. Paleognaths other than tinamous are commonly known as ratites from the Latin word for raft, ratis, because they have a breastbone shaped like a raft.

Anatomy & Characteristics of Paleognath Birds

Ratite birds are the easiest birds to distinguish - they are mostly large, flightless, have extended necks, a breastbone shaped like a raft (with the exception of tinamous), they have a simpified wing bone structure, strong legs, and no feather vanes, making it unnecessary to oil their feathers. And as a direct result of this they have no preen gland that contains preening oil. Ratite sizes range from 10 inches (25 centimeters) to 9 feet (2.7 meters) and weight can be from 2.86 pounds (1.3 kilograms) to 345 pounds (155.25 kilograms). Ostriches are the largest struthioniforms (members of the Struthioniformes order), with long legs and neck. They range in height from 5.7 to 9 feet (1.8 to 2.7 meters) and weigh from 139 to 345 pounds (63 to 157 kilograms). They have loose-feathered wings. Males have black and white feathers while the female has grayish brown feathers. Emus are about 6.5 feet in height and weigh 51 to 120 pounds (23 to 55 kilograms). They have long, strong legs and can run up to 30 miles per hour (48 kilometers per hour). They have short wings and the adults have brown feathers. Rheas are 4.5 to 5.6 feet (1.3 to 1.7 meters) and weigh 55 to 88 pounds (24.75 to 40 kilograms). Their feathers are gray or spotted brown and white. Cassowaries are 3.3 to 5.6 feet (1 to 1.7 meters) in height and weigh 30 to 130 pounds (14 to 59 kilograms). They have tiny wings with black feathers. Kiwis are the smallest of ratites, ranging in height from 14 to 22 inches (35 to 55 centimeters) and weight 2.6 to 8.6 pounds (1.2 to 3.9 kilograms). They have brown and black hair-like feathers. The tinamous have a keeled breastbone (shaped like a wishbone) and can fly. They range in size from 8 to 21 inches (20 to 53 centimeters) and weigh 1.4 ounces to 5 pounds (43 grams to 2.3 kilograms). The basic anatomy of ratites is simple in principle and there are few exceptions to the rules stated.

Evolution

Paleognaths probably descended from a common ancestor in the late Cretaceous period on the supercontinent of Gondwana. As the continents separated several forms of paleognaths reached different parts of the world, to support this idea all the continents where living (ostrich, cassowary, kiwi, tinamous, emu, and the rhea) and various fossil forms are found were connected during the late Cretaceous. Recent genetic evidence seems to also point towards this theory, DNA shows that they may not be descended from one common ancestor, but two or more. The results also show that ratites probably didn't lose the ability to fly until the middle Eocene. Many similarites in morphology and genetic analysis show that there is probably no convergent evolution between ratite birds. Currently, there is no exact way to way where, why, or when the paleognaths diverged.

Taxonomy

As told in the Evolution section of this article, the exact evolutionary history of the paleognaths is unresolved presently. Despite this many taxonomists try to place them in the grand scheme of the Aves. As a result two main branches developed, they are shown in the classification part below.

Early Taxonomy

Initially the ratites were placed in a single group Ratita. Later it was realized that they were too different to be placed in a single order, so they were split into the several orders in the infobox at the top of this article. Then they were split into two orders, the Struthioniformes and the Lithioniformes. Although the first classification is adopted in this article, either one is commonly used in sources.

Cladistics

Evolutionary Cladogram

+ (Lithornithiformes and other Fossil Paleognaths) | '

---

+ (Living Ratites)+ (Struthioniformes) | '---+ (Palaeotis) | '--- (Struthio) | '

---

+--+ (Rheiformes) | |---+ (Opisthodactylidae) | | |-?- (Diogenornis) | | '--- (Opisthodactylus) | | | '---+ (Rheidae) | |--- (Heterorhea) | |--- (Hinasuri) | | | '---+--- (Rhea americana) | | | '--- (Pterocnemia pennata) |+---+ (Aepyornithiformes) | | '--- (Aepyornithidae) | | | '---+---+ (Dinornithiformes) | | |---+ (Dinornithidae) | | | '--- (Dinornis) | | | | | '---+ (Emeidae) | | |--- (Emeinae) | | '--- (Anomalopteryginae) | | | '---+ (Apterygiformes) | '---+ (Apterygidae) | |--- (Megapteryx) | '--- (Apteryx) | '---+ (Casuariformes) '---+---+ ("Emuwaries") | | | '--- (Dromaius) | '--- (Casuarius)

Regional Cladogram

+ (Oceanic paleognaths) | |

---

+ (Australian and Indonesian paleognaths) | | '

---

+ (Casuariformes) | | '---+ ("Emuwaries") | | '---+--- (Emus) | | '--- (Cassowaries) | | | '

---

+ (New Zealand paleognaths) | |

---

+ (Dinornithiformes) | | |--- (Dinornithidae) | | '--- (Emeidae) | | | '

---

+ (Apterygiformes) | '---+ (Apterygidae) | '--- (Kiwis) |+ (South American paleognaths) | |

---

+ (Tinamiformes) | | '---+ (Tinamidae) | | '--- (Tinamous) | | | '

---

+ (Rheiformes) | |--- (Opisthodactylidae) | '--- (Rheidae) | '

---

+ (Laurasian and African paleognaths)?---+ (Remiornithiformes) | '--- (Remiornithidae) |?---+ (Lithornithiformes) | '--- (Lithornithidae) |?---+ (Aepyornithiformes) | '--- (Aepyornithidae) |?---+ (Struthioniformes) | '---+ (Struthionidae) | '--- (African Ostrich |?---+ (Ambiornithiformes) | '--- (Ambiortidae) |?---+ (Gansuiformes) | '--- (Gansuidae) | '---?---+ (Paleocursornithiformes) '--- (Paleocursornithidae) -->

Classification

Order Lithornithiformes (fossil)

• Family Lithornithidae
• Genus Eurolimnornis
• Eurolimnornis corneti
• Genus Lithornis (Early Eocene)
• Genus Promusophaga (Early Eocene)
• Genus Paracathartes (Early Eocene of WC USA) - tentatively placed here
• Genus Pseudocrypturus - tentatively placed here

Order Dinornithiformes - moa (prehistoric/extinct)

• Family Dinornithidae
• Genus Dinornis - giant moa (2-4 species)
• Dinornis novaezealandiae - North Island giant moa
• Dinornis robustus - slender moa
• Dinornis giganteus
• Dinornis maximus
• Dinornis struthoides
• Dinornis torosus
• Dinornis hercules
• Dinornis gazelle
• Family Anomalopterygidae - lesser moas
• Subfamily Emeinae
• Genus Emeus - Eastern Moa
• Emeus crassus
• Emeus huttoni
• Genus Euryapteryx - broad-billed or turkey moas (2 species)
• Eurypteryx curtus - North Island broad-billed moa
• Eurypteryx geranoides - South Island broad-billed moa
• Eurypteryx gravis
• Genus Zelornis
• Zelornis exilis
• Zelornis haasti
• Subfamily Anomalopteryginae
• Genus Anomalopteryx - Bush Moas
• Anomalopteryx antiquus
• Anomalopteryx didiformis - lesser bush moa
• Anomalopteryx oweni
• Genus Megalapteryx - upland moas (2 species)
• Megalapteryx benhami - Benham's megalapteryx
• Megalapteryx didinus - lesser megalapteryx
• Genus Pachyornis - stout moas (3-5 species)
• Pachyornis australis
• Pachyornis elephantopus
• Pachyornis mappini
• Pachyornis murihiku
• Pachyornis septentrionalis

Order Aepyornithiformes - elephant birds (prehistoric/extinct)

• Family Aepyornithidae
• Genus Aepyornis (4 species)
• Aepyornis titan (synonym A. maximus)
• Aepyornis gracilis
• Aepyornis hildebrandti
• Aepyornis medius
• Genus Mullerornis (4 species)
• Mullerornis agilis
• Mullerornis gracilis
• Mullerornis hildebrandti
• Mullerornis titan (synonym M. titan)
• Genus Eremopezus
• Eremopezus eocaenus
• Genus Stromeria
• Stromeria fajumensis
• Genus Paraphysornis
• Paraphysornis brasiliensis

Order Struthioniformes - ostriches

• Family Struthionidae
• Genus Palaeotis (fossil: Middle Eocene) - includes Palaeogrus geiseltalensis
• Palaeotis wiegelti
• Genus Struthio (1 living species)
• Struthio coppensi
• Struthio linxiaensis
• Struthio karingarabensis
• Struthio kakesiensis
• Struthio daberasensis
• Struthio oldawayi
• Struthio anderssoni
• Struthio brachydactylus
• Struthio chersonensis
• Struthio mongolicus
• Struthio orlovi
• Struthio wimani
• Struthio asiaticus - Asian ostrich
• Struthio camelus - African ostrich
• Struthio camelus syriacus - Arabian ostrich
• Struthio camelus australis - Southern African ostrich
• Struthio camelus camelus - North African ostrich
• Struthio camelus massaicus - Masai ostrich
• Struthio camelus rothschildi
• Struthio camelus spatzi
• Struthio camelus molybdophanes - Somali ostrich

Order Rheiformes

• Family Opistodactylidae (fossil)
• Genus Opisthodactylus (Miocene of Argentina) - rheid?
• Opisthodactylus patagonicus
• Genus Diogenornis - tentatively placed here
• Diogenornis fragilis
• Family Rheidae - rheas
• Genus Heterorhea (fossil: Pliocene of Argentina)
• Heterorhea dabbenei
• Genus Hinasuri (fossil)
• Hinasuri nehuensis
• Genus Rhea (2 species, includes Pterocnemia)
• Rhea americana - common rhea
• Rhea americana americana
• Rhea americana intermedia
• Rhea americana albescens
• Rhea americana nobilis
• Rhea americana araneipes
• Rhea pennata - Darwin's rhea
• Rhea pennata pennata
• Rhea pennata garleppi
• Rhea pennata tarapacensis - puna rhea

Order Casuariiformes

• Family Casuariidae - cassowaries
• Genus Casuarius
• Family Dromaiidae
• Genus Emuarius - "emuwaries" (fossil: Late Oligocene - Late Miocene; formerly in Dromaius)
• Genus Dromaius - emus (1 living species, 2 recently extinct)

Order Apterygiformes - kiwis

• Family Apterygidae
• Genus Apteryx (about 6 living species, possibly 1 recently extinct)

Order Tinamiformes - tinamous

• Family Tinamidae
• Subfamily Tinaminae
• Genus Crypturellus
• Genus Tinamus
• Genus Nothocercus
• Genus Taoniscus
• Genus Tinamotis
• Subfamily Rhynchotinae
• Genus Eudromia - crested tinamous
• Genus Rhynchotus
• Genus Nothoprocta
• Genus Nothura - nothuras

Alternate classification

• Superorder Paleognathae
• Order Struthioniformes
• Order Lithioniformes
• Order Tinamiformes

Sometimes placed here

• Ambiortus
• Eremopezus - includes Stromeria
• Gansus
• Limenavis
• Palaeocursornis
• Wyleyia

Paraphysornis is a phorusrhacid.

Ootaxa

• Gobioolithus (Late Cretaceous) - paleognath?
• Incognitoolithus (Eocene of North America) - ratite?
• Type A ("aepyornithoid") eggs (Tsondab Early Miocene of Namibia - Pliocene of Asia)
• Namornis (Middle Miocene of Namibia - Late Miocene of Kenya) - ratite?
• Diamantornis (Middle Miocene of Namibia - Late Miocene of UAE and Kenya) - ratite?
• Psammornis - may be from Eremopezus

Locomotion

Many of the larger ratite birds have extremely long legs and the largest living bird, the ostrich, can run at speeds over 60 km/h. And cassowaries, emus, and rheas show a similar likeness in agility and some extinct forms may have reached speeds of 75 km/h. Moas, the largest birds, had legs over 3 feet high and may have been the fastest land animals to live outrunning even the cheetah.

Paleognaths and humans

Paleognaths probably first interacted with Australopithecines about 3 million or so years ago in the middle pliocene in the form of an ancient ostrich or elephant bird. As Homo erectus evolved and left Africa for other countries and continents not much contact was made with ratites, until the Maori and Aborigines arrived in New Zealand and Australia. This was very bad for native ratites that had no adaptations to keep from being killed off by humans, as a result many of the Oceanian flora and fauna, not just ratites, became extinct during this period, like the moa for example. Natives used every part of a ratite for something- meat was eaten, bones and eggs were decorated, even the feces served a religious purpose. Most giant birds became extinct by the end of the 18th century, and quite a number of these were ratites. A lot of paleognaths are now considered endangered species. Ratites such as the Ostrich are farmed and sometimes even kept as pets. Today ratites play a large part of human culture- they are farmed, eaten, raced, protected, cheered, and kept in zoos. As likely as the giraffe or elephant- the ratites may not survive until the next decade.

See also

• Fossil birds
• Late Quaternary prehistoric birds
• Extinct birds
• Flightless birds

External links

• Page On the classification of Paleognaths of Animal Diversity Web
• Regional Cladogram of Paleognaths
• Evolutionary Cladogram of Paleognaths
• Spanish Page on Ratites
• Info on How to Prepare Ratites as Food
• Avibase
• Introduction to the Paleognathae
• Oxford Journal on the Molecular Biology and Evolution of Aves
• Paleognath Monophyly
• Ornithology and Natural History
• Avian Biotech
• Paleognathae on the Tree of Life Web Project
• Page on Ratites as Livestock

References

Harrison, C.J.O. (1979). Birds: their life, their ways, their world. Pleasantville, New York: The Reader's Digest Association, Inc., 8-412. ISBN 0-89577-065-2. 

Lambert, David (2001). Dinosaur Encyclopedia: from dinosaurs to the dawn of man. New York, New York: DK publishing, inc., 138-145. ISBN 0-7894-7935-4. 

(2005) Animal: the definitive visual guide to the world's wildlife. New York, New York: DK publishing, inc., 260-265. ISBN 0-7894-7764-5. 

Monroe, Burt (1993). A World Checklist of Birds. New Haven: Yale University Press. ISBN 0-3000-5547-1. 

Elwood, Ann (1991). Ostriches, emus, rheas, kiwis, & cassowaries. Mankato, Minnesota: Creative Education. ISBN 0-8868-2338-2. 

Wexo, John (2000). Zoobooks: Ostriches and other Ratites. Poway, California: Wildlife Education. ISBN 1-8881-5357-1. 

(1996) The Ratite Encyclopedia. Charley Elrod. ISBN 0-0642-9402-8. 

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