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'Charadriiformes' is a diverse order of small to medium-large birds. It includes about 350 species and has members in all parts of the world. Most Charadriiformes live near water and eat invertebrates or other small animals; however, some are pelagic (sea birds), some occupy deserts and a few are found in thick forest.

Contents

Systematics

Families in taxonomic order

Evolution

References

Systematics

The order was formerly divided into three suborders:

★ The 'waders' (or "Charadrii"): typical shorebirds, most of which feed by probing in the mud or picking items off the surface in both coastal and freshwater environments.

★ The 'gulls' and their allies (or "Lari"): these are generally larger species which take fish from the sea. Several gulls and skuas will also take food items from beaches, or rob smaller species, and some have become adapted to inland environments.

★ The 'auks' (or "Alcae") are coastal species which nest on sea cliffs and "fly" underwater to catch fish.
The Sibley-Ahlquist taxonomy, which has been widely accepted in America, lumps all the Charadriiformes together with the seabirds and birds of prey into a greatly enlarged order Ciconiiformes. However, the resolution of the DNA-DNA hybridization technique used by Sibley & Ahlquist was not sufficient to properly resolve the relationships in this group, and indeed it appears as if the Charadriiformes consititute a single large and very distinctive lineage of modern birds of their own (Fain & Houde 2004).
The auks, usually considered distinct because of their peculiar morphology, are more likely related to gulls, the "distinctness" being a result of adaptation for diving. Following recent research (Ericson ''et al.'', 2003; Paton ''et al.'', 2003; Thomas ''et al.'', 2004a, b; van Tuinen ''et al.'', 2004; Paton & Baker, 2006), a better arrangement may be as follows:

Families in taxonomic order

★ 'Suborder Scolopaci': snipe-like waders

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★ Family Scolopacidae: snipe, sandpipers, phalaropes, and allies

★ 'Suborder Thinocori': aberrant charadriforms

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★ Family Rostratulidae: painted snipe

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★ Family Jacanidae: jacanas

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★ Family Thinocoridae: seedsnipe

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★ Family Pedionomidae: Plains Wanderer

★ 'Suborder Lari': gulls and allies

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★ Family Laridae: gulls

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★ Family Rhynchopidae: skimmers

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★ Family Sternidae: terns

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★ Family Alcidae: puffins, guillemots, murres, and allies

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★ Family Stercorariidae: skuas

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★ Family Glareolidae: pratincoles and coursers

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★ Family Dromadidae: Crab Plover

★ 'Suborder Turnici': buttonquails

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★ Family Turnicidae: buttonquails

★ 'Suborder Chionidi': thick-knees and allies

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★ Family Burhinidae: thick-knees

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★ Family Chionididae: sheathbills

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★ Family Pluvianellidae: Magellanic Plover

★ 'Suborder Charadrii': plover-like waders

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★ Family Ibidorhynchidae: Ibisbill

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★ Family Recurvirostridae: avocets and stilts

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★ Family Haematopodidae: oystercatchers

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★ Family Charadriidae: plovers and lapwings
More conservatively, the Thinocori could be included in the Scolopaci, and the Chionidi in the Charadrii, or the Glareolidae could be placed in a tribe of their own. The buttonquails are of indeterminate, quite basal position in the Lari-Scolopaci ''sensu lato'' group. The arrangement as presented here is basically the consensus of the recent studies (see van Tuinen ''et al.'', 2004; Paton & Baker, 2006).

Evolution

That the Charadriiformes are an ancient group is also borne out by the fossil record. Much of the Neornithes' fossil record around the Cretaceous–Tertiary extinction event is made up of bits and pieces of birds which resemble this order. In many, this is probably due to convergent evolution brought about by semi-aquatic habits. Specimen VI 9901 (López de Bertodano Formation, Late Cretaceous of Vega Island, Antarctica) is probably a basal charadriiform somewhat reminiscent of a thick-knee. However, more complete remains of undisputed charadriiforms are known only from the mid-Paleogene onwards. Present-day orders emerged around the Eocene-Oligocene boundary, roughly 35-30 mya. Basal or unresolved charadriiforms are:

★ ''Jiliniornis'' (Huadian Middle Eocene of Huadian, China) - charadriid?

★ ''Boutersemia'' (Early Oligocene of Boutersem, Belgium) - glareolid?

★ ''Turnipax'' (Early Oligocene) - turnicid?

★ ''"Larus" desnoyersii'' (Early Miocene of SE France) - larid? stercorariid?

★ ''"Larus" pristinus'' (John Day Early Miocene of Willow Creek, USA) - larid?

★ Charadriiformes gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary: Gál ''et al'' 1998-99)

★ ''"Totanus" teruelensis'' (Late Miocene of Los Mansuetos, Spain) - scolopacid? larid?
The "transitional shorebirds" ("Graculavidae") are a generally Mesozoic form taxon formerly believed to constitute the common ancestors of charadriiforms, waterfowl and flamingos. They are now assumed to be mostly basal taxa of the charadriiforms and/or "higher waterbirds", which probably were two distinct lineages 65 mya already, and few if any are still believed to be related to the well-distinct weaterfowl. Taxa formerly considered graculavids are:

★ 'Laornithidae' - charadriiform? gruiform?

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★ ''Laornis'' (Late Cretaceous?)

★ '"Graculavidae"'

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★ ''Graculavus'' (Lance Creek Late Cretaceous - Hornerstown Late Cretaceous/Early Palaeocene) - charadriiform?

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★ ''Palaeotringa'' (Hornerstown Late Cretaceous?) - charadriiform?

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★ ''Telmatornis'' (Navesink Late Cretaceous?) - charadriiform? gruiform?

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★ ''Scaniornis'' - phoenicopteriform?

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★ ''Zhylgaia'' - presbyornithid?

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★ ''Dakotornis''

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★ "Graculavidae" gen. et sp. indet. (Gloucester County, USA)
Other wader- or gull-like birds ''incertae sedis'', which may or may not be Charadriiformes, are:

★ ''Ceramornis'' (Lance Creek Late Cretaceous)

★ ''"Cimolopteryx"'' (Lance Creek Late Cretaceous)

★ ''Palintropus'' (Lance Creek Late Cretaceous)

★ ''Torotix'' (Late Cretaceous)

★ ''Volgavis'' (Early Paleocene of Volgograd, Russia)

★ ''Eupterornis'' (Paleocene of France)

★ ''Fluviatitavis'' (Early Eocene of Silveirinha, Portugal)

References

★ 'Ericson', P. G. P.; Envall, I.; Irestedt, M. & Norman, J. A. (2003): Inter-familial relationships of the shorebirds (Aves: Charadriiformes) based on nuclear DNA sequence data. ''BMC Evol. Biol.'' '3': 16. PDF fulltext

★ 'Fain', Matthew G. & 'Houde', Peter (2004): Parallel radiations in the primary clades of birds. ''Evolution'' '58'(11): 2558-2573. PDF fulltext

★ 'Gál', Erika; Hír, János; Kessler, Eugén & Kókay, József (1998-99): Középsõ-miocén õsmaradványok, a Mátraszõlõs, Rákóczi-kápolna alatti útbevágásból. I. A Mátraszõlõs 1. lelõhely [Middle Miocene fossils from the sections at the Rákóczi chapel at Mátraszőlős. Locality Mátraszõlõs I.]. ''Folia Historico Naturalia Musei Matraensis'' '23': 33-78. [Hungarian with English abstract] PDF fulltext

★ 'Paton', Tara A. & 'Baker', Allan J. (2006): Sequences from 14 mitochondrial genes provide a well-supported phylogeny of the Charadriiform birds congruent with the nuclear RAG-1 tree. ''Molecular Phylogenetics and Evolution'' '39'(3): 657–667. (HTML abstract)

★ 'Paton', Tara A.; Baker, Allan J.; Groth, J. G. & Barrowclough, G. F. (2003): RAG-1 sequences resolve phylogenetic relationships within charadriiform birds. ''Molecular Phylogenetics and Evolution'' '29': 268-278. (HTML abstract)

★ 'Thomas', Gavin H.; Wills, Matthew A. & Székely, Tamás (2004a): Phylogeny of shorebirds, gulls, and alcids (Aves: Charadrii) from the cytochrome-''b'' gene: parsimony, Bayesian inference, minimum evolution, and quartet puzzling. ''Molecular Phylogenetics and Evolution'' '30'(3): 516-526. (HTML abstract)

★ 'Thomas', Gavin H.; Wills, Matthew A. & Székely, Tamás (2004b): A supertree approach to shorebird phylogeny. ''BMC Evol. Biol.'' '4': 28. PDF fulltext Supplementary Material

★ 'van Tuinen', Marcel; Waterhouse, David & Dyke, Gareth J. (2004): Avian molecular systematics on the rebound: a fresh look at modern shorebird phylogenetic relationships. ''Journal of Avian Biology'' '35'(3): 191-194. PDF fulltext