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 (seabirds), some occupy deserts and a few are found in thick forest.

Temporal range: Late Cretaceous-Present, 75–0 Ma
Several members of the order
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Infraclass: Neognathae
Clade: Neoaves
Clade: Aequorlitornithes
Order: Charadriiformes
Huxley, 1867

See text.

Taxonomy, systematics and evolution

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 lumps all the Charadriiformes together with other 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 constitute a single large and very distinctive lineage of modern birds of their own.[1]

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,[2] a better arrangement may be as follows:

Families in taxonomic order

This is a list of the charadriiform families, presented in taxonomic order.

More conservatively, the Thinocori could be included in the Scolopaci (this combined sub-order is called Limicoli), and the Chionidi in the Charadrii. The suborders Thincori, Scolopaci, Chionidi, and Charadri are commonly referred to collectively as waders. Some taxonomy sources place the family Glareolidae in its own suborder, instead of being classified under suborder Lari.[3] The buttonquails are of indeterminate or basal position in the Lari-Scolopaci sensu lato group. The arrangement as presented here is a consensus of the recent studies.[4]














































































Dromas ardeola


















































Cladogram based on Baker, A.J. et al. (2012)[5] and Boyd, J. H. et al. (2016) [3]

Evolution history

That the Charadriiformes are an ancient group is also borne out by the fossil record. Much of the Neornithes' fossil record around the Cretaceous–Paleogene 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 semiaquatic 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.[6] 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:

  • "Morsoravis" (Late Paleocene/Early Eocene of Jutland, Denmark) - a nomen nudum?
  • Jiliniornis (Huadian Middle Eocene of Huadian, China) - charadriid?
  • Boutersemia (Early Oligocene of Boutersem, Belgium) - glareolid?
  • Turnipax (Early Oligocene) - turnicid?
  • Elorius (Early Miocene Saint-Gérand-le-Puy, France)
  • "Larus" desnoyersii (Early Miocene of SE France) - larid? stercorarid?
  • "Larus" pristinus (John Day Early Miocene of Willow Creek, USA) - larid?
  • Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid?[7]
  • Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid?[8]
  • Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - larid?[9]
  • Charadriiformes gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary[10]
  • "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 waterfowl. Taxa formerly considered graculavids are:

  • Laornithidae - charadriiform? gruiform?
  • "Graculavidae"
    • Graculavus (Lance Creek Late Cretaceous - Hornerstown Late Cretaceous/Early Palaeocene) - charadriiform?
    • Palaeotringa (Hornerstown Late Cretaceous?) - charadriiform?
    • Telmatornis (Navesink Late Cretaceous?) - charadriiform? gruiform?
    • Scaniornis - phoenicopteriform?
    • Zhylgaia - presbyornithid?
    • Dakotornis
    • "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)
  • Neornithes incerta sedis (Late Paleocene/Early Eocene of Ouled Abdoun Basin, Morocco)[11]
  • Fluviatitavis (Early Eocene of Silveirinha, Portugal)

Evolution of parental care in Charadriiformes

Shorebirds pursue a larger diversity of parental care strategies than do most other avian orders. They therefore present an attractive set of examples to support the understanding of the evolution of parental care in avians generally (as reviewed in Thomas et al. 2007). The ancestral avian most likely had a female parental care system (Tullberg et al. 2002). The shorebird ancestor specifically evolved from a bi-parental care system, yet the species within the clade Scolopacidae evolved from a male parental care system. These transitions might have occurred for several reasons. Brooding density is correlated with male parental care. Male care systems in birds are shown to have a very low breeding density while female care systems in birds have a high breeding density. (Owens 2005). Certain rates of male and female mortality, male and female egg maturation rate, and egg death rate have been associated with particular systems as well (Klug et al. 2013). It has also been shown that sex role reversal is motivated by the male-biased adult sex ratio (Liker et al. 2013). The reason for such diversity in shorebirds, compared to other birds, has yet to be understood.

See also


  1. ^ Fain & Houde (2004)
  2. ^ Ericson et al. (2003), Paton et al. (2003), Thomas et al. (2004a,b), van Tuinen et al. (2004), Paton & Baker (2006)
  3. ^ a b John, Boyd. "Charadriiformes". Retrieved 2017-07-16.
  4. ^ van Tuinen et al. (2004), Paton & Baker (2006)
  5. ^ Baker, A.J. et al. (2012) Eight independent nuclear genes support monophyly of the plovers: The role of mutational variance in gene trees.
  6. ^ Case, J. A. and C. P. Tambussi. 1999. Maastrichtian record of neornithine birds in Antarctica: comments on a Late Cretaceous radiation
  7. ^ Proximal right humerus (MNZ S42416) and proximal left carpometacarpi (MNZ S42415, S42435) of a bird the size of a red-necked stint: Worthy et al. (2007)
  8. ^ Several wing and thorax bones of a bird the size of a double-banded plover: Worthy et al. (2007)
  9. ^ Premaxillae (MNZ S42681, S42736) and proximal right scapula (MNZ S41058) of a bird apparently similar to the black-billed gull but almost the size of a kelp gull: Worthy et al. (2007)
  10. ^ Gál et al. (1998-99)
  11. ^ A wading bird the size of a white stork (Ciconia ciconia): Bourdon (2005)


  • Bourdon, Estelle (2006): L'avifaune du Paléogène des phosphates du Maroc et du Togo: diversité, systématique et apports à la connaissance de la diversification des oiseaux modernes (Neornithes) ["Paleogene avifauna of phosphates of Morocco and Togo: diversity, systematics and contributions to the knowledge of the diversification of the Neornithes"]. Doctoral thesis, Muséum national d'histoire naturelle [in French]. HTML abstract
  • Ericson, Per 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. doi:10.1186/1471-2148-3-16 PDF fulltext
  • Fain, Matthew G. & Houde, Peter (2004): Parallel radiations in the primary clades of birds. Evolution 58(11): 2558-2573. doi:10.1554/04-235 PMID 15612298 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
  • Klug, H., M. B. Bonsall, and S.H Alonzo. 2013. Sex differences in life history drive evolutionary transitions among maternal, paternal, and bi‐parental care. Ecology and Evolution. 3: 792–806.
  • Liker, A., R. P. Freckleton, and T. Székely. 2013. The evolution of sex roles in birds is related to adult sex ratio. Nature Communications. 4: 1587.
  • Owens, I.P. 2002. Male–only care and classical polyandry in birds: phylogeny, ecology and sex differences in remating opportunities. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 357: 283-293.
  • 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. Mol. Phylogenet. Evol. 39(3): 657–667. doi:10.1016/j.ympev.2006.01.011 PMID 16531074 (HTML abstract)
  • Paton, T.A.; Baker, A.J.; Groth, J.G. & Barrowclough, G.F. (2003): RAG-1 sequences resolve phylogenetic relationships within charadriiform birds. Mol. Phylogenet. Evol. 29: 268-278. doi:10.1016/S1055-7903(03)00098-8 PMID 13678682 (HTML abstract)
  • Székely, T and J.D. Reynolds. 1995. Evolutionary transitions in parental care in shorebirds. Proceedings of the Royal Society of London. Series B: Biological Sciences. 262: 57-64.
  • Thomas, G. H., T. Székely and J.D. Reynolds. 2007. Sexual conflict and the evolution of breeding systems in shorebirds. Advances in the Study of Behavior. 37: 279-342.
  • 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. Mol. Phylogenet. Evol. 30(3): 516-526. doi:10.1016/S1055-7903(03)00222-7 (HTML abstract)
  • Thomas, Gavin H.; Wills, Matthew A. & Székely, Tamás (2004): A supertree approach to shorebird phylogeny. BMC Evol. Biol. 4: 28. doi:10.1186/1471-2148-4-28 PMID 15329156 PDF fulltext Supplementary Material
  • Tullberg, B. S., M. Ah–King and H. Temrin. 2002. Phylogenetic reconstruction of parental–care systems in the ancestors of birds. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 357: 251-257.
  • van Tuinen, Marcel; Waterhouse, David & Dyke, Gareth J. (2004): Avian molecular systematics on the rebound: a fresh look at modern shorebird phylogenetic relationships. J. Avian Biol. 35(3): 191-194. doi:10.1111/j.0908-8857.2004.03362.x PDF fulltext
  • Worthy, Trevor H.; Tennyson, A.J.D.; Jones, C.; McNamara, J.A. & Douglas, B.J. (2007): Miocene waterfowl and other birds from central Otago, New Zealand. J. Syst. Palaeontol. 5(1): 1-39. doi:10.1017/S1477201906001957 (HTML abstract)

Aequorlitornithes is a clade of waterbirds recovered in a compressive genomic systematic study using nearly 200 species in 2015. It contains the clades Charadriiformes (waders and shorebirds), Mirandornithes (flamingos and grebes) and Ardeae (Eurypygimorphae and Aequornithes). Previous studies have found different placement for the clades in the tree.


Aequornithes (from Latin aequor, expanse of water + Greek ornithes, birds), or core water birds are defined as "the least inclusive clade containing Gaviidae and Phalacrocoracidae".The monophyly of the group is currently supported by several molecular phylogenetic studies.Aequornithes includes the clades Gaviiformes, Sphenisciformes, Procellariiformes, Ciconiiformes, Suliformes and Pelecaniformes. It does not include several unrelated groups of aquatic birds such as flamingos and grebes (Mirandornithes), shorebirds and auks (Charadriiformes), or the Anseriformes.

Based on a whole-genome analysis of the bird orders, the kagu and sunbittern (Eurypygiformes) and the three species of tropicbirds (Phaethontiformes) together styled as the Eurypygimorphae are the closest sister group of the Aequornithes in the clade Ardeae.

Cladogram based on Burleigh, J.G. et al. (2015)

Black-naped tern

The black-naped tern (Sterna sumatrana) is an oceanic tern mostly found in tropical and subtropical areas of the Pacific and Indian Oceans. It is rarely found inland.


The coursers are a group of birds which together with the pratincoles make up the family Glareolidae. They have long legs, short wings and long pointed bills which curve downwards. Their most unusual feature for birds classed as waders is that they inhabit deserts and similar arid regions.

They have cryptic plumage and crouch down when alarmed to avoid detection by predators.

Like the pratincoles, the coursers are found in warmer parts of the Old World. They hunt insects by running.

Their 2–3 eggs are laid on the ground.

Egyptian plover

The Egyptian plover (Pluvianus aegyptius), also known as the crocodile bird, is a wader, the only member of the genus Pluvianus. Formerly placed in the pratincole and courser family, Glareolidae, it is now regarded as the sole member of its own monotypic family Pluvianidae.

The species is one of several plovers doubtfully associated with the "trochilus" bird mentioned in a supposed cleaning symbiosis with the Nile crocodile.


Gruimorphae is a clade of birds that contains the orders Charadriiformes (plovers, gulls, and allies) and Gruiformes (cranes and rails) identified in 2014 by genome analysis. This grouping has had historical support, as various charadriiform families such as the families Pedionomidae and Turnicidae were classified as gruiforms. The relationship between these birds is due similar anatomical and behavioral characteristics. A morphological study went further to suggest that the gruiforms might be paraphyletic in respect to the shorebirds, with the rails being closely related to the buttonquails.


The jacanas (sometimes referred to as Jesus birds or lily trotters) are a group of tropical waders in the family Jacanidae. They are found in the tropical regions around the world. They are noted for their elongated toes and toenails that allow them to spread out their weight while foraging on floating or semi-emergent aquatic vegetation. They are also among the rare groups of birds in which females are larger and several species maintain harems of males in the breeding season with males solely responsible for incubating eggs and taking care of the chicks.

Large-billed tern

The large-billed tern (Phaetusa simplex) is a species of tern in the family Laridae. It belongs to the monotypic genus Phaetusa.

It is found in most of South America (east of the Andes and north of the Pampas). It has occurred as a vagrant in Aruba, Bermuda, Cuba, Panama and the United States.

Its natural habitats are rivers and freshwater lakes.

These birds are rarely found in the UK or in Canada, however, they do appear to enjoy flying individually.


The suborder Lari is the part of the order Charadriiformes that includes the gulls, terns, skuas and skimmers; the rest of the order is made up of the waders and snipes. The auks are now placed into the Lari too, following recent research. Sometimes, the buttonquails are also placed here, but the molecular data and fossil record rather suggests they are a quite basal offshoot along with the snipe-like and aberrant waders.The larids are generally larger species that 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.


Laridae is a family of seabirds in the order Charadriiformes that includes the gulls, terns and skimmers. It includes around 100 species arranged into 22 genera. They are an adaptable group of mostly aerial birds found worldwide.

Lesser jacana

The lesser jacana (Microparra capensis) is a species of bird in the Jacanidae family. It is monotypic within the genus Microparra.It is found in Angola, Botswana, Burkina Faso, Burundi, Cameroon, Central African Republic, Chad, Democratic Republic of the Congo, Ivory Coast, Ethiopia, Kenya, Malawi, Mali, Mauritania, Mozambique, Namibia, Niger, Nigeria, Rwanda, Sierra Leone, South Africa, Sudan, Swaziland, Tanzania, Uganda, Zambia, and Zimbabwe. Its primary habitats are coastal and inland wetlands and waterways.

Long-billed dowitcher

The long-billed dowitcher (Limnodromus scolopaceus) is a medium-sized shorebird. The genus name Limnodromus is Ancient Greek from limne, "marsh" and dromos, "racer". The specific scolopaceus is New Latin for "snipe-like", from Latin scolopax, scolopacis, a snipe or woodcock. The English name is from Iroquois and was first recorded in 1841.Adults have yellowish legs and a long straight dark bill. The body is dark brown on top and reddish underneath with spotted throat and breast, bars on flanks. The tail has a black and white barred pattern. The winter plumage of both an adult and a juvenile is largely grey.

Their breeding habitat is wet tundra in the far north of North America and eastern Siberia. They nest on the ground, usually near water.

They migrate to the southern United States and as far south as Central America. Long-billed dowitcher is a rare but regular visitor to western Europe, with some individuals staying for long periods.

These birds forage by probing in shallow water or on wet mud. They mainly eat insects, mollusks, crustaceans and marine worms, but also eat some plant material.

They are more likely to be seen near fresh water than the short-billed dowitcher.

Long-billed plover

The long-billed plover (Charadrius placidus) is a species of bird in the family Charadriidae.

It breeds in East Asia (mainly China, Manchuria and Japan).


Quail is a collective name for several genera of mid-sized birds generally placed in the order Galliformes.

Old World quail are placed in the family Phasianidae, and New World quail are placed in the family Odontophoridae. The species of buttonquail are named for their superficial resemblance to quail, and form the family Turnicidae in the order Charadriiformes. The king quail, an Old World quail, often is sold in the pet trade, and within this trade is commonly, though mistakenly, referred to as a "button quail". Many of the common larger species are farm-raised for table food or egg consumption, and are hunted on game farms or in the wild, where they may be released to supplement the wild population, or extend into areas outside their natural range. In 2007, 40 million quail were produced in the U.S.The collective noun for a group of quail is a flock, covey, or bevy.

Saunders's tern

The Saunders's tern (Sternula saundersi) is a species of tern in the family Laridae.

It is found in Bahrain, Iran, Israel, Kenya, Madagascar, Pakistan, Oman, Saudi Arabia, Sri Lanka, Tanzania, United Arab Emirates, and Yemen.

Senegal lapwing

The African wattled lapwing (Vanellus senegallus) is sometimes called Senegal wattled plover.

The Senegal lapwing or lesser black-winged lapwing (Vanellus lugubris) is a species of bird in the family Charadriidae.

It is found in Angola, Burundi, Cameroon, Republic of the Congo, Democratic Republic of the Congo, Ivory Coast, Gabon, Gambia, Ghana, Guinea, Kenya, Liberia, Malawi, Mali, Mozambique, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Africa, Swaziland, Tanzania, Togo, Uganda, Zambia, and Zimbabwe.


Sterna is a genus of terns in the bird family Laridae. Sterna is derived from Old English "stearn" which appears in the poem The Seafarer; a similar word was used to refer to terns by the Frisians. It used to encompass most "white" terns indiscriminately, but mtDNA sequence comparisons have recently determined that this arrangement is paraphyletic. It is now restricted to the typical medium-sized white terns occurring near-globally in coastal regions.

Temminck's courser

Temminck's courser (Cursorius temminckii) is a bird in the pratincole and courser family, Glareolidae. It is a wader which lives in sub-Saharan Africa. It is noted for laying its eggs in the burnt bushes and grass of the African savannah.


Waders are birds commonly found along shorelines and mudflats that wade in order to forage for food (such as insects or crustaceans) in the mud or sand. They are called shorebirds in North America, where the term "wader" is used to refer to long-legged wading birds such as storks and herons. Waders are members of the order Charadriiformes, which includes gulls, auks and their allies.

There are about 210 species of wader, most of which live in wetland or coastal environments. Many species of Arctic and temperate regions are strongly migratory, but tropical birds are often resident, or move only in response to rainfall patterns. Some of the Arctic species, such as the little stint, are amongst the longest distance migrants, spending the non-breeding season in the southern hemisphere.

Many of the smaller species found in coastal habitats, particularly but not exclusively the calidrids, are often named as "sandpipers", but this term does not have a strict meaning, since the upland sandpiper is a grassland species.

The smallest member of this group is the least sandpiper, small adults of which can weigh as little as 15.5 grams and measure just over 13 cm (5.1 in). The largest species is believed to be the Far Eastern curlew, at about 63 cm (25 in) and 860 grams (1.90 pounds), although the beach thick-knee is the heaviest at about 1 kg (2.2 lb).

In the Sibley-Ahlquist taxonomy, waders and many other groups are subsumed into a greatly enlarged Ciconiiformes order. However, the classification of the Charadriiformes is one of the weakest points of the Sibley-Ahlquist taxonomy, as DNA–DNA hybridization has turned out to be incapable of properly resolving the interrelationships of the group. Formerly, the waders were united in a single suborder Charadrii, but this has turned out to be a "wastebasket taxon", uniting no less than four charadriiform lineages in a paraphyletic assemblage. However, it indicated that the plains wanderer actually belonged into one of them. Following recent studies (Ericson et al., 2003; Paton et al., 2003; Thomas et al., 2004a, b; van Tuinen et al., 2004; Paton & Baker, 2006), the waders may be more accurately subdivided as follows:

Suborder Scolopaci

Family Scolopacidae: snipe, sandpipers, phalaropes, and allies

Suborder Thinocori

Family Rostratulidae: painted snipe

Family Jacanidae: jacanas

Family Thinocoridae: seedsnipe

Family Pedionomidae: plains wanderer

Suborder Chionidi

Family Burhinidae: thick-knees

Family Chionididae: sheathbills

Family Pluvianellidae: Magellanic plover

Suborder Charadrii

Family Ibidorhynchidae: ibisbill

Family Recurvirostridae: avocets and stilts

Family Haematopodidae: oystercatchers

Family Charadriidae: plovers and lapwingsIn keeping more in line with the traditional grouping, the Thinocori could be included in the Scolopaci, and the Chionidi in the Charadrii. However, the increasing knowledge about the early evolutionary history of modern birds suggests that the assumption of Paton et al. (2003) and Thomas et al. (2004b) of 4 distinct "wader" lineages (= suborders) already being present around the Cretaceous–Paleogene boundary is correct.

Birds (class: Aves)
Fossil birds
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