Charadriiformes (, from ''

Charadrius ''Charadrius'' is a genus of plovers, a group of wading birds. The genus name ''Charadrius'' is a Late Latin word for a yellowish bird mentioned in the fourth-century Vulgate. They are found throughout the world. Many ''Charadrius'' species are ...

'', the

type genus In biological taxonomy, the type genus (''genus typica'') is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearin ...

of family

Charadriidae The bird family Charadriidae includes the plovers, dotterels, and lapwings. The family contains 69 species that are divided into 10 genera. Taxonomy The family Charadriidae was introduced (as Charadriadæ) by the English zoologist William El ...

) is a diverse order of small to medium-large

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

s. It includes about 390

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

and has members in all parts of the world. Most charadriiform birds live near water and eat

invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...

s or other small animals; however, some are

pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...

(seabirds), others frequent deserts, and a few are found in dense forest. Members of this group can also collectively be referred to as shorebirds.

Taxonomy, systematics and evolution

The order was formerly divided into three suborders: * The

wader 245px, A flock of Red_knot.html" ;"title="Dunlins and Red knot">Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflats in order to foraging, ...

s (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

gull Gulls, or colloquially seagulls, are seabirds of the subfamily Larinae. They are most closely related to terns and skimmers, distantly related to auks, and even more distantly related to waders. Until the 21st century, most gulls were placed ...

s and their allies (or "

Lari Lari may refer to: Currency * Georgian lari, the currency of Georgia * Maldivian laari, or lari, a coin denomination of the rufiyaa of the Maldives Places * Lari, Ardabil, or Lahrud, a city in Iran * Lari, East Azerbaijan, a village in Ir ...

"): these are generally larger species which take fish from the sea. Several gulls and

skua The skuas are a group of predatory seabirds with seven species forming the genus ''Stercorarius'', the only genus in the family Stercorariidae. The three smaller skuas, the Arctic skua, the long-tailed skua, and the pomarine skua, are called ...

s will also take food items from beaches, or rob smaller species, and some have become adapted to inland environments. * The

auk Auks or alcids are birds of the family Alcidae in the order Charadriiformes. The alcid family includes the Uria, murres, guillemots, Aethia, auklets, puffins, and Brachyramphus, murrelets. The family contains 25 extant or recently extinct speci ...

s (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

seabird Seabirds (also known as marine birds) are birds that are adaptation, adapted to life within the marine ecosystem, marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent ...

s and

birds of prey Birds of prey or predatory birds, also known as (although not the same as) raptors, are hypercarnivorous bird species that actively predation, hunt and feed on other vertebrates (mainly mammals, reptiles and smaller birds). In addition to speed ...

into a greatly enlarged order

Ciconiiformes Storks are large, long-legged, long-necked wading birds with long, stout bills. They belong to the family Ciconiidae, and make up the order Ciconiiformes . Ciconiiformes previously included a number of other families, such as herons and ibise ...

. 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. 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.

Families

The order Charadriiformes contains 3

suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...

s, 19 families and 391 species. * Suborder

Charadrii FIle:Vadare - Ystad-2021.jpg, 245px, A flock of Dunlins and Red knots Waders or shorebirds are birds of the order Charadriiformes commonly found wikt:wade#Etymology 1, wading along shorelines and mudflats in order to foraging, forage for food c ...

** Family

Burhinidae The stone-curlews, also known as dikkops or thick-knees, consist of 10 species within the family Burhinidae, and are found throughout the tropical and temperate parts of the world, with two or more species occurring in some areas of Africa, Asia, ...

– stone-curlews, thick-knees (10 species) ** Family Pluvianellidae – Magellanic plover ** Family Chionidae – sheathbills (2 species) ** Family Pluvianidae – Egyptian plover ** Family

– plovers (69 species) ** Family

Recurvirostridae The Recurvirostridae are a family of birds in the wader suborder Charadrii. It contains two distinct groups of birds, the avocets (one genus) and the stilts (two genera). Description Avocets and stilts range in length from and in weight from ; ...

– stilts, avocets (10 species) ** Family Ibidorhynchidae – ibisbill ** Family Haematopodidae – oystercatchers (12 species) * Suborder

Scolopaci Scolopaci is a suborder of wading birds within the order Charadriiformes containing the following families: * Family Rostratulidae – painted-snipes (3 species) * Family Jacanidae – jacanas (8 species) * Family Pedionomidae – plains-wan ...

** Family

Rostratulidae The Rostratulidae, commonly known as the painted-snipes, are a family of wading birds that consists of two genera: '' Rostratula'' and '' Nycticryphes''. Description The painted-snipes are short-legged, long-billed birds similar in shape to the ...

– painted-snipes (3 species) ** Family

Jacanidae The jacanas (sometimes referred to as Jesus birds or lily trotters) are a group of tropical waders in the family (biology), family Jacanidae. They are found in the tropical regions around the world. They are noted for their elongated toes and to ...

– jacanas (8 species) ** Family

Pedionomidae The plains-wanderer (''Pedionomus torquatus'') is an atypical species of wading bird, the only representative of family Pedionomidae and genus ''Pedionomus''. It is endemic to Australia. Its historic range included much of eastern Australia, ...

– plains-wanderer ** Family

Thinocoridae The seedsnipes are a small family, Thinocoridae, of small gregarious waders which have adapted to a herbivorous diet. The family is divided into two genera, ''Attagis'' and ''Thinocorus'', each containing two species. The family has a South Amer ...

– seedsnipes (4 species) ** Family

Scolopacidae Scolopacidae is a large family of shorebirds, or waders, which mainly includes many species known as sandpipers, but also others such as woodcocks, curlews and snipes. Most of these species eat small invertebrates picked out of the mud or soil ...

– sandpipers, snipes (98 species) * Suborder

** Family

Turnicidae Buttonquail or hemipodes are members of a small family of birds, Turnicidae, which resemble, but are not closely related to, the quails of Phasianidae. They inhabit warm grasslands in Asia, Africa, Europe, and Australia. There are 18 species in ...

– buttonquails (18 species) ** Family

Dromadidae The crab-plover or crab plover (''Dromas ardeola)'' is a coastal wader (shorebird). It is the only member of the genus ''Dromas'' and the family Dromadidae. It is unique among waders in making use of ground warmth to aid the incubation of its e ...

– crab-plover ** Family

Glareolidae Glareolidae is a family of birds in the wader suborder Lari. It contains two distinct groups, the pratincoles and the coursers. The atypical Egyptian plover (''Pluvianus aegyptius''), traditionally placed in this family, is now known to be o ...

– coursers, pratincoles (17 species) ** Family

Laridae Laridae is a family of seabirds in the order Charadriiformes that includes the gulls, terns (including white terns), noddies, and skimmers. It includes around 100 species arranged into 22 genera. They are an adaptable group of mostly aerial bird ...

– gulls, terns, skimmers (103 species) ** Family Stercorariidae – skuas (7 species) ** Family

Alcidae Auks or alcids are birds of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The family contains 25 extant or recently extinct species that are divided into ...

– auks (25 species)

Evolutionary history

That the Charadriiformes are an ancient group is also borne out by the fossil record. Alongside the

Anseriformes Anseriformes is an order (biology), order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest f ...

, the Charadriiformes are the only other order of modern bird to have an established fossil record within the late Cretaceous, alongside the other dinosaurs. Much of the

Neornithes Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweight ...

' fossil record around the

Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...

is made up of bits and pieces of birds which resemble this order. In many, this is probably due to

convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...

brought about by

semiaquatic In biology, being semi-aquatic refers to various macroorganisms that live regularly in both aquatic and terrestrial environments. When referring to animals, the term describes those that actively spend part of their daily time in water (in ...

habits. Specimen VI 9901 (

López de Bertodano Formation The Lopez de Bertodano Formation is a geological Formation (geology), formation in the James Ross Island, James Ross archipelago of the Antarctic Peninsula. The strata date from the end of the Late Cretaceous (upper-lower Maastrichtian stage) to t ...

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...

Vega Island Vega Island () is an island in Antarctica, long and wide, which is the northernmost of the James Ross Island group and lies in the west part of Erebus and Terror Gulf. It is separated from James Ross Island by Herbert Sound and from Trinit ...

, Antarctica) is probably a basal charadriiform somewhat reminiscent of a thick-knee. However, more complete remains of undisputed charadriiformes are known only from the mid-

Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...

onwards. Present-day orders emerged around the

Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

boundary, roughly 35–30 mya. Basal or unresolved charadriiforms are: *''"

Morsoravis ''Morsoravis'' is an extinct genus of neoavian bird from the Early Eocene Fur Formation of Denmark. It contains a single named species, ''Morsoravis sedilis''. Fossils of Morsoravis have also been found in the Green River Formation of Wyoming and ...

"'' (Late Paleocene/Early Eocene of Jutland, Denmark) - a ''

nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...

''? *'' 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, US) - larid? * Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid? * Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid? * Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - larid? *Charadriiformes gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary *''"Totanus" teruelensis'' (Late Miocene of Los Mansuetos, Spain) - scolopacid? larid? The "transitional shorebirds" (" Graculavidae") are a generally

Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...

form taxon Form classification is the classification of organisms based on their morphology (biology), morphology, which does not necessarily reflect their biological relationships. Form classification, generally restricted to palaeontology, reflects uncer ...

formerly believed to constitute the common ancestors of charadriiforms,

waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which i ...

and

flamingo Flamingos or flamingoes () are a type of wading bird in the family Phoenicopteridae, which is the only extant family in the order Phoenicopteriformes. There are four flamingo species distributed throughout the Americas (including the Caribbe ...

s. 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? **''

Laornis __NOTOC__ ''Laornis'' is a genus of a prehistoric neornithine birds, known only from Specimen YPM 820, a single tibiotarsus leg bone discovered in the late 19th century. Consequently, the genus is monotypic, containing only the species ''Laorni ...

'' (Late Cretaceous?) *" Graculavidae" **''

Graculavus ''Graculavus'' is a prehistoric bird genus that was described in the 19th century by American paleontologist O. C. Marsh. Its remains were found in the Late Cretaceous Austin Chalk of Texas, USA, and Lance Formation (late Maastrichtian faunal s ...

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

Palaeotringa ''Palaeotringa'' is a prehistoric bird genus that was discovered by O. C. Marsh during the late 19th century American bone wars. Its remains were found in the controversial Hornerstown Formation of New Jersey (United States) which straddles the ...

'' (Hornerstown Late Cretaceous?) - charadriiform? **''

Telmatornis ''Telmatornis'' is a valid prehistoric bird genus which has been placed in Charadriiformes. It apparently lived in the Late Cretaceous; its remains were found in the early Maastrichtian (c.71-68 million years ago) Navesink Formation of New Jerse ...

'' (Navesink Late Cretaceous?) - charadriiform? gruiform? **''

Scaniornis ''Scaniornis'' is a prehistoric bird genus. The only known species, ''Scaniornis lundgreni'', lived in the MP 1–5 (Early Paleocene, perhaps Middle Paleocene: c. 65–59 million years ago). It is known from a partial fossil skeleton of a righ ...

'' - phoenicopteriform? **''

Zhylgaia ''Zhylgaia'' is a genus of fossil bird. Its remains, consisting of two partial humeri, were recovered from an upper Paleocene deposit in Kazakhstan. The relationships of this genus are unknown; it was initially placed in the Presbyornithidae, ...

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

Ceramornis ''Ceramornis'' is a genus of ornithuran dinosaurs from the Late Cretaceous. It lived shortly before the Cretaceous–Paleogene extinction event in the Maastrichtian, some Longrich, N. (2009). "An ornithurine-dominated avifauna from the Belly Rive ...

'' (Lance Creek Late Cretaceous) * ''"

Cimolopteryx ''Cimolopteryx'' (meaning "Cretaceous wing"Holtz, Thomas R. Jr. (2008) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages'Winter 2010 Supplementary Information) is a prehistoric bird genus from the Late Creta ...

"'' (Lance Creek Late Cretaceous) * '' Palintropus'' (Lance Creek Late Cretaceous) * ''

Torotix ''Torotix'' is a Late Cretaceous genus of aquatic birds. They lived along the shores of the Western Interior Seaway, but it is not clear whether they were seabirds or freshwater birds, as the genus is only known from a humerus. Consequently, the ...

'' (Late Cretaceous) * '' Volgavis'' (Early Paleocene of Volgograd, Russia) * '' Eupterornis'' (Paleocene of France) * Neornithes incerta sedis (Late Paleocene/Early Eocene of

Ouled Abdoun Basin The Oulad Abdoun Basin (also known as the Ouled Abdoun Basin or Khouribga Basin) is a phosphate sedimentary basin located in Morocco, near the city of Khouribga. It is the largest in Morocco, comprising 44% of Morocco's phosphate reserves, and at ...

, Morocco) * '' 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. The ancestral avian most likely had a female parental care system. 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. It has also been shown that sex role reversal is motivated by the male-biased adult sex ratio.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. The reason for such diversity in shorebirds, compared to other birds, has yet to be understood.

Footnotes

References

* 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 The French National Museum of Natural History ( ; abbr. MNHN) is the national natural history museum of France and a of higher education part of Sorbonne University. The main museum, with four galleries, is located in Paris, France, within the Ja ...

n French N, or n, is the fourteenth letter of the Latin alphabet, used in the modern English alphabet, the alphabets of other western European languages, and others worldwide. Its name in English is ''en'' (pronounced ), plural ''ens''. History ...

br>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. PDF fulltext
* Fain, Matthew G. & Houde, Peter (2004): Parallel radiations in the primary clades of birds. ''

Evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...

'' 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 iddle Miocene fossils from the sections at the Rákóczi chapel at Mátraszőlős. Locality Mátraszõlõs I. ''Folia Historico Naturalia Musei Matraensis'' 23: 33–78. 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. ''Molecular Phylogenetics and Evolution">Mol. Phylogenet. Evol.'' 39(3): 657–667. (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. (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, 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. (HTML abstract) * Thomas, Gavin H.; Wills, Matthew A. & Székely, Tamás (2004): A supertree approach to shorebird phylogeny. '' BMC Evol. Biol.'' 4: 28. PDF fulltextSupplementary 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. 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. (HTML abstract) {{Authority control Seabirds Wading birds Bird orders Extant Lutetian first appearances Eocene taxonomic orders Oligocene taxonomic orders Miocene taxonomic orders Pliocene taxonomic orders Pleistocene taxonomic orders Holocene taxonomic orders

Taxonomy, systematics and evolution

Families

Evolutionary history

Evolution of parental care in Charadriiformes

See also

Footnotes

References