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 a ...

'', the

type genus In biological taxonomy, the type genus 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-bearing type of a nominal ...

of family

Charadriidae The bird family Charadriidae includes the plovers, dotterels, and lapwings, about 64 to 68 species in all. Taxonomy The family Charadriidae was introduced (as Charadriadæ) by the English zoologist William Elford Leach in a guide to the con ...

) is a diverse order of small to medium-large

bird 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 lightweig ...

s. It includes about 390

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

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

invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...

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 (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or w ...

(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 Dunlins and Red knots">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 mudflat ...

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 family Laridae in the suborder Lari. They are most closely related to the terns and skimmers and only distantly related to auks, and even more distantly to waders. Until the 21st century ...

s 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

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

s and

birds of prey Birds of prey or predatory birds, also known as raptors, are hypercarnivorous bird species that actively hunt and feed on other vertebrates (mainly mammals, reptiles and other smaller birds). In addition to speed and strength, these predat ...

into a greatly enlarged order

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

. 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. Following recent research,

Families

The order Charadriiformes contains 3 suborders, 19 families and 390 species. * Suborder Charadrii ** Family Burhinidae – stone-curlews, thick-knees (10 species) ** Family Pluvianellidae – Magellanic plover ** Family Chionidae – sheathbills (2 species) ** Family Pluvianidae – Egyptian plover ** Family

– plovers (68 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 fro ...

– stilts, avocets (10 species) ** Family Ibidorhynchidae – ibisbill ** Family Haematopodidae – oystercatchers (12 species) * Suborder Scolopaci ** Family Rostratulidae – painted-snipes (3 species) ** Family Jacanidae – jacanas (8 species) ** Family Pedionomidae – plains-wanderer ** Family Thinocoridae – seedsnipes (4 species) ** Family Scolopacidae – sandpipers, snipes (98 species) * Suborder Lari ** Family

Turnicidae Buttonquail or hemipodes are members of a small family of birds, Turnicidae, which resemble, but are unrelated to, the quails of Phasianidae. They inhabit warm grasslands in Asia, Africa, Europe, and Australia. There are 18 species in two genera ...

– buttonquails (18 species) ** Family Dromadidae – crab-plover ** Family Glareolidae – coursers, pratincoles (17 species) ** Family

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

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

Alcidae An auk or alcid is a bird of the family Alcidae in the order Charadriiformes. The alcid family includes the murres, guillemots, auklets, puffins, and murrelets. The word "auk" is derived from Icelandic ''álka'', from Old Norse ''alka'' (a ...

– auks (25 species)

Evolution history

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

' fossil record around the

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

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 com ...

brought about by

semiaquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semi aquatic animals include: * Ve ...

habits. Specimen VI 9901 ( López de Bertodano Formation,

Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger 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 ''creta'', ...

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

- 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? * 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 ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...

form taxon Form classification is the classification of organisms based on their morphology, which does not necessarily reflect their biological relationships. Form classification, generally restricted to palaeontology, reflects uncertainty; the goal of s ...

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 Caribbea ...

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'' (Late Cretaceous?) *" Graculavidae" **'' Graculavus'' (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'' (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 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 a ...

, 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, known in French as the ' (abbreviation MNHN), is the national natural history museum of France and a ' of higher education part of Sorbonne Universities. The main museum, with four galleries, is loc ...

n Frenchbr>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 change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

'' 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.html" ;"title="ungarian 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.___(HTML_abstract) *_Paton,_T.A.;_Baker,_A.J.;_Groth,_J.G._&_Barrowclough,_G.F._(2003):_RAG-1_sequences_resolve_phylogenetic_relationships_within_charadriiform_birds._''Molecular_Phylogenetics_and_Evolution.html" ;"title="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._''Molecular_Phylogenetics_and_Evolution.html" ;"title="Molecular Phylogenetics and Evolution">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._''[ 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._''Journal_of_Avian_Biology">J._Avian_Biol.''_35(3):_191–194._PDF_fulltext
*_ 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._''Journal_of_Avian_Biology">J._Avian_Biol.''_35(3):_191–194._PDF_fulltext
*_Trevor_H._Worthy">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._''Journal_of_Systematic_Palaeontology.html" ;"title="Trevor_H._Worthy.html" ;"title="ournal_of_Avian_Biology.html" ;"title="Molecular Phylogenetics and Evolution">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
*_Trevor_H._Worthy">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._''Journal_of_Systematic_Palaeontology">J._Syst._Palaeontol.''_5(1):_1-39.__(HTML_abstract) {{Authority_control Charadriiformes.html" ;"title="Journal of Avian Biology">J. Avian Biol.'' 35(3): 191–194. PDF fulltext
* Trevor H. Worthy">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. ''Journal of Systematic Palaeontology">J. Syst. Palaeontol.'' 5(1): 1-39. (HTML abstract) {{Authority control Charadriiformes"> 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