Charadriiformes (/kəˈrædri.ɪfɔːrmiːz/, fromCharadrius, thetype genus of familyCharadriidae) is a diverse order of small to medium-largebirds. It includes about 390species and has members in all parts of the world. Most charadriiform birds live near water and eatinvertebrates or other small animals; however, some arepelagic (seabirds), others frequent deserts, and a few are found in dense forest. Members of this group can also collectively be referred to asshorebirds.
The order was formerly divided into three suborders:
Thewaders (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.
Thegulls and their allies (or "Lari"): these are generally larger species which take fish from the sea. Several gulls andskuas will also take food items from beaches, or rob smaller species, and some have become adapted to inland environments.
Theauks (or "Alcae") are coastal species which nest on sea cliffs and "fly" underwater to catch fish.
TheSibley-Ahlquist taxonomy lumps all the Charadriiformes together with otherseabirds andbirds of prey into a greatly enlarged orderCiconiiformes. However, the resolution of theDNA-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.[6]
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.[7]
Phylogeny of the Charadriiformes based a study by Heiner Kuhl and collaborators published in 2020.[4] The families and the number of species are from the list maintained byFrank Gill,Pamela Rasmussen and David Donsker on behalf of theInternational Ornithologists' Union.[8] The suborders are those defined by Joel Cracraft in 2013.[9]
That the Charadriiformes are an ancient group is also borne out by the fossil record. Alongside theAnseriformes, the Charadriiformes are the only other order of modern bird to have an established fossil record within the late Cretaceous, alongside the other dinosaurs.[10] Much of theNeornithes' fossil record around theCretaceous–Paleogene extinction event is made up of bits and pieces of birds which resemble this order. In many, this is probably due toconvergent evolution brought about bysemiaquatic habits. Specimen VI 9901 (López de Bertodano Formation,Late Cretaceous ofVega Island, Antarctica) is probably abasal charadriiform somewhat reminiscent of athick-knee.[11] However, more complete remains of undisputed charadriiformes are known only from the mid-Paleogene onwards. Present-day orders emerged around theEocene-Oligocene boundary, roughly 35–30mya. Basal or unresolved charadriiformes are:
"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?[12]
Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid?[13]
Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - larid?[14]
Charadriiformes gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary[15]
"Totanus" teruelensis (Late Miocene of Los Mansuetos, Spain) - scolopacid? larid?
The "transitional shorebirds" ("Graculavidae") are a generallyMesozoicform taxon formerly believed to constitute the common ancestors of charadriiforms,waterfowl andflamingos. They are now assumed to be mostly basal taxa of the charadriiforms and/or "higher waterbirds", which probably were two distinct lineages 65mya already,[16] and few if any are still believed to be related to the well-distinct waterfowl. Taxa formerly considered graculavids are:
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.[18] The ancestral avian most likely had a female parental care system.[19] 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.[20] It has also been shown that sex role reversal is motivated by the male-biased adult sex ratio.[21] The reason for such diversity in shorebirds, compared to other birds, has yet to be understood.
^Mayr, Gerald (2016).Avian evolution: the fossil record of birds and its paleobiological significance. Topics in Paleobiology. Wiley-Blackwell. p. 306.ISBN978-1-119-02076-9.
^Černý, David; Natale, Rossy (2021-07-16). "Comprehensive taxon sampling and vetted fossils help clarify the time tree of shorebirds (Aves, Charadriiformes)".bioRxiv10.1101/2021.07.15.452585.
^abCracraft, Joel (2013).Dickinson, E.C.;Remsen, J.V. Jr. (eds.).The Howard & Moore Complete Checklist of the Birds of the World. Vol. 1: Non-passerines (4th ed.). Eastbourne, UK: Aves Press. pp. xxxvii–xxxviii.ISBN978-0-9568611-0-8.
^A wading bird the size of awhite stork (Ciconia ciconia): Bourdon (2005)
^Thomas, Gavin H.; Székely, Tamás; Reynolds, John D. (2007). "Sexual Conflict and the Evolution of Breeding Systems in Shorebirds".Advances in the Study of Behavior. Vol. 37. Elsevier. pp. 279–342.doi:10.1016/s0065-3454(07)37006-x.ISBN9780120045372.ISSN0065-3454.
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
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 Matraensis23: 33–78. [Hungarian with English abstract]PDF
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.; Székely, Tamás; Reynolds, John D. (2007). "Sexual Conflict and the Evolution of Breeding Systems in Shorebirds".Advances in the Study of Behavior. Vol. 37. Elsevier. pp. 279–342.doi:10.1016/s0065-3454(07)37006-x.ISBN9780120045372.ISSN0065-3454.
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.
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