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{{DISPLAYTITLE:Shorebirds, auks and gulls}}{{automatic taxobox
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{{DISPLAYTITLE:Charadriiformes: Shorebirds, auks and gulls}}{{automatic taxobox
−
|color={{Taxobox_colour|[[Animalia]]}}
  
 
|name = Charadriiformes
  
|name = Charadriiformes
 
|fossil_range = [[Late Cretaceous]]-Recent, {{fossilrange|75|0}}
  
|fossil_range = [[Late Cretaceous]]-Recent, {{fossilrange|75|0}}
−
|image = Vanellus miles novaehollandiae.jpg
 +
|image = Charadriiformes diversity.png
 
|image_width = 250px
  
|image_width = 250px
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|image_caption = [[Masked Lapwing]] (''Vanellus miles'')
 +
|image_caption =
 
|taxon = Charadriiformes
  
|taxon = Charadriiformes
 
|authority = [[Thomas Henry Huxley|Huxley]], 1867
  
|authority = [[Thomas Henry Huxley|Huxley]], 1867
 
|subdivision_ranks = Suborders
  
|subdivision_ranks = Suborders
  +
|subdivision = {{Linked taxon list
−
|display_children=1
  
  +
| Chionidi |
  +
| Charadrii |
  +
| Limicoli |
  +
| Turnici |
  +
| Lari |
  +
}}
 
}}
  
}}
   
−
'''Charadriiformes''' is a diverse order of small to medium-large [[bird]]s. It includes about 350 [[species]] and has members in all parts of the world. Most Charadriiformes live near water and eat [[invertebrate]]s or other small animals; however, some are [[pelagic]] (sea birds), some occupy deserts and a few are found in thick forest.
 +
'''Charadriiformes''' is a diverse order of small to medium-large [[bird]]s. It includes about 350 [[species]] and has members in all parts of the world. Most Charadriiformes live near water and eat [[invertebrate]]s or other small animals; however, some are [[pelagic]] (seabirds), some occupy deserts and a few are found in thick forest.
   
  +
==Taxonomy, systematics and evolution==
−
==Systematics==
  
  +
The order was formerly divided into three suborders:
−
{{clade|style=font-size:90%;line-height:100%
  
−
|label1=Charadriiformes (shorebirds)
  
−
|1={{clade
  
−
|1=[[Chionidi]]
 
−
|2={{clade
  
−
|1=[[Charadrii]]
 
−
|2={{clade
  
−
|1=[[Limicoli]]
  
−
|2={{clade
  
−
|1=[[Turnici]]
  
−
|2={{clade
  
−
|1=[[Lari]] }} }} }} }} }} }}
 
   
  +
* The '''[[wader]]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 order was formerly divided into these suborders:
  
  +
* The '''[[gull]]s''' and their allies (or "[[Lari]]"): these are generally larger species which take fish from the sea. Several gulls and [[skua]]s will also take food items from beaches, or rob smaller species, and some have become adapted to inland environments.
  +
* The '''[[auk]]s''' (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,{{citation needed|date=September 2013}} lumps all the Charadriiformes together with other [[seabird]]s and [[bird of prey|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.<ref>Fain & Houde (2004)</ref><!-- morphological refs should be locatable via family books etc -->
−
* The '''[[Magellanic Plover]], [[sheathbill]]s and [[thicknee]]s''': [[Chionidi]]. These birds are found in the [[Southern Hemisphere]].
  
−
* The '''[[Egyptian Plover]], ''[[Pluvialis]]'' plovers, [[Recurvirostridae|slilts and avocets]], [[Ibisbill]], [[oystercatcher]]s, and [[plovers]]''': [[Charadrii]].
 
−
* The '''[[Plains-wanderer]], [[seedsnipe]]s, [[painted-snipe]]s, [[jacana]]s and [[sandpiper]]s''': [[Limicoli]].
  
−
* The '''[[buttonquail]]''': [[Turnici]].
  
−
* The '''[[Crab plover (family)|Crab plovers]], [[Glareolidae|coursers and pratincoles]], [[skua]]s, [[auk]]s and [[Laridae|gulls, terns and skimmers]]''': [[Lari]].
  
   
  +
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,<ref>Ericson ''et al.'' (2003), Paton ''et al.'' (2003), Thomas ''et al.'' (2004a,b), van Tuinen ''et al.'' (2004), Paton & Baker (2006)</ref> a better arrangement may be as follows:
−
The [[Sibley-Ahlquist taxonomy]], which has been widely accepted in America, lumps all the Charadriiformes together with other [[seabird]]s and [[bird of prey|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.<ref>Fain & Houde (2004)</ref><!-- morphological refs should be locatable via family books etc -->
  
   
  +
===Families in taxonomic order===
−
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,<ref>Ericson ''et al.'' (2003), Paton ''et al.'' (2003, Thomas ''et al.'' (2004a,b), van Tuinen ''et al.'' (2004), Paton & Baker (2006)</ref> a better arrangement may be as follows:
  
−  
−
==Families in taxonomic order==
  
 
This is a list of the charadriiform [[family (biology)|families]], presented in [[taxonomic order]].
  
This is a list of the charadriiform [[family (biology)|families]], presented in [[taxonomic order]].
−  
 
* '''Suborder [[Chionidi]]''': thick-knees and allies
  
* '''Suborder [[Chionidi]]''': thick-knees and allies
−
** Family [[stone-curlew|Burhinidae]]: thick-knees
 +
** Family [[Pluvianellidae]]: Magellanic Plover
−
** Family [[sheathbill|Chionididae]]: sheathbills
 +
** Family [[Chionididae]]: sheathbills
−
** Family [[Magellanic Plover|Pluvianellidae]]: Magellanic Plover
 +
** Family [[Burhinidae]]: thick-knees
   
 
* '''Suborder [[Charadrii]]''': plover-like waders
  
* '''Suborder [[Charadrii]]''': plover-like waders
−
** Family [[Egyptian Plover|Pluvianidae]]: Egyptian plover
 +
** Family [[Pluvianidae]]: Egyptian plover
−
** Family [[Pluvialidae]]: Golden plovers
 +
** Family [[Pluvialidae]]: golden-plovers
 
** Family [[Recurvirostridae]]: avocets and stilts
  
** Family [[Recurvirostridae]]: avocets and stilts
−
** Family [[Ibisbill|Ibidorhynchidae]]: Ibisbill
 +
** Family [[Ibisbill|Ibidorhynchidae]]: ibisbill
 
** Family [[oystercatcher|Haematopodidae]]: oystercatchers
  
** Family [[oystercatcher|Haematopodidae]]: oystercatchers
 
** Family [[Charadriidae]]: plovers and lapwings
  
** Family [[Charadriidae]]: plovers and lapwings
   
−
* '''Suborder [[Limicoli]]''': sandpipers and allies
 +
* '''Suborder [[Limicoli]]''': snipe-like waders
−
** Family [[Plains Wanderer|Pedionomidae]]: Plains Wanderer
 +
** Family [[Pedionomidae]]: plains wanderer
−
** Family [[seedsnipe|Thinocoridae]]: seedsnipe
 +
** Family [[Thinocoridae]]: seedsnipe
 
** Family [[painted snipe|Rostratulidae]]: painted snipe
  
** Family [[painted snipe|Rostratulidae]]: painted snipe
−
** Family [[jacana|Jacanidae]]: jacanas
 +
** Family [[Jacanidae]]: jacanas
 
** Family [[Scolopacidae]]: snipe, sandpipers, phalaropes, and allies
  
** Family [[Scolopacidae]]: snipe, sandpipers, phalaropes, and allies
   
Line 67: Line 56:
   
 
* '''Suborder [[Lari]]''': gulls and allies
  
* '''Suborder [[Lari]]''': gulls and allies
−
** Family [[Crab Plover|Dromadidae]]: Crab Plover
 +
** Family [[Dromadidae]]: crab plover
 
** Family [[Glareolidae]]: pratincoles and coursers
  
** Family [[Glareolidae]]: pratincoles and coursers
 
** Family [[Skua|Stercorariidae]]: skuas
  
** Family [[Skua|Stercorariidae]]: skuas
 
** Family [[auk|Alcidae]]: puffins, guillemots, murres, and allies
  
** Family [[auk|Alcidae]]: puffins, guillemots, murres, and allies
−
** Family [[gull|Laridae]]: gulls
 +
** Family [[Laridae]]: gulls, terns, and skimmers
−
** Family [[Rhynchopidae]]: skimmers
  
−
** Family [[tern|Sternidae]]: terns
  
   
  +
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 [[wader]]s. Some taxonomy sources place the family Glareolidae in its own suborder, instead of being classified under suborder Lari.<ref name=":0">{{Cite web|url=http://jboyd.net/Taxo/List9a.html|title=Charadriiformes|last=John|first=Boyd|date=|website=jboyd.net|language=en|access-date=2017-07-16}}</ref> The buttonquails are of indeterminate or [[Basal (phylogenetics)|basal]] position in the Lari-Scolopaci ''sensu lato'' group. The arrangement as presented here is a consensus of the recent studies.<ref>van Tuinen ''et al.'' (2004), Paton & Baker (2006)</ref>
−
List is from:{{TiF8}}
  
  +
{{clade| style=font-size:100%;line-height:80%
  +
|label1=Charadriiformes
  +
|1={{Clade
  +
|label1=[[Charadrii]]
  +
|1={{Clade
  +
|label1=[[Chionidi|Chionida]]
  +
|1={{Clade
  +
|label1=[[Burhinidae]]
  +
|1={{Clade
  +
|1=''[[Burhinus]]''
  +
|2=''[[Esacus]]''
  +
}}
  +
|2={{Clade
  +
|label1=[[Chionidae]]
  +
|1=''[[Chionis]]''
  +
|label2=[[Pluvianellidae]]
  +
|2=''[[Pluvianellus]]''
  +
}}
  +
}}
  +
|label2=[[Charadrii|Charadriida]]
  +
|2={{Clade
  +
|label1=[[Pluvianidae]]
  +
|1=''[[Pluvianus]]''
  +
|2={{Clade
  +
|1={{Clade
  +
|label1=[[Pluvialidae]]
  +
|1=''[[Pluvialis]]''
  +
|2={{Clade
  +
|1={{Clade
  +
|label1=[[Ibidorhynchidae]]
  +
|1=''[[Ibidorhyncha]]''
  +
|label2=[[Haematopodidae]]
  +
|2=''[[Haematopus]]''
  +
}}
  +
|label2=[[Recurvirostridae]]
  +
|2={{Clade
  +
|1=''[[Recurvirostra]]''
  +
|2={{Clade
  +
|1=''[[Cladorhynchus]]''
  +
|2=''[[Himantopus]]''
  +
}}
  +
}}
  +
}}
  +
}}
  +
|label2=[[Charadriidae]]
  +
|2={{Clade
  +
|label1=[[Charadriinae]]
  +
|1={{Clade
  +
|1=''[[Oreopholus]]''
  +
|2={{Clade
  +
|1={{Clade
  +
|1=''[[Phegornis]]''
  +
|2=''[[Zonibyx]]''
  +
}}
  +
|2={{Clade
  +
|1=''[[Eudromias]]''
  +
|2={{Clade
  +
|1=''[[Afroxyechus]]''
  +
|2={{Clade
  +
|1=''[[Charadrius]]''
  +
|2=''[[Thinornis]]''
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
|2={{Clade
  +
|label1=[[Vanellinae]]
  +
|1=''[[Vanellus]]''
  +
|label2=[[Anarhynchinae]]
  +
|2={{Clade
  +
|1=''[[Erythrogonys]]''
  +
|2={{Clade
  +
|1=''[[Peltohyas]]''
  +
|2={{Clade
  +
|1=''[[Eupoda]]''
  +
|2={{Clade
  +
|1=''[[Anarhynchus]]''
  +
|2=''[[Ochthodromus]]''
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
|2={{Clade
  +
|label1=[[Limicoli]]
  +
|1={{Clade
  +
|label1=[[Thinocori|Jacanida]]
  +
|1={{Clade
  +
|label1=Thincoroidea
  +
|1={{Clade
  +
|label1=[[Pedionomidae]]
  +
|1=''[[Pedionomus]]''
  +
|label2=[[Thinocoridae]]
  +
|2={{Clade
  +
|1=''[[Attagis]]''
  +
|2=''[[Thinocorus]]''
  +
}}
  +
}}
  +
|label2=Jacanoidea
  +
|2={{Clade
  +
|label1=[[Rostratulidae]]
  +
|1={{Clade
  +
|1=''[[Nycticryphes]]''
  +
|2=''[[Rostratula]]''
  +
}}
  +
|label2=[[Jacanidae]]
  +
|2={{Clade
  +
|1={{Clade
  +
|1=''[[Hydrophasianus]]''
  +
|2=''[[Jacana (genus)|Jacana]]''
  +
}}
  +
|2={{Clade
  +
|1=''[[Actophilornis]]''
  +
|2={{Clade
  +
|1=''[[Metopidius]]''
  +
|2={{Clade
  +
|1=''[[Microparra]]''
  +
|2=''[[Irediparra]]''
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
|label2=[[Scolopaci|Scolopacida]]
  +
|2={{Clade
  +
|label1=[[Scolopacidae]]
  +
|1={{Clade
  +
|label1=[[Numeniinae]]
  +
|1={{Clade
  +
|1=''[[Bartramia (bird)|Bartramia]]''
  +
|2=''[[Numenius (bird)|Numenius]]''
  +
}}
  +
|2={{Clade
  +
|label1=[[Limosinae]]
  +
|1=''[[Limosa]]''
  +
|2={{Clade
  +
|label1=[[Arenariinae]]
  +
|1={{Clade
  +
|1=''[[Limicola]]''
  +
|2=''[[Ereunetes]]''
  +
|3=''[[Calidris]]''
  +
|4={{Clade
  +
|1=''[[Arenaria (bird)|Arenaria]]''
  +
|2=''[[Prosobonia]]''
  +
}}
  +
}}
  +
|2={{Clade
  +
|label1=[[Tringinae]]
  +
|1={{Clade
  +
|1={{Clade
  +
|1=''[[Xenus]]''
  +
|2=''[[Phalaropus]]''
  +
}}
  +
|2={{Clade
  +
|1=''[[Actitis]]''
  +
|2=''[[Tringa]]''
  +
}}
  +
}}
  +
|label2=[[Scolopacinae]]
  +
|2={{Clade
  +
|1={{Clade
  +
|1=''[[Lymnocryptes]]''
  +
|2=''[[Limnodromus]]''
  +
}}
  +
|2={{Clade
  +
|1=''[[Scolopax]]''
  +
|2={{Clade
  +
|1=''[[Gallinago]]''
  +
|2={{Clade
  +
|1=''[[Chubbia]]''
  +
|2=''[[Coenocorypha]]''
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
|label2=[[Lari]]
  +
|2={{Clade
  +
|label1=[[Turnici|Turnicida]]
  +
|1={{Clade
  +
|1=''[[Ortyxelos]]''
  +
|2=''[[Turnix]]''
  +
}}
  +
|label2=[[Larida]]
  +
|2={{Clade
  +
|label1=Glareoloidea
  +
|1={{Clade
  +
|label1=[[Dromadidae]]
  +
|1=''[[Dromas ardeola]]''
  +
|label2=[[Glareolidae]]
  +
|2={{Clade
  +
|1=''[[Stiltia]]''
  +
|2=''[[Rhinoptilus]]''
  +
|3={{Clade
  +
|1=''[[Cursorius]]''
  +
|2=''[[Glareola]]''
  +
}}
  +
}}
  +
}}
  +
|2={{Clade
  +
|label1=Alcoidea
  +
|1={{Clade
  +
|label1=[[Stercorariidae]]
  +
|1=''[[Stercorarius]]''
  +
|label2=[[Alcidae]]
  +
|2={{Clade
  +
|label1=[[Fraterculinae]]
  +
|1={{Clade
  +
|1={{Clade
  +
|1=''[[Cerorhinca]]''
  +
|2=''[[Fratercula]]''
  +
}}
  +
|2={{Clade
  +
|1=''[[Ptychoramphus]]''
  +
|2=''[[Aethia]]''
  +
}}
  +
}}
  +
|label2=[[Alcinae]]
  +
|2={{Clade
  +
|1=''[[Brachyramphus]]''
  +
|2={{Clade
  +
|1=''[[Cepphus]]''
  +
|2={{Clade
  +
|1=''[[Synthliboramphus]]''
  +
|2={{Clade
  +
|1={{Clade
  +
|1=''[[Uria]]''
  +
|2=''[[Alle (genus)|Alle]]''
  +
}}
  +
|2={{Clade
  +
|1=''[[Alca (bird)|Alca]]''
  +
|2=†''[[Pinguinus]]''
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
|label2=Laroidea
  +
|2={{Clade
  +
|label1=[[Laridae]]
  +
|1={{Clade
  +
|label1=[[Gyginae]]
  +
|1=''[[Gygis]]''
  +
|label2=[[Rynchopinae]]
  +
|2=''[[Rynchops]]''
  +
|label3=[[Anoinae]]
  +
|3={{Clade
  +
|1=''[[Anous]]''
  +
|2=''[[Procelsterna]]''
  +
}}
  +
|label4=[[Sterninae]]
  +
|4={{Clade
  +
|1=''[[Onychoprion]]''
  +
|2={{Clade
  +
|1=''[[Sternula]]''
  +
|2={{Clade
  +
|1=''[[Phaetusa]]''
  +
|2={{Clade
  +
|1={{Clade
  +
|1=''[[Gelochelidon]]''
  +
|2=''[[Hydroprogne]]''
  +
}}
  +
|2={{Clade
  +
|1=''[[Larosterna]]''
  +
|2={{Clade
  +
|1=''[[Chlidonias]]''
  +
|2={{Clade
  +
|1=''[[Thalasseus]]''
  +
|2=''[[Sterna]]''
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
|label5=[[Larinae]]
  +
|5={{Clade
  +
|1=''[[Creagrus]]''
  +
|2={{Clade
  +
|1=''[[Hydrocoloeus]]''
  +
|2=''[[Rhodostethia]]''
  +
}}
  +
|3={{Clade
  +
|1=''[[Rissa (genus)|Rissa]]''
  +
|2={{Clade
  +
|1=''[[Pagophila]]''
  +
|2=''[[Xema]]''
  +
}}
  +
}}
  +
|4={{Clade
  +
|1=''[[Saundersilarus]]''
  +
|2={{Clade
  +
|1=''[[Chroicocephalus]]''
  +
|2={{Clade
  +
|1=''[[Leucophaeus]]''
  +
|2={{Clade
  +
|1=''[[Larus]]''
  +
|2=''[[Ichthyaetus]]''
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
  +
}}
   
  +
Cladogram based on Baker, A.J. ''et al''. (2012)<ref>Baker, A.J. ''et al.'' (2012) [http://www.sciencedirect.com/science/article/pii/S1055790312002758 Eight independent nuclear genes support monophyly of the plovers: The role of mutational variance in gene trees].</ref> and Boyd, J. H. ''et al.'' (2016) <ref name=":0" />
−
More conservatively, the Thinocori could be included in the Scolopaci, and the Chionidi in the Charadrii, or the Glareolidae could be placed in a suborder of their own. The buttonquails are of indeterminate, quite [[Basal (phylogenetics)|basal]] position in the Lari-Scolopaci ''sensu lato'' group. The arrangement as presented here is a consensus of the recent studies.<ref>van Tuinen ''et al.'' (2004), Paton & Baker (2006)</ref>
  
   
  +
The order was formerly divided into these suborders:
−
==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 (phylogenetics)|basal]] charadriiform somewhat reminiscent of a [[thick-knee]]{{Citation needed|date=April 2007}}. 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 (unit)|mya]]. Basal or unresolved charadriiforms are:
  
  +
* The '''[[Magellanic Plover]], [[sheathbill]]s and [[thicknee]]s''': [[Chionidi]]. These birds are found in the [[Southern Hemisphere]].
  +
* The '''[[Egyptian Plover]], ''[[Pluvialis]]'' plovers, [[Recurvirostridae|slilts and avocets]], [[Ibisbill]], [[oystercatcher]]s, and [[plovers]]''': [[Charadrii]].
  +
* The '''[[Plains-wanderer]], [[seedsnipe]]s, [[painted-snipe]]s, [[jacana]]s and [[sandpiper]]s''': [[Limicoli]].
  +
* The '''[[buttonquail]]''': [[Turnici]].
  +
* The '''[[Crab plover (family)|Crab plovers]], [[Glareolidae|coursers and pratincoles]], [[skua]]s, [[auk]]s and [[Laridae|gulls, terns and skimmers]]''': [[Lari]].
  +
  +
The [[Sibley-Ahlquist taxonomy]], which has been widely accepted in America, lumps all the Charadriiformes together with other [[seabird]]s and [[bird of prey|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.<ref>Fain & Houde (2004)</ref><!-- morphological refs should be locatable via family books etc -->
  +
  +
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,<ref>Ericson ''et al.'' (2003), Paton ''et al.'' (2003, Thomas ''et al.'' (2004a,b), van Tuinen ''et al.'' (2004), Paton & Baker (2006)</ref> a better arrangement may be as follows:
  +
  +
===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 [[List of semiaquatic tetrapods|semiaquatic]] habits. Specimen VI 9901 ([[López de Bertodano Formation]], [[Late Cretaceous]] of [[Vega Island]], Antarctica) is probably a [[Basal (phylogenetics)|basal]] charadriiform somewhat reminiscent of a [[thick-knee]].<ref>Case, J. A. and C. P. Tambussi. 1999. Maastrichtian record of neornithine birds in Antarctica: comments on a Late Cretaceous radiation</ref> 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 (unit)|mya]]. Basal or unresolved charadriiforms are:
 
*''"[[Morsoravis]]"'' (Late Paleocene/Early Eocene of Jutland, Denmark) - a ''[[nomen nudum]]''?
  
*''"[[Morsoravis]]"'' (Late Paleocene/Early Eocene of Jutland, Denmark) - a ''[[nomen nudum]]''?
 
*''[[Jiliniornis]]'' (Huadian Middle Eocene of Huadian, China) - charadriid?<!-- *Condor104:896 -->
  
*''[[Jiliniornis]]'' (Huadian Middle Eocene of Huadian, China) - charadriid?<!-- *Condor104:896 -->
Line 88: Line 412:
 
*''"Larus" desnoyersii'' (Early Miocene of SE France) - larid? stercorarid?<!-- JSystPaleontol5:1. -->
  
*''"Larus" desnoyersii'' (Early Miocene of SE France) - larid? stercorarid?<!-- JSystPaleontol5:1. -->
 
*''"Larus" pristinus'' (John Day Early Miocene of Willow Creek, USA) - larid?<!-- JSystPaleontol5:1. -->
  
*''"Larus" pristinus'' (John Day Early Miocene of Willow Creek, USA) - larid?<!-- JSystPaleontol5:1. -->
−
* Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid?<ref>[[Proximal]] right [[humerus]] ([[MNZ]] S42416) and proximal left [[carpometacarpus|carpometacarpi]] (MNZ S42415, S42435) of a bird the size of a [[Red-necked Stint]]: Worthy ''et al.'' (2007)</ref>
 +
* Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid?<ref>[[Anatomical terms of location#Proximal and distal|Proximal]] right [[humerus]] ([[Museum of New Zealand|MNZ]] S42416) and proximal left [[carpometacarpus|carpometacarpi]] (MNZ S42415, S42435) of a bird the size of a [[red-necked stint]]: Worthy ''et al.'' (2007)</ref>
−
* Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid?<ref>Several wing and thorax bones of a bird the size of a [[Double-banded Plover]]: Worthy ''et al.'' (2007)</ref>
 +
* Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid?<ref>Several wing and thorax bones of a bird the size of a [[double-banded plover]]: Worthy ''et al.'' (2007)</ref>
−
* Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - larid?<ref>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)</ref>
 +
* Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - larid?<ref>Premaxillae ([[Museum of New Zealand|MNZ]] S42681, S42736) and [[Anatomical terms of location#Proximal and distal|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)</ref>
 
*Charadriiformes gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary<ref>Gál ''et al.'' (1998-99)</ref>
  
*Charadriiformes gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary<ref>Gál ''et al.'' (1998-99)</ref>
 
*''"Totanus" teruelensis'' (Late Miocene of Los Mansuetos, Spain) - scolopacid? larid?
  
*''"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 [[flamingo]]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 (unit)|mya]] already{{Citation needed|date=April 2007}}, and few if any are still believed to be related to the well-distinct weaterfowl. Taxa formerly considered graculavids are:
 +
The "transitional shorebirds" ("[[Graculavidae]]") are a generally [[Mesozoic]] [[form taxon]] formerly believed to constitute the common ancestors of charadriiforms, [[waterfowl]] and [[flamingo]]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 (unit)|mya]] already,{{Citation needed|date=April 2007}} and few if any are still believed to be related to the well-distinct waterfowl. Taxa formerly considered graculavids are:
 
*'''[[Laornithidae]]''' - charadriiform? gruiform?
  
*'''[[Laornithidae]]''' - charadriiform? gruiform?
 
**''[[Laornis]]'' (Late Cretaceous?)
  
**''[[Laornis]]'' (Late Cretaceous?)
Line 113: Line 437:
 
* ''[[Volgavis]]'' (Early Paleocene of Volgograd, Russia)
  
* ''[[Volgavis]]'' (Early Paleocene of Volgograd, Russia)
 
* ''[[Eupterornis]]'' (Paleocene of France)
  
* ''[[Eupterornis]]'' (Paleocene of France)
−
* Neornithes incerta sedis (Late Paleocene/Early Eocene of Ouled Abdoun Basin, Morocco)<ref>A wading bird the size of a [[White Stork]] (''Ciconia ciconia''): Bourdon (2005)</ref>
 +
* Neornithes incerta sedis (Late Paleocene/Early Eocene of [[Ouled Abdoun Basin]], Morocco)<ref>A wading bird the size of a [[white stork]] (''Ciconia ciconia''): Bourdon (2005)</ref>
 
* ''[[Fluviatitavis]]'' (Early Eocene of Silveirinha, Portugal)
  
* ''[[Fluviatitavis]]'' (Early Eocene of Silveirinha, Portugal)
   
  +
==Evolution of parental care in Charadriiformes==
−
==Footnotes==
  
−
{{Reflist}}
  
   
  +
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.
−
==References==<!-- Auk93:587 -->
  
−
* {{aut|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]. [http://bcmnts1.mnhn.fr/ipac20/ipac.jsp?session=12C9G373T0897.297548&profile=mq&source=~!muscat&view=&uri=full=3100001~!405665~!2 HTML abstract]
  
−
* {{aut|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 journals|BMC Evol. Biol.]]'' '''3''': 16. <!-- BMC ejournals use article numbers, not page numbers --> <small>{{doi|10.1186/1471-2148-3-16}}</small> [http://www.biomedcentral.com/content/pdf/1471-2148-3-16.pdf PDF fulltext]
  
−
* {{aut|Fain, Matthew G. & Houde, Peter}} (2004): Parallel radiations in the primary clades of birds. ''[[Evolution (journal)|Evolution]]'' '''58'''(11): 2558-2573. <small>{{DOI|10.1554/04-235}} PMID 15612298</small> [http://biology-web.nmsu.edu/houde/Parallel_radiations.pdf PDF fulltext]
  
−
* {{aut|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] [http://www.matramuzeum.hu/Personal/folia/23/03MATSZOLOS.PDF PDF fulltext]
  
−
* {{aut|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. <small>{{DOI|10.1016/j.ympev.2006.01.011}} PMID 16531074</small> (HTML abstract)
  
−
* {{aut|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|Mol. Phylogenet. Evol.]]'' '''29''': 268-278. <small>{{doi|10.1016/S1055-7903(03)00098-8}} PMID 13678682</small> (HTML abstract)
  
−
* {{aut|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|Mol. Phylogenet. Evol.]]'' '''30'''(3): 516-526. <small>{{doi|10.1016/S1055-7903(03)00222-7}}</small> (HTML abstract)
  
−
* {{aut|Thomas, Gavin H.; Wills, Matthew A. & Székely, Tamás}} (2004): A supertree approach to shorebird phylogeny. ''[[BMC journals|BMC Evol. Biol.]]'' '''4''': 28. <small>{{doi|10.1186/1471-2148-4-28}} PMID 15329156</small> [http://www.pubmedcentral.org/picrender.fcgi?artid=515296&blobtype=pdf PDF fulltext] [http://www.pubmedcentral.org/articlerender.fcgi?artid=515296#supplementary-material-sec Supplementary Material]
  
−
* {{aut|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. <small>{{doi|10.1111/j.0908-8857.2004.03362.x}}</small> [http://www.stanford.edu/group/hadlylab/images/Lab%20Members/Marcel/JAB2004.PDF_1.pdf PDF fulltext]
  
−
* {{aut|[[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. <small>{{doi|10.1017/S1477201906001957}}</small> (HTML abstract)
  
   
  +
==See also==
  +
* [[List of Charadriiformes by population]]
   
  +
==Footnotes==
  +
{{Reflist}}
   
  +
==References==<!-- Auk93:587 -->
  +
* 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]. [https://web.archive.org/web/20110721012025/http://bcmnts1.mnhn.fr/ipac20/ipac.jsp?session=12C9G373T0897.297548&profile=mq&source=~!muscat&view=&uri=full=3100001~!405665~!2 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 journals|BMC Evol. Biol.]]'' '''3''': 16. <!-- BMC ejournals use article numbers, not page numbers --> <small>{{doi|10.1186/1471-2148-3-16}}</small> [http://www.biomedcentral.com/content/pdf/1471-2148-3-16.pdf PDF fulltext]
  +
* Fain, Matthew G. & Houde, Peter (2004): Parallel radiations in the primary clades of birds. ''[[Evolution (journal)|Evolution]]'' '''58'''(11): 2558-2573. <small>{{DOI|10.1554/04-235}} {{PMID|15612298}}</small> [https://web.archive.org/web/20130407204436/http://biology-web.nmsu.edu/houde/Parallel_radiations.pdf 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] [http://www.matramuzeum.hu/Personal/folia/23/03MATSZOLOS.PDF 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. <small>{{DOI|10.1016/j.ympev.2006.01.011}} {{PMID|16531074}}</small> (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|Mol. Phylogenet. Evol.]]'' '''29''': 268-278. <small>{{doi|10.1016/S1055-7903(03)00098-8}} {{PMID|13678682}}</small> (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. ''[[Molecular Phylogenetics and Evolution|Mol. Phylogenet. Evol.]]'' '''30'''(3): 516-526. <small>{{doi|10.1016/S1055-7903(03)00222-7}}</small> (HTML abstract)
  +
* Thomas, Gavin H.; Wills, Matthew A. & Székely, Tamás (2004): A supertree approach to shorebird phylogeny. ''[[BMC journals|BMC Evol. Biol.]]'' '''4''': 28. <small>{{doi|10.1186/1471-2148-4-28}} {{PMID|15329156}}</small> [http://www.pubmedcentral.org/picrender.fcgi?artid=515296&blobtype=pdf PDF fulltext]{{dead link|date=July 2017 |bot=InternetArchiveBot |fix-attempted=yes }} [http://www.pubmedcentral.org/articlerender.fcgi?artid=515296#supplementary-material-sec Supplementary Material]{{dead link|date=November 2016 |bot=InternetArchiveBot |fix-attempted=yes }}
  +
* 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. <small>{{doi|10.1111/j.0908-8857.2004.03362.x}}</small> [http://www.stanford.edu/group/hadlylab/images/Lab%20Members/Marcel/JAB2004.PDF_1.pdf 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. <small>{{doi|10.1017/S1477201906001957}}</small> (HTML abstract)
   
 
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[[Category:Seabirds]]
  
[[Category:Seabirds]]
 
[[Category:Wading birds]]
  
[[Category:Wading birds]]
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[[Category:Extant Lutetian first appearances]]
  +
[[Category:Eocene taxonomic orders]]
  +
[[Category:Oligocene taxonomic orders]]
  +
[[Category:Miocene taxonomic orders]]
  +
[[Category:Pliocene taxonomic orders]]
  +
[[Category:Pleistocene taxonomic orders]]
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[[Category:Holocene taxonomic orders]]

Latest revision as of 19:05, 15 March 2018

Charadriiformes
Temporal range: Late Cretaceous-Recent, 75–0 Ma
PreЄ
Є
Є
O
O
S
S
D
D
C
C
P
P
T
T
J
J
Pg
Pg
N
N
File:Charadriiformes diversity.png
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Clade: Aequorlitornithes
Order: Charadriiformes
Huxley, 1867
Suborders

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.

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, which has been widely accepted in America,[citation needed] 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.

  • Suborder Turnici: buttonquails

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]

Charadriiformes
Charadrii
Chionida
Burhinidae

Burhinus



Esacus




Chionidae

Chionis


Pluvianellidae

Pluvianellus




Charadriida
Pluvianidae

Pluvianus




Pluvialidae

Pluvialis




Ibidorhynchidae

Ibidorhyncha


Haematopodidae

Haematopus



Recurvirostridae

Recurvirostra




Cladorhynchus



Himantopus






Charadriidae
Charadriinae

Oreopholus





Phegornis



Zonibyx





Eudromias




Afroxyechus




Charadrius



Thinornis








Vanellinae

Vanellus


Anarhynchinae

Erythrogonys




Peltohyas




Eupoda




Anarhynchus



Ochthodromus












Limicoli
Jacanida
Thincoroidea
Pedionomidae

Pedionomus


Thinocoridae

Attagis



Thinocorus




Jacanoidea
Rostratulidae

Nycticryphes



Rostratula



Jacanidae


Hydrophasianus



Jacana





Actophilornis




Metopidius




Microparra



Irediparra








Scolopacida
Scolopacidae
Numeniinae

Bartramia



Numenius




Limosinae

Limosa



Arenariinae

Limicola



Ereunetes



Calidris




Arenaria



Prosobonia





Tringinae


Xenus



Phalaropus





Actitis



Tringa




Scolopacinae


Lymnocryptes



Limnodromus





Scolopax




Gallinago




Chubbia



Coenocorypha












Lari
Turnicida

Ortyxelos



Turnix



Larida
Glareoloidea
Dromadidae

Dromas ardeola


Glareolidae

Stiltia



Rhinoptilus




Cursorius



Glareola






Alcoidea
Stercorariidae

Stercorarius


Alcidae
Fraterculinae


Cerorhinca



Fratercula





Ptychoramphus



Aethia




Alcinae

Brachyramphus




Cepphus




Synthliboramphus





Uria



Alle





Alca



†Pinguinus









Laroidea
Laridae
Gyginae

Gygis


Rynchopinae

Rynchops


Anoinae

Anous



Procelsterna



Sterninae

Onychoprion




Sternula




Phaetusa





Gelochelidon



Hydroprogne





Larosterna




Chlidonias




Thalasseus



Sterna









Larinae

Creagrus




Hydrocoloeus



Rhodostethia





Rissa




Pagophila



Xema






Saundersilarus




Chroicocephalus




Leucophaeus




Larus



Ichthyaetus














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

The order was formerly divided into these suborders:

The Sibley-Ahlquist taxonomy, which has been widely accepted in America, 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 consititute 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. Following recent research,[7] a better arrangement may be as follows:

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.[8] 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?[9]
  • Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - charadriid? scolopacid?[10]
  • Charadriiformes gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) - larid?[11]
  • Charadriiformes gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszõlõs, Hungary[12]
  • "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,[citation needed] 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 (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)[13]
  • 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

  • List of Charadriiformes by population

Footnotes

  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". jboyd.net. 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. ^ Fain & Houde (2004)
  7. ^ Ericson et al. (2003), Paton et al. (2003, Thomas et al. (2004a,b), van Tuinen et al. (2004), Paton & Baker (2006)
  8. ^ Case, J. A. and C. P. Tambussi. 1999. Maastrichtian record of neornithine birds in Antarctica: comments on a Late Cretaceous radiation
  9. ^ 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)
  10. ^ Several wing and thorax bones of a bird the size of a double-banded plover: Worthy et al. (2007)
  11. ^ 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)
  12. ^ Gál et al. (1998-99)
  13. ^ A wading bird the size of a white stork (Ciconia ciconia): Bourdon (2005)

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 [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 Template:PMID 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 Template:PMID (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 Template:PMID (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 Template:PMID PDF fulltextThis link is dead. Supplementary MaterialThis link is dead.
  • 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)
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