Ruff
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Panellet
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{{otheruses}}
{{Taxobox
| name = Ruff
| status = LC
| status_system = IUCN3.1
| image = Kampfläufer 2007-06-08 118.jpg
| regnum = [[Animal]]ia
| phylum = [[Chordate|Chordata]]
| classis = [[bird|Aves]]
| ordo = [[Charadriiformes]]
| familia = [[Scolopacidae]]
| genus = '''''Philomachus'''''
| genus_authority = [[Paul Möhring|Moehring]], 1758
| species = '''''P. pugnax'''''
| binomial = ''Philomachus pugnax''
| binomial_authority = ([[Carolus Linnaeus|Linnaeus]], 1758)
}}
[[Image:Kampfläufer weiß 070608.jpg|thumb|Satellite male with white ruff]]
The '''Ruff''' (''Philomachus pugnax'') is a medium-sized [[wader]]. It is usually considered the only member of its [[genus]] ''Philomachus'', but more recent research (Thomas ''et al'', 2004) indicates that the [[Broad-billed Sandpiper|Broad-billed]] and [[Sharp-tailed Sandpiper]] may belong there too.
[[Image:Ruffs I IMG 9501.jpg|thumb|left| In non-breeding plumage near [[Hodal]] in [[Faridabad]] District of [[Haryana]], [[India]]. ]]
Their breeding habitat is bogs, marshes and wet meadows with short vegetation in northern Europe and [[Russia]]. Ruff are [[bird migration|migratory]], wintering in southern and western Europe, Africa and [[India]]. They are highly gregarious, with a wintering flock of 1 million birds reported in [[Senegal]]. This species is a rare migrant to North America, but has nested (but is not known to have hatched eggs) in [[Alaska]].
These birds nest in a well-hidden location on the ground, laying 3-4 eggs. The males display at a [[Lek (animal behavior)|lek]] during the breeding season, standing erect, crouching or taking a variety of postures with the ruff erected. They may jump into the air. They are silent while displaying and when not breeding. These birds forage in wet grassland and soft mud, probing or picking up food by sight. They mainly eat insects and earthworms.
The Ruff is one of the species to which the ''Agreement on the Conservation of African-Eurasian Migratory Waterbirds'' ([[AEWA]]) applies.
==Description==
[[Image:Ruffs (Philomachus pugnax) at Bharatpur I IMG 5580.jpg|thumb|left| In non-breeding plumage at [[Keoladeo National Park]], [[Bharatpur]], [[Rajasthan]], [[India]].]]
Ruffs have a distinctive "gravy boat" appearance, with a small head and medium [[beak|bill]], longish neck and pot-bellied profile. Males are larger than females. They have longish usually yellowish legs, and show white ovals on the tail sides in flight. Males are 29-32 cm long with a 54-60 cm wingspan. Breeding males grow the most individually distinctive breeding [[plumage]]s of any wild bird, and recognize each other as individuals by plumage differences. They can have black, chestnut or white "neck ruffs" and "head tufts", made up of solid, barred or irregularly colored feathers (see images below). Their backs are grey-brown and the underparts white except for the black breast.
Females, also called reeves, are 22-26 cm long with a 46-49 cm wingspan. They are grey-brown above and white below. Females nest alone and provide all parental care. In winter, both sexes are a paler grey above and white below and can be differentiated only by size. Young birds have browner more scaly upperparts and are buffy below.
==Breeding behaviour==
There are two main behavioral types of males (but see end of article), which are fixed by a simple [[gene|genetic]] [[Polymorphism (biology)|polymorphism]]: territorial ("resident") and satellite males. These types can be distinguished by their appearance and behaviour: Resident or independent males (84%) have intensely-colored ruffs (black or chestnut) and stake out and occupy small (ca. 1x1 m) mating territories in the [[lek]]. They actively court females and display a high degree of aggression towards other resident males. Satellite males (16%) have light (white or mottled) ruffs and do not occupy own territories, but rather use those occupied by resident males and attempt to mate with females that visit the territory.
The relationship between resident and satellite males can be described as an uneasy alliance, which persists because having both kinds of males on a territory appears to attract additional females to that territory. This encourages partial cooperation between the two males, but they remain competitors to mate with visiting females.
[[Image:Philomachus pugnax (Marek Szczepanek).jpg|thumb|left|Non-breeding plumage]]
Notably, while satellite males are on average slightly smaller and lighter, the nutrition of the chicks does not influence mating strategy, but rather, the inherited mating strategy heavily influences body size: Resident-type chicks will, if provided with the same amount of food, grow heavier than satellite-type chicks. Therefore, the mating strategies constitute an environmentally stable simple polymorphism.
Moreover, the genetic [[Locus (genetics)|locus]] relevant for the mating strategy is located on an [[autosome]]. That means that both sexes carry 2 [[allele]]s, not only males (whereas [[mammal]]ian males have the [[gonosome]]s XY and females have XX, in birds it is the other way around, males having the gonosomes ZZ and females WZ. W, X, Y and Z are technical terms, with X- and Z-chromosomes or Y- and W-chromosomes being analogous in male- and female-determining [[Sex linkage|heterogame]] systems, respectively). If ruff females (which are always non-territorial) are given [[testosterone]] implants, they display the male territorial behaviour according to their genotype.
In 2006, a third type of male was described: permanent female mimics, the first such reported for birds. About 1 in 100 males are small, intermediate in size between males and females, and do not grow the elaborate breeding plumages of territorial and satellite males. These "female mimics" apparently obtain access to mating territories together with females, and then "steal" matings when females crouch to solicit matings.
==References==
* {{IUCN2006|assessors=BirdLife International|year=2004|id=49092|title=Philomachus pugnax|downloaded=[[11 May]] [[2006]]}} Database entry includes justification for why this species is of least concern
*D B Lank, C M Smith, O Hanotte, T Burke & F Cooke: ''Genetic polymorphism for alternative mating behaviour in lekking male ruff Philomachus pugnax''. [[Nature (journal)|Nature]] 378, p.59-62 (1995)
*J Jukema & T Piersma: ''Permanent female mimics in a lekking shorebird''. [[Biology Letters]] doi:10.1098/rsbl.2005.0416.
* '''Thomas''', Gavin H.; Wills, Matthew A. & Székely, Tamás (2004): A supertree approach to shorebird phylogeny. ''[[BMC journals|BMC Evol. Biol.]]'' '''4''': 28. {{doi|10.1186/1471-2148-4-28}} [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]
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==Gallery==
<gallery>
Image:Ruffs, Spotted Redshanks, Black-winged Stilts etc near Hodal I Picture 2124.jpg| Ruffs, Spotted Redshanks, Black-winged Stilts etc. near [[Hodal]] in [[Faridabad]] District of [[Haryana]], [[India]].
Image:Ruffs, Spotted Redshanks, Black-winged Stilts etc near Hodal I .jpg| Ruffs, Spotted Redshanks, Black-winged Stilts etc. near [[Hodal]] in [[Faridabad]] District of [[Haryana]], [[India]].
Image:Ruffs, Redshanks, Black-winged Stilts etc near Hodal I Picture 2133.jpg| Ruffs, Redshanks, Black-winged Stilts etc. near [[Hodal]] in [[Faridabad]] District of [[Haryana]], [[India]].
Image:Ruffs I IMG 9504.jpg| near [[Hodal]] in [[Faridabad]] District of [[Haryana]], [[India]].
Image:Ruffs (Philomachus pugnax) with a Sandpiper at Bharatpur I IMG 5557.jpg| with a Sandpiper at [[Keoladeo National Park]], [[Bharatpur]], [[Rajasthan]], [[India]].
Image:Ruffs (Philomachus pugnax) at Bharatpur I IMG 5580.jpg| At [[Keoladeo National Park]], [[Bharatpur]], [[Rajasthan]], [[India]].
Image:Ruff (Philomachus pugnax) at Bharatpur I IMG 5554.jpg| At [[Keoladeo National Park]], [[Bharatpur]], [[Rajasthan]], [[India]].
Image:Ruff90.jpg|Breeding birds, mainly males
</gallery>
== External links ==
*[http://www.ebepe.com/html/ruff.html Ruffs] Image documentation
*[http://www.sfu.ca/biology/wildberg/ruff.html] Genetics and behavioral research
*[http://www.ebc.uu.se/zooeko/FredrikW/Ruff.html#Strategies] Mating strategy introduction and research
*[http://www.journals.royalsoc.ac.uk/openurl.asp?genre=article&id=doi:10.1098/rsbl.2005.0416] Cryptic males publication
{{Commons|Philomachus pugnax}}
[[Category:Philomachus]]
[[Category:Wading birds]]
[[Category:Birds of Pakistan]]
[[Category:Birds of Europe]]
[[Category:Birds of South Australia]]
[[Category:Birds of Western Australia]]
[[Category:British Isles coastal fauna]]
[[bg:Бойник (животно)]]
[[ca:Philomachus pugnax]]
[[cs:Jespák bojovný]]
[[cy:Pibydd Torchog]]
[[da:Brushane]]
[[de:Kampfläufer]]
[[et:Tutkas]]
[[es:Combatiente]]
[[eo:Duelbirdo]]
[[fr:Combattant varié]]
[[gl:Combatente]]
[[ko:목도리도요]]
[[it:Philomachus]]
[[lb:Ambaraskréimer]]
[[lt:Gaidukas]]
[[hu:Pajzsoscankó]]
[[nl:Kemphaan]]
[[ja:エリマキシギ]]
[[no:Brushane]]
[[nn:Brushane]]
[[pl:Bojownik batalion]]
[[sk:Pobrežník bojovný]]
[[sl:Togotnik]]
[[fi:Suokukko]]
[[sv:Brushane]]
[[tr:Dövüşken kuş]]