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Plumage Colour and Pattern in Waders

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Plumage Colour and Pattern in Waders Plumage colour and pattern in waders P.N. FERNS School of Biosciences,Cardiff University,PO Box 915, Cardiff CF10 3TL, UK, e-marl: [email protected]. uk Ferns,P.N. 2003. Plumagecolour and pattern in waders.Wader Study Group Bull. 100: 122-129. At least 16 of the hypothesesthat have beenproposed to explainthe evolutionof plumagecolour and pattern in birdsmay be applicableto waders- five straightforwardphysical explanations, and the restinvolving some form of communicationbetween animals. Eight of the latterinvolve communication between different species (suchas predator and prey) and the rest involve communication between individuals of the samespecies (such aspotential mates). These hypotheses are reviewed and an additionalone based on socialselection proposed. INTRODUCTION Tyndall blue is one of the coloursproduced by physical effects.It is alsoreferred to as incoherentRayleigh or Nie At least30 hypotheseshave been proposed to explainadap- scattering,though it is actuallyproduced by coherentinter- tive colorationin birds(Savalli 1995). The evidenceusually ference (Prum et al. 1998). This involves the reflection of consistsof little more than a few examples,and it is often shorterwavelengths of light by minute air bubblesin the possibleto cite just as many contradictoryones (Baker & feathersand the absorptionof longerwavelengths by under- Parker 1979). We tend to rely on our own judgementabout lying melanindeposits. No wadersshow pure blue but the what constitutes,for example,good camouflage, overlook- Ibisbill Ibidorhynchastruthersii has bluish-grey on the head ing how easyit is to make the facts fit our preconceptions and neck, and the Egyptian Plover Pluvianusaegyptius has (see Fig. 1). Most mammalian and avian predatorshave blue-greywing covertsand tail feathers.These probably owe visionthat is very differentfrom our own,probably better in their colorationto Tyndall effectssince bluish pigments are the case of birds and worse in the case of mammals (Endler extremelyrare in birdfeathers. Iridescence occurs more com- 1978, Jacobs1993, Finger & Burkhardt1994). This makes monlyin waders.For example,it is responsiblefor theglossy it difficult to designgood experimentsto test many of the sheenon the back of many oystercatchers,the Red-kneed ideasabout animal coloration, but this shouldnot discourage Dotterel Charadrius cinctus and Black Stilt Himantopus wader workersfrom trying to do so. novaezelandiae,as well as the iridescent blue, green and brown of the mantle and wings in NorthernLapwings and TYPES OF PLUMAGE COLOUR SouthernLapwings V. chilensis.Typically, iridescenceis producedby severalevenly spacedand parallel layersof Plumagecoloration can be producedby physicaleffects as melaningranules, separated by keratin.Light is reflectedby well asby pigments,and there are examples of bothof these the top of eachlayer andthe interactionbetween the refrac- in waders.Melanin is perhapsthe mostfamiliar and wide- tive indexof keratinand the spacingof the layersdetermines spreadpigment occurring in feathers.It is derivedfrom the the colourproduced (Lucas & Stettenheim1972). aminoacid tyrosineand occursin at leasttwo quite differ- Carotenoidsare the othermain groupof pigmentsfound ent forms- eumelaninand phaeomelanin. Eumelanin is re- widely in bird feathers.They are responsiblefor a rangeof sponsiblefor the browns,greys and blacksthat are present brightred, orangeand yellow colours.Most of the yellows in at leastsome of thefeathers in nearlyevery species of bird. and reds in wader plumageare producedby phaeomelanin The pigmentoccurs in granulesthat are restricted to the out- (Harrison1965, Lucas & Stettenheim1972). Indeed, the only ermostlayers of the featherkeratin. publishedevidence of carotenoidpigment in the feathersof Phaeomelaninproduces various shades of brown, chest- a wader, refers to an unidentified one responsiblefor the nut and red, includingthe bright red breedingplumage of greenish-yellowwing feathers of theNorthern Jacana Jacana many wader species,such as Red Knot Calidris canutus, spinosa(Auber 1957). In waders,therefore, carotenoids are Dunlin C. alpina, Curlew SandpiperC. ferrugineaand Bar- largelyrestricted to the iris, orbitalring, legsand beak. The tailed Godwit Limosa lapponica (Harrison 1965, Lucas & red colourof all of theseareas in oystercatchersHaematopus Stettenheim1972). Severalother species have orange-brown spp.is probablydue to carotenoids. or chestnutareas of feathering that are probably due to phaeomelaninbut in whichthe precise nature of thepigment TYPES OF PLUMAGE PA'I-rERN remainsto be established,e.g. NorthernLapwing Vanellus vanellus(vent), American Avocet Recurvirostraamericana The transitionbetween differently colouredareas of plum- (head and breast),Red-necked Avocet A. novaehollandiae agecan be dividedinto threedifferent types - (i) gradual, (headand neck), BandedStilt Cladorhynchusleucocephalus (ii) abruptand irregular,and (iii) abruptand regular. In the (breast)and many plovers(breast and/or belly). Another caseof a gradualtransition, the colours merge into one another, form of melaninis responsiblefor the yellowishplumage of andit is impossibleto definea preciseborder. For example, manyyoung birds (Lucas & Stettenheim1972) including the the orange-brownneck of the AmericanAvocet fades gradu- chicksof Pluvialis plovers. ally into the white of the rest of the body. Irregular,abrupt Bulletin 100 April 2003 122 Ferns: Plumage colour and pattern •n waders 123 Fig. 1. Curlewdrawing from Thayer (1909). Althoughthe Curlewdoes have plumagethat is crypticagainst a backgroundof grass, it is possibleto infer that almostany plumagepattern is crypticby presentingthe bird againstan appropriatebackground. transitionsare not especiallycommon, but are seenin the PHYSICAL FUNCTIONS OF COLOUR AND black wing tips of the Pied AvocetRecurvirostra avosetta, PATTERN and the white wing patches of the White-tailed Plover Vaneflus leucurusand Long-toedLapwing V. crassirostris. Thermoregulation Transitions that are abrupt and regular, i.e. that involve a suddenchange in colour along a straightor smoothlycurv- Dark coloursincrease the absorptionand emission of energy ing borderare commonin waders;for examplemost of the from solarradiation, whilst light onesdecrease it (Campbell head and neck bandsof plovershave bordersof this type. & Norman 1998). For this reason,Fjeldsfi (1977) has sug- Indeed,of the ninefamilies of waders,this type of transition gestedthat the dark dorsalplumage of somechicks from is commonon the headand body of eight of them (Rostra- damphabitats may be an adaptationto fasterwarming in the tulidae, Dromadidae,Haematopodidae, Ibidorhynchidae, sun,whilst the pale dorsalcolours of othersfrom arid envi- Recurvirostridae,Glareolidae, Burhinidae (head only) and ronmentsmay protectthem by reflectinglight andheat. Red- Charadriidae).Only in the largestfamily - the Scolopacidae neckedPhalarope Phalaropus lobatus chicks may be in the - are abrupt,regular borders largely absent(with the excep- former category, and Stone Curlew Burhinus oedicnemus tion of a few eye stripes).The overallpatterns that are sepa- chicks in the latter. rated by bordersof thesetypes may dependsimply on the shapeof a particulararea of the bodythat they highlight, e.g. Physical protection therump, or theymay have a specificshape - notablystripes and lines on the head and wings, and curved bandson the Melaninimproves the resistance of feathersto wearin a wide chin, breastand belly. range of species(Barrowclough & Sibley 1980, Bergman Otherpatterns, including those involving streaks and spots 1982).This probablyexplains why theflight feathersof most cannoteasily be categorisedin the aboveway sincethey do wadersare dark, even when the rest of the plumageis pale, not involve uniform blocksof colour and do not have very e.g. most coursersCursorius spp. In the caseof variegated preciseborders. These patterns are usually associatedwith feathers,the paler edgeswear fasterin waders(Hayman et crypsis. al. 1986). Bufferin 100 April 2003 124 Wader Study Group Bulletin Radiation protection WoodcocksScolopax spp. and snipesGallinago spp.pro- vide classicexamples of blendingcoloration in which their Dark colours,especially those produced by melanin,protect mottledplumage makes them virtually invisibleagainst the feathersfrom damagingUV radiationthat might otherwise ground(Darwin 1871). Protectiveresemblance is no more acceleratethe rate of feather wear (Burtt 1978, Hayman et than blendingcoloration in whichthe resemblanceto some al. 1986). Suchwear is especiallylikely in the exposedtips specificfeature of thebackground environment is particularly of the primaries(Burtt 1978, 1986) and so theseare often notable.The resemblanceof WoodcockScolopax rusticola protectedby heavydeposits of melanin,and there is usually plumageto deadleaves, sticks and otherground vegetation a slightdecrease in the amountof melaninproximally, e.g. (Thayer 1909, McKelvie 1986), andRinged Plover Charad- mostlapwings and ploversof the genusVanellus. rius hiaticula plumageto stones(Cott 1940) are examples. The Ringed Plover is a classiccase of disruptivecamou- Sighting lines flage, with both the chick andthe adult cited (Thayer 1909, Cott 1938), and drawingsof the chick featuringin the clas- It has been suggestedthat eye stripes,with their sharpbor- sic work on animal coloration(Cott 1940). Both have black ders,can act as sightlines and henceimprove the accuracy and white bands acrossthe neck, with abrupt and regular with which birds can strike at mobile prey (Ficken & Wil-
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