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BIO554/754
Ornithology
LectureNotes3BirdFlightII

Birdsflyinavarietyofways,rangingfromglidingtosoaringtoflappingflighttohovering.Ofthese,thesimplesttypeofflightisgliding.

Aglidingbirdusesitsweight(mass)toovercomeairresistancetoitsforwardmotion.Todothiseffectively,ofcourse,requiresacertainmass&,asaresult,
onlylargebirds,suchasvultures,glideonaregularbasis.Whengliding,abirdlosesaltitudeatsome'sinkingspeed'(Vs)whiletravelingforwardatsome
'flightspeed'(V).Abird'sglideratioequalsV/Vs(thedistancetraveledforwardperunitofaltitudelost).Someofthebest'birdgliders'(suchasBlack
Vultures)maytravelupto20metersforeverymeterofaltitudelost(or,aglideratioof20).
Asoaringbird(e.g.,TurkeyVultures)maintainsorincreasesitsaltitudewithoutflappingitswings.Onewaytodothisistotakeadvantageofrisingair,e.g.,
updraftsaregeneratedwhenasteadywindstrikesahill,cliff,orbuilding,&thisisreferredtoasobstructionlift:

thermals,orupdraftscausedbytheunevenheatingofairneartheearth'ssurface.Airoverfieldsheatsfasterthanairoveraforestorlake.Thewarmer
airoverafieldislighterthanthesurroundingcoolerair&,therefore,rises.However,athighaltitudesthewarmairbeginstocool&sink.Asaresult,
birdsusingthermalsforlifttypicallyflyincircles(tostayintheareaofrisingair).

Source:kosetal.(2008)

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TrajectoryorflightpathofaPeregrineFalconsuperimposedonablackandwhitesatellitemapofthearea(southeastHungary).
Colorindicatesverticalvelocity,withmorereddishcolorindicatingclimbingwithinthermalsandbluishcolorindicatingsinking
(i.e.,periodsofglidingbetweenthermals)(Source:kosetal.2008 ).

DorsalwingprofileinsilhouetteofArgentavisiscomparedforscalingwiththoseofaBaldEagle.

ArgentavismagnificensfromtheupperMiocene(6millionyearsago)ofArgentina,withanestimatedmassof7072kgandawingspanof7m,wastheworld's
largestknownflyingbird.BecausethefossilsofArgentavisarefoundinthefoothillsoftheAndestothepampas,itislikelythatitusedprimarilyslopesoaringover
thewindwardslopesoftheAndesandthermalsoaringovertheopenpampas.Inslopesoaring,abirdfliesinaregionofrisingaircausedbyupwarddeflectionof
windoveraridgeoracliff.Ifthesinkingspeedoftheanimalislessthanthevelocityoftherisingair,thebirdisabletoremainairborneindefinitelywithoutflapping
itswings.CranialmorphologyindicatesthatArgentavis,likeotherteratorns,wasanactivepredatorratherthanascavenger.Itwasprobablyadiurnalpredator,
dependentonthermalsforflightactivityformuchofthetimemuchaslarge,broadwingedcarnivorousbirdsweseetoday.Strongthermalsoccurbymiddayand
disappearintheevening,sothermalsoaringforArgentaviswouldhavebeenpossibleonlybetweenthosetimes.Withaskull>55cmlongand15cmwide,
ArgentaviswascapableofcatchingsizeablepreywithitsformidablebeakFrom:Chatterjeeetal.(2007).

Overtheopenocean,largebirdsliketheWanderingAlbatrossshownbelowtakeadvantageofwindvelocitygradientsinatypeofsoaringcalleddynamic
soaring.

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WanderingAlbatross
PaulWardandCoolAntarctica

Soaringalbatross

DynamicsoaringAlbatrossesperformafascinatingand
complicatedflightmaneuvercalleddynamicsoaring,inwhich
energycanbeextractedfromhorizontallymovingairandtransferred
tothebirdsothatanenergygainisachievedwhichenablesittofly
continuouslywithoutflapping.Dynamicsoaringispossiblewhen
thewindspeedchangeswithaltitude.Thistypeofwind,whichis
calledshearflow,existsintheboundarylayerabovetheocean
surfaceinareasinwhichalbatrossesarefound.Dynamicsoaring
consistsofperiodicallyrepeatedcycles,withonecycleillustratedto
theleft:1climb(windwardflight)2uppercurve(changeof
flightdirectiontoleeward)3descent(leewardflight)&4lower
curve(changeofflightdirectiontowindward)(Sachs2005).

Dynamicsoaringisenergeticallyefficient.TheheartrateofaWanderingAlbatrosswasrecordedoveratwodayperiod
anditsheartratewasjustaboverestingrateswhensoaring,suggestingthatdynamicsoaringrequireslittlemoreenergythan
restingonland(Weimerskirchetal.2000).

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AntipodeanAlbatrosswithtubularnasalpassages

Procellariiformbirds:thetubeknowsairspeed?AlbatrossesandsmallerProcellariiformbirdslikepetrels
andshearwaterscantravellongdistancesovertheoceanbydynamicsoaring.Thismethodofsoaringrequiresthat
thesebirdsbeabletodetectvariationinwindspeedatvariousdistancesabovetheoceansurface.Pennycuick
(2002)proposedthatalbatrossesandtheirrelativescanusetheirtubenosesasapitottubetoveryaccurately
determineairspeed.Pitottubesonairplaneshavetwoholes,onepointingforward(inthedirectionoftheplaneis
flying)tomeasurewhatiscalledthestagnation(orpitot)pressure,andasideholethatmeasuresstaticpressure(the
ambientpressureofthesurroundingair).Thedifferencebetweenthestagnationpressureandstaticpressureis
calledthedynamicpressure,whichcanbeusedtodetermineaplanesairspeed.Forexample,inthediagrambelow,
asaplaneincreasesitsairspeed,thepressuregeneratedatthestagnationpointwillincreaserelativetothestatic
pressureandthefluidinthedifferentialmanometerwillbeforceddownward,outofthemanometer,andupwardin
thetubeleadingoutofthemanometer.Pitottubesarecalibratedsothatdynamicpressurereadingsaretranslated
intoairspeedreadings.

Inthisdiagram,thedifferentialmanometermeasuresdynamicpressure.

ThetubenosesofalbatrossesandotherProcellariiformbirdsresemblepitottubesandmayfunctioninthesame
way.Theforwardpointingnostrilsortubenoses(pointinginthedirectionthebirdsfly),thatcouldmeasure
stagnationorpitotpressure,leadintonasalchambersalsoconnectedtothemouth,ororal,cavity,wherepressure
wouldcorrespondtostaticpressure.Mangold(1946)identifiedanexpandablepocketorcapsuleoneithersideof
thenasalseptumofpetrelsandproposedthattheseweresenseorgansthatcouldmeasuredynamicpressure,the
differencebetweenstagnationandstaticpressure,andprovideinformationaboutairspeed.Additionalstudyis
neededtotestPennycuicks(2002)andMangolds(1946)hypothesis.

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(A)TransversesectionthroughaFulmarsskullshowingnasalchamberswithexpandablepocketsorcapsules.The
insideofthecapsulesareconnectedtotheforwardpointingnostrils(possiblydetectingstagnationorpitot
pressure),whereasthechamberoutsidethecapsulesisconnectedtotheoralormouthcavity(possiblydetecting
staticpressure).(B)Diagramofatypicalpitottubeusedtomeasureairspeedofairplanes.Theexpandablecapsule
registersthedifferencebetweenthestagnationorpitotpressurecomingfromtheforwardpointingtubeandthe
staticpressurecomingfromholesonthesidesofthetube.(From:Pennycuick2008).

Ofcourse,mostbirdsflaptheirwingswhentheyfly.Flappingflightinvolvesupanddownmovementofthewingsand,duringsuchflight,
differentpartsofawinghavedifferentfunctions:
theproximalpartofthewing(basicallythehalfclosesttothebody)movesless&providesmostofthelift
thedistalpartofthewingmovesthroughawidearcandgeneratesthethrustthatpropelsabirdforward.
Duringthedownstroke(powerstroke),awingmovesdownward&forward.Asaresult,theleadingedgeofthewing,particularlyfor
thedistalportionofthewing,islowerthanthetrailingedge.Asaresult,asshowninthefigurebelow,theresultantforce(R)isangledforward,
producingforwardthrust(checkoutaBaldEagletakingflight!).
Duringtheupstroke(recoverystroke),thetipsoftheprimariesseparate(lookattheCanadaGooseimagebelow)&these'slots'allowpassageof
airthroughthem(whichreducesfrictionasthewingcomesup).Also,thewingispartiallyfoldedatthewrist&elbowanddrawnintowardthe
bodytoreducedrag.

Differentforcesalongaflappingwing.(A)Thereislittleverticalmovementofthewingclosetothebird'sbody,but
thedistalportionofthewingisangleddownward(withtheleadingedgelowerthanthetrailingedge)andairmovingpastthedistalwing
ismovingfaster,andatadifferentangle,becauseofthewingsflappingmotion.(B)AtcrosssectionX,theliftisalmostvertical.(C)At
crosssectionY,becauseoftheangle,theliftforceistiltedforwardandproducesforwardthrust.D=dragforce,L=liftforce,R=resultantforce
(From:Alexander2002Fig.4.6).

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Theseimages,takenfromahighspeedrecordingofacockatielflyingat1meter/sec,showthetipreversalupstroke.
Inthefirstframe,thewinghasalreadyreverseddirectionandthehumerushasbeenelevated.Inthesecondframe,
theprimaryfeathershaverotatedslightlytocreategapsbetweensuccessivefeathers.Betweenthesecondandthirdframes,
therotatedprimariessweepupwardasthewristjointextends.Bythethirdframe,theprimarieshavebeenrotatedbackinto
theirstandardorientationandthewinghasbeguntomoveforwardaswellasupward(Hedricketal.2004).

Measuring magpie ight energetics

ABlackbilledMagpieflyinginawindtunnelattwodifferentspeeds
wearingcustomrespirometrymasks.

Cockatielflyinginawindtunnel

Gullinslowmotion

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RockPigeonsinflight

Birdstakingoffslowmotion

Gullsinslowmotion

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Mostspeciesofbirdsdonotflaptheirwingscontinuouslyduringflight.Rather,theyexhibitoneoftwointermittentflightpatterns:flapglidingandflap
bounding.Mathematicalmodelspredictthatflapboundingisenergeticallycheaperthancontinuousflappingflightathighspeeds,whileflapglidingismore
efficientthancontinuousflappingatlowspeeds.However,fewspeciesofbirdexhibitbothtypesofintermittentflight,soflapboundingmaybeacompromise
betweentheneedtomaintainmusclecontractionsatanoptimalvelocityandtheneedtovarypoweroutputandflightspeed.Inaddition,theprimaryflight
muscle,thepectoralis,ofmanysmallbirdsiscomposedofasinglemusclefibertype,furtherlimitingtherangeofusefulstrainratesforthesespecies.Thus,a
"fixedgearhypothesis"suggeststhattheonlyeconomicalmethodforsmallbirdstovarypoweroutputistointermittentlybound.However,investigatorsatthe
FlightLaboratoryattheUniversityofMontanahavefoundthatsomesmallbirds,suchasBudgerigarsandEuropeanStarlings,doexhibitbothtypesof
intermittentflight,withflapglidingbeingusedatlowerspeeds,andflapboundingathigherspeeds.Thissuggeststhatsomesmallbirdsarecapableof
optimizingtheirflightstylesdespitethetheoreticalconstraintsoftheirmusclecomposition.

Flapglidingflightpath(left)andflapboundingflightpath(right)
Source:http://www.biology.leeds.ac.uk/staff/jmvr/Flight/PWV/index1.htm

404NotFound
nginx

FlapglidingEuropeanSparrowhawk(Accipiternisus)

Timingofaflapboundingcyclerelativetowingmovements,altitude,andbodyangle(relativetohorizontal)fora
ZebraFinchflyinginawindtunnel(Videler2005,basedonTobalskeetal.1999).

DarkeyedJuncoflapboundingflight

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Source:http://biology.umt.edu/flightlab/Intermittent.htm

Asflightspeedincreasedinawindtunnel,budgerigarsthatexhibitedintermittentflightatallspeedstendedtoflextheir
wingsduringintermittentnonflappingperiods,apparentlyinresponsetoincreasedprofiledrag(TobalskeandDial1994).

Boundingflightisnotseeninbirdsofgreaterthan300grams,thuslikelytobeconstrainedbysize.Thesmallbirdswhichmostfrequentlyutilizeboundingtendto
haveshortroundedwings(lowaspectratio),poorlysuitedforgliding,makingundulatingflightlessaerodynamicallyattractive.
(Source:www.biology.leeds.ac.uk/staff/jmvr/Flight/PWV/index1.htm)

Goshawk Flies Through Tiny Spaces in Slo-Mo! - The Animal's Guide to ...

Afewbirdsusinghoveringflight.Somebirds,likeAmericanKestrels,'hover'orremaininplacebyflyingintothewindata
speedequaltothatofthewind,andotherbirds,likeOspreys(totheright)hovermomentarilywhileforaging.But
hummingbirds(checkthisshortvideo)areabletoremaininthesameplaceinstillairaslongastheywishtheyaretrue
hoverers.Ahoveringhummerkeepsitsbodyatabouta45degreeangletothegroundandmovesitswingsinmoreorlessa
figureeightpattern,withthe"eight"lyingonitsside.Hummingbirds,unlikeotherbirds,canalsoflybackwards(videoofa
hummingbirdinslowmotion).

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TurbulencecreatedasaRubythroatedHummingbirdhoverswithwingsbeatingabout40timespersecond.Thehummingbirddownstrokegeneratesabout2.5times
as
muchverticalforceastheupstroke.Associatedwithliftproductionisthesimilarpowerimbalancebetweenthetwohalfstrokes.Analysisindicatesthatinadditionto
theangleof
attack,wingvelocityandsurfacearea,dragbasedforceandwingwakeinteractionalsocontributesignificantlytotheliftasymmetry(From:Songetal.2014).Also,
checkthis
linkSciencegraphicoftheweek:hummingbirdwingaerodynamics.

Source:http://www.njsouth.com/leamingsrun.htm

Clickonthisphototoseeahummingbirdinslowmotion.

FemaleRubythroatedHummingbird(about45wingbeats/second)

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FemaleRubythroatedHummingbird(slowmotion)

BroadtailedHummingbirds(slowmotion)

Source:http://www.ae.utexas.edu/design/humm_mav/theory.html

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Ananalysisshowsairvorticesatthetip
Credit:NicolleRagerFuller,NSF ofahummingbird'swingsasitflies.
DouglasR.Warrick,BretW.Tobalske
andDonaldR.Powers

Doesahummingbirdflylikeaninsectorabird?Abitlikeboth.Manyexpertshadarguedthathummingbirds'skillat
hovering,ofwhichinsectsaretheundisputedmasters,meansthatthetwogroupsmaystayaloftinthesameway:by
generatingliftfromawing'supstrokeaswellasthedown.Thisturnsouttobeonlypartiallytrue.Otherbirdsgetalloftheir
liftfromthedownstroke(duringslowflightandwhenhoveringnotduringfasterflight),andinsectsmanagetogetequallift
frombothupanddownbeats(checkthisshortvideo),buthummingbirdsliesomewhereinbetweenandgettabout75%of
theirliftfromthedownstrokeand25%fromtheupstroke.

Credit:NicolleRagerFuller,NSF

Todeterminethis,Warricketal.(2005)trainedRufousHummingbirds(Selasphorusrufus)tohoverinplacewhilefeeding
fromasyringefilledwithsugarsolutionandlookedattheswirlsofairleftintheirwake.Theyfilledtheairwithamistof
microscopicoliveoildroplets,andshoneasheetoflaserlightinvariousorientationsthroughtheairaroundthebirdstocatch
twodimensionalimagesofaircurrents.Acoupleofquickphotographstakenaquartersecondapartcaughttheoildropletsin
theactofswirlingaroundawing.Althoughhummingbirdsdoflaptheirwingsupanddowninrelationtotheirbody,theytend
toholdtheirbodiesuprightsothattheirwingsflapsidewaysintheair.Togainliftwitheachstrokethebirdspartiallyinvert
theirwings,sothattheaerofoilpointsintherightdirection.Theirflightlooksalittlelikethearmandhandmovementsused
byaswimmerwhentreadingwater,albeititatamuchfasterpace.
Insectsattainthesameliftwithbothstrokesbecausetheirwingsactuallyturninsideout.Ahummingbird,withwingsof
boneandfeathers,isn'tquitesoflexible.Butthebirdsarestillveryefficient."Theirwingsareamarvellousresultofthe
considerabledemandsimposedbysustainedhoveringflight,"Warricksays."Providedwithenoughfood,theycanhover
indefinitely."Theresearchersaddthatthehummingbird'sflappingbearsastrikingresemblancetothatoflargeinsectssuchas
hawkmoths,anexampleofhowevolutioncanproducesimilarengineeringsolutionsinhugelydistantanimalgroups.
MichaelHopkin,NatureNews

Incontrasttootherbirds,thehummingbirdwingisfreetorotateinalldirectionsattheshoulder
becauseit'saballandsocketjoint(uniquetohummingbirdsandswifts).
(Source:http://www.ae.utexas.edu/design/humm_mav/theory.html)

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Hummingbird hovering under no rain, light r...

HummingbirdsflyingintherainFlightinrainrepresentsagreaterchallengeforsmalleranimalsbecausetherelativeeffectsofwaterloadinganddropimpact
aregreateratreducedscalesgiventheincreasedratiosofsurfaceareatomass.Nevertheless,itiswellknownthatsmallvolanttaxasuchashummingbirdscan
continueforagingeveninextremeprecipitation.OrtegaJimenezandDudley(2012)evaluatedtheeffectoffourrainintensities(i.e.zero,light,moderateandheavy)
onthehoveringperformanceofAnna'sHummingbirds(Calypteanna)underlaboratoryconditions.Lighttomoderaterainhadonlyamarginaleffectonflight
kinematicswingbeatfrequencyofindividualsinmoderaterainwasreducedby7%relativetocontrolconditions.Bycontrast,birdshoveringinheavyrainadopted
morehorizontalbodyandtailpositions,apositionthatmayreducethenumberofdropshittingabird'swingsandkeepitmorestableintheair.Inheavyrain,
hummingbirdsalsoincreasedwingbeatfrequencysubstantially,whilereducingstrokeamplitudewhencomparedwithcontrolconditions.Theratiobetweenpeak
forcesproducedbysingledropsonawingandonasolidsurfacesuggeststhatfeatherscanabsorbassociatedimpactforcesbyuptoapproximately50%.
Remarkably,hummingbirdshoveredwellevenunderheavyprecipitation(i.e.,270mmh1)withnoapparentlossofcontrol,althoughmechanicalpoweroutput
assumingperfectandzerostorageofelasticenergywasestimatedtobeabout9%and57%thigher,respectively,comparedwithnormalhovering.

Links:

Howhummingbirdsweatherthestorm

Secretofhummingbirds'abilitytoflyintherain

HoveringishardworkformostbirdsEverseenasongbirdhoveroveracrowdedfeedingstation,waitingforaperchtoopenupsoitcanlandandeat?
Lookslikehardwork,doesn'tit?Itis,whichiswhyhoveringissomethingmostbirdsdon'tliketodoorcan'tdoforverylong.Dialetal.(1997)surgically
implantedstraingaugesinthewingsofthreeBlackbilledMagpies.Thedevicesmeasuredtheforceexertedbythemainflappingmusclewitheachwingbeat.
Thebirdsthenflewinawindtunnelatarangeofspeeds.Thestraingaugeallowedthescientiststocalculatethepower(theamountofworkdoneperunittime)
requiredtomaintainagivenspeed.Hoveringtooknearlytwiceasmuchpowerasflyingataveragespeed,theresearchersfound.Evenwhenthemagpiesflew
attopspeed,theyexpendedfarlesspowerthantheydidwhentheyhovered.Evidencesuggestedthatwhentheyhovered,thebirdswereworkingattheir
physicallimits.Theirwingmusclesappearedtobeemployinganaerobicmetabolism,asourceofenergythatcan'tbesustainedforlong.Thereareclearly
exceptionstothis.Hummingbirds,theauthorsnote,haveanunusualshoulderdesignthatallowsthemtogenerateliftonbothdownbeatandupbeat.Butbirds
withabodydesignsimilartomagpiesarelikelytohavestrictlimitsontheirabilitiestoflystandingstill.

Kestrelhovering

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WhitetailedKitehovering

FormationFlying

Somebirds,likegeese&cranes,areoftenobservedflyinginVformation.Thereasoniswingtipvortices.Thebirdstakeadvantageoftheupwindsideofthe
vortexsheddingoffthebirdinfrontofthem.Thisupdraftactuallyliftsthebirdup,makingtheflightalittleeasier.

Airmovesfromtheareaofhighpressure(underthewing)totheareaoflowpressure(topofthewing)atthewingtips.Thisisnicelyillustratedinthephotooftheplanepassingthroughclouds.
BirdsflyinginVformationusethesevorticesofrisingair.

Avortexisformedinthewakeofeachwingtip,creatingdownflowbehindthewingandupliftoutside
thewake,asindicatedatthetipoftherightwingoftherighthandbird.Atrailingbirdcantakeenergeticadvantage
ofthisupliftbyflyingatasuitablylateralpositionrelativetothebirdahead.Theorysuggeststhattheoptimalwingtip
overlapforthetrailingbirdisaboutonetenthofthewingspanb.Adistanceofabout0.78bseparatesthecenters
ofthetwotrailingvorticesfromabirdoraircraft(AnderssonandWallander2004).

See"Mysteryofbird'V'formationsolved"(BBCNews)

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WingedMigration

FlyinginaflockcomesatacostFlyingbirdsoftenformflocks,withsocial,navigational,andantipredatorimplications.Further,flyinginaflockcanresultin
aerodynamicbenefits,thusreducingpowerrequirements,asdemonstratedbyareductioninheartrateandwingbeatfrequencyinpelicansflyinginaVformation.
Buthowgeneralisanaerodynamicpowerreductionduetogroupflight?Vformationflocksarelimitedtomoderatelysteadyflightinrelativelylargebirds,andmay
representaspecialcase.Whataretheaerodynamicconsequencesofflyinginthemoreusualclusterflock?Usherwoodetal.(2011)useddatafrombackmounted
GlobalPositioningSystem(GPS)andinertialsensorstoshowthatpigeons(1)maintainpowered,bankedturnslikeaircraft,imposingdorsalaccelerationsofupto2
g,effectivelydoublingbodyweightandquadruplinginducedpowerrequirements,(2)increaseflapfrequencywithincreasesinallconventionalaerodynamicpower
requirements,and(3)increaseflapfrequencywhenflyingnear,particularlybehind,otherbirds.Therefore,unlikeVformationpelicans,pigeonsdonotgainan
aerodynamicadvantagefromflyinginaflock.Indeed,theincreasedflapfrequency,whetherduetodirectaerodynamicinteractionsorrequirementsforincreased
stabilityorcontrol,suggestsaconsiderableenergeticcosttoflightinatightcluster.

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Foodandformationhelpbirdsflyefficiently.Swimmingafteraheavymealmaynotbewisebutflyingisanothermatter.Birdsflymoreefficientlywhen
loadedwithfood,recentresearchsuggests,helpingtoexplainhowtheycanmigratethousandsofkilometreswithoutstopping(Kvistetal.2001).Andasecond
studyhasconfirmedthecenturyoldsuspicionthatbirdsflyinaVformationtosavesubstantialamountsofenergy(Weimerskirchet
al.2001).AndersKvistatLundUniversityinSwedenandhiscolleagueslookedatflyingefficiencyinRedKnots(showntothe
right),smallwadersthatdoubleinsizefortheirannualmigrationfromSiberiatoAfrica.Fullyfed,RedKnotsflyinginawindtunnel
for610hoursextractedsignificantlymorepowerfromeachunitoffood.Thismighthelptoexplainwhybirdsoftenmakelongnon
stopflightsevenwhentheydon'thavetocrossanoceanordesert,saysKvist."Sinceefficiencyincreaseswhenthebirdsareheavy,it
mightnotbeasbadtomakelongflightsaspeoplethought."Theresearchfliesinthefaceofcomputerpredictionsthatbirdsareless
efficientwhenfull.SaysbirdaerodynamicsspecialistJeremyRayneroftheUniversityofLeeds:"It'samajoradvance,becauseithas
disprovedsomethingwe'veheldontoforalongtime."Thefindingis"extremelyunexpected",agreesJohnSpeakmanwhoworkson
animalenergyuseattheUniversityofAberdeen."Thischangesourwholeviewofmigrationalstrategiesintermsofhowmuchfat
birdsshoulddeposittocross,say,theSaharaDesert."Understandingtherelationshipbetweenfoodandflightmighthelpecologiststo
measuretheimpactofhabitatchangeonmigratorybirds,Speakmansays."Ifyou'redecidingwhethertofloodanestuary,for
example,thiscouldhelpyoumakemoresensiblepredictionsabouthowitwillaffectbirdsthatusetheestuaryasastopover."Itis
unclearhowbirdsincreasetheirefficiencywhenmigrating,Kvistsays.Puzzlingly,theydon'tadoptthemosteconomicalstrategyatalltimes.Kvistspeculatesthat
whenbirdsarebreedingtheymaykeepreservesofstrengthforsuddenmanoeuvressuchasspeedinguporswervingtoavoidapredator.
BirdsalsoconservefuelbyflyinginVformations.Bymeasuringheartrates,researchersinFrancenowhaveproofthatpelicansuse1114%lessenergyflying
together,evenwhentheyarenotperfectlypositionedtotakeadvantageofthewakefromthoseinfrontofthem.Configuredflightmaycreateastreamofairthat
allowsbirdstoglidelonger,suggestsHenriWeimerskirch,thebiologistattheNationalCentreofScientificResearchatVilliersenBois,wholedthestudy."Ifyou
lookclosely,youseethatthebirdsatthebackareglidingmorethantheleader."PeoplehavebeenaskingwhetherVformationsaremoreefficientformorethan100
years,Speakmansays,butnoonehadmeasuredenergysavingsbefore."Theytookacenturyoldproblemandwenttotheheartofit,"hesays.WrittenbyErica
Klarreich.

FlightMetabolism

Allbirdshavehighmetabolicrates,andflyingbirdshaveevenhigherrates.Themetaboliccostofflightdependsonthetypeofflight(gliding,soaring,flapping,
orhovering),wingshape,andspeed.Ofcourse,flappingflightandhoveringarethemostcostlytypesofflight.Laboratorystudiesofbirdstrainedtoflyinwind
tunnels(liketheonebelow)indicatethatthemetabolic'cost'offlappingflightcanbeanywherefromabout7to15timesabird'sbasalmetabolicrate.

Source:http://www.swe.org/iac/LP/wind_03.html

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Cockatielinawindtunnel

Speedinfluencesthecostofflight,withlowspeedflight(suchaswhentakingofforlanding)requiringmoreenergy.Someinformationalsosuggeststhatbird's
flyingatmaximumspeedsalsousemoreenergythanat'medium'speeds.Forexample,inthegraphbelow,notethatEuropeanStarlingsusemuchmoreenergyat
lowspeeds(02meters/second)thanathigherspeeds.TherelationshipbetweenflightspeedandenergyconsumptionisalsoveryapparentforBudgerigars(below).
Lowspeedflightismorecostlybecausethereismoredrag(induceddrag).Atlowspeeds,airflowoverwingsisrelativelyslowand,tomaximizelift,birdsmust
maximizetheangleofattackandflaptheirwingsfasttoincreaseairspeedandthisrequireslotsofpower.Highspeedflappingflight(asillustratedforBudgerigars
andEuropeanStarlingsbelow)ismorecostlybecause,atgreaterspeeds,frictiondragandparasitedragincrease,requiringanincreaseinwingbeatfrequencyand/or
anincreaseintheproportionofmusclecells(inthepectoralismuscles)contracting.Thegraphsbelowclearlyrevealsthatflightismostefficientat'medium'speed.

Totalmechanical(aerodynamicPaero )poweroutputduringflappingflight
atdifferentflightspeedsinCockatiels(Nymphicushollandicus).DatarepresentsmeansSE.
Pind =inducedpower,Ppar=parasitepower,andPpro =profile(friction)power(Brighton2007).

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Superficial(left)anddeep(right)flightmusclesofaBlackbilledMagpie(Tobalskeetal.1997).

Distributionofenergyduringflight(Bejan2005).

A10,000kmnonstopflightFourBartailedGodwits
(Limosalapponica)flewtheirwayintotherecordbooks
withnonstopflightsofmorethan10,000kmfromNew
ZealandtotheYellowSea.Thegodwits,trackedby
satellitetransmittersinMarch2007,didnotstoptoeator
drinkonthefirstlegoftheirnorthernmigrationthatends
inAlaskainMay.PhilBattley,anecologistatMassey
University,saidithadbeensuspectedthatthebirdscould
flysuchdistancesbutnowithadbeenproved.Noother
animalhasshownsuchendurance,hesaid.Thefemale
godwitstook6to7daystocovertheroute,flyingat
altitudesupto2kmandatanaveragespeedof56km/h.
WhenthegodwitsleaveNewZealand,theyareclinically
obese,butloseabouthalftheirbodyweightduringeach
portionoftheirmigratoryflight.Afterarrivinginthetidal
flatsoftheYellowSea,offChinaandSouthKorea,they
stayforamonthortwotorefuel.Battleysaid"It'sthe
equivalentofridingtheTourdeFrancebutkeepingitup
foraweeknonstop." CheckthePacificShorebirdMigrationProjectWebpageforup
todateinformation.

Birds,ofcourse,getaroundinwaysotherthanflying.Infact,somebirdsareflightlessanddependentirelyonwalking,running,or

swimmingtogetfromplacetoplace.Somebirdsspendmostoftheirtimeonorinwater.Birdshavespecialadaptationsofthelegs,feet,&
wingsforterrestrialandaquatic(swimminganddiving)locomotion.

Walking,running,hopping,&waddlingbirdsthattravelalongthegroundregularlyoftenhaverelativelylonglegs.Amongtheratites,
suchasOstrichesandEmus(orcheckthisvideo),therehasbeenareductioninthenumberoftoes(lessweightatendofthelimb=more
efficientlocomotion).

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Withshort,lighttails(duetoreductioninthenumberofcaudalvertebrae)andlargeflightmuscles,naturalselectionhasfavored
positioningofbirdfeetunderamorecranialpositionedcenterofmass.Thisisachievedbyasubhorizontalorientationofthefemur
and,whenwalkingandrunning,thekneeactingasthemainfulcrumnearthebird'scenterofmass(From:Nyakaturaetal.2012).

How fast does an Ostrich run?.mov

WesternGrebes

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Runningshorebirds

WoodDuckswalking

Waddlingmakesmostofpenguin'sshortlegsItmaynotbegraceful,butthepenguin'swaddlemakesperfectsensetoscientists,whofoundthatthebird'ssideto
sidegaitconservesenergy.UniversityofCaliforniaresearchersfoundthatthegaitworkslikeapendulum,withenergystoredattheendofeachswingforthebird's
nextstep."Ourfindingsindicatethatwalkingisexpensiveforpenguinsnotbecauseoftheirwaddling,butbecausetheyhavesuchshortlegsthatrequiretheirleg
musclestogenerateforceveryquicklywhentheywalk,"saidTimothyGriffin,aUCBerkeleygraduatestudent.GriffinandKram(2000)decidedtostudypenguins
becausetheyseemtobedoingeverythingwrong.Anearlierstudyshowedpenguinswereburningtwiceasmanycalorieswhenwalkingasotheranimalsofsimilar
size.Butresearchersfoundtheproblemwasthepenguins'legs,nottheirjerkysidetosidemovements.TheEmperorpenguinsstudiedatSanDiego'sSeaWorld,for
instance,wereatleast3feettallbuthadlegsonlyabout10incheslong.Penguinsburnaboutthesameamountofcaloriesasanimalswithsimilarleglengths,Griffin
said.Theresearcherscoaxedpenguinsacrossaforceplatform"kindofafancybathroomscale,"saysGriffinwithbitsoffish.Usingscalemeasurementsand
videos,thescientistsmeasuredthesidetosideandforeandaftforcesthepenguinsexertwhilewalking,aswellastheverticalforcessupportingtheirweight.The
fivepenguinsstudiedhadawalkingspeedofabout1.5feetpersecond.Thepercentageofenergyretainedduringtwostepsiscalledtherecoveryrate.Humanshave
arecoveryrateofabout65percent.ThepenguinsstudiedbyGriffinandKramhadanimpressiverecoveryrateofupto80percent.

HeadmovementsinwalkingWhoopingCranes.(A)Oneframeofavideorecordingofawalkingcrane,showingmethod
ofmeasurementofhead,body,andlegposition.Theheadisfitwithagraphicalmodeloftheeyeandbill,thebodywithacirclescaledto
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headandlegsizeandcenteredoverthepelvisofthebird,andeachlowerlegwithalinesegmentextendingfromtheankletothefoot
(green,rightlegred,leftleg).(B)Onesequenceofmeasurements,atintervalsof33ms,ofaspontaneouslyforagingWhoopingCrane
throughseveralcompletesteppingcycles.Thebirdwalkedatanaveragespeedofabout0.46ms1.Duringthissequence,theright
footcompletednearly3stepsandtheleftfoot,about2.5steps.Theheadwasstabilizedthroughoutmostofeachfootsstep,withits
positionsateachofthesetimesindicatedbythearrows.(Watchacranewalk,clickhere!videobyThomasCronin).

AvianheadbobbingManyspeciesofbirdsmovetheirheadsforwardthroughaseriesofsuccessive,fixedpositionswhenwalking.Thisuniqueheadbobbing
behaviorstabilizesvisualfieldsduringbodymovement,preventingmotionbluroftheretinalimage.Gazestabilizationcouldberequiredforsuccessfulvisual
search,particularlyformovingobjects,butthetimeavailableforstabilizationvarieswithwalkingspeed.Nodirectevidencehasbeenpublishedshowingthatbirds
favorthestabilizationphasewhileforagingeitherformovingorimmobilefood.Croninetal.(2005)examinedheadbobbingbehaviorinforagingWhoopingCranes
(Grusamericana)astheysearchedthegroundforfood,andfoundthattheywalkatspeedsthatallowtheheadtobeimmobilizedatleast50%ofthetime.Thestable
phaseofbirdheadbobbingmovementsisparticularlyinterestingbecausethebehavior,uniquetobirds,clearlycontributestovisualgazestabilization.Pigeonshead
bobwhenlanding,andheronsstabilizetheirheadsrigidlywhenwalkingorwhentheirperchmoves,almostcertainlyforvisualfunction.Headmovements
neverthelessplayessentialrolesinvision,givingvisualcuesfordistancesandrelativelocationsofobjects,providinganopportunityforchangesinheadangle,and
permittingbirdstofixatenewobjectsofvisualinterest.

Climbingbirdsthatclimb,likewoodpeckers,nuthatches,BlackandwhiteWarblers,andPiedMonarchshavesharplyrecurvedclawstohelpgripthe
substrate(e.g.,barkofatree)

WhitebreastedNuthatch

Source:http://animalpicturesarchive.com/animal/APAsrch3.cgi?qt=nuthatch

BlackandwhiteWarbler

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Acomplete'hop'ofatreecreeperclimbingupwardsonaverticaltrunk.Thesequenceisfromlowerlefttoupperright.

Birdsadaptedforclimbing,likewoodpeckersandtreecreepers,havesharplyrecurvedclawsandtoes,sometimesrelativelylong,thatcanbespreadaparttohelp
firmlygripthesubstrate(typicallytreebark).Otheradaptationsforclimbingdifferwithforaginghabits.Climbersthattypicallymoveuptrees,likewoodpeckers
(Picidae)andtreecreepers(Certhidae),haverelativelyshortlegs(particularlythetibotarsus)thatkeeptheircenterofmassclosetothesubstrateandalong,stifftail
thatprovidessupportagainsttheforceofgravity.Aswoodpeckersandtreecreepersmoveupatree,theyhopupwardandinward(tocounteracttheforceofgravity
thattendstopullthemawayfromaverticaltreetrunk),movingbothfeetinunison.Tailsupportprovidestwoadvantages:(1)thelongtailcreatesalongbaseline
betweenthepointsofattachment(feetandtail)and,thelongerthisbaseline,thesmallerthehorizontalforcebetweenfeetandbarkagainstwhichthebirdmustwork
whenpullingitselftowardsthetrunkwhenhoppingupward,and(2),whennotmoving,thetail,ratherthanthelegmuscles,supportspartorallofthebirdsweight
(Norberg1981).

Climbingbirdsthatusetheirtailforsupportalmostalwaysmoveupwardswhenforaging.Foragingwoodpeckersandtreecreepersapproachingthetopofonetree,
typicallyflydownwardtoalowerpositiononanothertreethenagainclimbupwardsand,whenapproachingthetop,repeattheprocess.Notonlydoessucha
foragingstrategymakesenseenergetically(becauseflyingdownwardislesscostly),butattemptingtomovedownwardwhenforagingwouldcreateatleastthree
problems(Norberg1981):(1)difficultyinseeingwheretograspthebarkafterahop,(2)thestifftailcouldgetcaughtontheirregularsurfaceofthebark,and(3)
potentialpreywouldbealertedtothepresenceofapossiblepredatorbeforethebirdcouldgetinapositiontocapturethem.

Nuthatches(Sittidae)areadaptedforclimbingdownwardaswellasupwards.Theirrelativelyshorttailsarenotusedforsupportand,ratherthanhopping,nuthatches
walkupanddowntreetrunksandbrancheswithalternatinglegmovements.Nuthatcheshaverelativelylonglegs(particularlythetibiotarsus),allowingarelatively
longbaselinebetweenthefeetandreducingthehorizontalforcebetweenfeetandbarkandtheenergeticcostoflocomotion(Norberg1981).

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WhitethroatedTreecreeper(Cormobatesleucophaeus)

Swimmingaquaticbirds(likeTuftedDucks)typicallyhave:
lowspecificgravity(lightweightsotheyareverybuoyant)featherswithlotsofbarbules&hooklets(lesspermeabletowater)welldevelopeduropygial
gland(secretionshelpkeepfeathersingoodcondition)
webbedfeetthatactlikeoars

Source:http://www.oaklandzoo.org/atoz/video22.html

WoodDuck

AswimminganddivingAnhinga

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CommonLoon

Divingbirdsthatfrequentlydiveunderwater,suchasgrebes,cormorants,&loons,have:
relativelyhighspecificgravities(heavierandlessbuoyant)
feetlocatedwellbackonthebodytopermitbetterpropulsionandmaneuveringunderwaterand/orsmallerwingsthatpermit'flying'underwater(e.g.,
scoters,petrels,murres,and,ofcourse,penguinsliketheAdeliePenguinbelow)

Source:http://www.bionik.tuberlin.de/intseit2/xs2pinfi.html

WingbonesofaJackassPenguin(Spheniscusdemersus).
Forwingpropelledswimmingunderwater,thepaddlemustbehighlymobileattheshoulder
(humeruspectoralgirdlearticulation),buttheremainingjointsneedtoberelativelyfixedtominimize
themusclecontractionneededtomaintaintheproperposition(Louw1992).

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EmperorPenguins

BubblesinthewakeofaPigeonGuillemot(Cepphuscolumba)swimminghorizontallyunderwater,
indicatingpatternsofintermittentthrustmainlyonthedownstroke.B)Wingpositionsduringhorizontal
swimmingbyaCommonMurre,asdrawnfromfilmstakenat32frames/sec.Sequenceisfrom
lefttorightandtoprowtobottomrow.Angleofattackofthewingssuggestssubstantialliftduringtheupstroke
(From:Lovvorn2001).

Guillemotsdiving

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RapidascentofEmperorPenguinsTojumpoutofwaterontoseaice,EmperorPenguinsmustachievesufficientunderwaterspeedtoovercometheinfluenceof
gravitywhentheyleavethewater.Therelevantcombinationofdensityandkinematicviscosityofairismuchlowerthanforwater.Injectionofairintoboundary
layers(airlubrication,i.e.,anairfilmseparatesthewaterfromthesurfaceofastructureorabirdthusreducingfriction)hasbeenusedbyengineerstospeed
movementofvehicles(ships,torpedoes)throughseawater.Basedonanalysisofpublishedandunpublishedunderwaterfilm,Davenportetal.(2011)hypothesized
thatfreerangingEmperorPenguinsemployairlubricationinachievinghigh,probablymaximal,underwaterspeeds(mean=5.3meters/sec),priortojumps.
Penguinsdiveto15to20meterswithairintheirplumageandthatcompressedairisreleasedasthebirdssubsequentlyascendwhilemaintainingdepressedfeathers.
Finebubblesemergecontinuouslyfromtheentireplumage,formingasmoothlayeroverthebodyandgeneratingbubblywakesbehindthepenguins.Inseveral
hoursoffilmofhundredsofpenguins,nonewereseentoswimrapidlyupwardswithoutbubblywakes.Penguinsdescendandswimhorizontallyatabout2
meters/sec.Davenportetal.(2011)hypothesizedthatasignificantproportionoftheenhancedascentspeedisduetoairlubricationreducingfrictionalandformdrag
andthatbuoyancyforcesalonecannotexplaintheobservedspeeds.

EmperorPenguinscreatebubbletrails(image:BluePlanet,BBC)

Links:

Penguinstaketotheair

Theflightofthepenguins

Why Divers Have Small Wings Many researchers believe that small
wings reduce drag underwater and, therefore, are bettersuited for diving.
But until recently, there was no concrete evidence for the supposed
benefits of small wings. Studying the effects of wing area on diving is
difficultcrossspeciesstudiesnevergivefaircomparisons.Bridge (2004)
decided to study the effect of altered wing size on Common Guillemots
(Uria aalge) and Tufted Puffins (Fratercula cirrhata) during their brief
moltingperiods.

Bridge (2004) used video cameras to film the bird's diving activity at
SeaWorldCaliforniabymountingonecamerainfrontofthepool'sviewing
window, and the other above the
pool pointing straight down. This
way, he could plot the bird's
movement in three dimensions and
calculate diving parameters such as
dive speed and angle of descent.
Bridge (2004) found that instead of
improving the bird's diving
performance, wing molt had an
unexpectedly adverse effect. During
molt, the birds dived a shorter distance with each flap of the wings, and
energyoutputfromthewingmovement,asmeasuredbyworkperflap,was
also reduced, especially when both primary and secondary feathers were WingmoltstagesofaTuftedPuffin
missing. wing.Approximationsofthe
percentageofintactwingareawith
Butifreducedwingareasdonotimprovedivingability,whyhasnatural thewinglooselyextendedarelisted
selectionfavoredsmall,pointedwingsinmanyaquaticbirds?Apparently foreachmoltstage(Bridge2004).
birds with small, pointed wings are adept at highspeed, longdistance
flight,essentialforrapidmovementbetweenhabitats.But,small,pointed

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wings cannot generate lift at low speed, so rapid vertical takeoffs are
impossible.Thisisnotabigproblemformostdivingbirdsbecausetheir
openaquatichabitatspreventcloseapproachbyundetectedpredators.In
addition,whenthebirdsslowdowntoland,theirsmallwingsstalleasily
andloselift.Fortunately,highspeedhardlandingsaremoreacceptableon
water than on land. Thus, aquatic habitats relax the constraints on the
evolution of small, pointed wings. Jane Qiu, Journal of Experimental
Biology

GreatCormorantfeathershavearegularandhighlywaterproofcentralpartwhereasthedistalregionisirregularandwettable.
Aventralfeatherisshown(From:Grmilletetal.2005).

UnusualfeatherstructureofGreatCormorantsWaterhasveryhighspecificheatandthermalconductivity,sothatdivingendothermscanpotentially
losemuchheattosurroundingwater.Todealwiththischallengingenvironment,mostwarmbloodeddivershavehighlyefficientbodyinsulation.
Marinemammalshaveevolvedthickskinsandextensiveperipheralfatlayers,whiledivingbirdshavedense,highlywaterproofplumagethattrapsan
insulatinglayerofair.Afewdivingbirdspecies,includingAnhingasandcormorantsarepuzzlingexceptionstothispattern,havingplumagethatisapparently
penetratedbywaterduringsubmersion.TheGreatCormorant(Phalacrocoraxcarbo)isthoughttohaveawettableplumage,providinglow
bodyinsulationduringforaging.GreatCormorantsshouldthusbeconstrainedbywatertemperatures,andshowhighenergyrequirements.
Surprisingly,thisspecieshasoneofthewidestbreedingdistributionsofalldivingbirds,anddoesnotrequiremorefoodthantheseotherspecies.
Grmilletetal.(2005)exploredthisapparentparadoxbycomparingtheinsulativepropertiesofbodyplumageinfoursubspeciesofgreatcormorants
rangingfromtropicaltopolarregions.Theauthorsfoundthatallsubspeciesretainedaninsulatingairlayerintheirplumage,whichwas,however,much
thinnerthanforotherspeciesofdivingbirds.Detailedexaminationoftheplumageshowedthateachcormorantbodyfeatherhasaloose,
instantaneouslywet,outersectionandahighlywaterproofcentralportion.Thisindicatesthattheplumageofgreatcormorantsisonlypartlywettable,
andthatbirdsmaintainathinlayerofairintheirplumage.Thesefindingssuggestanunusualmorphologicalfunctionaladaptationtodivingwhichbalances
theantagonistconstraintsofthermoregulationandbuoyancy.

DoublecrestedCormorant

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ImperialCormorantdiving

Photosource:http://www.copyrightfreepictures.org.uk/

FootpropelledlocomotionWhensubmerged,
GreatcrestedGrebes(Podicepscristatus)swimwith
synchronizedfootstrokes,keepingtheirwingsclosely
foldedagainstthebody.Duringthepowerstroke,the
feetmovefromacranialandventrolateralpositionto
acaudalanddorsomedialpositionrelativetothebody.
Themeanswimmingspeedvariedfrom0.71.2
meters/sec(JohanssonandNorberg2001).
Dorsal(left)andlateral(right)videoframesofadivinggrebe.
Thedorsalviewwasrecordedafterreflectionfromamirror.

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WesternGrebe

Next:BirdBiogeographyI

LectureNotes:
IIntroductiontoBirds

IIBirdFlightI

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Alexander,D.E.2002.Nature'sflyers:birds,insects,andthebiomechanicsofflight.JohnsHopkinsUniversityPress,Baltimore,MD.

Andersson,M.andJ.Wallander.2004.Kinselectionandreciprocityinflightformation?BehavioralEcology15:158162.

Bejan,A.2005.Theconstructallawoforganizationinnature:treeshapedflowsandbodysize.JournalofExperimentalBiology208:16771686.

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Chatterjee,S.,R.J.Templin,and
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Pennycuick,C.J.2008.Informationsystemsforflyinganimals.In:TheoreticalEcologySeries,vol.5.Modellingtheflyingbird(C.J.Pennycuick,ed.),pp.305
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Tobalske,B.W.,N.E.Olson,andK.P.Dial.1997.FlightstyleoftheBlackbilledMagpie:variationinwingkinematics,neuromuscularcontrol,andmuscle
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Tobalske,B.W.,W.L.Peacock,andK.P.Dial.1999.KinematicsofflapboundingflightintheZebraFinchoverawiderangeofspeeds.JournalofExperimental
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Warrick,D.R.,B.W.Tobalske,andD.R.Powers.2005.Aerodynamicsofthehoveringhummingbird.Nature435:10941097.

Weimerskirch,H.,T.Guionnet,J.Martin,S.A.Shaffer,andD.P.Costa.2000.Fastandfuelefficient?Optimaluseofwindbyflyingalbatrosses.Proceedingofthe
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Usefullinks:

DynamicSoaring

FlappingFlightWebSite

HummingbirdFlight

TheintermittentflightofZebraFinches:Unfixedgearsandbodylift

"UnderwaterFlight"ofthePenguin

BacktoBIO554/754syllabus

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