19/09/2016 Flightinslowmotion:aerodynamicsofthepterosaurwing|ProceedingsoftheRoyalSocietyofLondonB:BiologicalSciences

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Flightinslowmotion:aerodynamicsofthe
pterosaurwing
ColinPalmer
Published24November2010.DOI:10.1098/rspb.2010.2179

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Abstract

Theflightofpterosaursandtheextremesizesofsometaxahavelongperplexedevolutionarybiologists.
Pastreconstructionsofflightcapabilitywerehandicappedbytheavailableaerodynamicdata,whichwas
unrepresentativeofpossiblepterosaurwingprofiles.Ireportwindtunneltestsonarangeofpossible
pterosaurwingsectionsandquantifythelikelyperformanceforthefirsttime.Thesesectionshave
substantiallyhigherprofiledragandmaximumliftcoefficientsthanthoseassumedbefore,suggestingthat
largepterosaurswereaerodynamicallylessefficientandcouldflymoreslowlythanpreviouslyestimated.In
ordertoachievehigherefficiency,thewingbonesmustbefaired,whichimpliesextensiveregionsof
pneumatizedtissue.Whetherfairedornot,thepterosaurwingswereadaptedtolowspeedflight,unsuitedto
marinestyledynamicsoaringbutadaptedforthermal/slopesoaringandcontrolled,lowspeedlanding.
Becausetheirthinwalledbonesweresusceptibletoimpactdamage,slowflightwouldhavehelpedtoavoid

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injuryandmayhavecontributedtotheirattainingmuchlargersizesthanfossilorextantbirds.Thetradeoff
wouldhavebeenanextremevulnerabilitytostrongorturbulentwindsbothinflightandontheground,akin
tomoderndayparagliders.

1.Introduction

PterosaurswereflyingreptilesthatlivedalongsidethedinosaursthroughoutmostoftheMesozoic(ca220
65Ma)[1].Unliketheirlivingfunctionalcounterparts,birdsandbats,pterosaurspossessedaflexible,
membranouswingthatwassupportedbyasingle,superelongatewingfinger[13].Asfarasisknown,no
otherflyingvertebrateshaveeveradoptedthisgrossmorphologyandtherearenodirectanaloguesin
mechanicalaerodynamics,theclosestbeingthemainsailofasailboat[4].

Althoughweknowfromfossilswithpreservedsofttissuesthatthepterosaurwingmembranewasthin(most
likelyofvaryingelasticity),withinternalreinforcingfibresperhapssurfacefibres(pycnofibres)[3]and
possiblypneumatizedintheregionsclosesttothewingbones[5],itremainsdifficulttodeterminethenature
ofthemembraneattachmenttothebonyskeleton.Itwasprobablyeithermidwaybetweenthedorsaland
ventralsidesofthewingbones[6],asinbats[7],orattachedentirelytotheirdorsalsurfaces[8].

Reconstructionsofpterosaurflightcapabilitieshavevariedhugelybecauseofstructuralandbiomechanical
unknowns(e.g.[917]).Mosthavebeenbasedonassumptionsofwingbonemorphologyfromfossil
evidence,combinedwithwingsectiondatadrawnfromthepre1950saerodynamicliterature,whichare
necessarilyunrepresentativeofactualpterosaurwingprofiles.Resultsfrommorerecenttheoreticaland
experimentalworkonlowspeedaerofoils[18]andsailboats[19,20]canimproveourunderstandingofthe
relevantairflowphenomena(seeelectronicsupplementarymaterial),butstilldonotprovideresultsthatcan
bedirectlyappliedtopossiblepterosaurwingsections.Onlytwospecificinvestigations[11,21]ofsuch
sectionshavebeenpublished,butbothcontainanomalousresults(seeelectronicsupplementarymaterial).
Consequently,itisnotpossibletoquantifytheeffectsofvaryingthelocationofthewingbonerelativetothe
liftingsurface,varyingthecamberofthewingsectionorofdifferentwingbonecrosssections.Inthe
absenceofsuchdataitisnotpossibletopredicttheoverallflightperformancewithanycertainty.Toaddress
thisdeficit,twodimensionalmodelsofarangeofwingsectionsweremadeandtestedinalowspeedwind
tunnelandtheresultsusedtoproducecomparativeflightperformancecurvesforlarge,generic
ornithocheiridomorph(Pteranodontidae,Istiodactylidae,Anhangueridae,etc.)pterosaurs.Thesourcesof
informationaredescribedinmoredetailintheelectronicsupplementarymaterial,butthemodelsweremade
intentionallygenericsothattheresultswouldbewidelyapplicableandnotspeciesspecific.

2.Methods

(a)Modeltests

Twodimensionalrigidandflexiblemodelsofwingsectionsrepresentativeofdifferentcrosssectionlocations
(figure1)alongthewingofageneric,5.8mwingspanderivedpterosaurwereconstructedandtestedina

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windtunnelatappropriatevaluesofReynoldsnumber(Reseeelectronicsupplementarymaterial).Therigid
modelsweremadefromthin,curvedsheetsofepoxyresin/carbonfibrecomposite(figure2).Sincethe
actualcamberofthepterosaurwingmembranescannotbeknownwithanyprecision,modelsweremade
withthreedifferentcambervaluestoprovidearangeofresults.Pterosaurwingscomprisedtwodistinct
regionstheproximalregionwhereapropatagiumwaspresentandthedistalregionwhereitwasabsent
[1,2].Thedistalregionofthemainwingmembranewassupportedbywingbonessituatedalongtheanterior
margin,whereasintheproximalregionthewingboneslaywithinthemarginsofthemembrane(figure1).
Thelocalwidthofthepropatagiumdependsupontheassumedangleoftheelbowjoint,andorientationand
pointoflocationofthepteroid.Theproximalregionwasthereforemodelledwiththewingbone(ulna)onthe
ventralsideat20and40percentofthewingchordfromtheanteriormargin,torepresentdifferent
propatagiumwidthreconstructions.Fairedandunfairedgeometrieswereusedtomodelpossiblemuscle
tissue.Thefirstwingphalanx(WP1)wasmodelledwithovalsectionsoftwodifferentsizes,withandwithout
asofttissuefairingonitsposteriorface[8].Thesecondwingphalanx(WP2)modelsweresubtriangularin
section.Onewasfairedextensively.

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Abstract
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1.Introduction

2.Methods
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3.Results

Figure1. 4.Discussion

Generalizedshapeofthewingofalargeornithocheiridpterosaur.Redrawnfromrecentreconstructions
Acknowledgements
[16,17,46],andshowingthelocationsofthewingsectionsthatweretestedandtheextentofthewing
References
membraneassumed(crosshatchedarea).
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Figure2.
Wingsectionsthatweretested.(a)Highcamber(14.6%)proximalregionwithulnafairedandunfaired,
Article
20%and40%ofchordfromanteriormargin.(b)Mediumcamber(11.5%)sectionrepresentingcentral
regionofthewingfittedwithWP1andWP2phalanxsectionsoftwodifferentsizes(nominallywithdepth
Abstract
of7.5%and10%ofsectionchord)fittedonventralsideofthewingsection.TheWP1sectionwasalso
testedonthedorsalsideandwithasmallaerodynamicfairing[8].(c)Lowcambersection(8.5%)fitted
1.Introduction
withWP1andWP2sections.WP1sectionfairedaccordingtoPadian&Rayner[8]andWP2section
2.Methods
fairedtotheleastextentrequiredtominimizeseparation,aspredictedfromXFOILanalysis[47].(d)
Flexiblesectionwiththreedifferentdegreesofslackness(intheunloadedconditiontheactualcamber
3.Results
increasedwithaerodynamicload).
4.Discussion

Acknowledgements
Themembraneoftheflexiblemodelwasmadefromlatexrubber,reinforcedintheproximodistaldirection
References
withthincottonfibrestorepresenttheanisotropicpropertiesofthemembraneconferredbyinternal
Figures&Data
reinforcingfibres[3,22].Themembraneelasticitywasscaledtobeequivalentto2mmthickskinwiththe
Info&Metrics
samepropertiesasbatwings[23].Theposterioredgeofthemembranecouldbemovedinthe

eLetters
anteroposteriordirectiontochangetheslacknessinthemembrane,andthusitscamber.Threelocations
weretested,giving0,9.5and18percentcamberwhenslack.Alltheresultswerecorrectedforwindtunnel
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blockageeffects[24].

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Tovalidatethetests,thecamberedplatemodelresultswerecomparedwithresultspublishedbyMilgram
[25],theonlycomprehensivedataavailableforthincamberedplateaerofoilsatlowRe(<500000)overa
rangeofcamberratios.Theoverallshapeandtrendsoftheresultswereverysimilar,givingconfidencethat
theresultsbeingreportedherearereliable.Detaileddifferencesareprobablyowingtodifferencesbetween
thesectionshapesandcharacteristicsoftheairflowinthewindtunnels,wellknownproblemswhenresults
arecomparedbetweenfacilities[26,27].Thesedifferencesdonotinvalidatedetailedcomparisonsbetween
theresultsobtainedinthesametestfacilityormoregeneralcomparisonsacrossfacilities.

(b)Analysisofresults

Resultswereanalysedbyreferencetotherelationshipsbetweenclandcd(sectionliftcoefficientanddrag
coefficient).Onetypicalsetofresultsisshowninfigure3.Inordertocompareandquantifytheeffectsof
thedifferentwingsectioncharacteristics,thelift:dragrelationshipswereusedtocalculatetheflight
performanceofanotionalthreedimensionalpterosaurwithawingareaof2.2m2,wingspanof5.8mand
massrangingbetween13.9and32kg[10,12,28,29],usingthestandardtechniquesofaircraftdesign[30].
Thedragwascalculatedasthesumoftheprofiledragmeasuredinthetwodimensionaltests,theparasitic
dragofthebodyandtheinduceddrag(dragowingtolift).Theresultingliftanddragvalueswerethenused
tocalculateaglidepolarcurve(thevariationofsinkspeedwithforwardspeed[31,32]).

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Article
Abstract

1.Introduction

2.Methods

3.Results

4.Discussion

Acknowledgements
Downloadfigure
References

Figures&Data Openinnewtab

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eLetters
Figure3. PDF
TypicalwindtunneltestresultsatRe=200000.(a)Variationofsectionliftcoefficient(cl)withangleof
attackforWP2wingbonesection.Opencircles:resultswithincreasingangleofattack.Filledcircles:
resultsfordecreasingangleofattack.Theclosenessoftheseresultsdemonstratesgoodtest
repeatabilityandabsenceofhysteresiseffects.(b)EffectofsectionfairingonclcdfortheWP2wing

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section.Theextensivefairingsignificantlyreducestheminimumdragcoefficientbuthasnoeffecton
maximumliftcoefficient.

TheparasiticdragofthebodywascalculatedusingthemethodologyofBramwell&Whitfield[10]inorder
thatcomparisonscouldbemadewiththeirwork.Theinduceddragwascalculatedusingthestandard
aerodynamicformulation[30]: ,whereeisaconstantdependentuponthegeometryofthe
wingplanform,ARtheaspectratioofthewingsandCLthewingliftcoefficient.Avalueofe=0.9wasused,
applicabletoahighlytaperedwing[4].ARwascalculatedfromAR=B2/S(whereB=thetotalwingspanand
Sthewingarea).

Thepolarcurvewascalculatedusingtheidentity ,whereCTistheresultantofCLand
CD(thewingliftanddragcoefficients),Wtheweightoftheanimal,themassdensityofairandVathe
airspeed.Thisissolvedtogivetheairspeedvector,fromwhichthehorizontalandverticalspeed
componentscanbederived.

TheresultantcurveisaninvertedUshape,andthemaximumofthecurveisthepointofminimumsink.The
pointwhereitistangentialtoalinethroughtheaxisisthemaximumaerodynamicefficiency(L/Dmax)and
alsothemaximumrangeinstillair.Itisimportanttobeawareofthisdistinction.Atminimumsink,timeinthe
airismaximized,whereasatL/Dmaxtherangeismaximized.Theformermattersmoreforbehaviourthat
reliesonsoaringinrisingair[31].Minimumsinkcanbeimprovedintwoways:byincreasingtheL/Dmax
and/orbyreducingtheflightspeed,asmovingtowardsthetopleftofthegraphimprovesthesinkrate
withoutanincreaseinaerodynamicefficiency.Indeed,itispossibletoimproveminimumsinkevenasL/Dmax
reduces.

Oneotherregionofthepolarcurvedeservesattentionthebottomleftofthegraph.Thisiswherespeed Close
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becomesverylowandflightcanonlybesustainedbyachievinghighvaluesofliftcoefficient.Aprecipitous
dropinthepolarcurvereflectsasuddenstallandprobablelossofcontrol.Amoreroundedshapereflectsa
Article
moregradualandcontrolledtransitionfromflighttostall.
Abstract

1.Introduction

3.Results 2.Methods

3.Results
Theulnapositionedat40percentofwingchord(fairedorunfaired)reducedmaximumliftbutdidnot
4.Discussion
increaseminimumdrag,resultinginonlyasmallreductioninaerodynamicefficiency.Amoreanterior(20%
Acknowledgements
chord)locationforthiswingboneresultedinagreaterreductioninperformance,butinbothcasestheeffect
References
ofthefairingwasverysmall.
Figures&Data
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TestswiththeWP1andWP2phalangesgavethefollowingresults.(i)Withthesectionsontheventralside,
dragincreasedsubstantially(whencomparedwithacamberedsectionalone),withlittledifferencebetween
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thetwoshapes.Therewasalmostnoeffectonthemaximumlift(infactitincreasedalittlewiththeWP2
PDF
phalanx).(ii)Aphalanxlocatedonthedorsalsideofthewingreducedthesectionperformancesubstantially,
increasingdraganddecreasingthemaximumliftcoefficient.(iii)Thelargerthebonesectionrelativetothe
widthofthewingsection,thegreaterthedrag.(iv)Fairingthephalanxsectionstotheextentpreviously

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suggested[8]didnotreducethedragorinfluencethemaximumlift.(v)Amoreextensivefairing,designedto
minimizeseparation,reducedthedragby35percentwithnochangeinmaximumlift.(vi)Theflexible
sectionhadsimilarminimumdragbutgreatlyincreasedmaximumlift(>25%)(atthecostofhighdrag).(vii)
Themaximumliftcoefficientsproducedbytherigidsectionsapproached2.0andwerestillhigherforthe
flexiblesection,valuesconsiderablyinexcessofthosereportedforbirds[32,33].

4.Discussion

Pterosaurwingsectionperformanceissensitivetowingbonelocationandsizebecausethebonestrigger
flowseparation(whichcreatesdrag)butthiseffectisreducedwhentheyarepositionedposteriortothewing
margin(ontheventralside).Thehumerusandulnaarethelargestdiameterbonesinthepterosaurwing
andwouldhavebeensurroundedbysofttissue,furtherincreasingtheirsizesoproximalregionsofthewing
wouldhavepotentiallysufferedsubstantiallossofperformanceowingtothesebones.However,thiseffect
wasmitigatedbythepropatagium,whichpositionedthebonesposteriortotheanteriormarginofthe
membrane,andsoactedasadragreductiondevice(contra[21]).Suchathinleadingedgesectionisvery
sensitivetothelocalangleofincidentflow[34],whichmightexplaintheroleofthepterosaurpteroidtovary
thelocalincidenceangleofthepropatagiumastheoverallangleofattackofthewingchanged,andthus
maintaintheoptimumflowconditions.

Whenaphalanxwaspositionedontheanterodorsalwingmargin,theperformancewashighlydegraded.
Similarresultshavebeenseeninsail/masttests[20,35,36],togetherprovidingstrongsupportforventral
wingfingerpositioning[8].

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Theglidepolarsconstructedfromthesewingsectionresultsshowtheinfluencesontheoverallflight Close

performance(figure4).Flightefficiencyissignificantlyinferiortopreviousestimates[10,12],andsomewhat
lowerthansomeextantsoaringbirds[3740].However,owingtothelowflightspeed,theminimumsinkrate
Article
(approx.1.0ms1)wascomparabletoextantbirds[32,3740]andbats[41].Aswingbonesizeincreases
Abstract
(relativetothewingchord),thesinkrateincreaseswithlittleeffectuponflightspeed.
1.Introduction

2.Methods

3.Results

4.Discussion

Acknowledgements

References

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Figure4.
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Polarglidecurves.(a)Comparisonwithresultsfrompreviousstudies((i)[10](ii)[12]),withresultsfrom Close
presenttests(WP1mediumcambersection)attwoextremesofmass((iii)and(iv)).TheWP1winghas
onlyhalftheaerodynamicefficiency(L/Dratio)andalmosttwicethesinkrateoftheearlierestimates.(b)
Effectofdifferentwingbonesizesandshapes:(i)sectionwithnobone(ii)smallWP1(iii)largeWP1(iv)
Article
smallWP2and(v)largeWP2.TheWP1phalangesmovethepolarcurvedownwardswithincreasing
size,resultinginlittlechangeintheoptimumflightspeedbutalargeincreaseinthesinkrate.TheWP2
Abstract
phalangeshavesimilareffects,butbecausetheyincreasebothdragandthemaximumliftcoefficient,the
1.Introduction
glidepolarmovesalittletotheleft,extendingthelowspeedflightenvelope.(c)Potentialperformanceof
anoptimizedwingsection:(i)417asectionusedinearlierreconstructions(ii)optimizedmodernS1223
2.Methods
laminarflowaerofoilsection[48](iii)WP2fairedwithXFOILdesignedfairingand(iv)smallWP2only.
3.Results
Thetwoaerofoilsectionsgivesimilarpeakefficienciesof20:1,buttheS1223sectionmaintainsgood
performancetohighervaluesofliftcoefficient,shiftingthepolarcurvetotheleft.(d)Effectofflexibility:(i)
4.Discussion
envelopeofrigidsectionswithsamebonedepthasflexiblesection,and(ii)(iv)theflexiblesectionwith
increasingmembraneslackness.
Acknowledgements

References
Theflightperformancewasimprovedbyextensivefairingofthewingbones,and,whilethepresenceofsuch
Figures&Data
afairingisentirelyspeculative,itmayhavebeenprovidedbypneumatizedtissue[5].Thefairingincreased
Info&Metrics
theaerodynamicefficiencyandflightspeed,buthadonlyalimitedeffectupontheminimumsinkspeed,the
eLetters
parameterthatdeterminestheloiteringandthermal/slopesoaringcapability.
PDF
Withaflexiblemembrane,theflightenvelopewasextendedtolowerspeedsowingtotheenhancedhighlift
capabilityandprogressivestallofthesesections.Sincetheanimalspresumablyhadsomecontroloverthe
wingcamber[3,22],theenvelopecurvearoundtheresultswiththeflexiblemembranebestshowsthefull

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rangeofperformance.Whencomparedwiththerigidwingsectionresults,thelowspeedflightcapabilityis
extendedandcombinedwithasofterstall,whichwouldhaveenhancedcontrolduringlandingmanoeuvres
whenlowspeed,highdragandhighliftarerequired.

Inconclusion,thesetestshavequantifiedthetwodimensionalcharacteristicsofpossiblepterosaurwing
sectionsforthefirsttime.Theyshowthatthesecreaturesweresignificantlylessaerodynamicallyefficient
andmorecapableofflyingatlowerspeedsthanpreviouslyestimated.Inordertoachievehigherefficiency,
thewingbonesmustbefaired,whicheitherimpliesasubstantialweightpenaltyowingtotheadditional
tissuerequiredor,morelikely,ahighdegreeofpneumaticityinthefairingtissue.

Whetherfairedornot,thepterosaurwingsectionswereadaptedtoalowspeedflightregimethatminimizes
thesinkrate.Thisregimeisunsuitedtomarinestyledynamicsoaringadoptedbyprocellariiformbirds,which
requireshighflightspeedcoupledwithhighaerodynamicefficiency[42,43],butiswellsuitedto
thermal/slopesoaring.Thelowsinkratewouldhaveallowedpterosaurstousetherelativelyweakthermallift
foundoverthesea[44,45].Sincethebonesofpterosaurswerethinwalledandconsequentlyvery
susceptibletoimpactdamage,thelowspeedlandingcapabilitywouldhavemadeanimportantcontribution
toavoidinginjury,andsohelpedtoenablepterosaurstoattainmuchlargersizesthanextantbirds.Indeed,
themaximumliftcapabilityofapterosaurwingwassubstantiallyhigherthaninbirds,enablingslowflightand
mitigatingtheallometricconstraintonsizethatmanifestsinthefastlandingspeedsoflargeextantbirds.The
tradeoffforpterosaurswouldhavebeenextremevulnerabilitytostrongwindsandturbulence,notunlike
moderndayparagliders.

Acknowledgements
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IamgratefultoG.Dyke,M.Benton,E.Rayfield,P.Gillandtwoanonymousreviewersfortheirhelpful
commentsonearlierdraftsofthismanuscript,andtoMalachyO'RourkefortheuseoftheUCDSchoolof
Article
Electrical,ElectronicandMechanicalEngineeringwindtunnel.
Abstract

ReceivedOctober8,2010. 1.Introduction
2.Methods
AcceptedNovember3,2010.
3.Results
ThisJournalis2010TheRoyalSociety
4.Discussion

Acknowledgements

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