CHAPTER 6 ANIMAL FLIGHT P r e p a r e d f o r t h e C o u r s e T e a m b y D a v i d R o b i n s o n

6.1 Introduction

A U f i v e c ~ o f ~ ~ ~ t h t m ~ ~ ~ l a a r e d t h e s e , ~ i s ~ t o g l i d i q g ~ ~ p o a r a a d ~ Z n a d d i t i o n t o t h e living vmkbmk~, them are tBe flying d tbe W p r h 3 (24S2og d o n years k&m (BP)), tbt J& ptr ior l (SI46 millioP BP) and the Crcmcm million y t m BP), minds which, in t e t m s o f s i z e , h a v e m , ~ ~ . P h 6 . 1 sbowsaddondpesent - d a y @ ~ v e a e b r a t e s d b 6 2 ~ m M s i m p r t ~ i o n o f t b e ~ k r a o w n f o s s i l ~ , ~ ~ whkhmearwrcdsome 12m fromwingtiptowingtip.

As you h o w from tbe previous d q t m , any mimd that flies requires a o f d l e m c a t h a t c a n a c t a s anamfoil. Fwgliding, thwe mannmberof opcionsavadable, suchasanexpamkmofthe feetor of the thinmenhum m t i n g the limbs a d body. Aerofd mryst be ~~ to rwme extent, a d tbe involvement o f t h e ~ i n g l i d i m g d ~ e x e r t s ~ ~ w n t r o l . T h e 0 y i n g ~ d o w n o t & i t i a 9 0 ~ . .

of-wwitl ~ U U E qpm later (Sectioa 6.2.3).

The power required for I%@ is r rmja limiting h far dl nnimsls. Tk dividing line between having ar llot bnvigg orrfffcitne power ouqRlt from the m d e s t o f l y , i s a f t e n a f i n e ~ . F a r ~ , t h e t l i g e t d o f M canootpduwsuffircisntpawaudthewimnlisdupinkhg. Z f y o u e n c o u n t e r l ~ ~ y i n ~ m o a n i n g t b e y ~ a M e t o P l m p a w a y f r o m y o u a n d s t a r t ~ f l y , b n t ~ ~ ~ b o ~ ~ a o d ~ ~ o r ~ k k o n t o t h e ~ m w e u ~ ' f d m ~ ~ f l y b e c a u s e t h e p o w ~ ~ o f t h e ~ i e ~ t o ~ t b e ~ o f f t b e g r o u n d i s t r u e a t r r w > m ~ b t t t k P c r i v i t y o f t b e ~ E e s b e f ~ E a l m o f f w a r m s t h e m . A t a h i g h e r ~ t b e ~ ~ a ~ p ~ w e r wtputandso,owxwanned-up,tftebumblab#m~dstill'obey'tbela~ of physics.

T h e ~ ~ ~ ~ f w ~ a u d ~ . ' h flight i s o b v & l y o f ~ ~ t o ~ . F l i g h t i s ~ e x p e n s i v e i n a n e s g y , d ~ t h r t ~ t b e ~ o f ~ h a v t a n e v o l u t i ~ ~ . A t t x m w t i m e i n t f i t p s t , t f i c ~ a f ~ l b s d f h a d t o ~ t s e e a e r ~ y ~ f a I h s ~ t o f l y b g t o b n v e e v o l v e d a n d k n ~ C P l a h P t i e g a e o t a l ~ ~ f o a a ~ a n i m a r i s f a r f r r a n ~ ~ s y ~ h ~ a ~ o f ~ ~ a e a d t o b e ~ U n d e r ~ ~ ~ ~ ~ u c b & s ~ ~ ~ o f ~ i o e l r d

5324 Peek 4 Animal Pcrfmrrnancc

6.2 Gliding and parachuting

Cha)ter 4 Animal I

6.2. I Frogs

I t i s ~ c l ~ w h y ~ g l i d e a ~ h u t e . ~ u e a ~ o % ~ s ~ W h a v e w e l l & ~ ~ b e e n t b t ~ m t b e i r ~ , f o r example Rkacuphm m i w a d i asd -his s p m d 6.la d 6.3, respectively), w h i c h s u g g e s t s t h a t t h c ~ ~ i n ~ . T b c m o s t likelysekction~cmdfircrgnispdWh,bHtfaarhdhgstbe a W t y t o g l i d e b e t w e e s l ~ o f t r a s s m a y ~ P d ~ i n t h e ~ forfoodand~~Howevw,inolltspacSea,~io~crrty~ofthe t w e n t i e t h c e n t u r y , t k t d u m w h ~ o f ~ ~ w e t e f a u n d t o i n c l ~ r e ~ o f ~ ~ & ~ ~ ~ y c a t m ~ a a d ~ ~ f i w m d on the ma. It remaim a pddity, however, that for somc p i e s , gfiding is @ y m ~ t o f d i n g .

I n ~ t i c m t o t h c ~ t h s e g h & , ~ ~ ~ & a t ~ ~ t e a i r ~ a f descent i n a i r , h t ~ a c h i e v t a ~ w g I i & e m m d y cute.

6.2.2 Fish

m y i n g f i s h m o n e o f t b e ~ ~ f o r o c t a n t t s t v ~ I l a s i a t b e ~ ~ . Ttsefi&lqmtofthewatcrdglidtovcstbs*~yEorqHi8e longdistaraces.Flyingfishoocurtbmughouttb~dtWewtselrerrl s p e c i e s o f s u c h f i & i n o n e f a m i l y 4 ~ . ~ ~ ~ i s b t u s , ajishof about 2 0 c m ~ ~ ~ ~ i s sbowek P l a t e 6 . l b . T b e p e c t a r a l h , ~ ~ ~ C Z W d 0 0 h ~ d & , m g r e a t l y ~ f w m i n g ~ ~ b o d y i s ~ d t h e ~ t o b c o f t b e ~ o r d h i s ~ A ~ d f o r m o f 9 . i n g f i l i s e x e m p l i f i o d ~ t b c g c n u s C y ~ h n * ~ , w ~ W t h e p & d v l l d ~ ~ a r e ~ ~ ~

' w i n g s a a d t b e l o w e r l o b e o f t h e c a r d a l 6 a ~ ~ ~ ~ 6 ~ ) . T b t t a o typesof fisharcreferradbas t k m o n o p h r r e ( ~ ~ ~ a p d h i @ m w ~ w i n g s ) t y p , ~ v e l y . l k e a r 8 ~ ~ ~ ~ ~ m ~ n ~ a r m a g e m e n t o f ~ ~ ~ w h i l i e t h e ~ ~ ~ 5 m a x i m i z e s m T b e b i p b n e ~ b * v e ~ ~ n t i o ~ 9ad a lower wing loading for + v d ~ ~ & body mass, gMkg Iift at 1- speeds (Chapter 5).

k 4 Animal Perfarmancc

W h e n ~ ~ , t h t ~ c y p l w u a ~ w i a a l g e ~ l o r n ~ a l o b t u f ~ ~ 6 u i n ~ w h h t h t w g D g ; ~ ~ ~ ~ t b e ~ p e d f r o m l O r n ~ - ~ t o 1 & 2 0 m s - l . k h ~ d t 8 t f i l i s & w o f t b e ~ t h e p e c b o r P l f i n s o p e n w i t h a a r m l l p & h q $ e d ~ garerPtine~Tbe~htbeaopen,Zrfflae.thetailootd~u~.Tbe average kn@d the taxi !Am WwITis 9-

rds ad snakes

5324 Leek 4 Animal Petfmrmance

6.24 Mammalr

' Chapter 6 Animal Flight

5324 B0.k 4 Animal terformance

If~wing~ishigb,theglidtisEsst,so~t8csrir0mwatbcwing geeeaatessuf6cientl i f fTbegh&pathdtbe~issepto~withandit l-h@tatarateof6ms-'.AsspeedWup,k~WW shallower. Tbt b a l g r a d qwd is orormd 15 ms-I mgurc 6.5).

~ b e r e a r e ~ q m a i m u o f g e m r a l i n t e r e s t t b a t ~ ~ a t ~ gliding nwmds:

Chapter 4 Animal Flight

A l l f i v e v ~ ~ ~ ~ ~ m e m b e E s t h o t ~ ~ , ~ p o t R e r a d ~ i s ~ e ~ c t e d t o ~ ~ a s d m e ~ u f ~ h h , a s d d y m a n b g . s o f t h e f i r s t t W ~ ~ a m a @ k o f ~ ~ ~ ~ ~ t f r e 4

r e p C i l e s a n d ~ ~ i s a r r l n g c o f ~ t o ~ d O f ~ r r t c o ~ w i h ~ f h t s u r f a c e a r e a o n w h i c h r i r c a a ~ T b e ~ ~ ~ d ~ i ~ a a t h e s n g k o f ~ t o ~ ~ g f i d t + I f ~ i s g r e a t e r ~ 4 5 " , t b e ~ i s n o a a r r l l y c m s i d m d t o b e ~ l l g d ~ t o k v e J W g ~ ~ e c , ~ a distinction is a rather dAmy cmc.

6.3 Flight in bats Bats are the only mammals capable of hgpmg flight. 'Zbey are very diverse, with over 1 000 recophi @a, and thus almost onequarter of mammal s p e c i e s ~ ~ 1 ~ a r e ~ . ~ y r a a g c i a ~ ~ 2 g ~ 1 ~ ( ~ b o d y m a s r s ) a n d r n o s t f e e d o n ~ , a l ~ t b e r e a r e a n u m b e r o f ~ ~ W f e a d ~ ~ T h P . t ~ ~ s o m t ~ ~ b a t s t h a e ~ o n n e c t a r c a r pollen, fib, h g s or " M o o d

5324 Imok 4 Animal Perfermance

T b t ~ o f a b n t a r i n g i s s h o m i n F i g m e 6 . 8 . T b e b o c l e ~ i s a modifiedformofthempmnuli.nfadimb strumme. Tbamownmtintbe wrist a n d e l b o w ~ ~ ~ t o t b e p ~ o f t h e w i n g s o t h a t t b e w i n g ~ w t twistin~The~bomhverysmallincomp&mwiLthe~us~qurc & 9 c ~ p r e v ~ t h t ~ m w ~ t h a t i p s i b b i n w r o w n w r i s t joints.Thtwing~,ofcoursc,hastobtkqtundeatensionduringflight ortheairflowmitwOuldMbe~dliftwwldbeMFigure6.8 ~ m s ~ t h e b d t e n S i o n ( b l u t a m r w s ) d ~ w ~ r r m w s ) i n tbewingkeepittaut.misadinthc-Wmoacotbe d d i g i S p l l l i n g i t f ( ~ ( n d d a w a y f T o m t h e W ~ ( ~ b y t b e long, thin b k k arrow). Tht force c x m d by t i i s muscle, the extensor carpi radialis longus, keeps the third digit extend4 as the force the m d c exerts on t h e ~ d i g i t i s ~ t o t h e t h i r d b y a l i ~ T h e ~ p l l l s t o w ~ t h c b o d y , w ~ e x e t t P a ~ w h ~ ~ i n ~ i t i m t o t h a t pllling thc d d third digits f w d , maintaining Won in the wing.

FIgnm6.8 ThtrigMwingofaWsgreadasin~Tbeblaeaaomsbowkfo~cws o f ~ o n i n t b e ~ d t h e ~ ~ t h e f a r c o s o f ~ m i n ~ b a n a s . T h c t h a c k ~ ~ s b w v t b t ~ i n w h i c h p b e f ~ m w c k d t b e ~ exert a forrx w th wing.

Chapter 6 Animal Flight

In flight, the dciwnmk is tbe pow- ns ht& majority of birds (Section 5.6).Duringtbe~,tbwhgDlcmbrotwi~~wr.isdic*tiag&at 1 i f i i s ~ ~ ~ P r i n g w a b o ~ a ~ w ~ d ~ t o t h e a i r f l o w ~ ~ P p s t r o k e , w b i e h ~ l i f f . ~ ~ i a ~ m l y ~ l i f t , ~ - "birdsdWcormbting~cxccpt ioas. ~ e s P e B p r m i c ~ b e t a t e e n b j r d s d ~ a r c ~ p p s ~ l ~ t f m n a c o m p a a i s o n e t f ~ t b s i r a p c t a t i o a n d ~ ~ ~ ~ & l U ) .

. 4 Animai Performance

~ y o u l ~ e o f l p r i t s m l i r & w U @ s ~ y m ~ f a the following bats: (a) P l c c o t r r s h , a - slow--bat; a d ( b ) N y c t a k r s - , ~ - ~ ~ ~ ~

Chapter 6 Animal Flight

Summary of Section 6.3

6.4 Flight muscles ~ ~ m u s c l e s ~ ~ ( b e ~ ~ ~ ~ t b e p c c t o r a l i s m l m a n d ~ ~ m w c h ( S m h 5 . 6 ) . h ~ ~ ~ itisthepectoralis~lesthrtdomastdtbe~h~of~,the pctmab nmcbare, mavlerage, 15-mof bodymass,lxrtthc s ~ s ~ l e s ~ m a c h s ~ ~ ~ ~ l 3 4 6 d t h e m a s s o f the pctmab &w.

I n s w r r e b i r d s t h e d i & r e n c e i n * ~ i k ~ ~ i r ~ ~ , w i ~ t h e s ~ & u s ~ ~ , a r o u n d ~ d ~ ~ o f ~ ~ muscle. Can y w suggest what habits such a blrd might have?

Birds that hover p o w oe W~I tbe downswoke and the up&, with b o t h s ~ d e v c l o p i n g ~ l i f t r ~ q p t t h e h i y ~ i n h ~ ~ . ~ s u ~ ~ l e i s ~ h r g e h ~ , w k i & i a related to lkir ability to swim under water.

T h e ~ t m u s c l e o f ~ c o n t a i n s a v a r i e t y o f ~ ~ , a s i n ~ ( S c c t i o n 2 . 2 ) a n d r r l . m f l l e l n . B & M a n d ~ t e h = ~ b u t t f i e ~ d i f k between specim, with the Mumm being related to WstyIc. T h e d o n m t k c h i c h r a d y ~ c a a n y p a t d i s c a n e t a t d t h i s ~ ~ d w i t h t h e ~ m o f t h e f i l x s ~ i n t t w : ~ ~ .

Whichdoyouthinkitbe~~brcintfaepectwalismuscEeof chicken?

Qlicken breast meat is white, with wbite, East glycolytic muscle fik compri~~67%of~W.Ele~apgcentMttSe~~led,withtbe remailing 22% bizkg ng. In co- ia* hmmh@fdA&kht# t h e m u s c ~ s t m ~ ~ 1 1 ~ o f d 6 b r e s .

The flight I n ~ l e s ' a m obviously major * of- xtivity 4 quire substantial ammts of oxygen to be d e f i d b them. The bearc is larger in r b i r d t h a n i n a m P m m a l 8 f t b e s ; r m e & , t ~ ~ y t h e ~ ~ i s l o w e r O w e r

5324 Beak 4 Animal Performance

6.5 Flight energetics

. Chapter C Animal I

6.5.1 Bounding flih

6.5.2 Flying in groups

. Chapter 6 Animal

6.5.3 The cost af m s p r t

Theminimumaddtmqmtfacthb birdi0.24W1Eg1m-1 (atr fligb- of 11.7 m s-I ), which camp^ with 0.26 W kg1rn-' at 10 ms-I aad 025 W ~ l m - l s t ; ? O m s - l , a # , t b e ~ ~ ~ ~ - I P O C B ~ ~ a ~ Q f ~ .

w h a t ~ c a n y o r l * f a t l m h t t b s t t k l a g e r a a d b e a v i m ~ have a lower minbm cost of m m p t ?

5324 Book 4 Animal Ferfmrmanct

6.5.4 Energetic costs in gliding mammals

Summary of Seaion 6.5

G l i d i n g h a s e v d v e d ~ a e v e r a l ~ i n ~ d t h e F e ~ ~ ~ i a t b e ~ h a b i t f o r ~ t h o t ~ * ~ h t Ws have pFobably e v d d from g l j b g mmm~

. Chapter C Animal Flight

6.6 Conclusion A p a r t f r o m t h c b h d ? 3 r s d m , t h e r e ~ o r d j r a k w ~ 9 f v ~ t h a t c a n f l y , b p r t ~ ~ a m t h m f r l l f i v e ~ o f ~ . M o a t o f ~ o t h e a s ~ g k k a r ~ d a = ~ t q m b k o f p w m d % i g b t A ~

. . . k m a d e b e t w c e n ~ d p m c h u t i n g o e t k ~ o f ~ a n g l e a t w h i c h t h animaldmcds. I f t b t m g k b e t w ~ e n t h e & s c a N ~ s d t b t ~ i s always g m t u than 45*, d m he animal is desgibad s -uting-situply s l o w i n g ~ r p ~ e o f ~ A t ~ s d h t b v a 4 5 " , ~ P a r i m a l i s ~ d is obtaining some lift d m d tbe Ldy QW.

A l t b o u g h ~ p r e ~ y f l y i n s m k h t c s , k w o f t b e ~ s p e c i e s a n d ~ h a v e h n ~ ~ ~ F l i g h t ~ ~ 0 n ~ o c c a s I o n s i n ~ ~ d t k e h a s b e F n a w a p r i s i n g ~ o f c m v q e w e . T h i s i s p @ ~ , f m m a k ~ * o f * , t B e 3 u d m a f ~ b ~ b u t i t m i g h t h b e ~ ~ h ~ v 6 d t k r o u g h a ~ p h a s e a d t h e glidingphase has -con- km Wy a- o n t h e e v o l u t i o ~ t y p a t b t o ~ ~ m ~ ~ ~ 4 A n ~ o f t h e ~ ~ o f s p e & ~ g l i & ~ ~ t h e r e a r e ~ s i n g l i ~ m t h e r e i s a w ~ d q . I t m i g h t d b e t h p t h M w d v m ~ ~ b y ~ ~ o o u l d ~ e v e n t u a l l y l e d t o t h e e v h t i o n o f t h e b a t s f m m a ~ g ~ . Studitsof flightofmanyoftbeglidings~ have bm limited became s w h animals m relatively bcwsshk in fa& areas. Birds dbatstoopresentpmbkms, f or~manyof themotrbmuch b t # e r k n O W n t f r a n g l i d i a g ~ t b E e c t t h a t t b e y a m d ~ t h e m k s s ~ y t o w o r k w i t h . I t i s ~ d e v e l ~ o f m o d w n ' ' - tionandwind ~ t h p e ~ m a d e w o r k o a b d s d b a t s i n i l i g h t p s i M e . M o d a r n ~ s h a s ~ i t p s i b k f m p h y ~ ~ t s t o b e m a d e s o d ~ t u r a t r i d ~ ~ c m r y ~ ~ w i f h a ~ - i n ~ o -.This-caabeusedforflying-toO,batmabird o r b a t w i t h a n i n s ~ ~ ~ ~ o o t ~ 1 ~ . * ~ , a r s o o v t r l o a d i t ~ i t c p D m t t a l c t o f f , i s s t i n s r p r o b l e m t h s t h a s n o t b m d v e d . F e w t e l c " ~ m ~ i l y i n g m r i m a l R h a v e b e e a ambduut,butwhcnthey pre,astbeysraely will beintfatntarfutllre, we may getanwchbettermkstmhqofthctnasycostsofflying.

5324 Leek 4 Animal Perfmrmance

Objectives for Chapter 6

Questions for Chapter 6

(d) 'hhm7m Voiitam

(el Hykr nzi-