The Auk 113(4):802-810, 1996

INFLUENCEOF THE TRAILING-EDGE

NOTCH ON FLIGHT
PERFORMANCE OF GALLIFORMS

SERGEI V. DROVETSKI •
BurkeMuseumandDepartment
of Zoology,Universityof Washington,
Seattle,Washington
98195,USA

ABsTRACT.--Trailing-edge notches,formed by shortenedfirst secondaries,characterizethe

wings of most galliforms. I investigated the function of these notcheswith comparative
measurementsof notch size taken from extended-wing specimensand with experimental
studiesof modelwings of four representative species.Pheasants, quail, and turkeys,all of
which useflight to escapepredators,have wide wings and deep notches.Grousewith dark
flight muscleshave long, narrow wings with small trailing-edge notchesand typically fly
relatively long distancesfrom one foraging site to another. Grousewith light coloredflight
muscleshave short,broadwings with large trailing-edgenotchesand mostlyfly from ground
to canopyor from branchto branchto reachtheir food.Model wings of two pairsof galliforms
with different wing shapeswere usedin the experiments.White-tailedPtarmigan(Lagopus
leucurus) and SageGrouse(Centrocercus urophasianus) have small notches,high aspectratios,
relatively heavy wing loadings,low maximumlift coefficients,and dark pectoralmuscles.In
contrast,Wild Turkey (Meleagrisgallopavo) and California Quail (Callipeplacalifornica)have
deep notches,low aspectratios,relatively light wing loadings,high lift coefficients,and light
coloredpectoralmuscles.Experimentsusing model wings in a water flow tunnel show that
the trailing-edgenotch increasesthe maximumlift-to-drag ratio and stabilizesairflow around
the wing, but reducesthe maximumlift coefficient.Thus, the trailing-edgenotch increases
performancein vertical and slow flight but reducesefficiencyin level flight. Sucha function
is consistentwith the suiteof differencesthesebirds show in musclecolor,wing shape,and
predominantmode of flight. Received 17 November 1995,accepted27 February1996.

BIRD WINGSEXHIBITmorphological adapta- ed. However, only Shestakova(1971) hasnoted

tions for different kinds of flight (Rayner 1988,
Norberg 1989).Regardlessof the species,slots
between the primariesincreasethe lift-to-drag
ratio by increasinglift and reducing drag (es-
peciallyinduceddrag)that resultsfrom deflec-
tion of an air streamarounda tilted wing. Such
conclusions aboutthe functionalsignificanceof
wing slotscomeprimarily from the established
correlation between deep slotting and slow,
level flight that characterizes
birdswith wings
that are broad at the tip. Wind-tunnel experi-
ments supportthese conclusions(Vinogradov
1951; Hofton 1978; Tucker 1993, 1994).
Galliformshave deepslotsbetweentheir pri-
mariesand a large alula, but they rarely have
been usedas subjectsfor aerodynamicstudies
(Shtegman 1953, Shestakova1971). Further-
more, no study of the aerodynamicsof galli-
forms hasaddressedthe functional significance
of the large notch on the trailing edge of the
wing that is formedby a shortfirst secondary.
This feature is exhibited by all galliformsand
is quite evident when the wing is fully extend-
• E-mail: svd@u.washington.edu
802
the existenceof this notch in variousgalliforms
and some gulls.
In this paper, ! presentcomparativedata on
the size of the trailing-edge notchfor 31 species
of galliforms. I also present resultsof experi-
mentsthat showhow this notchinfluencesglid-
ing performancein the wings of four species
with different wing shapes.
MATERIALs AND METHODS
Measurementsof wing length, wing breadth, and
trailing-edgenotchsizewere taken from 220 extend-
ed-wing specimensof 31 galliform species(Table 1).
I calculated notch size (in %) as:
(WS - WSs•)/WB, (1)
where WBs•is the distancebetween the leading edge
of the wing and the tip of the first secondary,and
WBis wing breadthmeasuredfrom the leadingedge
of the wing to the tip of the fifth secondary.I chose
the fifth secondaryfor these measurementsbecause
in somebirds (e.g. turkeys)the wing notch is formed
by severalshortenedsecondaryfeathers.Wing length
is the distancefrom the proximal end of the humerus
to the tip of the longest primary, measuredalong a
line parallel to the leading edge of the wing.
Throughout this paper, meansare reported + I SD.
October
1996] FlightPerformance
andtheWingNotch 803

Four speciesof galliformswere chosenfor flight- TABLE1. Trailing-edge notch size and wing length-
performanceexperimentsbasedon extremesof wing to-breadth ratio of galliforms. Values are means
shape(see Fig. 1) and body size within the order: (with SD in parentheses).
White-tailed Ptarmigan ([Lagopus leucurus];Tetraoni-
nae;subadultmale specimen);SageGrouse([Centro- Wing
cercusurophasianus]; Tetraoninae; adult male); Wild Notch size length-to-
(% of wing breadth
Turkey ([Meleagris gallopavo];
Meleagridinae;adult fe-
Species n breadth) ratio
male); and California Quail ([Callipeplacalifornica];
Turnicidae
Odontophorinae;adult male).I refer to themas"mod-
el" species.Other speciesof galliformsprovide more Turnixsylvatica 3 12.36 (4.93) 1.77 (0.04)
extreme examplesof wing-shapevariation, but my Phasianinae
choiceswere limited by the availability of frozen Tetraogallus
himalayensis 1 18.14 1.81
specimensat the University of Washington Burke Alectorischukar 3 36.62 (3.54) 1.69 (0.01)
Museum for use in creating modelsof wings. Francolinussephaena 3 35.45 (1.60) 1.62 (0.02)
I measuredflight performanceby steady-state (con- F. africanus 4 24.98 (2.29) 1.86 (0.05)
stantvelocity) lift and drag characteristicsof various F. swainsonii 2 47.34 (5.64) 1.57 (0.01)
wings.Active flight in thesespeciesmay not be char- Perdixperdix 6 30.17 (2.34) 1.91 (0.01)
P. dauurica 1 36.36 1.78
acterized by this particular dynamic condition, but
Coturnix coturnix 3 21.57 (2.04) 2.25 (0.05)
any differencesin the coefficientsassociatedwith wing
Lophuraignita 1 35.71 1.58
lift and drag are presumedto apply to flapping, at
Gallusgallus 3 38.11 (2.67) 1.48 (0.07)
least for fast forward flight. Lift (C•) and drag (Cd) Phasianuscolchicus 13 30.86 (3.20) 1.71 (0.10)
coefficientsare defined by the following relation-
ships: Odontophorinae
Colinusvirginianus 15 32.49 (3.76) 1.61 (0.05)
C, = L/(0.5 p SU2), and (2) Callipeplasquamata 2 35.06 (3.46) 1.63 (0.07)
C. californica 9 31.84(3.76) 1.66 (0.05)
cd = D/(0.5 p STY2), (3) C. gambelii 2 37.37 (0.58) 1.62 (0.01)
where L is the measured lift force, D is the measured Meleagridinae
drag force,• is the densityof the fluid medium, S is Meleagrisgallopavo 4 31.40 (2.04) 1.47 (0.10)
the plainform area of the wing, and U is the fluid
Tetraoninae(light and intermediate
velocity relative to the wing. Lift and drag coefficients flight muscles)
were measured for model wings. Models were cut
from 0.2-mm thick brass foil and soldered to a metal
Dendragapus
falcipennis 2 26.50 (3.23) 1.95 (0.11)
D. canadensis 10 21.82 (4.29) 1.95 (0.09)
rod 2.5 mm in diameter and about 130 mm long. All D. obscurus 19 23.24 (3.31) 1.90 (0.07)
modelshad a width of 30 mm (length varied from 46 Bonasabonasia 14 24.52 (2.67) 1.94 (0.07)
to 62 mm depending on species).Each model was B. umbellus 22 28.94 (2.49) 1.88 (0.10)
slightly bent around the axis of the metal rod, and
Tetraoninae(dark flight muscles)
the spacebetween the leading end of the modelwing
Lagopus
lagopus 21 22.09 (1.89) 2.09 (0.05)
and the rod wasfilled with glue.The bendingcreated L. mutus 14 20.68 (3.08) 2.35 (0.08)
camberin the models.The top of the curvaturewas L. leucurus 8 19.60 (2.86) 2.12 (0.09)
10 mm behind the leadingedgeof a modelwing, and Tetrao tetrix 1 18.87 2.08
the height of the curvature,measuredas the distance T. urogallus 3 20.67 (1.29) 2.05 (0.16)
from a line connectingthe leading and trailing edges T. parvirostris 10 15.17 (2.52) 2.24 (0.11)
of a model wing to the top of the curvature, was Centrocercus urophasi-
approximately4 mm. The patternsusedfor preparing anus 10 18.06 (3.54) 2.24 (0.16)
the modelswere images(reducedby photocopy)of Tympanuchus
phasianel-
lus 4 19.25 (2.49) 2.16 (0.05)
freshlythawedwingsthat had beenpinned in a fully
T. cupido 7 20.76 (3.17) 2.05 (0.10)
extendedposition.
Lift and drag coefficientsare functionsof the Reyn-
olds number:
tunnel. The flow tunnel provided a uniform flow of
Re = UL/•, (4) relatively low turbulence intensity (ca. 5%). The
working section of the tunnel was square in cross
where U is definedas above,L is the length of the section,with sides of 0.2 m and a length of 1.2 m.
wing chord, and v is the kinematicviscosityof the The flow in the working sectionhad all four bound-
fluid mediumof the wing. Consequently,scalemod- aries fixed. During the measurements I placed each
elswerepreparedsothatin the testingapparatus (see model such that its center was in the center of the
below),eachmodelhada Reynoldsnumberrelatively working section. Apart from the model, there was
similar to that of real wings. only a piece of rod approximately7 cm long in the
Lift and drag forceswere measuredin a water flow flow. This rod connected the model to the force trans-
 804 Sm•G•
V.DItOVETSKI [Auk,
Vol.113

FIG.1. Extended-wing specimens

usedasmodelsfor the experiments.White-tailed
Ptarmigan(Lagopus
leucurus;
upperleft);SageGrouse
(Centrocercus
urophasianus;
upperright);c31ifornia
Quail(Callipepla
californica;
lowerleft);andWild Turkey(Meleagris
gallopwoo;
lowerright).Scaleundereachwingis in inches(upper)
and cm (lower).

ducer.Useof a waterflow tunnelprovideshigh res- ficient remains stable and constant across Re values
olutionof lift and dragforces,which,aslongasthe of 5 x lip to 5 x l0 s(Vogel 1983).
Reynoldsnumberis similar(seebelow),givesuseful Forces weremeasured separatelywitha strain-gauge
datafor lift anddragcoefficients.
Waterspeedin the transducer(sensitivity2 x 10-3 N) as one arm of a
flow tunnel was 0.5 m/s. Wheatstone bridge(seeLanyon1976).Theoutputwas
If modelsareperfectlyscaled,then the Reynolds amplifiedand collectedby a PC computer.For lift
numbersof themodelsandthe wingstheyrepresent measurements,the wing was mounted to the force
will be equal.Due to the limited size of the flow tank, transducer with its sensitive axis normal to the flow;
constantflow velocity,and the needto keepmodel for drag,the sensitiveaxiswas parallelto the flow.
geometrysimilarto the real wing geometry,models For all models,lift and dragforceswere measuredat
were madesuchthat the greatestbreadthof eachwas sixanglesof attack:0, 5, 10, 15, 20, and 25 degrees.
30 mm. Thus,the Reynoldsnumberof all modelswas At eachangleof attack,2,000samples weremeasured
the same, viz. Re = 1.5 x l0 s.This is somewhat lower over 10 s and averaged.To reduceerror, five mea-
thantheReynolds numbers fortherealwings(White- surements were made before water flow was started,
tailedPtarmigan,Re= 12.5 x lOS;SageGrouse,Re= five during the stableflow, and five after the flow
19.5 x lOS;Wild Turkey,Re= 33.5 x lOS;California was stopped.The mean of the 10 electricpotential
Quail, Re= 9.0 x l0 s)at a presumedflight speedof measurements (five measurements each before the
15 m/s, a valueavailablefor the Ring-neckedPheas- flow wasstartedand after it wasstopped)wascon-
ant (Phasianuscolchicus;
Rayner1985).Despitethe dif- sidered to be the "0 force" condition. This mean value
ferencesbetweenthesefigures,the modelsare still was subtracted from the mean of the five measure-
useful for the experimentsbecauseRe = 1.5 x 1OSis ments that were taken when flow was stable, and the
within theperformance rangeof realwings.Reynolds difference between these two means was then con-
number increasesproportionallywith flight speed vetted to a force.By measuringthe "0 force"condi-
betweentakeoffandfastforwardflight.Also,aslong tionbeforeandaftermeasuring thelift anddragforc-
as geometricsimilarity is maintained,the drag coef- es, errors in measurements were reduced. Measure-
October
1996] Flight
Performance
andtheWingNotch 805

48-
mentswerefirstmadeonwing modelswithoutnotch-
es. Then, notches were cut in each model, and the
measurement cycleswere repeated.The areaof each NS = 74.33 - 25.29 (WL / WB)

cut piecewascalculatedby weighing it (+ 0.001mg)

and using foil thicknessand specificgravity of the
metal.
I used differencesin pectoral muscle color as an
index of differencesin power output and resistance
to fatigue among galliforms.White muscleshave
higher poweroutputbut are lessresistantto fatigue.
Galliform muscles that are white or intermediate
colorare composedpredominatelyof glycogen-load-
in 24'

20
Wild
Turkey
-• _
ed white fibers(Norberg 1989).For example,glyco-
gen-loadedwhite fibersconstitute> 80%of all muscle
fibersin the pectoralmusclesof Ring-neckedPheas-
antsand SpruceGrouse(Dendragapus canadensis).
This
16
12 ß
-e
tailed
Ptarmigan
e•e use
•.4 1:6 •:s 2:0 2:2 2:4
similaritysuggests that the proportionof red and in-
Wing length-to-breadth
ratio
termediatefibers varies among galliformsby about
only 20%.Because galliformsarethoughttobe mono- F]c. 2. Relationshipbetweennotchsizeand wing
phyletic, all specieswithin the order are likely to length-to-breadth
ratiofor 31 galliforms.Circlesare
sharesimilar types of red and intermediatemuscle tetraonidswith dark pectoralmuscles,diamondsare
fibers.When musclefiber typesare the same,differ- tetraonidswith light pectoralmuscles,and squares
encesin musclecolormight correlatewith differences are other galliforms.
in the proportionsof thesefiber typesand, thus,dif-
ferences in muscle function that are related to the
ecologyof the respecitvespecies. that undergoeslong, strait flights. The Hima-
While preparing museum specimensI recorded layan Snowcockis characterizedby downhill
pectoralmusclecoloraccordingto a visualscale.This gliding insteadof activeflapping flight. Despite
scalehad three grades:dark or red (e.g.Lagopus),in-
theseexceptions,I found a strongnegativecor-
termediate(e.g.Dendragapus),and light or white (e.g. relation between notch size and the ratio of
Bonasa).
wing length to wing breadth (r = -0.76, n =
31, P < 0.001; Fig. 2). Galliforms with short,
RESULTS
wide wings have deeper notches.
DIFFERENCES AMONG GALLIFORMS IN WING The differencesin wing shapeand notch size
GEOMETRY AND NOTCH SIZE between grouseand other galliforms could be
related to differencesin the frequencyand du-
Wing narrowness (length-to-breadth ratio) ration of flights performed by the two groups.
and notch size vary widely among the 31 spe- Flight is a critical element of foraging behavior
cies studied (Table 1). Ptarmigan and grouse in grouseand ptarmigan, especiallyin winter,
(Tetraoninae)have narrower wings and smaller when they feed on buds and foliage in trees.
notchesthan othergalliforms.The rangeof wing To reach these foods,forest grousemust make
length-to-breadth ratios for grouse and ptar- verticalflightsto limbsandfrequentshortflights
miganis 1.88to 2.35,comparedwith 1.47to 1.91 from limb to limb, and ptarmigan mustfly from
in all other speciesexceptCommonQuail (Co- one foraging site to another. Other galliforms
turnixcoturnix),which is migratory and haslong, fly lessoften, usingflight primarily to flee from
narrow wings (length-to-breadthratio of 2.25). predators.
Notch size varies between 15.17% and 28.94%
of wing breadth in the tetraoninaeand between MORPHOLOGICAL FLIGHT PARAMETF,RS OF THE
24.98%and 47.34%in mostother galliforms ex- MODEL BIRDS
cept three species:Small Buttonquail (Turnix
sylvatica), Common Quail, and Himalayan Several morphological measurementsare
Snowcock(Tetraogallus himalayensis;
Table1).The usefulin comparativeanalysesof flight perfor-
systematicposition of buttonquailsis not clear. mance in birds (Rayner 1988, Norberg 1989).
Most authorstreat them asgalliforms,but some Among the most important are wing loading
place them with Gruiformes and others in the and aspectratio. Wing loading (in N/m 2) is de-
Turniciformes.The Common Quail is a migrant fined as:
806 SERGEI
V. DROVeSKI [Auk,Vol. 113

125-
TABLE2. Wing loading, aspectratio, and trailing-
edge notch sizefor the specimensusedto make the
model wings. 115- ßSage
Grou•
Wing
Notch
size
(% of
m•'
105-
z

• Wing
loading
=91.4
M
0.201
Species
loading Aspect wing
(N/m 2) ratio area)
• 85 / Wild
Tunkey
,
Lagopusleucurus
Centrocercusurophasianus
74.12
121.52
9.36
5.79
1.57
1.64
75
//y/White-tailed
85

55
Ptarmigan
/. California
Quail
Meleagrisgallopavo 87.19 4.23 4.11 o.o ,.0 2:0 d.5 3.0 ;.0
Callipeplacalifornica 59.43 3.67 2.16
Bodymass (kg)

FIG.3. Plot of wing loading vs. body massfor the

wing loading = Mg,/S, (5) four model speciesrelative to expectedrelationship
(curve)basedon an allometricequationcalculatedfor
where M is body massin kg, g, is acceleration galliforms by J. M. V. Raynet (pets. comm.).
due to gravity (i.e. 9.81 m/s2), and S is the total
wing area (the planar area of both wings and
that part of body between the leading and trail- the first type are red, whereasfibers of the sec-
ing edges of the wings in m2). Wing loading ond type are white. These types of fibers have
determinesimportant flight characteristics suchdifferent biochemistry that results in differ-
asspeedand maneuverability.Birdswith heavy ences in contraction frequency and speed as
wing loadingfly at high speedand havea large well as resistanceto fatigue. Red fibers have
turning radius. Aspectratio is defined as: lower frequencyand speedof contraction,but
AR = b:/S, (6)
much higher resistanceto fatigue than white
fibers (Norberg 1989). Therefore, muscleswith
where b is wingspan(in m), and S is total wing a high proportion of red fibers are suited for
area (in m2). Aspect ratio determines cost of sustained(horizontal) flight, but white muscles
transport,or the ratio of power to speed.Birds are adaptedfor short, powerful bursts(take off,
with a high aspectratio uselessenergy per unit verticalflight). Wild Turkey and CaliforniaQuail
of distanceflown (i.e. less power) to generate have muchlighter coloredpectoralmusclesthan
the samehorizontal speedcomparedwith birds do White-tailed Ptarmigan and SageGrouse.
with low aspectratio (Raynet 1988). Grouseand ptarmiganwith red pectoralmus-
The White-tailed Ptarmigan has the highest cles(i.e. Lagopus, Tetrao,Tympanuchus, and Cen-
aspectratioof the four modelsand a fairly heavy trocercus)have smaller trailing-edge notches
wing loading (Table 2). The aspectratio for Sage than thosewith white or intermediatepectoral
Grouseis lower, but it still is much higher than muscles(i.e. Dendragapus and Bonasa;œ= 19.46
that of the Wild Turkey and Common Quail. + 2.01%, n = 9 vs. • = 25.04 + 2.79%, n = 5,
Wing loading of the SageGrouseis exception- respectively;t = 4.32, df = 13, P < 0.001) and
ally high, whereasthe CaliforniaQuail hasthe wings with larger length-to-breadthratios(œ=
lightestwing loading.Wild Turkeyshavesome- 2.15 + 0.10 vs. œ = 1.92 + 0.03; t = 4.76, df =
what greater wing loading than White-tailed 13, P < 0.0005).
Ptarmigan.However, when size is taken into The four model speciesform two pairs, each
account,wing loading changesproportionally with different wing characteristics and pectoral
to M ø':ø•in galliforms (J. M. V. Raynet pets. musclecolors.White-tailed Ptarmiganand Sage
comm.).Thus,Wild Turkeysactuallyhave much Grouse have high wing loading, high aspect
lower wing loadingthan expectedby allometry ratio, and red pectoralmuscles,which servetheir
(Fig. 3). long, straightflights at high speeds.In contrast,
Another important morphologicalparameter Wild Turkey and CaliforniaQuail havelow wing
that canbe usedfor comparativestudyof close- loading, low aspectratio, and white pectoral
ly related speciesis the color of the pectoral muscles that are better suited for slower, more
muscles.In galliforms it may indicate relative powerful and maneuverableflights.Their short,
abundanceof fast-twitchoxidative-glycolyticfi- broad wings are better suited to frequent short
bers and fast-twitch glycolytic fibers. Fibers of (and vertical)flightsbecausethey generatemore
October
1996] Flight
Performance
andtheWing
Notch 807

T^BLœ3. Maximumlift coefficients of the modelwings,maximumlift-to-dragratios(L/D), anglesof attack

at which they occur,and increasein lift-to-dragratioswith wing notching.

Max. lift coefficient Max. L/D Angle of attack'

Species No notch Notched No notch Notched No notchNotchedincrease
b
Lagopusleucurus 0.69 0.61 3.96 5.98 5 5 33.8
Centrocercus
urophasianus 0.64 0.56 2.40 2.61 10 10 8.0
Meleagris
gallopavo 0.80 0.76 4.57 5.23 5 10 12.6
Callipepla
californica 0.70 0.65 4.81 5.12 5 10 6.1
At maximumlift-to-drag ratio (L/D).
Increasein maximumL/D with wing notching.

power on the upstroke, facilitating takeoff, were the same as those for lift measurements.
landing,and steepclimbsat low forwardspeed. In addition,notchingdecreased the CV for mea-
The wing notch sizes of White-tailed Ptar- surementsof drag to a different extent in dif-
migan and SageGrouseare similar (Table2). ferent species(White-tailed Ptarmigan,70.2%;
This suggeststhat such a structureis lessim- SageGrouse,25.1%;CaliforniaQuail,21.3%;and
portantfor galliformbirdswith relativelyhigh Wild Turkey, 0.1%).
aspectratiosand heavy wing loading. In con- The inverserelationshipsbetweenangle of
trast, the wing notch sizesof California Quail attack and the CV for lift and drag measure-
and Wild Turkeys are much larger. Thesecom- mentsmaybe relatedto the meansof the forces
parisonssuggestthat birdswith short,rounded increasingwith an increasein angle of attack.
wingsshouldderivemorebenefitfrom the wing
notch than do longer-winged species. 1.0- 1,0 ÷
While-tailed ptarmigan Sage Grouse
0.8
LIFT-TO-DRAG COEFFICIENT RATIOS IN WINGS nn

WITH AND WITHOUT NOTCHES

0.8-
• ø'8
0.8-
0.4

••2n
The trailing-edge notch increasedthe maxi- 0.4-
mal lift-to-drag ratio in all the model wings 0.2- 0.2
(Fig. 4). However, this increasewas small in
0.0- 0.0
SageGrouse,Wild Turkey, and CaliforniaQuail
but reasonablylarge in White-tailed Ptarmigan -0.2 - -0.2 ß

(Table 3). The angle of attack of maximal lift- 0.0 0.2 0.4 0.6 0.0 0.2 0.4 0.6

Drag coefficient Drag coefficienl

to-drag ratio did not change in White-tailed
Ptarmigan and SageGrouse,but it increasedin
Wild Turkey and California Quail after notches 1.0
1.0-
were createdexperimentally. California Quail Wild Turkey

All wing models also had smaller standard 0.8-

deviationsof lift and drag coefficientsafter the

0.8
•n
trailing-edge notcheswere created (Fig. 4). I 0.6- • 0.6
testedfor the effectof the notch,angle of attack, 0.4- • .4
and species,and for the interactionof species 0.2- ..3 0.2
and notch, on coefficients of variation (CV) of
the lift and drag measurements(Table 4). Co- 0.0- 0.0

efficients of variation for measurements of lift

-0.2- -0.2
were significantlyaffectedby only two factors: 0.0 0.2 0.4 0.6 0.0 0.2 0.4 0.6

(1) increasingthe angle of attack from 0øto 25ø Drag coefficient Drag coefficient

progressivelyreducedthe CV, and (2) notches Fla.4. Plotof lift vs.dragcoefficients

for the mod-
reducedthe CV by 53.4%regardlessof species. el wings.Eachcurveis basedon independentmea-
Coefficients of variation for measurements of
surements
(indicatedby standarddeviations)madeat
drag were significantlyaffectedby all four fac- sixanglesof attack.Fromleft to right: 0ø,5', 10ø,15ø,
tors (Table 4). The relationships of CV to the 20%and 25ø;nn is model with no notch, wn is model
angle of attackand the presenceof the notch with notch.
808 SERGE•
V. DROVErrSK/ [Auk, Vol. 113

TABLE 4. Effectof wing notching and angle of attackon the CV of lift and drag measurements
of model
wings.

Notching Angle of attack Species Notchingand species

F P F P F P F P

Lift 7.77 0.008 30.43 <0.001 2.81 ns 0.23 ns

Drag 20.48 0.001 34.09 <0.001 3.91 0.016 8.94 <0.001

Because the CV is a ratio of standard deviation during take off. Therefore, notchesmay help
to the mean, increasing the means will lower such birds to ascend faster.
the CV if the SD doesnot change.Drag increas-
es through the whole range of anglesof attack, DISCUSSION
and lift increasesthrough most of the range,
and then levels off or slightly decreases. Also, My comparativedata show that galliforms
the rate of changein lift and dragis not constant displaya wide variety of trailing-edgenotch
through the range of angles of attack (Fig. 1). sizesand wing shapes(Fig. 2). In general, gal-
For example, a minor error in positioning the liforms of all groups except Tetraoninae have
model wing for a small angle of attack (closeto the shortestwings and deepest notches found
0ø)will have a large effecton lift measurements in the order. Bonasaand Dendragapus have in-
and a small effecton drag measurements.When termediate notch sizesand intermediate wing
the angle of attack is closeto 25ø, the oppositelength-to-breadthratios.Other grouse(Lagopus,
is true. Tetrao,Tympanuchus and Centrocercus)and the
Despite other factors,wing notches signifi- migratory Common Quail have the longest
cantly reduced the CV of lift and drag mea- wings and smallestnotchesfound in the order,
surements.This suggeststhat wings with trail- and all, except the quail, have dark pectoral
ing-edgenotchesshouldbe morestablein flight, muscles,whereasthe other 22 specieshave light
and that their performanceshouldbe more pre- or intermediate colored pectoral muscles.
dictable.Trailing-edgenotchesalsoreducedthe Although the flight performancedata were
maximum lift coefficients (Table 3). The maxi- collectedfor fixed wings in steadyflow, differ-
mum lift coefficients of models without notches encesin lift and drag characteristics
shouldap-
were inversely related to the amount of their ply to fast forward flight with reduced fre-
reduction (r = -0.84, n = 4, P = 0.16). This quencies(see Norberg 1989) less than 0.5 Hz.
indicatesthat long wings that producelesslift The reducedfrequencyparameter(2a-fL / U) for
(e.g. SageGrouseand White-tailed Ptarmigan) these birds in forward flight is about 0.65 (cal-
loseproportionatelymore lift with the addition culated for Ring-neckedPheasant,f = 9 Hz;
of a notchthan do short,round,high-lift wings Greenewalt 1962:table 13). Despitea rough ap-
(e.g. turkey and quail). However, they alsolose proximation (the pheasanthas an intermediate
proportionatelymore drag, so the lift-to-drag body mass,soits speedis probablyhigher than
ratio of long wings increasesmore than that of the speed of California Quail and White-tailed
short wings. Therefore, trailing-edge notches Ptarmigan but lower than that of Wild Turkey
make takeoff easier by improving the lift-to- and SageGrouse;the oppositeis true for wing-
drag ratio, but they also increasethe energetic beat frequency), the calculated reduced fre-
cost of forward flight by decreasinglift, thus quency parameter is only slightly higher than
forcing birds to fly faster. the generally acceptedcutoff. Thus, the results
The angle of attackat maximum lift-to-drag of the experimentsstill are likely to apply to
ratio is probably stable regardlessof the pres- the flight of model species.However, in take
enceor absenceof a notch in long-winged birds off or landing, somecautionshouldbe usedin
(Sage Grouse, White-tailed Ptarmigan), but it interpreting differencesin steady state aero-
increasesin short-winged birds (Wild Turkey, dynamic coefficients.
California Quail). The increasein angle of at- The differencesin wing geometryand notch
tackin birdswith short,roundedwings should size are related to the ecologyand behavior of
increaselift (at the expenseof some increasein differentspecies.Exceptfor the grouseandptar-
drag) and, thus, increasevertical acceleration migan, which use flight between bouts of for-
October
1996] Flight
Performance
andtheWing
Notch 809

aging, most galliforms use flight primarily to other branch in the same tree, or to a neigh-
flee from predators.Thus, becausethe main el- boring tree in the same canopy. During one
ement of their flight is vertical (takeoff and as- foraging bout a Hazel Grouse visits 6-12
cending), level flight is not so important. In branches.Flightswithin foragingboutsareshort
contrast,flight is an important part of the for- and slow, and a flock does not move more than
aging behavior of grouse and ptarmigan, and 300 m during a single bout of foraging. Thus,
the relative importance of level flight varies the majorityof the energy Hazel Grousespend
dramatically in specieswith dark versus light in foraging flights is investedin vertical move-
(or intermediate)pectoralmuscles.Grousewith mentsand in the frequent takeoffsand landings
a higher proportion of red fibers in the pector- required to move from branch to branch. Fur-
alismajorfly relatively long distancesfrom one thermore, Hazel Grouse land and feed on such
foraging site to another, but then they either small branchesthat they can not jump from
walk on the ground or perch on branchesto them with their feet to aid in takeoff.This places
feed. Thesegrouseuselevel flight primarily to even greater demands on their wings for ac-
reachfood patches.However, grousewith light celeration.For theseshort burstsof flight, white
or intermediate pectoral muscle fibers do not muscle fibers; short, broad wings; and deep
fly long distances.Rather,theyuseverticalflight trailing-edge notchesare the best compromise.
primarily from groundto canopy,or slowflight I usedmodelwings of two pairsof galliforms
from branch to branch, to reach their food. The with different flight characteristicsin my ex-
grouse with white and intermediate fibers and periments. White-tailed Ptarmigan and Sage
short, broad wings have deep trailing-edge Grouse represent specieswith small notches
notchesthat increaseboth their lift-to-drag ra- (Fig. 2). They have high aspectratios, heavy
tio and the angle of attack at which their wings wing loading for their size, low maximumlift
are more efficient. coefficients,and dark pectoral muscles(Tables
For example, the Willow Ptarmigan (Lagopus 1-3). Theseareall adaptationsfor relativelylong,
lagopus),which has red pectoral muscles,feeds fast, straightand efficient flight (Rayner 1988,
on willows (Salix schwerinii,S. rorida) and cho- Norberg 1989).In contrast,the Wild Turkey and
senia (Chosenia arbutifolia)twigs and buds dur- California Quail, which represent specieswith
ing winter in northeastern Russia (Andreev deep notches(Fig. 2), have low aspectratios,
1980, Drovetski 1992b). The willow and cho- relatively light wing loading, high lift coeffi-
senia thickets where ptarmigan forage are sit- cients,and light pectoralmuscles(Tables1-3).
uated on river islandsand along the banks of Thesefeaturesare adaptationsto powerful short
braided river channels.During foraging birds flightsand maneuverability(Rayner 1988,Nor-
walk along patchesof thickets and pick twigs berg 1989).
and buds from bushesand snow. After passing Short, wide wings can generateas much lift
through a patchof willows, a flockof ptarmigan aslong, narrow wings of the sameareaonly by
must fly to reach another willow patch, which increasingthe angle of attack (Vogel 1983).
usually will be situatedon another river island However, higher anglesof attackgeneratein-
or along another channel. Thus, Willow Ptar- creasesin both profile and induced drag. My
migan perform several flights of several hun- resultsshow that the trailing-edge notchesin
dred metersa day. They fly through openplaces galliform wings increasethe maximumlift-to-
at a fast pace and at low altitude. Becausethese drag ratio by improving air flow around the
flights are relatively long, the most expensive wing. Notcheshad a similar effect on all four
componentis not takeoffand landing, but rath- species,i.e. increasingthe maximumlift-to-drag
er level movement. ratio and stabilizingair flow around the wing.
In the sameregion, the Hazel Grouse(Bonasa However, the trailing-edge notch slightly re-
bonasia), which haswhite pectoralmuscles,feeds ducedthe maximumlift coefficient(Fig. 2). This
in trees but in denser, more continuous forests reduction in maximum lift is presumablymore
(Andreev 1980; Drovetski 1992a, 1992b). Unlike detrimental to speciescharacterizedby rela-
Willow Ptarmigan,Hazel Grouseforagemostly tively long flightsthan to speciesthat spenda
by perching on branchesin the canopy,and high percentageof their flight time in landings
extremely rarely by walking to branchesthat and takeoffs, times when the angle of attack
canbe reachedfrom the snow.Eachbird spends must be great.
3 to 8 rain on one branch and then flies to an- Theexperimentaldatashowa tradeoffin flight
810 s•a•i V. DRoversKI [Auk,VoL 113

performanceof galliforms associatedwith the with takingpicturesof modelwings.I am very grate-

trailing-edge notch. The notch reducesmaxi- ful to JeremyRayner, Carl Heine, Kenneth Dial, and
mum lift, but increasesthe lift-to-dragratio and an anonymousreviewer for very helpful comments
stabilizesairflow around the wing. This means on the manuscript.This studywassponsoredby Mr.
that an increasein the ability to take flight and GarretEddy.Thanksalsoto the BurkeMuseum'sEddy
Endowmentfor Excellencein Ornithology for sup-
climb fast,and in the ability to maintainheight
port during theseinvestigations
and to the University
during a slow flight, may be achievedat a cost of WashingtonBurkeMuseumand the Universityof
of reducedefficiencyin level flight. Galliforms Puget Sound Slater Museum of Natural History for
that experiencerelatively long and fast level useof specimens. The comparativepart of this study
flights shouldbenefit more from a high-lift (i.e. could not have been done without the Burke Mu-
high-efficiency)wing. In suchspeciesthe trail- seum'scollections
of extendedwingsfromAsia,North
ing-edge notch should be smaller. In short America, and Africa.
flights,when verticalmovementsbecomea large
componentof eachflight, increasingthe lift-to- LrrERXTUP, E CITED
drag ratio becomesmore advantageous. In this
circumstanceselectionshouldfavor larger trail- ANI)REEV,A. V. 1980. Adaptatsiyaptits k zimnim
ing-edge notches. usloviyamsubarktiki. "Nauka," Moscow.
Comparedwith othervolantbirds,galliforms DROVETSKI, S. V. 1992a. Materialy po ecologii Ry-
spend small amounts of time in the air, rarely abchika (TetrastesbonasiaL.) na Uge Magadan-
fly long distances,and have pronouncedtrail- skoy oblasti v zimniy period. Zoologicheski
Zhurnal 71:45-59.
ing-edge notches. The comparative analysis
DROV'a'TSKI,
S.V. 1992b. Novye dannyeo funktsion-
within the order supportsthe resultsof exper-
al'nomznacheniirogovoybakhromyna pal'tsakh
iments and suggeststhat wing geometry and teterevinykhptits.ZoologicheskiZhurnal 71:100-
size of the notch result from of a compromise 109.
between what is optimal for level flight and GREENEWALT, C.H. 1962. Dimensionalrelationships
what is optimal for frequent takeoffand vertical for flying animals. Smithsonian Miscellaneous
flight. Grouse that spend much time in level Collections 144:1-46.
flight have small trailing-edge notches,where- HOFTON, A. 1978. How sails can save fuel in the air.
New Scientist 78:146-147.
as those whose flight is characterizedby fre-
L•Yolq, L.E. 1976. The measurement of bone strain
quent takeoffs and landings, and by vertical
movement, have deep trailing-edge notches. in vivo. Acta Orthopaedica Belgica 42 (Supple-
ment 1):98-108.
One may ask why ptarmigan and other long-
NORI•ERC, U. M. 1989. Vertebrate flight. Zoophy-
winged galliformshaveany trailing-edgenotch. siology.Springer-Verlag,Berlin.
The answer, I think, lies in the fact that even
RAYNER, J. M.V. 1985. Speedof flight. Pages225-
thosegrousewith relativelylong, narrowwings 226 in A dictionary of birds (B. Campbell and E.
fly very little comparedwith other birds. Thus, Lack, Eds.).T. and A.D. Poyser,London.
relative to other species,long-winged galli- RAYIqER,J. M.V. 1988. Form and function in avian
forms spend a considerable amount of their flight. Current Ornithology 5:1-66.
SHEST.a,
flight time in takeoff,where the notch is help- KOV.% G.S. 1971. Stroeniekryl'ev i mekhan-
ful. ika polyota ptits. "Nauka," Moscow.
Whereas the trailing-edge notch improves SHTEC;MAN, B.K. 1953. Osobennostilyotnykh ka-
chestv seroy i kamennoykuropatok.Zoologi-
maximum lift-to-drag ratio of the extended cheski Zhurnal 37:677-683.
wing, the notch could be even more important TUCKER, V.A. 1993. Gliding birds:Reductionof in-
for a powered upstroke.The possibility that it duceddragby wing tip slotsbetweentheprimary
createsa separatewing should be examinedby feathers.Journalof ExperimentalBiology 180:
photography,and, if confirmed,be exploredby 285-310.
further work in wind tunnels or flow tanks. TUCKER, V.A. 1994. Drag reductionby wing tip slots
in a gliding Harris' Hawk, Parabuteounicictus.
ACKNOWLEDGMENTS Journalof ExperimentalBiology198:775-781.
VINOGRADOv, I. N. 1951. Aerodinamikaptits pari-
I thank Sievert Rohwer, Thomas Daniel, and Chris teley. DOSARM, Moscow.
Wood for their help with this study, and Dmitriy VotEr, S. 1983. Life in moving fluids. Princeton
Banin, Richard Droker, and Carol Spaw for the help University Press,Princeton,New Jersey.
Associate Editor: K. P. Dial