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Morphometrics and Flight Performance of Southern African Peregrine and Lanner Falcons
Author(s): Andrew R. Jenkins
Source: Journal of Avian Biology, Vol. 26, No. 1 (Mar., 1995), pp. 49-58
Published by: Wiley on behalf of Nordic Society Oikos
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JOURNAL OF AVIANBIOLOGY 26: 49-58. Copenhagen1995

Morphometrics and flight performance of southern African

Peregrine and Lanner Falcons
Andrew R. Jenkins

Jenkins,A. R. 1995. Morphometricsand flight performanceof southernAfrican

Peregrineand LannerFalcons.- J. AvianBiol. 26: 49-58

Twenty-fourmorphometric parameterswere measuredfromsamplesof live southern

AfricanPeregrineFalconsFalco peregrinusminorand LannerFalconsF. biarmicus
biarmicus.The two species were differentin most measurements,especiallythose
relevantto preycaptureandhandlingtechniques,andflightperformance (bill size, foot
size, wing span,wing area,tail lengthand wing loading).Flightperformanceparam-
eters calculatedfrom mensuraldata predictedsignificantdifferencesin the flying
abilitiesof the two species,notablythatPeregrinesshouldfly fasterin level powered
andglidingflight,butincurgreaterfuel costs in termsof bothtimeanddistanceflown.
Peregrinesshould glide less efficientlyand be restrictedin their ability to soar in
thermals.Thesepredictionswere comparedwith observationsof PeregrinesandLan-
nersunderuniformenvironmental conditions,andmostlywere confirmed.Peregrines
flew fasterbut for less time, flappedmore and soaredin thermalsless thanLanners.
Theoretically,Peregrinesshould tend more towardssedentaryperch huntingthan
Lanners,andbe morehabitatselectiveas a result.Observations anddistributional data
from South Africa corroboratethis. Form and functionaldifferencesin these two
falcons are relatableto differencesin foragingmode, distributionand abundance.I
suggestthatsimilarinferencesmaybe drawnfrommorphological comparisonsof other
largefalconsto provideproximalexplanationsfor broad-scalepatternsof distribution.

A. R. Jenkins,PercyFitz PatrickInstituteof AfricanOrnithology,Universityof Cape

Town,Rondebosch7700, SouthAfrica.

Morphometric datahave been used as both evidencefor sized raptors found widely throughout sub-Saharan
and predictorsof differencesin flight performanceand Africa. The basic habitat and food requirements of these
feedingecology of co-existingraptors.Intraspecificstud- two species are the same: they usually nest on cliffs and
ies have comparedbirds of differentraces (e.g. White feed on birds which are caught in aerial chases (Cramp
1982), sexes (e.g. AnderssonandNorberg1981) andage and Simmons 1980, Steyn 1982). In southern Africa,
classes (e.g. Muelleret al. 1981,Brown1989).Interspec- Peregrines and Lanners are sympatric in many areas, but
ific studieshavebeen concernedmainlywith nicheparti- Peregrines are more sparsely distributedand generally are
tioning(e.g. Barnard1986). Authorshave tendedto em- much rarer(Mendelsohn 1988). In South Africa, Lanners
phasizethe importanceof specific wing and tail param- outnumberPeregrines by 10:1 in most areas and occupy a
etersin predictingthe energeticcosts of flight andhence breeding range five times larger (Jenkins in press). This
the optimumhuntingmode (e.g. Jaksic and Carothers probably is because the Lanner is a more generalized
1985). Othermorphometriccharacters,such as toe and feeder and is less specific in its nest-site requirements,
tarsallengthsand beak sizes (Cade 1982, White 1982), enabling it to occupy a wider variety of habitats. Pe-
are thoughtto reflect differencesin the food handling regrine populations in southern Africa may be restricted
abilitiesof differentforms. either by the competitively superior Lanner (Tarboton
The Africanrace of the PeregrineFalcon Falco pe- 1984, Thomson 1984) or by food and/or habitat limita-
regrinus minor and the nominaterace of the Lanner tions (Jenkins 1991).
FalconE biarmicusbiarmicusareboth smallto medium Differences in the feeding ecologies and habitat prefer-



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ences of southernAfricanPeregrinesandLannersshould
be reflectedin theirrespectivemorphologiesand flying
Predicted flight performance
andhuntingabilities.In this paperI examinethe relation- The body mass, wing spanand wing areameasurements
ship between form and function and distributionand of each individualwere used to calculateflightperform-
abundancein thesetwo falconspecies.Mensuraldataare ance parameterswith Pennycuick's(1989) bird flight
used to predict flight performanceparameterswhich performance computerPrograms1 and2. Theparameters
might influenceforagingmode, food and habitatselec- calculatedfor horizontalflappingflight were: minimum
tivity, dispersabilityand ultimately populationstatus. power speed (Vmp)- the air speed at which power output
Fieldobservationsof huntingandflying behaviourof the for flight is least, maximum range speed (V,mr)- the air
two species underthe same conditionsare then used to speed at which the ratio of power to speed is lowest,
test these predictions.Intraspecificdifferencesbetween effective lift: drag ratio (L/D max) at Vmr, fuel consump-
adultsand immaturesalso are discussed. tion - gramsof fat consumedper km flown at Vmr, the
minimum aerobic scope - the minimum power required
to fly (Pmin)dividedby an estimateof the basalmetabolic
rate (Pmet) (see Pennycuicket al. 1994). The parameters
Methods calculated for gliding flight were: stall speed - the mini-
mumair speedrequiredto avoid stalling,minimumsink-
Morphometrics ing speed - the lowest rate of descent attainable, best
Body mass,wing area,wing loading,aspectratioand20 glide ratio - the best achievable ratio of forward to
linearparameterswere measuredfrom a total of 73 live downwardmovement,flying at the best glide speed,the
falcons and one recently dead specimen. The sample best glide speed (Vbg)- the air speed at which the ratio of
comprised33 Peregrines(15 adultfemales, five imma- power to speed is lowest, circling radius - the minimum
turefemales,eight adultmalesandfive immaturemales) radiusof a thermalin which the bird is able to climb,
and 41 Lanners(nine adult females, 12 immaturefe- banking at a standard angle of 240, and cross-country
males, seven adultmales and 13 immaturemales). Any speed (opt)) - the groundspeed of cross country
birdnot in full adultplumagewas consideredimmature. flights(Vxc
if inter-thermal glides aremadeat optimumspeed
Of the birds measured,14 Peregrinesand 28 Lanners and using thermalsrisingat a standardspeedof 5 m s-'.
werelive-trappedin theTransvaalandtheCapeProvince, Formoredetailson the derivationof theseparameters see
South Africa, between 1989 and 1994. The remainder Pennycuick(1989). The programs'default values for
werecaptivebirdsfromvarioussources(eitherinjured,in accelerationdue to gravity(9.81 m s-' - standardearth
captivebreedingfacilities or being flown by falconers). gravity)andairdensity(1.23 kg m-3)wereused through-
Measurementstaken and units used were those of out.
Biggs et al. (1977) and Mendelsohnet al. (1989) except Two-sampleStudent'st tests were used to test the
wherethey were relevantfor flightperformancecalcula- significanceof differencesbetweenthe means of each
tions, in which case they followed Pennycuick(1989). morphometric andflightperformancevariablein each of
Body mass was measuredcorrectto the nearest 10 g, the groupingscompared.For those variableswheresig-
using either a 1000 g or 1500 g springscale for wild- nificantdifferenceswerefoundbetweenadultandimma-
trappedbirds,or variousbalancescalesfor captivebirds. ture birds in within-speciescomparisons,only data for
Linearmeasurementswere takento the nearest0.1 mm adultbirdswere used in between-speciescomparisonsto
using Verniercalipersor to the nearest1 cm with a ruler preventthe proportionof immaturesfrominfluencingthe
or tapemeasure.Wingareawas measuredto the nearest1 result.
cm2from a tracingof the extendedwing, flatteneddor-
sally. The massof the cropcontentsof wild-trappedbirds
was estimatedaccordingto the degreeof cropdistension Observed flight performance
(10 g incrementsfrom 10 g to 50 g), andsubtractedfrom
the measuredmassto give an emptymassfor all individ- Two adult pairs of each species were observedin the
uals. Ten per cent was addedto the actualbody mass of AugrabiesFalls National Park (AFNP), South Africa
falconrybirdsto approximatewild condition. (28oS, 200E) between 25 April and 16 May 1993. The
Morphometricdatafor each species were groupedby birdsareresidentin the park,andbreedon the walls of a
sex. Data for Peregrinefemales and Lannermales were deep gorge alongthe lowerreachesof the OrangeRiver.
tested for differencesbetweenwild-trappedand captive The gorge is about 15 km long, and featuresopen ex-
birds. These two groupingshad the most comparable panses of sheerand semi-sheerrock on both sides. The
samplesof birdsfromeach source.Datafor bothsexes of cliffs vary fromabout80-120 m high, and the valley is
both species were tested for differencesbetweenadults about 150-500 m wide. The gorge runsthroughan area
and immatures. of undulating,rockyhills, about650 m above sea level.
The area generallyis dry, and vegetationis sparseand
mostlyconfinedto the watercourses.Thenestcliffs of the
fourpairsof falconsarespacedunevenlyalongthe first8

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Table1.Morphometric betweenadultandimmature the activitiesof both birds at once. Observationswere
southern AfricanPeregrines andLanners. Datagivenaremeans madefromone or two positionsalong the gorge wall at
? ISD, n.s. = not significant,*P < 0.05, **P < 0.01, ***P < each
0.001. site, andthe birdswere not activelyfollowed.Data
from the two sites for each sex of each species were
Adults Immaturest pooled.Timesspenton each activitycategoryfor eachof
(n) (n) value the observationperiodswere expressedas percentagesof
the timebirdswerein sight,or for flight activities,of the
Peregrines time the birds were observedflying. These percentages
males were then grouped by species, or within species by
secondary length 124?4 137?4 5.43***
(mm) (6) (5) weatherconditions,and differenceswere tested for sig-
outerrectrix 132?6 141?4 2.97* nificanceusing non-parametric Mann-Whitney U tests.
length(mm) (6) (5) Wheneverpossible,huntsandlongerflightswereplot-
innerrectrix 134?7 142?3 2.33* ted on 1:5000 aerialphotographs,using landscapefea-
length(mm) (6) (5)
claw 1 length 19.2?0.8 18.1?0.7 2.33* tures to estimatethe routesfollowed. These plots were
(mm) (6) (5) measuredand providedindices of strikeand flight dis-
females tances.Indicesof strikeand flight groundspeeds could
secondarylength 149?4 155?4 2.73* thenbe generatedin cases whereflights were both mea-
(mm) (11) (5)
outerrectrix 152?5 159?7 2.65* suredandtimed.The observedhomerangeof eachof the
length(mm) (10) (5) four pairswas estimatedby outliningthe minimumarea
covered by all the flights plotted at each site. When
males observingbouts of flappingflight, I tried to count the
outerrectrix 159?5 167?7 2.70* numberof beatscompleted.In flappingboutslongerthan
length(mm) (7) (13) 5 s in duration,I dividedthe numberof beatscountedby
females the whole numberof seconds elapsed, to result in an
bill length 22.3?1.5 21.1?0.9 2.42* index of wingbeat frequency (see Pennycuick 1990).
(mm) (9) (12)
bill width 16.5?0.7 17.9?1.6 2.38* These variousindices and estimatesall are prone to a
(mm) (9) (12) substantialdegreeof error,but this was consideredto be
claw2 length 18.6?1.3 17.3?0.7 2.86** equal for both species. The data are intendedonly to
(mm) (9) (12) reflectthe relativeflightperformanceof the two species.
Meanvalues for flight and striketimes, and distance,
km of the gorge, downriverfrom the falls. Althoughthe each species, and for males and females within each
gorge is narrower,shallowerand steeper-sidedat its up- species. Differenceswere tested for statisticalsignifi-
per end thanits lower end, overallit providesa structur- cance using Student'st tests.
ally uniformhabitatfor foragingfalcons,ideallysuitedto Localmaximumtemperatures for eachdayof the study
a study comparingthe huntingand flying behaviourof wereprovidedby the SouthAfricanWeatherBureau,and
the two species. wind speed was ranked 0-4 (0 = calm, 1 = light, 2 = light
PeregrinesandLannersdo not breedat the sametime to moderate,3 = moderateto strongand 4 = strong)for
in southernAfrica (Steyn 1982). This study was con- each observationperiod.
ductedin the non-breedingseason of both species, and
was timedto excludeas muchas possiblethe influences
of moult,courtshipandbreedingon maintainance activity
budgets. Results
Falconswere observedusing 10 x 40 binocularsanda
20-60x spottingscope, at distancesof 100-1500 m. All Morphometrics
observationswere recordedon a dictaphone,and sub- No significant differenceswere found between wild-
sequentlywere timed correctto the nearestsecond,and trappedand captive birds in any of the measurements
transcribed.Bird activitieswere divided into four cate- taken.
gories:perched,gliding (all flying withoutflapping,in- ImmaturePeregrineshad longer secondaryfeathers
cluding soaring in orographicupdrafts (Pennycuick and longerrectrices(only outerfeathersin females)than
1989) but excludingthermalsoaring),flapping and ther- adultsin both sexes (Table1), and immaturemale Lan-
maling(circlingand gainingheightin a risingpocketof ners had longer outer rectricesthan adults. Generally,
differentiallyheatedair). immaturesof both species tendedto be lighter and to
A focal bird (Altmann 1974) was selected and its have largerflight surfaceareasand lower wing loadings
activitieswere recordedfor as long as the bird was in than adults,althoughthese differenceswere not signif-
sight, or until the end of the observationperiod.When icant. Male Peregrinesand female Lannersshowed sig-
one or bothof the pairat a site were perchedin an easily nificant differencesbetween adults and immaturesin
observableposition,it was sometimespossibleto record some bill and claw measurements(Table1).

4* JOURNAL OF AVIAN BIOLOGY 26:1 (1995) 51

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Table2. Comparative morphometric datafor southernAfricanPeregrinesandLanners.Datafor adultsandimmatureswerepooled,
exceptwheretheseweresignificantlydifferent,whenonly adultswerecompared.Datagivenaremeans? 1SD,n.s. = not significant,
*P < 0.05, **P < 0.01, ***P < 0.001.

Peregrine Lanner t Peregrine Lanner t

males males value females females value
(n) (n) (n) (n)

Body mass 0.528?0.029 0.506?0.035 2.01 n.s. 0.771?0.072 0.726?0.063 2.13*

(kg) (13) (20) (20) (21)
Body length 360?11 390?11 6.98*** 403?11 444?16 8.44***
(mm) (11) (19) (15) (20)
Bill length 19.5?1.0 19.3?1.0 0.52 n.s. 22.7?1.0 22.3?1.5 0.84 n.s.
(mm) (11) (20) (11) (9)
Bill width 15.5?1.8 15.2?1.9 0.43 n.s. 15.2?1.2 16.5?0.7 1.90 n.s.
(mm) (11) (20) (11) (9)
Bill depth 14.4?0.5 14.7?0.5 1.63 n.s. 16.5+0.5 16.2?0.7 1.68 n.s.
(mm) (11) (20) (16) (21)
Wing span 0.873?0.022 0.964?0.034 9.17*** 0.984?0.052 1.065?0.036 6.52***
(m) (13) (20) (20) (21)
Wing length 284?7 315?11 8.31*** 321?7 354?10 11.27***
(mm) (11) (19) (16) (21)
Wing area 0.091?0.005 0.118?0.008 6.85*** 0.119?0.008 0.145?0.008 10.31***
(m2) (13) (20) (20) (21)
Ulna length 80?6 90?5 5.08*** 95?6 98?5 1.75 n.s.
(mm) (11) (20) (16) (21)
Secondary 129?8 156?5 12.27*** 149?4 177?7 8.07***
length(mm) (11) (20) (11) (9)
Outerrectrix 132?6 159?5 9.24*** 152+5 183+6 8.88***
length(mm) (6) (7) (10) (9)
Innerrectrix 138?7 165?7 10.04*** 153?6 189?9 13.57***
length(mm) (11) (20) (15) (21)
Tarsus 45.4?4.5 48.3?3.7 1.94 n.s. 53.0?3.2 52.2?3.8 0.71 n.s.
length (mm) (11) (20) (16) (21)
Tarsus 6.1+0.3 6.0?0.3 0.79 n.s. 7.2?0.4 6.8?0.4 3.23**
width (mm) (11) (20) (16) (21)
Toe 1 27.3?3.9 25.7?3.2 1.23 n.s. 32.0?3.3 25.8?4.0 4.92***
length (mm) (11) (20) (15) (21)
Toe 2 46.4?2.8 43.4+2.1 3.40** 51.1?3.0 45.3?2.8 5.94***
length(mm) (11) (20) (15) (21)
Toe 3 34.0?3.3 29.2?3.3 3.85*** 37.7?3.5 32.0?3.5 4.83***
length (mm) (11) (20) (15) (21)
Toe 4 23.2?2.9 20.4?2.3 2.96** 26.1?3.0 22.3?1.6 5.09***
length (mm) (11) (20) (15) (21)
Claw 1 18.6?0.9 16.7?1.3 4.24*** 21.2?1.2 18.5+1.0 7.36***
length (mm) (10) (20) (15) (21)
Claw 2 17.4?0.9 16.2+0.6 4.30*** 20.1+1.1 18.6?1.3 3.10*
length (mm) (10) (20) (15) (21)
Claw 3 16.3?0.9 15.2?0.9 3.22** 19.9?5.9 16.5?1.1 2.64*
length (mm) (10) (20) (15) (21)
Claw 4 19.6?0.9 18.3?0.8 4.33*** 22.9?1.9 20.0?1.0 6.04***
length(mm) (10) (20) (15) (21)
Wing loading 57.2?4.3 42.1?3.9 10.38*** 63.3?3.9 49.1?4.4 10.96***
(N m-2) (13) (20) (20) (21)
Aspect 8.49?0.37 7.92?0.42 3.77*** 8.18?0.56 7.83?0.54 1.95 n.s.
ratio (11) (20) (16) (21)

Female Peregrines were significantly different from (Fig. 1), and had heavier wing loadings, and higher aspect
female Lanners in 18 of the 24 measurements compared ratios.
(Table 2). Bill size, ulna length, tarsus length and aspect
ratio were the same. Male Peregrines were significantly
different from male Lanners in 17 of the measurements
Predicted flight performance
compared (Table 2). Body mass, bill size, tarsus length
and width, and toe 1 length were the same. Pennycuick (1989) recommends that information gener-
Overall, Peregrines tended to be heavier and to have ated by his programs be compared in terms of percentage
longer bills (not significant), were shorterbodied, smaller change rather than absolute values. In this study, Pe-
winged, shorter tailed and bigger footed than Lanners regrines and Lanners were significantly different in all


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terms of fuel consumed per unit time - Vmp) and the speed
at which the power to speed ratio is lowest (i.e. fuel
consumption per unit distance flown is minimized - Vmr)
(Pennycuick 1989) are 3-5% higher for Peregrines than
for Lanners, (ii) the effective lift:drag ratios for flight at
Peregrine Vmr are 3-7% lower for Peregrines than for Lanners, (iii)
fuel consumption (grams of fat consumed per km flown
at Vmr) is 8-9% higher for Peregrines than for Lanners
and (iv) the minimum aerobic scope required is 12-15%
greater. In gliding flight (i) Peregrines must fly 11-15%
Lannermale faster than Lanners to avoid stalling, (ii) Peregrines
achieve a 3-5% lower best glide ratio at a best glide
speed 6-9% higher than Lanners, (iii) the minimum rate
of vertical sink is 10-12% higher for Peregrines than for
Lanners, (iv) Peregrines require thermals 20-26% larger
than Lanners in order to soar and (v) when flying cross-
Peregrine female
country, gliding at optimum speed between thermals ris-
ing at a standard5 m s-', Peregrines fly 1-3% faster than
Overall, optimum flight speeds for Peregrines are
higher, but fuel consumption for powered flight is greater
and gliding performanceis inferior in terms of its sustain-
Lanner female
ability in calm conditions. The cross-country flying abil-
ity of Peregrines is impaired by their dependence on
larger thermals. In strong winds Peregrines should be
able to maintain sufficiently high air speeds and glide
relatively more efficiently (Tucker and Parrot 1970).
Fig. 1. Simplifiedflightoutlinesof southernAfricanPeregrines
and Lanners,based on the morphometric datain Table2. Although differences were not significant, immatures
tended to have slower optimum flight speeds, higher
lift:drag ratios and glide ratios, slower stall speeds and
the flight performance parameters compared (Table 3). narrowercircling radii than adults, and flight tended to be
These differences predicted that in horizontal flapping less energetically expensive.
flight (i) the speeds at which flight is least strenuous (in

Table3. Comparative
flightperformance of southernAfricanPeregrinesandLanners,calculatedusingPrograms1 and2
fromPennycuick(1989). Data for adultsand immatureswere pooled,except wherethese were significantlydifferent,when only
adultswerecompared.Samplesizes werePeregrinemalesn = 13, Peregrinefemalesn = 20, Lannermalesn = 20, Lannerfemalesn
= 21. Data given are means ? 1SD, n.s. = not significant, *P < 0.05, **P < 0.01, ***P < 0.001.

Peregrine Lanner t Peregrine Lanner t

males males value females females value

Vmp (m s-1) 9.1?0.2 8.5?0.2 8.03*** 9.8?0.4 9.3?0.3 4.80***

VmrS-') 15.0?0.3 14.2?0.3 6.79*** 16.2?0.5 15.3?0.5 5.95***
Max. L/D 7.35?0.18 8.12?0.24 9.82*** 7.64?0.25 8.28?0.25 8.07***
Fat at Vm, 0.079?0.005 0.068?0.005 5.37*** 0.113?0.012 0.098?0.012 3.04**
(g km-')
Aerobic 15.35?0.84 12.91?0.77 8.61*** 15.20?1.20 6.58***
Stall speed 7.62?0.29 6.52?0.29 10.58*** 8.02+0.20 7.06?0.30 10.88***
(m s-')
Best glide 13.3?0.3 13.9?0.3 5.78*** 13.5?0.4 14.0?0.4 3.74***
Best glide 10.2+0.3 9.2?0.3 9.51*** 10.9?0.4 10.0?0.4 7.82***
speed (m s-')
Min. sinkingspeed 0.669+0.020 0.578?0.026 9.17*** 0.708?0.040 0.627?0.040 6.40***
(m s-')
Circling 16.7?1.3 12.2?1.1 10.92*** 18.4?1.1 14.3?1.3 10.993***
radius (m)
Vxc (opt) (m s-') 12.5?0.1 12.2?0.2 5.23*** 13.2?0.2 12.9?0.2 4.84***


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Table 4. Comparativeactivitybudgetdata for Peregrinesand 50.5%of the observationtime. Includingtime whentwo
Lannersat the AFNP,from observationsof two pairsof each birdswereunderobservationat once, andexcludingtime
species. Data for each sex of each species were pooled. Pe- when no birdswere in sight, 119.8 h of activitybudget
regrinesn=17 observationperiods,Lannersn= 11 observation
periods,n.s. = not significant,*P < 0.05, **P < 0.01, ***P < data were collected at Peregrinesites (males 58.9 h,
0.001. females60.9 h), and68.0 h at Lannersites (males34.8 h,
females33.2 h). Becauseobservationsamplesweresmall
Peregrines Lanners U value
for the sexes withineach species, datawere pooled and
activity budgetswere comparedbetween species only.
Flying' 4 21 182*** Observationperiodswhere birds were in sight for less
(range 1-9) (range7-62)
Gliding2 55 53 98 n.s. than 60 min (n = 1 at Peregrinesites, n = 2 at Lanner
(range29-87) (range43-73) sites) wereconsideredunrepresentative samplesandwere
Flapping2 40 14 172*** excluded from the analysis. Of the total samples, Pe-
(range13-71) (range5-29)
Thermaling2 5 33 177*** regrinesflew for 4.7 h (3.9%) and Lannersfor 11.4 h
(range0-31) (range4-48) (16.8%).Seventy-threeflights and 25 strikeswere mea-
suredat Peregrinesites, covering94.2 km. Eighty-five
'Average% of time observed. flights and 41 strikes were measuredat Lannersites,
2Average% of time observedflying.
totalling 123.4 km of flying. Mean maximumtemper-
atures for Peregrine(26.5 ? 6'C, n = 17) and Lanner
(25.5 ? 3oC, n = 11) observationperiods used in the
Observed flight performance activitybudgetanalyseswere not significantlydifferent
(Student'st = 0.77, df = 26, P > 0.2), andthe modalwind
Periods of observationranged from 1.6 h to 11.5 h.
speed rankswere the same (mode = 1: light wind).
Eighteen observationperiods were completedat Pere- Peregrines flew significantly less than Lanners
grine sites and 13 at Lannersites, totalling104.3 h and (Table4), and when flying, flapped significantlymore
100.9h of observationrespectively.Peregrinesremained andsoaredin thermalssignificantlyless. Temperature did
almostexclusivelywithinthe confinesof the gorge, and not significantlyinfluencethe flight behaviourof either
were in sight for 78.5% of the total observationtime,
whereasLannersfrequentlyflew out of sight over the species. On average,Peregrineflights were shorterin
durationand flight speeds were faster than Lanners
countryadjacentto the gorge, andwere in sight for only

Table5. Comparative flighttimesanddistance,speedandwingbeatfrequencyindicesof maleandfemalePeregrinesandLannersat

the AFNP,fromobservationsof two pairsof each species.Datafor each sex for each specieswerepooled.Datagiven aremeans+
1SD, n.s. = not significant,*P < 0.05, **P < 0.01, ***P < 0.001.

Peregrines Lanners t
(n; range) (n; range) value

Flight distance 1297?1078 1469?1083 0.69 n.s.
index (m) (33; 200-5750) (43; 100-4625)
Flighttime (s) 126?134 269?274 2.39*
(24; 16-585) (26; 11-1149)
Flight speed 13.5?6.3 7.3?5.6 3.55***
index (m s-') (24; 3.6-32.7) (26; 2.4-25.0)
Strikedistance 398?268 179?137 2.92**
index (m) (11; 150-1100) (18; 50-600)
Strikespeed 26.7?7.9 18.9?9.4 2.20*
index (m s-') (10; 16.7-40.3) (16; 6.7-35.0)
Wingbeatfrequency 4.23?0.76 3.59?0.60 8.52***
index (beatss-') (176; 1.33-7.00) (170; 1.67-5.20)
Flightdistance 1079?827 1263?1202 1.43 n.s.
index (m) (40; 150-3175) (41; 200-7650)
Flighttime (s) 88?68 167?114 3.23**
(32; 24-334) (25; 26-390)
Flight speed 12.1?1.3 7.0?3.2 5.20***
index (m s-') (32; 4.7-21.6) (25; 3.4-14.4)
Strikedistance 279?190 234?94 1.10 n.s.
index (m) (14; 70-675) (22; 100-400)
Strikespeed 24.9?9.1 19.5?7.8 1.65 n.s.
index (m s-') (11; 10.0-40.0) (16; 8.6-31.8)
Wingbeatfrequency 3.17?-0.64 11.29***
index (beatss-') 3.94_?0.56
(182; 2.13-5.33) (127; 1.60-4.75)


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Table6. Comparative activitybudgetdatarelative
to windspeed Peregrinesactively flushed prey before chasing it (see
for PeregrinesandLanners at theAFNP,fromobservations of Jenkinsandvan Zyl 1994)in 13 (28.9%)of the observed
twopairsof eachspecies.Dataforeachsexof eachspecieswere
n = 10 observation hunts, and pairs hunted together on four occasions
pooled.Peregrines periodswithcalm/light
winds(ranks0 and 1) and n = 7 observation periodswith (8.9%).Lannersonly flush-huntedfour times, but pairs
moderate to strong(ranks3 and4), Lanners n = 4 andn = 7 huntedtogether27 times (6.5% and 43.5% respectively
respectively,n.s. = not significant,*P<0.05, **P<0.01, of the huntsobserved).Both species huntedsmall pas-
***P< 0.001. serines (Red-eyed Bulbuls Pycnonotus nigricans and
Calm/light Moderate/strongU value Redbilled Queleas Quelea quelea), aerial insectivores
wind wind (Alpine Swifts Apus melba and Rock MartinsHirundo
fuligula) and columbids (Streptopelia sp. and Rock Pi-
Peregrines geons Columba guinea). Forty-three (95.5%) of the Pe-
Flying' 2 6 60** regrine strikes observed apparentlywere attemptsto
(range1-5) (range2-9) make clean, aerial catches, once a small bird was
Gliding2 50 62 51 n.s.
snatchedoff therockwall of thegorge,andonce a female
(range29-87) (range28-79)
Flapping2 49 27 62** caughta Rock Pigeon while perched,as it tried to dis-
(range13-71) (range13-40) lodge her from a ledge (Jenkinsand van Zyl 1994).
Lanners Lannersattemptedto catchpreyin the air in 56 (90.3%)
Flying' 28 17 15 n.s. of the strikesobserved,andon six occasionseitherstruck
(range7-62) (range7-42) and retrievedprey, chased it out of cover on foot or
Gliding2 53 54 12 n.s. caughtit on the cliff face.
(range29-87) (range28-79) Overall,Peregrinesperchedmore/flewless thanLan-
Flapping2 16 12 16 n.s.
ners, flew faster and tendedto flap and glide whereas
(range9-28) (range5-25)
Lannersglided and soaredin thermals.Peregrineswere
%of timeobserved.
'Average significantlymoreaerialin windyconditions,were more
%of timeobservedflying.
2Average thanLanners,andhuntedover
longerdistancesat higherspeeds.
(Table5). Peregrines(especially males) made strikes
over longerdistancesandat greaterspeedsthanLanners,
and had higher wingbeat frequencies.Peregrinesflew
significantlymore,andof the timeflying, flappedsignif- Discussion
icantlyless in windy conditionsthanin calm conditions Possible differences between adults and
(Table6). Windspeeddid not significantlyinfluencethe
activitybudgetsof Lanners.Mean Peregrineflight dis-
tance indices were significantlygreaterin windy condi- Age-relateddifferencesin morphologyand flight per-
tions than in calm conditions(calm = 790 m, windy = formance have been reportedfor falcons previously
1743 m, t = 3.63, df = 53, P < 0.001) and their flights (Beebe 1960, Amadon1980, Cade 1982). Longerflight
were significantlylongerin duration(calm= 64 s, windy feathersand lighterwing loadingsin immaturebirdsare
= 141 s, t = 2.54, df = 41, P < 0.02). Lanner flight
thoughtto functionin reducingthe risk of injury(Ama-
distanceswere not significantlygreaterin windy condi- don 1980),increasingmanoeuvrability at low speedsand
tions, and flight durationswere not significantlylonger reducingthe energeticcosts of flight in inexperienced
(calm = 154 s, windy = 172 s, t = 0.36, df = 29). Observed birds(Muelleret al. 1981). Brown(1989) has suggested
home rangesof the two Peregrinepairs (0.85 km2and thatlowerwing loadingsalso may facilitatethe dispersal
0.81 km2)weresmallerthanthoseof thetwo Lannerpairs of independentyoung away from natal territories,over
(1.29 km2and 1.13 km2). habitatswhereflying conditionsare less favourablethan
Peregrineshuntedalmostexclusivelyin the gorge.On in areasoccupiedby residentadults.In this study such
two occasions they were seen slope soaringalong the age relateddifferenceswereevident(Table1) butmostly
edge of the gorge in the late evening,makingnumerous, were not statisticallysignificant,perhapsbecausesample
fast flightsat andthroughloose aggregationsof insectiv- sizes were too small and the degree of individualvar-
orousbats.Five batswerecaughtin thisway andeatenon iationwas too large.
the wing beforehuntingwas resumed.Otherwise,of 45
discretehunts observedat Peregrinesites, 11 (24.4%)
were successful,44 (97.8%)were perch-huntsand only
one (2.2%)was initiatedfromthe air. Lannersmay have Morphometrics,catching and killing
huntedaway fromthe gorge duringthe periodsthatthey Preyhandlingabilitiesarerelatedto foot andbill size in
were out of sight. All the Lannerhunting attemptsI falcons(Cade1982).The largerfeet of Peregrines(Table
observedweremadein the gorge,andtheywere success- 2) indicate that they are better equippedto catch and
ful in 14 out of 62 (22.6%)strikes.Of these 53 (85.5%) control prey in aerial hunts. Thomson(1984) suggests
were perch-huntsand nine (14.5%) were aerial hunts. that Peregrinesin Zimbabwe,southernAfrica are more

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exclusively aerial in their huntingmethodsthan sym- Peregrineswatchedat coastal sites on the Cape Penin-
patric Lanners.The sample of hunts observed in this sula,SouthAfricawereabsentfromthe immediatevicin-
study suggest that Peregrinesare more likely to catch ity of theirhome cliffs for 40-60% of the time observed
preyin open,aerialhuntsthanLanners,andthatLanners (butwereperchedfor about90%of the timetheywerein
are less stereotypedin their capturetechniques.Pere- sight) (Jenkins1987 and unpubl.data).
grines tend to have largerbills (Table2), and probably Calculatedoptimumflight speeds are faster for Pe-
have more massive bill musculature(e.g. Hull 1991), regrinesthanfor Lanners,and observedflight speed in-
suggestingthat they are more efficient than Lannersat dices, for both flights and strikes,also were higherfor
quicklykillingpreywhile still in the airaftera successful Peregrines(Table5). Speed indices of Peregrineswere
aerialhunt(Cade 1982). similarto absolutevalues measuredfor level-flyingand
A higher wing loading in Peregrines(Table2) may stoopingPeregrinesin otherstudies(about15 m s-' and
meanthatthey encountermoredifficultywhen tryingto 40 m s-' respectively - Cochran and Applegate 1986,
take-offfromthe groundthanLanners,especiallyif car- Alerstam1987, White and Nelson 1991), althoughmy
ryingprey.Certainlyliftingan equivalentweightinto the strikespeedestimatesprobablywere too low, since they
airwitha smallerflight-surfaceareais moreenergetically did not accountfor verticaldistancecovered.Although
demanding.This also may contributefurtherto the Pe- Peregrinestendedto fly fasterthanpredictedandLanners
regrine'stendencyonly to huntandcatchpreyin the air. slower,flight speedindiceswere similarto the predicted
cross-countryspeeds (Tables3 and 5), especiallygiven
that air density at the AFNP (not measured)probably
differed from the default value used to calculate the
Predicted vs observed flight performance
The flightperformanceinformationgeneratedby Penny- by Peregrinesthanby Lannersare attributable mainlyto
cuick's (1989) programs(Table3), whichpredictedsub- differencesin the wing loadingsof the two species(Cade
tle but significantdifferencesbetween the two species 1982, Norberg1986). Lanners,with lighterwing load-
based on consistentmorphometricdifferences,largely ings, shouldbe moreagile andcapableof fasteracceler-
was confirmedby field observations(Tables4-6). In ation than Peregrines(Anderssonand Norberg 1981,
orderto minimizethe energyexpenditureof flight,birds Norberg1986). Observationssuggestthatthis is not the
should use sources of atmosphericenergy as much as case. In most of the huntsI recordedat the AFNP,Pe-
possible to stay in the air. Lannersare able to circle in regrinesappearedto be moreagileandquickerto respond
smaller thermalsthan Peregrines,and apparentlywere to the evasive tacticsof prey, andusuallyhuntedsingly.
able to use this source of lift to a greaterextent at the Lannershuntedin pairs relativelyfrequently,and this
AFNP,andfly for longerperiods,over greaterdistances, may have enabledthem to catch birds which were too
and to foragefrom the air to a greaterextent (Table4). agile for individualsto easily catch on their own (e.g.
Peregrinesmostlywere restrictedto glidingandflapping Alpine Swifts). The higherwingbeatfrequenciesof Pe-
flight. Energetic constraintsshould require that birds regrines(Table5), presumablyattainablethrougha com-
glide wherepossibleratherthanflap. Tuckerand Parrot binationof smaller wings and more massive pectoral
(1970) define two goals of gliding flight: coveringdis- musculaturethan Lanners,probablyaccount for their
tance over the groundfrom one point to another,and to visibly superioraerialdexterity(AnderssonandNorberg
stay in the air by static or slope soaring.Under most 1981, Cade 1982).
conditions,a bird with a higherlift:dragratio (e.g. the Wingbeatfrequencyindices were about 25% lower
Lanner)will achieve these goals more easily than one thanvalues calculatedusing Pennycuick's(1990) equa-
with a lower lift:dragratio(e.g. the Peregrine).In calm tion (Peregrinemales 5.33 beats s-', Peregrinefemales
conditions,Peregrinesthereforeshouldbe forcedto cover 5.15 beats s-', Lannermales 4.56 beats s-' and Lanner
groundby flappingflight moreoften thanLanners.This females 4.34 beats s-'), and probablyeven lower than
apparentlywas the case at the AFNP (Table4). In terms actualfrequencies(see Pennycuicket al. 1994).
of fuel consumptionper unit distanceand per unit time
flown, flappingflightis moreenergeticallyexpensivefor
Peregrinesthanfor Lanners.HencePeregrinesshouldbe
far less aerial than Lanners,which was the case at the
AFNP.Also, theorypredictsthat a bird with relatively
Flight performance,foraging mode and habitat
high optimumglide speedsand a relativelyhigh sinking
speed (e.g. the Peregrine)glides over distancerelatively JaksicandCarothers(1985) foundthathigherwing load-
efficiently into a stronghead wind (Tuckerand Parrot ings in raptorspredicteda sit-and-waitratherthanactive
1970). Peregrinesat the AFNP were significantlymore search hunting mode. This suggests that Peregrines
aerialin windy conditions(Table6). Both species prob- shouldbe more sedentaryhuntersthanLanners,prefer-
ably were more sedentaryin the rivergorge situationat ringto huntfroma perchratherthansearchfor preyfrom
the AFNPthantheymightbe in otherconditions,perhaps the air.This was confirmedby the smallsampleof hunts
whereprey are less spatiallyconcentrated.For example, observedin this study,andby the significantdifferencein


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the time the two species spent flying. Given the relatively grines are 17.0 and 19.3 suggesting that in relative terms,
high cost:benefit ratio of foraging for bird-eating raptors powered flight is about 20% more strenuous.
(Temeles 1985), additional energetic constraints on flying Differences in the morphology and flight performance
for Peregrines are likely to restrict foraging mode to the of southern African Peregrine and Lanner Falcons corre-
optimum, and increase habitat selectivity. Hence, Pe- spond to differences in their feeding ecologies, distribu-
regrines should be more restricted than Lanners to areas tion and abundance. The greater degree of morphological
where suitably high vantage points for perch-hunting are specialization in Peregrines means that, although they are
abundant (e.g. mountainous areas where high cliffs are better adapted to the high speed, open aerial pursuit of
frequent). This is the case in South Africa, where Pe- birds than Lanners, they are correspondingly less effi-
regrine distribution follows the distribution of cliff avail- cient in other hunting and flying modes and less able to
ability more closely than sympatric Lanners, and Pe- exploit other food sources. Thus, in poor food areas the
regrine habitat selectivity indices are higher (Jenkins in Peregrine Falcon may either be absent or restricted to
press). habitats which particularlyfavour their specialized hunt-
Differences in the primary foraging modes of Pere- ing mode. Lanner Falcons are less likely to be restricted
grines and Lanners should result in corresponding differ- in this way.
ences in their feeding rates and diets (Huey and Pianka In a morphometricstudy of the genus Falco, Kemp and
1981). Although no comprehensive data on the diets of Crowe (1991) found that smaller, tropical races of Pe-
the two species are available, provisioning rates at nests regrines (e.g. F p. minor and F. p. peregrinator) are
of Peregrines and Lanners in South Africa are signif- relatively stocky, short tailed, short winged and large
icantly different (Jenkins 1992). Also, as suggested by footed, and group with the morphologically extreme Or-
Norberg (1977), Peregrines, as sit-and-wait predators, angebreasted F deiroleucus and Taita Falcons E fascii-
may be expected to be more habitat selective and more nucha, whereas larger, northern races (e.g. F. p. pealei)
sedentary in their hunting methods as food availability have relatively longer wings and tails, and are more
decreases. Such a trend has been suggested for Peregrine closely allied to the desert or hierofalcons (Cade 1982)
populations globally, in relation to food availability at (e.g. F biarmicus). Hence, morphological differences of
different latitudes (Jenkins 1991). a similar nature to those illustrated here for southern
The difference in the two species' abilities to utilize African Peregrines and Lanners, and with similar func-
thermals has furtherimplications for their respective hab- tional, behavioural and ecological implications, may oc-
itat preferences and dispersability. Since Lanner Falcons cur between different races of F peregrinus, as well as
are relatively efficient users of thermals, they may be between Peregrines and congeners in other parts of the
resident in low relief areas where thermals are the pri- world. This may partly explain broad-scale patterns of
mary source of lift for active search hunting and cross- distribution and abundance of large falcons.
country flying. In contrast, Peregrines are relatively poor
users of thermals and may prefer areas of higher relief Acknowledgements - Thanksto Tim Wagner,Ian Hoffmanand
because of the slope soaring opportunities they provide the TransvaalFalconryClub, and Tom Davidsonof the Natal
FalconryClub,for allowingme to measuretheircaptivebirds.
(see also Pennycuick and Scholey 1984). Barnard(1986) Greg McBey and Tim Wagnerprovidedassistanceand advice
found that a montane raptor with a higher wing loading with the trappingof wild birds, and Zelda Bate helped and
was significantly more dependent on slope lift generated accompaniedme in thefield.TheNationalParksBoardof South
Africakindlyallowedme workin theAFNP,andPeterNovellie
by wind for prolonged, energetically efficient foraging andNico vanderWaltof ParksBoardwerehelpfulandcooper-
than a sympatric raptorwith a lower wing loading. Active ative. Roy Siegfriedand two anonymousrefereesmadeuseful
search hunting in Peregrines probably is similarly con- comments on drafts of this manuscript,and GerardMalan
strained. helped with the statisticalanalyses. This project was partly
There is evidence for local migratory movements of fundedby the Foundationfor ResearchDevelopment.
southern African Lanners (Steyn 1982, Van Zyl et al. in
press), but none has been reported for Peregrines. This
may be because African Peregrines are less efficient
flyers over low relief areas, and these may act as barriers References
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