Regional variation in body size of the cheetah

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Regional variation in body size of the cheetah (Acinonyx jubatus

)

Author(s): Lorraine K. Boast, Ann Marie Houser, Kyle Good, and Markus Gusset

Source: Journal of Mammalogy, 94(6):1293-1297. 2013.

Published By: American Society of Mammalogists

DOI: http://dx.doi.org/10.1644/13-MAMM-A-076.1

URL: http://www.bioone.org/doi/full/10.1644/13-MAMM-A-076.1

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Journal of Mammalogy, 94(6):12931297, 2013

Regional variation in body size of the cheetah (Acinonyx jubatus)

LORRAINEK. BOAST,* ANN MARIE HOUSER, KYLEGOOD, AND MARKUSGUSSET

Cheetah Conservation Botswana, Private Bag 0457, Gaborone, Botswana (LKB, AMH, KG)

Animal Demography Unit, Department of Biological Sciences, University of Cape Town, Rondebosch 7701, South Africa

(LKB)

Centre for Wildlife Management, University of Pretoria, Pretoria 0028, South Africa (AMH)

Wildlife Conservation Research Unit, Department of Zoology, University of Oxford, Abingdon OX13 5QL, United

Kingdom (MG)

* Correspondent: [email protected]

Body size affects almost every aspect of the biology of a species, with considerable intraspecific variation.

Cheetahs (Acinonyx jubatus) reportedly vary in body size across their geographical range. However, because

morphometric measurements were not taken in a standardized manner, it is impossible to rule out differences in

measurement protocols as the cause. Our study differed from previous ones in that we made use of a standardized

methodology for taking morphometric measurements in cheetahs. Free-ranging cheetahs in Namibia were shorter(3.54.1%) and slimmer (4.07.0%) than those in neighboring Botswana. Cheetah density was more than 3 times

higher and home-range sizes were more than 3 times smaller in Botswana compared to Namibia. This suggests

that variation in resource availability may be the main driver of the fine-scale spatial differences in morphometric

measurements. Overall, our study promotes the use of standardized protocols for measuring morphological traits

in free-ranging animals.

Key words: Acinonyx jubatus, body size, Botswana, cheetah, resource rule, sexual size dimorphism

2013 American Society of Mammalogists

DOI: 10.1644/13-MAMM-A-076.1

Body size affects almost every aspect of the biology of a

species, from physiology and life history to ecology (Roy

2008). There is considerable intraspecific variation in body size

with respect to geography and time (Yom-Tov and Geffen

2011). According to the resource rule (McNab 2010), the

abundance, availability, and size of resources (e.g., prey

biomass) are the main drivers of such spatial and temporal

variation. For example, free-ranging African wild dogs

(Lycaon pictus) decreased in body size by up to 17% over a

20-year period, concurrent with a significant decline in the

density of their main prey species (McNutt and Gusset 2012).

Other recent studies also linked a reduction in predator body

size to a possible decrease in prey availability (Yom-Tov et al.

2007, 2010; Rode et al. 2010).

In free-ranging animals, intraspecific comparison of body

size across study sites and periods is often confounded, because

morphometric measurements are rarely taken in a standardized

manner (De Waal et al. 2004). Cheetahs (Acinonyx jubatus)

show considerable variation in body size across their

fragmented geographical range spanning throughout Africa

and into Asia (Caro 1994). Klein (1986) found carnassial

length to vary with latitude, adhering to Bergmanns rule that

individuals tend to be larger in cooler climates. Cranial

measurements suggest that individuals from North Africa are

smaller than those from sub-Saharan Africa (Saleh et al. 2001).

Within sub-Saharan Africa, Marker and Dickman (2003) found

various morphological traits to vary in size across study sites.

However, whether this geographical variation in body size is

real and related to, for example, resource availability or is

merely an artifact of using different measurement protocols is

unknown. In our study, we made use of a standardized

methodology developed for taking 19 morphometric measure-

ments from more than 200 free-ranging cheetahs in Namibia

(Marker and Dickman 2003). For the 1st time, this allowed us

to assess regional variation in cheetah body size without the

confounding variable of measurement protocol, as exemplified

by a comparison between our study sites in Botswana and

those in neighboring Namibia.

In the absence of comparative data on prey biomass from the

respective study sites in Botswana and Namibia, we used 2

w w w . m a m m a l o g y . o r g

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proxies for resource availability: cheetah density and home-

range size. Prey biomass strongly predicts cheetah density

(Hayward et al. 2007) and is significantly negatively related to

the size of a cheetahs home range (Marker 2002). Although a

direct comparison between resource availability and body size

would have been preferable, the scaling relationships of prey

biomass, body size, population density, and home-range sizeamong carnivores (Gompper and Gittleman 1991; Carbone and

Gittleman 2002; Ferguson and Lariviere 2008) justify the use

of proxies. We hypothesized that if there are country-specific

differences in the 2 proxies for resource availability, the body

size of cheetahs in Botswana and Namibia will differ as well.

MATERIALS AND METHODS

Capture and care.A total of 64 cheetahs were captured in

the Southern, Kgalagadi, and Ghanzi districts of Botswana

between October 2003 and July 2012. Cheetahs were live-

captured in locally manufactured double-ended box traps (2.0

3 0.8 m), using limited access or bait trap sets. Limited access

traps used Acacia spp. cuttings to block access to waterholes,

marking trees, or along fence lines. Cheetahs were transported

in wooden squeeze boxes (1.2 3 0.8 m) to Cheetah

Conservation Botswana research bases and kept temporarily

in holding pens (20 3 40 m) until a medical workup could be

arranged. Cheetahs were held in captivity for 4.7 6 5.0 (range

121) days between capture and medical workup (animals held

in captivity for . 30 days were not included in the study

[Marker and Dickman 2003]).

Morphometric measurements also were taken from 5 dead

cheetahs (3 of which were euthanized after being hit by

vehicles). The absence of rigor mortis or bloating enabled allmeasurements to be collected on 4 of the 5 animals.

Postmortem changes on the 5th animal prevented the accurate

measurement of abdomen girth, total foreleg length, and total

hind-leg length. In total, morphometric measurements from 40

males and 29 females were taken. Handling procedures

followed guidelines of the American Society of Mammalogists

(Sikes et al. 2011).

Medical workup .Cheetahs were anesthetized using

medetomidine (Domitor; Pfizer Inc., New York City, New

York; 3040 lg/kg) and tiletaminezolazepam (Zoletil; Virbac,

Carros, France; 1 mg/kg), using a hand syringe in the squeeze

boxes or by dart gun in the holding pens. A physical health

check was performed and cheetahs were deemed to be inexcellent, good, fair, or poor health, using methods adapted

from Marker (2002). Superficial trap cage injuries were not

considered, because they do not reflect on wild cheetahs health

status. Cheetahs were assigned to 1 of 8 age classes based on

body mass, teeth wear and discoloration, gum recession, coat

condition, social grouping of animals caught together, and

reproductive condition (Marker and Dickman 2003). Cheetahs

. 30 months old were collectively referred to as adults ( n20

males and 15 females).

Morphometric measurements.Body mass was measured

using a hanging balance, with the animal placed in a stretcher,

to the nearest 0.5 kg. Another 18 measurements were taken as

described by Marker and Dickman (2003). In addition,

shoulder to point of the elbow (most dorsal point of scapula

to high olecranon) and point of the elbow to heel (high

olecranon to base of foot), as well as most dorsal point of hip to

knee (top of ilium to patella) and knee to heel (patella to base

of foot) were measured.Canine length, as well as skull, foot, and testis length and

width were measured with vernier calipers to the nearest 0.1

cm. All other measurements were taken using a 200-cm

measuring tape to the nearest 0.5 cm. Leg measurements were

taken with the leg in a normal walking position. A mean was

calculated for measurements taken on both sides of the body.

To test for reliability, measuring the same 2 animals 4 times in

succession showed a small dispersion of each measurement. To

avoid possible measuring errors from influencing data analysis,

any measurement outside of 2 standard deviations of the mean

was removed (3.8% of adult measurements); a new mean and

standard deviation were then calculated.

Data analysis.Sexual size dimorphism in adults was tested

for using all morphometric measurements. Development of

body mass, body length, chest girth, and total foreleg length in

males and females was compared over the age classes.

Correlation between chest girth and body mass was

calculated for all age classes combined and for adults only.

Measurements of Botswana cheetahs were compared with

data from 99 male and 39 female live or recently deceased

adult cheetahs from Namibia. All Namibian cheetahs had been

in captivity for, 30 days and 86% were in excellent or good

physical condition (Marker and Dickman 2003). Morphometric

measurements between Botswana and Namibian cheetahs were

compared with cheetah density (based on known individuals)and home-range sizes (based on 95% fixed kernels in Botswana

and 95% adaptive kernels in Namibia) using available data

from the respective study sites (Cheetah Conservation

Botswana, pers. comm.; Marker 2002; Marker and Dickman

2003; Marker et al. 2008a, 2008b; Houser et al. 2009a, 2009b;

Kent 2011).

Data are presented as mean 6 SD. KolmogorovSmirnov

tests revealed a normal distribution of the datas residuals. All

statistical tests (unpairedt-test and Pearson correlation) were 2-

tailed, with the significance level set atP , 0.05, and were run

in SYSTAT, version 12 (Systat 2007).

RESULTS

The majority of live cheetahs (n 64) were in excellent

(26.6%) or good (65.6%) health. Only 1 cheetah was in poor

health, suffering from an unknown respiratory infection. All

but 1 of the 23 morphometric measurements were significantly

larger in adult males compared to adult females (Table 1).

Males were generally larger than females with regard to body

mass, body length, and chest girth, but not total foreleg length,

from . 12 months old (Fig. 1). Females reached their full body

size at 3148 months old, whereas males did not reach their

full size until . 48 months old. There was a strong positive

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correlation between chest girth and body mass for all age

classes combined (n 68,r 0.95,P , 0.0001), with a linear

regression equation of body mass 1.213 chest girth 46.84

(r2 0.90). This correlation remained significant when only

adults were considered (n 34, r 0.84, P , 0.0001), in

which case the linear regression equation became body mass

1.263 chest girth 49.84 (r2 0.70).

When comparing the body size of adult cheetahs between

Botswana and Namibia, 13 (68.4%) of the 19 morphometric

measurements for both males and females were larger in

Botswana. Compared to Namibia (Marker and Dickman 2003),

cheetahs in Botswana had a significantly larger chest girth in

males and females, tail length in males, total length in males,

and testis length and width, but had a significantly smaller total

hind-leg length in females (Table 2). Cheetah density (3.2 6

3.1 versus 0.9 6 0.7 individuals/100 km2) was more than 3

times higher in Botswana, but due to small samples sizes this

difference only approached statistical significance (nBotswana

8,nNamibia 9,t 2.07 [Welch corrected], P 0.077). Home-range sizes (517.9 6 263.4 versus 1,651.1 6 1,594.2 km2)

were more than 3 times smaller in Botswana (nBotswana 5,

nNamibia 27, t 3.45 [Welch corrected], P 0.002).

DISCUSSION

We found sexual size dimorphism in cheetahs from

Botswana, as well as regional variation in cheetah body size

between Botswana and Namibia. Sexual size dimorphism, with

males being larger than females, has been reported for cheetahs

across their geographical range (Caro 1994; Marker and

Dickman 2003). In our study, sexual dimorphism in body

mass, chest girth, and body length was evident from . 12

months old; however, total foreleg length showed more

developmental variability. This contrasts with the aging scale

detailed by Caro (1994), which states that it is possible to

distinguish male and female cubs by shoulder height from 7

months old onward. We found females reached full body sizeas young adults (3148 months old), compared to as prime

adults (. 48 months old) in males. This coincides with the

findings of Caro (1994) that females 1st reproduce at 37

months of age, whereas males are unable to hold and defend a

territory, and presumably mate, until . 48 months old.

Similar to polar bears (Ursus maritimusDurner and

Armstrup 1996), Marker and Dickman (2003) stated that chest

girth can be used to extrapolate body mass in cheetahs when

weighing is not feasible. We also found a significant

correlation between chest girth and body mass. This correlation

was strongest when all age classes were considered; however,

age may influence the chest girthbody mass relationship

(Cook et al. 2003). Therefore, it may be advisable to use the

regression equation derived for adults. Nevertheless, because

chest girth in Botswana was larger than in Namibia, with

similar body mass, applying this equation to Namibian

cheetahs would underestimate their body size. Similar

geographical variation has been observed in polar bears

(Durner and Armstrup 1996), which emphasizes that the

appropriate regression equation for age and population should

be applied when estimating body mass from chest girth.

Cheetahs were previously reported to vary in body size

across their geographical range (Caro 1994; Marker and

Dickman 2003). However, because morphometric measure-

ments were not taken in a standardized manner, it wasimpossible to rule out differences in measurement protocols as

the cause. Our study differed in that we made use of a

previously developed standardized methodology for taking

morphometric measurements in cheetahs (Marker and Dick-

man 2003). We found regional variation in body size,

especially in males. Testes are difficult to measure reliably

(De Waal et al. 2004) and measuring total hind-leg length relies

on the subjective positioning of the leg in a walking stance

(Marker and Dickman 2003), which are likely to have

contributed to the observed differences. Because tail length is

a part of total length (Marker and Dickman 2003), differences

in these 2 measurements are interrelated. Together with the

observed difference in chest girth, cheetahs in Namibia wereshorter (3.54.1%) and slimmer (4.07.0%) than those in

Botswana. Similarly, wild dogs too became shorter and

slimmer with decreasing resource availability (McNutt and

Gusset 2012).

Concurrent with regional variation in cheetah body size

between Botswana and Namibia, our 2 proxies for resource

availability showed country-specific differences. Considerably

higher cheetah density and smaller home-range sizes in

Botswana are indicative (Marker 2002; Hayward et al. 2007)

of higher prey biomass in Botswana. As a caveat, it should be

remembered that these data were collected at the respective

TABLE1.Morphometric measurements taken from adult cheetahs

(Acinonyx jubatus) in Botswana ( X6 SD) and test values for sexual

size dimorphism (n 1620 for males and 1215 for females).

Measurement (unit) Males Females t-value P-value

Body mass (kg) 46.7 6 8.7 38.5 6 3.5 3.77a 0.0008

Upper-canine length (cm) 2.3 6 0.2 2.1 6 0.2 2.57 0.015

Lower-canine length (cm) 1.7 6 0.2 1.5 6 0.2 2.98 0.005Skull length (cm) 23.0 6 1.2 22.6 6 1.7 0.62 0.543

Skull width (cm) 14.6 6 0.8 13.1 6 0.9 4.77 , 0.0001

Muzzle girth (cm) 27.2 6 1.5 25.1 6 1.7 3.88 0.0005

Chest girth (cm) 77.1 6 4.7 70.1 6 3.3 4.81 , 0.0001

Abdomen girth (cm) 58.6 6 5.0 55.2 6 3.6 2.14 0.041

Body length (cm) 128.9 6 7.2 122.2 6 6.2 2.79 0.009

Tail length (cm) 80.0 6 3.6 73.5 6 7.2 3.23a 0.004

Total length (cm) 209.5 6 10.3 195.7 6 9.5 4.02 0.0003

Total foreleg length (cm) 77.5 6 3.7 73.3 6 4.0 3.18 0.003

Shoulderelbow length (cm) 41.0 6 2.4 38.0 6 1.6 4.16 0.0002

Elbowheel length (cm) 42.0 6 2.0 40.4 6 2.1 2.32 0.027

Total hind-leg length (cm) 79.3 6 4.4 74.2 6 5.5 2.98 0.006

Hipknee length (cm) 37.1 6 2.6 34.4 6 3.6 2.55 0.016

Kneeheel length (cm) 46.7 6 2.0 44.2 6 2.1 3.36 0.002

Front-foot length (cm) 8.1 6 0.4 7.8 6 0.4 2.17 0.038Front-foot width (cm) 6.2 6 0.3 5.9 6 0.3 3.27 0.003

Hind-foot length (cm) 9.3 6 0.5 8.9 6 0.3 2.33 0.026

Hind-foot width (cm) 6.3 6 0.4 5.9 6 0.3 2.62 0.013

Testis length (cm) 3.3 6 0.3

Testis width (cm) 2.5 6 0.3

aWelch corrected (variances are not equal).

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study sites where the morphometric measurements were taken,

and thus may not be representative of the 2 countries as a

whole. Nevertheless, in accordance with the predictions of the

resource rule (McNab 2010), our findings suggest that greater

resource availability in Botswana (with no indication of

stronger intraspecific competition) may be causal to the

generally larger body size of cheetahs in Botswana compared

to Namibia. Our findings also are consistent with the pattern

found in mammals for which fluctuations in food supply

differentially constrain growth patterns of the sexes (Isaac

2005), because male body size regionally differed more

substantially than that of female cheetahs (also see McNutt

and Gusset 2012).

Furthermore, there is fine-scale genetic substructuring in the

southern African cheetah subspecies (A. j. jubatus), including

differences indicative of limited gene flow between Botswana

and Namibia (Kotze et al. 2008; Charruau et al. 2011). These

genetic differences were found in neutral markers that do not

code for morphological traits. Nevertheless, phenotypically

unique lions (Panthera leo) from Ethiopia, with smaller body

size and mass, also differed genetically at neutral markers from

all other lions investigated (Bruche et al. 2013). Therefore, in

addition to possible differences in resource availability, there

might be a genetic component to the regional variation in

cheetah body size we found between Botswana and Namibia.

In conclusion, the use of standardized methodology allowed

us to assess regional variation in cheetah body size without the

confounding variable of measurement protocol, as exemplified

by a comparison between our study sites in Botswana and

those in neighboring Namibia. Variation in resource availabil-

ity may be the main driver of the observed fine-scale spatial

differences in morphometric measurements (McNab 2010;Yom-Tov and Geffen 2011), but a direct comparison between

resource availability and body size would be desirable. Overall,

our study promotes the use of standardized protocols for

measuring morphological traits in free-ranging animals (e.g.,

Marker and Dickman 2003; De Waal et al. 2004), to enable

intraspecific comparisons of body size across study sites and

periods.

ACKNOWLEDGMENTS

We are grateful to Botswanas Ministry of Environment, Wildlife

and Tourism and the Department of Wildlife and National Parks for

TABLE2.Morphometric measurements ( X6 SD) taken from adult

cheetahs (Acinonyx jubatus) in Botswana and Namibia that showed

significant regional variation (Botswana: n 1820 for males and 14

15 for females; Namibia: n 8194 for males and 38 for females).

Measurement

(cm) Sex Botswana Namibia a t-value P-value

Chest girth Male 77.1 6 4.7 71.7 6 3.9 5.39 , 0.0001

Tail length Male 80.0 6 3.6 76.7 6 5.2 3.41b 0.002

Total length Male 209.5 6 10.3 202.2 6 9.4 3.04 0.003

Testis length Male 3.3 6 0.3 2.8 6 0.5 5.74b , 0.0001

Testis width Male 2.5 6 0.3 2.0 6 0.2 7.08b , 0.0001

Chest girth Female 70.1 6 3.3 67.3 6 3.8 2.44 0.019

Total hind-leg

length Female 74.2 6 5.5 77.8 6 2.8 2.42b 0.028

aFrom Marker and Dickman (2003).

bWelch corrected (variances are not equal).

FIG. 1.Development of a) body mass, b) body length, c) chest girth, and d) total foreleg length in male and female cheetahs (Acinonyx

jubatus) from Botswana ( X6 SD) over the age classes (n 212 per age class).

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permission to conduct the study and for continuing assistance. The

study was supported by Cheetah Conservation Botswana, especially

R. Klein, and the farming community of Ghanzi. We thank Debswana

for allowing us to work and collect data in Jwana Game Reserve. The

manuscript benefitted from comments provided by L. Marker and an

anonymous referee.

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Associate Editor was I. Suzanne Prange.

December 2013 1297BOAST ET AL.CHEETAH BODY SIZE

Regional variation in body size of the cheetah

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