Extra Supplementary Material

Heat transfer model of emperor penguin

Surface area and characteristic dimensions
A simple geometric model of an emperor penguin Aptenodytes forsteri
was represented by main body (prolate spheroid), head (sphere), beak
(cone), flippers and feet (flat plates) (Fig. 1). A specimen (Hunterian
Museum, Catalogue: Z1125) mounted in a realistic upright stance was
used to provide representative body dimensions and body surface areas
for heat transfer calculations.

Atrunk
(m2) was determined by:

The surface area of the main body trunk

Atrunk 2

ab 1
sin e 2b 2
e

e
where

b2

and a and b are the semi-major

and minor axes lengths, respectively [1]. a was taken to be half the body
length from the black neck collar to the base of abdominal feathers (0.34
m), b was half the maximum diameter of the body trunk (0.16 m)
determined from a measurement of girth taken under the flippers. The

Abeak rs
beak area was a cone

where the radius r was half the maximum

width at the base of the beak (0.01 m) and hypotenuse s was

approximated by beak length (0.11m). The area of the head was
approximated by a sphere minus the base of a cone (beak):
Ahead d 2 r 2

where the diameter d was taken to be the mean (

standard error) of head height and width (0.110.004 m) measured from
the bottom of black head plumage and r as above. Flippers were
A flipper l w

approximated as flat rectangles of length l and width w,

and

were determined by tracing outlines onto 1mm squared graph paper. The
flipper length was the maximum diagonal length from top to bottom of
flipper (0.280.011 m) and mean flipper width (0.0650.0002 m).
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The surface area of each foot was obtained by tracing its outline onto 1
mm2 graph paper. The mean surface area of a single foot in contact with
the ground was 0.00360.0004m2. The total surface area of a foot was
therefore approximated as twice the measured area. As emperor penguins
often rest on their tarsometatarsus joint, this area was also traced from
the museum specimen and averaged 0.00060.00008 m2 or 17% of the
lower surface area of the foot. For radiation and convective calculations
the characteristic dimension of the foot was taken to be maximum foot
width, f (0.0560.0025 m). The thickness of the foot, t was determined
from the mean thickness of metatarsi (mean = 0.014 m SE = 0.001). The
total surface area of an average emperor penguin was calculated to be
0.56 m2 which was within previously measured values [1, 2].

Surfac % Total
e

surface

Area

area

d (m)

Nusselt Relationship

(m2)
Trunk (excluding flippers)

0.471

83.8

0.32

Prolate Nu =0.24Re0.6

Head and beak

0.040

7.2

0.11

Sphere Nu =0.34Re0.6

Flippers (outside surface

0.036

6.5

0.065

Flat plate Nu
=0.032Re0.8

only)
Feet

0.014

2.5

0.056

Flat plate Nu =
0.032Re0.8

Total

0.562

Table 1. Calculated surface area, percentage of total surface area,

characteristic dimensions for heat transfer calculations of emperor
penguin and relationship between Nusselt and Reynolds numbers [7].

Heat transfer
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A distributed parameter heat transfer model was used to estimate total

qtot
heat exchange,

(W) for an emperor penguin by summing heat transfer

from each body region (head, trunk, flippers and feet) assuming that the
penguin was in thermal equilibrium with its surroundings [3]:
q tot q head qtrunk q flippers q feet

(1)
Radiation
As measurements were undertaken during mid-winter (1h50-2h50 light/24h)
solar heat gain was assumed to make a trivial contribution to heat transfer.

q rad
Radiative heat loss

was determined by solving the radiation balance at

the surface. Heat loss by radiation was the difference between radiation

qbird
emitted from the penguins surface,

and radiation gained from the

q env
environment,

such that:

q rad qbird q env

(2)
Radiation emitted from each body part of surface area, A with surface
temperature Ts (K) was determined according to:
qbird ATs

(3)
Where

is emissivity of bird plumage (=0.98, [4]) and

is the Stefan-

Boltzmann constant (5.67 x10-8 Wm-2K-1). We assumed that each part of the
body exchanged radiation equally with sky and surroundings such that
amount of radiation absorbed was equal to the mean flux from sky and
ground surface:

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Ld Lu

qenv Aal

(4)

where A (m) is the radiative area and, al is the long wave absorptivity
(=emissivity) of the penguin. Ld (Wm-2) and Lu (Wm-2) are the downward
and upward radiative heat fluxes from sky and snow surface, respectively.
The downward radiative flux was estimated using the empirical
relationship measured in Antarctica [5]:

Ld 0.665Ta 18.175h 8.003c 14.088

(5)
where Ta is air temperature (K), h the specific humidity (gkg-1) and c the
cloud cover (oktas).
The upward radiative flux was determined by:

Lu g Tg

(6)

g
where

is the emissivity of ground surface (snow = 0.97, [6]) and Tg (K)

the ground ice temperature.

Convection

qconv
Heat transfer by convection,

from each region of the body was

calculated by:

q conv hA Ts Ta
(7)
Forced convection is the dominant mode of heat transfer in wind ( 0.5 ms-1
in this study) such that the heat transfer coefficient was determined by:

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h Nu

k
d

(8)

where k is the thermal conductivity of air (0.0225 Wm -1oC-1 at -20oC), d (m)

is the characteristic dimension of each body part in the direction of air flow
and Nu is the dimensionless Nusselt number. The Nusselt number is a
measure of the ratio of buoyant to viscous forces. It depends on shape and

can be related to the dimensionless Reynolds number,

Re

Re ud
from

where u is the wind speed (ms-1), d the characteristic dimension (m) and

the kinematic viscosity of air (11.6 x 10-6 m2s-1 at -20oC). The relationship
between Nu and Re has been determined empirically for a range of
geometric shapes and flow regimes (Table 1).
Conduction
Emperor penguins commonly cover the upper surface of their feet by
abdominal feathers and therefore a bird will lose heat by conduction, qcon
from the lower surface of its feet to the snow surface such that:

q con Ak

(T feet Tg )
x

(9)

where k is the thermal conductivity of the foot tissue (Wm-1oC-1) of

thickness x (m). The thermal conductivity of foot tissue was taken to equal
skin conductivity (0.502 Wm-1oC-1) [8]. Emperor penguin feet remain above
freezing and the minimum heat loss by conduction was estimated from
minimum foot temperature of 3.3C [9]. The temperature of ground
surface underlying the foot was assumed to equal the surface
temperature of surrounding snow surface. If feet were not visible it was
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assumed birds were resting on their tarsometatarsus where area in

contact with ground was 17% of foot area. When standing or walking, the
foot will also lose heat by radiation and convection from the upper surface
(as above).
Latent heat loss
Latent heat loss for emperor penguins was estimated from previous
measurements of the evaporative water loss that remained constant
between -47 and 20 oC and averaged 5.85 g h-1 [1]. The vaporisation of 1g
water requires 2.43 kJ, therefore latent heat loss for an emperor penguin
is equivalent to 4.0 W.
Data
Input data for the model was taken from surface temperature
measurements of emperor penguins and meteorological data recorded at
the breeding colony of Pointe Gologie in Terre Adlie (66o40S 140o 01E),
Antarctica in June 2008 (ESM2). Where surface temperature data were
missing for a particular body part for an individual, the missing value was
computed using regression with air temperature from GLM models (see
paper).
References
1.

Pinshow, B., Fedak, M.A., Battles, D.R., Schmidt Nielsen, K. 1976

Energy-expenditure for thermoregulation and locomotion in emperor

penguins. Am. J. Physiol. 231,903-12.
2.

Le Maho, Y., Delclitte, P., Chatonnet, J. 1976. Thermoregulation in

fasting emperor penguins. Am. J. Physiol. 231,913-22.

3.

McCafferty, DJ, Gilbert, C, Paterson, W, Pomeroy, PP, Thompson, D,

Currie, J, Ancel, A. 2011 Estimating metabolic heat loss in birds and

mammals by combining infrared thermography with biophysical
modelling. Comp. Biochem. Physiol.A-Molec. Integ. Physiol. 158, 337-345.
4.

Hammel, H.T. 1956 Infrared emissivities of some arctic fauna. J.

Mammal. 37, 375-8.

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5.

Cho, H.K., Kim, J., Jung, Y., Lee, Y.G., Lee, B.Y. 2008 Recent changes

in downward longwave radiation at King Sejong Station, Antarctica. J.

Climate. 21, 5764-76.
6.

Kondo, J., Yamazawa, H. 1986. Measurement of snow surface

emissivity. Boundary-Layer Met. 34,415-6.

7.

Monteith, J.L., Unsworth, M.H. 1990 Principles of environmental

physics. London: Edward Arnold.

8.

Gates, D.M. 1980 Biophysical Ecology. Berlin: Springer Verlag.

9.

Prvost, J, Sapin-Jaloustre, J. 1964 A propos des premieres mesures

de topographie thermique chez les Spheniscides de la Terre Adelie.

Oiseau. 34, 52-90.

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Fig.1. Geometric model of emperor penguin. Definition of terms given in text.