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Camp. Biochem. Physid. Vol. 80A, No. 3, pp. 355-358, 1985 0300-9629/U $3.00 + 0.

00
Printed in Great Britain 0 1985 Pergamon Press Ltd

FOOD AND ENERGY REQUIREMENTS OF CAPTIVE


BARN OWLS TYTO ALBA
KIRK L. HAMILTON*
Department of Biology and the Ecology Center, UMC 53, Utah State University, Logan, UT 84322, USA

(Received 5 June 1984)

Abstract-I. Food and energy ~quirements of captive barn owls were determined at 5. 15 and 25°C.
2. Food consumption, gross energy intake, dry matter intake, and existence metabolism increased as
ambient temperature decreased.
3. Barn owl energy assimilation efficiencies at the three test temperatures were approximately 78%.
4. An increase in bioenergetic parameters might be anticipated when a bird is cold-stressed and must
increase its metabohsm for ~e~ore~lation.

INTRODUCTION determined with a Phillipson microbomb calorimeter


(Gene-Wiege~ Instrument, Inc., South Carolina).
Many raptor food and energy requirement studies Live mice, food scraps, pellets and excreta were collected
have been conducted to determine food and gross separately at the end of each food trial and dried m a
energy intake, existence metabolism and energy as- convection oven (SO-60°C). Pellets were measured (to the
similation efficiencies (Graber, 1962; Ligon, 1969; nearest 0.1 mm, vernier caliper) as described by Hamilton
Gessaman, 1972; Duke et al., 1973; Marti, 1973; (1978). The dry material was weighed to the nearest 0.1 g,
Johnson, 1974; Barrett and Mackey, 1975; Wallick .around in a Wiley_ mill, homogenized
_ (wig-I-hue
\., uamalpam-
I

and Barrett, 1976; Kirkwood, 1979; Koplin et al., ator, Model 5AR, Crescent Dental Mfg. Co., Chicago,
Illinois), and energy content determined (in triplicate) by
1980; Hamilton and Neill, 1981; Postler and Barrett, microbomb calorimetry. An owl’s gross energy intake
1982). None have examined all four bioenergetic (GEI), during a feeding trial, was determined by subtracting
parameters in the barn owl (Gyro a&a). the energy content of the food scraps’ from the energy of the
In the present study, the food and energy require- food provided (Kendeigh et al., 1977). Existence metabolism
ments of captive barn owls have been examined. (EM) was derived as GE1 - pellet energy (PE) - excretory
energy (EE). Energy assimilation eficiency (EAE) was
MATERJALS AND METHODS calculated by EM/GE1 x 100 (Gessaman, 1972, 1973; Stal-
master and Gessaman, 1982).
Two adult owls (7j~to &a) were captured in April 1980 Statistical analysis of data presented here include analysis
at Welder Wildlife Foundation (Sinton, San Patricia Co., of variance and Student-Newman Kuels Test.
Texas) and a third owl was obtained from a local raptor
rehabilitator (MS S. ,Ure), and transported to the Environ-
mental Physiology Laboratory at Utah State University. In
1981, three adult owls were captured post-incubation RESULTS AND DISCUSSION
(April-May, Brigham City, Box Elder Co., Utah), and
Food consumption trials were conducted on six
likewise were transported to the laboratory facilities. All
owls were housed in separate 3 x 3 x 2.5 m walk-in environ- captive barn owls (l-6) to determine the gross food
mental chambers. Owls were maintained on a 12L:12D and energy intake, existence metabolism, and energy
photoperiod during all feeding trials. assimilation at 5, 15 and 25°C. Table 1 shows food
Food and energy consumption and energy assimilation by trial duration and weights of barn owls used during
barn owls were determined for one diet (laboratory mouse), the trials.
at 3 temperatures (5, 15 and 25°C). The temperatures were
repre~ntative of those experienced during the incubation Water a& catoric content of diet
period in Texas and Utah (Hamilton, 1978). Owls were Energetic characteristics of the mouse diet used for
acclimated to the test temperature for 1 week prior to the
experiment. Preweighed live mice (Mus musculus) (weighed barn owl consumption trials are shown in Table 2.
to the nearest 1.0 g analytical balance) were provided ad Mouse water content comprised 69.7% of the actual
kibitz. Food trials lasted 4-5 days (d) if the owl’s weight weight. Mouse diet caloric values (5.046 kcal/g DM,
did not deviate + 2% (Kendeigh et al., 1977) from the initial and 1.53 1 kcal/g wet wt) are similar to other reported
test weight (measured Cothe nearest 1.Og platform balance). values for Mus musculus (1.7 kcal/g wet wt) (Golley,
Composition of the laboratory mouse diet was measured 1958) and are representative of barn owl prey
in triplicate. Dry matter (DM) was determined by drying species observed in the field [&‘igmodon hispidus,
samples at 60°C in a convection oven. After grinding 4.91-5.49 kcaljg DM and 1.74 kcal!g wet wt; Rei-
carcasses in a Wiley mill, the caloric content of the DM was throdontomys megalotis, 5.10-X I5 kcal/g DM and
1.65 kcal/g; Microtus ochrogaster, 4.91-5.01 kcal/g
*Present address and to whom all correspondence should DM and I.42 kcal/g; Peromyscus maniculatus,
be addressed: Dept. of Physiology and Biophysics, 5.05-5.14 kcal/g DM and 1.61 kcai/g; (Fleharty et at.,
University of Texas Medical Branch, Galveston, TX 1973). Perognuth~ purvus, 1.55 kcal/g wet wt
77550, USA. Telephone: (409) 761-1826. (Schreiber and Johnson, 1975)].
356 KIRK L. HAMILTON

Table 1. Food consumotion duration and weiehts for 6 cautwe barn owls
Total
Trial period (days) period Body weight (g)
Bird Sex N ii &SD (days) X +SD
Owl 1 F 3 4 00 12 579.5 10.8
Owl 2 F 3 4 0.0 12 529.1 10.8
Owl 3 F 3 4.7 0.6 14 572 0 7.8
Owl 4 F 3 6.3 0.6 19 584.8 12.7
Owl 5 F 3 5.7 1.5 17 575.0 1.3
Owl 6 F 3 5.0 1.0 15 527.0 28.7
Total 18 4.9 1.1 89 561.3 27.8

Pellet and excreta production sumption trials held at 5, 15 and 25”C, respectively)
The number of pellets regurgitated/day for the are similar to that reported by Graber (1962)
three test temperatures was not significantly different (2.74 kcal/g DM).
(0.05 < P < 0.1); however, pellet dry matter pro- Food consumption
duction (g/day) at 25°C was significantly higher than
at 15°C (P < 0.05), or 5°C (P < 0.05) (Table 3). Pellet Daily food consumption (wet matter) increased
weights at 5°C (2.72 + 1.13 g/pellet, N = 78), were significantly (P < 0.001) as air temperature decreased
greater than at 25°C (2.19 f 0.84 g/pellet, N = 61, (Fig. la). Food intake at 5°C (116.9 + 7.6 g/day) was
P < 0.05), but not different at 15°C significantly greater than at 15°C (96.8. &-8.7 g/day,
(2.19 + 1.14 g/pellet, N = 75, P > 0.05). There was P < 0.05) or 25°C (76.9 f 16.2 g/day, P < 0.001).
no difference (P > 0.05) between pellet weights from Food consumption at 15°C was higher than at 25°C
15°C and 25°C food trials.
Pellet length (mm) was significantly different
(P < 0.01) for the test temperatures. Pellets regur- Table 3. Results of the 18 barn owl food consumption trials
gitated at 5°C (33.6 f 6.9mm, N = 78) were comrAeted at 3 temperatures, and a laboratory mouse diet
significantly longer (P < 0.001) than pellets expelled Total
at 15°C (29.9 + 6.1 mm, N = 75) or 25°C T, (“C) x -
&SD
(29.1 + 5.6 mm, N = 61) food trials. Pellet length was Wet matter intake/day (g/d)
not different for food trials conducted at 15 and 25°C. 5 116.9 7.6
There was a significant difference (P < 0.001) in 15 96.8 8.7
25 76.9 16.2
pellet width among the food trial temperatures. Pel- Dry matter intake/day (g/d)
lets regurgitated at 5°C (22.1 + 3.8 mm, N = 78) were 5 35.4 2.3
significantly wider (P < 0.001) than pellets of 15°C 15 29.5 2.7
trials (20.4 f 4.Omm, N = 75) or 25°C trials 25 23.5 4.9
Gross energy intake/day (kcal/d)
(20.4 + 3.3 mm, N = 61). Pellet widths from 15 and 179.0 11.6
5
25°C food trials were not different. 15 148.9 13.8
The amount of feces excreted/d (g dry wt) was not 25 117.7 24.8
different (P > 0.05) for 5, 15 and 25°C food trials Pellet energy loss/day (kcal/d)
5 23.2 3.5
(Fig. lb). Neither pellet nor excretory energy content 15 18.6 2.6
(kcal/g DM) were significantly different as ambient 25 13.7 3.1
temperature changed. Pellet energy content Excretory energy loss/day (kcal/d)
(3.20 + 0.13 kcal/g DM, 3.24 f 0.23, and 3.23 f 0.25 5 15 6 4.3
15 13.6 3.5
at 5, 15 and 25”C, respectively) was in agreement with 25 12.2 3.9
values reported by Graber (1962) (3.15 kcal/g DM) Existence metabolism (k&/d)
and Wallick and Barrett (1976) (3.01 + 0.13 kcal/g 5 140.3 6.7
DM). Excretory energy content (2.73 + 0.07 kcal/g 15 116.7 10.6
25 91.7 20.9
DM, 2.65 f 0.12, and 2.65 + 0.12 at food con-
Dry matter excretion/day (g/d)
5 51 1.6
15 5.3 1.3
25 4.8 1.4
Table 2. Energetic characteristics of the mouse (MU Dry matter pellets/day (g/d)
musculus) diet used for barn owl food trials 5 6.2 1.5
15 5.7 0.7
Diet type
25 4.3 0.9
Mus musculus
Parameter x SD Eneigy assimilation efficxncy (%)
78.4 2.1
Dry matter 30.3 1.36 15 78.3 1.4
(A as fed) (5) 25 77.6 51
Dry matter energy* 5.046 0.215 Dry5matter assimilation efficiency (%)
(5) 66.1 6.4
Wet matter energy* 1.531 0.112 15 62.7 3.2
(5) 25 61.1 2.1
Prey weight (X)t 30.4 5.9 Pellets regurgitated/day
(322) 5 2.6 0.4
15 2.4 0.4
*kcal/g; sample size in parentheses.
7ci 19 04
tMean weight in grams; sample we in parentheses.
Barn owl food and energy requirements 357

Rate of food consumption of barn owls at 25°C


(76.9 g/day) is very similar to other published values
for wild and captive barn owls (Marti, 1973; Johnson,
1974; Wallick and Barrett, 1976; Hamilton and Neill,
1981). However, Kirkwood (1979) reported that barn
owls maintained at 14°C on a diet of day-old cock-
erels, consumed 10.7 g of DM/day while birds in our
study consumed 29.5 g/day DM at 15°C (Table 3).
Some of this discrepancy lies in the energetic nutritive
values of the food sources used in the separate
studies. The mean DM content of day-old cockerels
and mouse diets were 25:3 and 30.3x, respectively,
and the energy values of the diets were 5.870 and
5.046 kcal/g DM, respectively (Table 2). Therefore, in
the present study, barn owls maintained on a mouse
diet must consume more biomass/day than when fed
day-old cockerels. Stalmaster and Gessaman (1982)
noted parallel results for bald eagles (Huliueetus
leucocephalus) maintained on different nutritional
diets.
Daily pellet energy loss (kcal/day) increased
I . (P < 0.001) as ambient temperature decreased, but
5 15 25 daily excretory energy loss was not significantly
Temp. (“Cl different (15.6kcal/day at 5°C 13.6 at 15°C and
Fig. 1. (a) Daily food consumption (wet weight) and (b) 12.2 at 25°C; P > 0.05) at any temperature (Fig. 2,
excretion (dry weight) of 6 captive barn owls fed on a Table 3). Pellet energy loss at 5°C (23.1 kcal/day) was
laboratory mouse diet at 3 temperatures. Mean (horizontal higher than at 15°C (18.5 kcal/day, P < 0.05) and
lines), i SD (boxes), ranges (vertical lines), and sample sizes 25°C (13.7 kcal/day, P < O.OOl), and at 15°C was
are presented. higher (P < 0.05) than at 25°C (Fig. 2). Wallick and
Barrett (1976) reported barn owl pellet energy loss of
16.1 kcal/day and excretory loss of 8.9 kcal/day,
(P < 0.05). During the food consumption trials, at which agree with results presented here. Likewise,
25, 15 and 5°C owls consumed 13.7, 17.2 and 20.8% Johnson (1974) found that excretory energy loss
of mean body weight (561.3 g) per day, respectively. of barn owls weighing 409.5-588.4 g was
Marti (1973) reported food consumption values of 12.7-15.4 kcal/day.
46.474.0 g/day for barn owls subjected to ambient Barn owl existence metabolism (EM) paralleled
temperatures (outside aviary) of -1 to +21”C. GEZ results in that EM increased (P < 0.001) as
Marti’s values are 3&35x lower than consumption ambient temperature decreased (Fig. 2). Existence
rates reported in the present study, and this might be metabolism at 5°C (140.2 kcal/day) was higher than
explained because Marti could not maintain the test at 15°C (116.7 kcal/day, P < 0.05) or at 25°C
temperature and did not state if the owls’ weights (91.7 kcal/day, P < O.OOl), and EM determined at
remained + 2% of the initial test weights (Kendeigh 15°C was higher (P < 0.05) than at 25°C (Fig. 2).
et al., 1977). Wallick and Barrett (1976) reported a Kendeigh et al. (1977) described the following two
food consumption rate of 74.1 g/day for barn owls equations to predict existence metabolism for non-
(experiment conducted in August) which was only 3%
lower than the value determined for birds held at
25°C in this study. GEl
EM
PE
Energy requirements EE

In the present study, food consumption, gross


energy intake (GEI), dry matter intake (DMI), and -I
G
EM increased significantly as ambient temperature
decreased (Figs 1 and 2, Table 3).
G
Gross energy intake (kcal/day) and DMI (g/day) ’ f
6
increased (P < 0.001) as ambient temperature de-
creased. Energy intake and DMI at 5°C t
G

(179.0 kcal/day, and 35.4 g/day, respectively) were


significantly higher than at 15°C (148.9 kcal/day and 20 ik
29.4g/day, P < 0.01) (Fig. 2, Table 3). The GEI G T, = %,
%- 7 9, 1

values at 5 and 15°C food trials agree with values 5 15 25


reported for long-eared (Asio otus) (159.0 kcal/d) and Ambient Temp. (%I
short-eared (Asio JEammeus) (188.0 kcal/day) owls Fig. 2. Daily gross energy intake (GEZ), existence metabo-
held at 18°C (Graber, 1962). Gross energy intake at lism (EM), pellet energy (PE), and excretory energy (EE)
25°C (117.7 kcal/day) is very similar to Wallick and of 6 captive barn owls fed on a laboratory mouse diet at 3
Barrett’s (1976) reported value (111 .Okcal/day) for temperatures. Means, *SD, ranges, and sample sizes are
captive barn owls held from 7 to 17 August. presented.
358 KIRK L. HAMILTON

passerines at 0 and 30°C: Duke G. E., Ciganek J. G. and Evanson 0. A. (1973) Food
At O”C, M = 4.142 Wo54” consumption and energy, water and nitrogen budgets in
(1)
captured great-horned owls (Bubo vzrginianus). Camp.
At 30°C M = 1.068 W” 6637 (2) Biochem. Physiol. 44A, 283-292.
Fleharty E. D., Krause M. E. and Stinnett D. P. (1973)
where M = existence metabolism (kcal/day) and
Body composition, energy content and lipid cycles of four
W = weight of bird (g). Equations of Kendeigh et al.
species of rodents. J. Mammal. 54, 42&438.
(1977) predict EMS of 129.9 kcal/day and Gessaman J. A. (1972) Bioenergetics of the snowy owl
71.3 kcal/day for barn owls maintained at 0 and (Nyctea scandiaca). Arctic Alp. Res. 4, 223-238.
30°C respectively, which are 7 and 22% lower than Gessaman J. A. (1973) Methods of estimating the energy
those determined in this study. Johnson’s (1974) cost of free existence. In Ecological Energetics of Homeo-
estimation of EM fl = 54.0 kcal/day) (conducted at therms (Edited by Gessaman J. A.), pp. 3-34. Utah State
ambient temperatures between 16 and 2lC) is University Press, Logan.
below the EM value for 25°C presented in this study, Golley F. B. (1958) Energy dynamics of a food chain of the
old-field community. Ph.D. Thesis. Michigan State Uni-
and the cause may be due to smaller owls used in
versity, East Lansing.
Johnson’s experiments fl owl weight of 484g vs
Graber R. R. (1962) Food and oxygen consumption in three
561 g for the present study). species of owls (Strigidae). Condor 64, 473-487.
Barn owl energy assimilation efficiency was not Hamilton K. L. (1978) Food habits and bioenergetics of a
significantly different (P > 0.05) at the 3 test tem- natural population of barn owls (Tyto alba). M.A. Thesis.
peratures (78.4, 78.3 and 77.6% at 5, 15 and 25°C University of Texas Arlington, Arlington.
respectively) (Table 3). Likewise, dry matter assimi- Hamilton K. L. and Neil1 R. L. (1981) Food habits and
lation efficiency was not different (P > 0.05) among bioenergetics of a pair of barn owls and owlets. Am. Mid/.
the test temperatures (Table 3). Energy assimilation Nat. 106, 1-9.
Johnson W. D. (1974) Bioenergetics of the barn owl, Tyto
did not increase with decreased temperatures (a trend
alba. M.A. Thesis. California State University, Long
was noted) (Table 3). Johnson (1974), Wallick and Beach.
Barrett (1976), and Kirkwood (1979) reported barn Kendeigh S. C., Dolink V. R. and Gavrilov V. M. (1977)
owl EAE’s commensurate to those determined in this Avian energetics. In Granivorous Birds in Ecosystems
study (79, 77 and 72x, respectively). (Edited by Pinkowski J. and Kendeigh S. C.), pp.
Acknowledgements-This paper is in partial fulfillment of a 127-204. Cambridge University Press, New York.
Ph.D. degree in the Department of Biology at Utah State Kirkwood J. K. (1979) The partition of food energy for
University. I would like to thank Dr J. A. Gessaman, my existence in the kestrel (Falco tinnunczdus) and barn owl
major professor, for his assistance and editorial comments (Tyto alba). Comp. Biochem. Physiol. 63A, 495-498.
on this manuscript. I thank Drs K. L. Dixon, R. T. Sanders, Koplin J. R., Collopy M. W., Bammann A. R. and
L. C. Ellis and R. P. Sharma for their assistance on my Levenson H. (1980) Energetics of two wintering raptors.
graduate committee. This research was funded in part by an Auk 97, 795-806.
Edwards H. and Winnie H. Smith Fellowship- from- the Ligon J. D. (1969) Some aspects of temperature relations in
Welder Wildlife Foundation, a Sigma Xi Grant-in-Aid of small owls. Auk 86, 458-472.
Research award, and the Ecology Center-Utah. State Uni- Marti C. D. (1973) Food consumption and pellet formation
versity. I would like to express my sincere thanks to Dr A. rates in four owl species. Wilson Bull. 85, 178-181.
B. Hamilton for her field assistance, research discussions, Postler J. L. and Barrett G. W. (1982) Prey selection and
and editorial comments of this manuscript. I thank MS P. bioenergetics of captive screech owls. Ohio J. Sci. 82,
Hornbeck for secretarial assistance and Ms. E. Tamayo for 55-58.
drawing the figures. This is the Welder Wildlife Foundation Schreiber R. K. and Johnson D. R. (1975) Seasonal changes
Contribution No. 164. in body composition and caloric content of great basin
rodents. Acta Theriol. 20, 343-364.
REFERENCES Stalmaster M. V. and Gessaman J. A. (1982) Food con-
sumption and energy requirements of captive bald eagles.
Barrett G. W. and Mackey C. V. (1975) Prey selection and J. Wildl. Mgmt 46, 64&654.
calorie ingestion rate of captive American kestrels. Wilson Wallick L. G. and Barrett G. W. (1976) Bioenergetics and
Bull. 87, 574-579. prey selection of captive barn owls. Condor 78, 1399141.

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