Biological Conservation 170 (2014) 120–129

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Biological Conservation
journal homepage: www.elsevier.com/locate/biocon

Amur tiger (Panthera tigris altaica) energetic requirements: Implications

for conserving wild tigers
C.S. Miller a,b,⇑, M. Hebblewhite a, Y.K. Petrunenko c, I.V. Seryodkin c, J.M. Goodrich b, D.G. Miquelle b
a
Wildlife Biology Program, Department of Ecosystem and Conservation Sciences, College of Forestry and Conservation, University of Montana, Missoula, MT 59812, USA
b
Wildlife Conservation Society, Bronx, NY 10460, USA
c
Pacific Geographical Institute, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690041, Russia

a r t i c l e i n f o a b s t r a c t

Article history: The Global Tiger Recovery Program has identified increasing prey populations as a crucial component of
Received 24 June 2013 recovering wild tiger numbers because prey density is a key determinant of tiger density, especially for
Received in revised form 12 October 2013 Amur tigers (Panthera tigris altaica) living at the northern limit of tiger distribution, where both tiger and
Accepted 6 December 2013
prey densities are the lowest range wide. To understand potential prey requirements for Amur tigers, we
developed a predictive energetics model based on allometric scaling and literature values for the costs of
reproduction to estimate energetic requirements. We parameterized the energetics model with empirical
Keywords:
activity budgets and movement rates estimated from 5 GPS radiocollared tigers between 2009 and 2013.
Amur tiger
Energetics
To avoid starvation, adult male tigers must consume a minimum of 5.2 kg/day, non-reproducing adult
Panthera tigris altaica females 3.9 kg/day, and an adult female raising 4 cubs to independence needs 200% more energy, or
Prey requirements an average of 11.4 kg/day over the 651 day reproduction period. Given knowledge of Amur tiger diet
Russian Far East composition and prey biomass in the Russian Far East, male tigers would consume 24.8 prey per year,
Tiger non-reproducing females 18.2 prey per year, and a female raising 4 cubs 53.8 prey per year. However,
our energetics model underestimated empirical consumption rates, emphasizing that while the relative
costs of reproduction are valid, wild tigers are likely consuming more prey than predicted by basal met-
abolic demands alone. Quantifying the energetic requirements of tigers allows resource managers to esti-
mate nutritional carrying capacity, estimate the impact of tigers on prey, and develop science-based
conservation recommendations.
Ó 2013 Elsevier Ltd. All rights reserved.

1. Introduction poaching is a primary threat to tiger persistence across Southeast

Asia, where conservation efforts are challenged by some of the
Recent estimates suggest there are fewer than 3500 wild tigers densest and fastest growing human populations in the world (Kar-
(Panthera tigris) remaining in the world (Walston et al., 2010). In anth and Stith, 1999; Miquelle et al., 1999). Recent efforts to en-
November 2011, in an unprecedented meeting, leaders of the 13 ti- hance prey density in Malaysia emphasize the challenge, but also
ger-range countries met in St. Petersburg, Russia, and committed viability, of recovering prey for tigers (Kawanishi et al., 2013). In
to the ambitious goal of doubling wild tiger numbers by 2022. Like Russia, the combination of intensive poaching, legal subsistence
other large carnivores, one of the main ecological drivers of tiger hunting, and predation by a spectrum of large predators including
density across their range is the density of preferred prey (Carbone both Amur tigers (P. t. altaica) and wolves (Canis lupus; Miquelle
and Gittleman, 2002; Karanth et al., 2004). For the Global Tiger et al., 2005) has likely reduced prey densities below carrying
Recovery Program (a collaborative effort between the 13 tiger- capacity, potentially limiting tiger densities.
range countries) to achieve this goal of doubling tiger numbers Prey depletion is even more important for Amur tigers because
within a decade, increasing prey populations will be crucial (Global lower primary productivity at the northern limit of tiger distribu-
Tiger Recovery Program, 2011). Prey depletion resulting from tion is hypothesized to be constrained by densities of key prey spe-
cies (Miquelle et al., 2010). Amur tigers exist at the lowest ungulate
⇑ Corresponding author at: Wildlife Biology Program, Department of Ecosystem densities (Miquelle et al., 1999; Stephens et al., 2006) and have the
and Conservation Sciences, College of Forestry and Conservation, University of largest home ranges (Goodrich et al., 2010) of any tiger subspecies.
Montana, Missoula, MT 59812, USA. Tel.: +1 (406) 243 5236. One of the primary challenges to conservation of Amur tigers is to
E-mail addresses: [email protected] (C.S. Miller), mark.hebblewhite@ manage ungulate harvest and reduce poaching to achieve a neces-
umontana.edu (M. Hebblewhite), [email protected] (Y.K. Petrunenko), sary minimum prey density to ensure not only the energy require-
[email protected] (I.V. Seryodkin), [email protected] (J.M. Goodrich),
ments for survival, but also for reproduction. The long-term
[email protected] (D.G. Miquelle).

0006-3207/$ - see front matter Ó 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.biocon.2013.12.012
 C.S. Miller et al. / Biological Conservation 170 (2014) 120–129 121

persistence of a tiger population is dependent on the availability of 2. Materials and methods

preferred prey species, which are generally similar to tigers in size
(Hayward et al., 2012; Miquelle et al., 2010), and of course limiting 2.1. Study area
human-caused mortality. Can tiger habitat composed of only 10 kg
musk deer (Moschus spp.) or 14 kg muntjac (Muntiacus spp.; prey Our research was conducted in the Sikhote-Alin Mountains of
weights from Hayward et al. (2012)) support viable tiger popula- the Russian Far East, focusing in and around the 4000 km2 Sikh-
tions? Quantifying the energetic needs of tigers provides a more ote-Alin Biosphere Zapovednik (SABZ). As an IUCN Class I protected
precise means of estimating their nutritional carrying capacity area, SABZ is closed to the public and access is limited to Zapoved-
(sensu Hobbs, 1989; Laundre, 2005), and consequently, the neces- nik (Reserve) staff and visiting scientists, resulting in low poaching
sary kill rates to successfully rear young. Given the challenge of rates and no legal hunting, whereas prey populations outside pro-
empirically relating field consumption rates to tiger demography, tected areas sustain legal hunting and high poaching rates (Miquelle
an energetics model could provide a useful basis for conservation et al., 2005). The Sikhote-Alin Mountains are characterized by
recommendations (Odden and Wegge, 2009). dry, cold winters (mean = 12.9 °C, January at Reserve headquar-
Most studies of wildlife energetics have relied on laboratory ters) and warm, humid summers (mean = 15 °C, July at Reserve
experiments that directly estimated the energetic expenditure of headquarters; Gromyko, 2010). Vegetation communities vary from
captive animals and then use these models to predict potential oak (Quercus mongolica) forests along the coast to mixed conifer–
energetic costs or requirements in the wild (Barboza et al., 2009; deciduous forests inland, including Korean pine (Pinus koraiensis),
Oftedal and Gittleman, 1989). Because of the inherent difficulties larch (Larix komarovii), birch (Betula spp.), spruce (Picea ajanensis)
facing field studies of large carnivores, few studies have attempted and fir (Abies nephrolepis). The preferred prey species of tigers in
to translate estimated energetic requirements of wild carnivores to the region are red deer (Cervus elaphus) and wild boar (Sus scrofa),
prey requirements (but see Powell, 1979). Calculating prey with sika deer (Cervus nippon) and roe deer (Capreolus pygargus)
requirements for survival and reproduction requires knowledge becoming more important in areas of diminished preferred prey
of energetic requirements and food consumption in the wild populations (Miquelle et al., 2010).
(Ackerman et al., 1986; Aldama et al., 1991; Laundre, 2005). Advances
in Global Positioning System (GPS) collars have provided an oppor- 2.2. General field methods
tunity to obtain empirical estimates of activity and movement
costs (Eriksen et al., 2011) and to estimate annual kill and con- We captured adult tigers in and around SABZ from 2009 to 2011
sumption rates for many carnivores (Knopff et al., 2010; Metz using Aldrich foot snares and anaesthetized with ZoletilÒ (Univer-
et al., 2012; Miller et al., 2013). There have been, however, few sity of Montana IACUC # AUP 043-09; Lewis and Goodrich, 2009).
attempts to understand energetic requirements by combining Tigers were fitted with LOTEK (Newmarket, Ontario, Canada) or
empirical data from GPS collars with models of energetic costs. VECTRONIC (Berlin, Germany) GPS collars, which were pro-
The primary energetic activities of adult tigers are resting, trav- grammed to obtain locations every 90, 120, or 180 min.
eling, eating, hunting, and the periodic costs associated with repro-
duction and thermoregulation. Time spent in different activities 3. Calculation
multiplied by the energetic costs of each activity results in an esti-
mate of daily energy requirements (Laundre, 2005; Powell et al., We estimated the daily energetic requirements of adult male,
1985). Sunquist et al. (1999) suggested that a reproductive tigress non-reproductive female, and reproductive female tigers raising
with two large cubs needs approximately 50% more food than sol- 1–4 cubs and an average litter (2.4 at birth but decreasing to 1.3
itary females because of the increased energetic requirements of when 1 year old; Kerley et al., 2003), during summer (21 April–
gestation, lactation, and hunting, but these estimates have not 30 November) and winter (Hojnowski et al., 2012).
been verified. Translating tiger energetic requirements into con-
sumption rates could provide a more precise means of defining 3.1. Tiger energetics model
minimum prey requirements for successful reproduction, benefit-
ting conservation efforts throughout tiger range and give quantita- We estimated the total daily energetic requirements (Ctotal) of
tive criteria to aim to restore ungulate populations to promote an adult tiger using the following equation (Aldama et al., 1991;
tiger recovery. Powell, 1979):
We developed the first energetics model to estimate prey
requirements for tigers to survive and reproduce in the wild using C total ¼ C r þ C tr þ C h þ C e þ C rep þ C th ð1Þ
a combination of empirical data and modeling. We had two broad which includes the energetic costs of resting (Cr), traveling (Ctr),
research objectives, first to estimate tiger energetic requirements hunting (Ch), eating (Ce), reproduction (Crep), and thermoregulation
and then to determine the consequences of these caloric demands (Cth). Eq. (1)requires an estimate of the proportional time allocated
in terms of predicted prey requirements in single and multi-prey to each activity. Daily time budgets of tigers were calculated by
communities. We developed this field-based tiger energetics mod- dividing the number of locations (at a fixed GPS fix interval/tiger)
el using information from the literature to estimate the energy from known activities (eating and digesting at clusters of locations
requirements of tigers, and tested the model using GPS-based, at predicted and known kill sites, clusters of locations while resting
empirically-derived movement rates and kill/consumption rates between kill sites, and single, solitary locations while traveling/
on Amur tigers in the Russian Far East (Miller et al., 2013). In addi- searching for prey) by the total number of received locations. We
tion, we validated our model against independent data from a used a logistic regression model to assign clustered locations to eat-
radiocollared reproductive tigress, addressing the most important ing at predicted kill sites and resting at day beds (Miller et al.,
predictions of the energetics model. Although we developed the 2013). We considered activity budgets constant throughout the
energetics model based on data from Amur tigers, we also demon- year. We assumed any biases from missed locations were low be-
strate its broad applicability using previously collected GPS data cause of our high overall fix rates (91.3%; Frair et al., 2010).
from Bengal tigers (P. t. tigris) in the Bangladesh Sundarbans The energetic cost of resting (Cr), also called the basal metabolic
(Barlow, 2009). rate (BMR), was estimated as:
122 C.S. Miller et al. / Biological Conservation 170 (2014) 120–129

C r ¼ 70W 0:75 t kcal=day ð2Þ urfaces. However, neither of these previous studies provided rigor-
ous or transferable relationships to extrapolate energy costs of
where t represents the daily time spent resting and W represents
movement in snow to large carnivores. Therefore, while we include
the weight of the tiger in kilograms when captured, which we as-
costs of thermoregulation, our inability to estimate the additional
sumed stayed constant (Kleiber, 1961). To apply our model to wild
costs of movement in snow for Amur tigers means our estimated
tigers, we used mean weight estimates from Slaght et al. (2005) for
energy requirements will be conservative of wild tiger needs dur-
wild adult Amur tiger males (176.4 kg) and wild adult females
ing winter. We return to this important caveat in the discussion.
(117.9 kg).
Reproduction is among the most energetically expensive activ-
Metabolic rates usually increase linearly with movement
ities in mammals (Harvey, 1986). The cost of reproduction in fe-
(Taylor et al., 1974) and are estimated using an allometric
males is the combined energetic costs of gestation (Cgest),
relationship to body weight. From Taylor et al. (1974), we
lactation, and post-weaning costs associated with the growth of
estimated energetic costs of traveling (Ctr) in tigers using:
cubs (Cgrow). To estimate the daily energetic costs of nursing and
C tr ¼ ð5:8W 0:75 tÞ þ ð2:6W 0:6 dÞ ð3Þ post-weaning dependence on a tigress, we estimated the daily
energetic cost of growth, travel, and maintenance for each cub as:
where W is defined above as tiger weight, t represents the time
spent traveling in hours, and d represents the total daily distance C cub ¼ C grow þ C r þ C tr þ C e ð6Þ
traveled (km). We used a combination of snow tracking and GPS As cubs are born throughout the year (Kerley et al., 2003), we
collar data to calculate seasonal movement rates. Straight-line dis- could not predict the age and weight of cubs during winter, and
tances between consecutive locations were calculated using Haw- therefore, did not include their costs of thermoregulation, which
th’s Tools (Beyer, 2004) in ArcMap 9.3 (Environmental Systems would also lead to conservative estimates of energy requirements.
Research Institute, Redlands, California, USA) and represent the The gestation period in tigers averages 103 days (Nowell and
minimum distance traveled between fixes. To represent travel rates, Jackson, 1996), and the birth weight of a tiger cub averages 1255 g
or distances covered while a tiger is actually moving, we excluded (Oftedal and Gittleman, 1989). The litter energy per metabolic
consecutive locations associated with a cluster of locations (which body size was calculated from litter weight, assuming energy
usually represented resting or kill sites). We estimated actual travel content of 0.98 kcal/g, multiplied by gestation length (Oftedal
distance by comparing straight-line distance between consecutive and Gittleman, 1989). Using these parameters, we calculated the
locations to real movement distances determined by snow tracking additive energetic cost of gestation using the following formula:
(Musiani et al., 1998; Pepin et al., 2004). We assumed the correction
factor was constant for both winter and summer. C gest ¼ ððDW g=day  28:624 kcal=day  0:98 kcal=g  LitterÞ
Tigers are solitary, stalk and ambush carnivores that capture  103Þ  1:27 ð7Þ
prey on average during 1 of 5 attempts (Matyushkin, 1977), using
a quick burst of energy in a very short time span. Due to these where DW was the age-specific weight gain in g/day, 28.624 kcal/
hunting techniques, we considered 0.1% of a tiger’s time budget, day is the daily energetic costs per metabolic body size in kg,
or 1.44 min each day, to be time spent attacking and subduing 0.98 kcal/g was the average caloric content of mammals, 103 was
prey. The maximum energetic output of 32 mammal species was the gestation period, in days, and the constant 1.27 accounts for
estimated using the formula: energetic costs of producing the placenta and fetal fluids (Kaczmar-
ski, 1966; Oftedal and Gittleman, 1989). We divided the birth
C h ¼ 191W 0:942 t ð4Þ
weight of a tiger cub by the gestation period to standardize the daily
where t represents the total time, in hours, spent attacking and sub- weight gain (DW) per cub across the 103 day gestation period to
duing prey (Weibel et al., 2004). estimate daily costs. We estimated reproductive costs across a
The energetic costs of feeding and digestion, while difficult to range of litter sizes from 1 to 4 cubs, as well as at the average litter
estimate, are higher than BMR (Barboza et al., 2009). We assumed of 2.4 cubs at birth and 1.3 cubs from 1 year to dispersal. We as-
the cost of eating to be equal to the cost of traveling at mean travel sumed a 50:50 sex ratio of cubs at birth. We assumed that repro-
speed (Ackerman et al., 1986; Aldama et al., 1991). Using mean tra- ductive costs for adult male tigers were included in overall
vel speeds, we replaced d in Eq. (3)with the product of speed and activity budgets.
time (Ts) and estimated the energetic costs of eating as: Lactation costs to the tigress were calculated assuming a 60%
conversion efficiency of the female’s metabolized energy to milk
C e ¼ ðð5:8W 0:75 Þ þ ð2:6W 0:6 T s ÞÞt ð5Þ and subsequent assimilation by the cubs (Moen, 1973). Following
where Ts represents the average travel speed in kilometers per hour Ackerman et al. (1986) and Laundre (2005), we estimated the ener-
and t represents the time spent eating in hours. getic cost of growth in wild tiger cubs from birth to dispersal as:
An additional energetic challenge faced by Amur tigers com- C grow ¼ ðDW  1430Þ=e ð8Þ
pared to other tiger subspecies is the harsh winter environment
of the Russian Far East. To account for thermoregulation (Cth), we where DW was the age-specific weight gain in kg/day, 1430 kcal/kg
increased energetic costs (Ctotal) during the winter season by 15%, wet weight was the energetic value of cougar (Puma concolor) flesh
as estimated under laboratory conditions for bobcats (Lynx rufus; (assumed to be the same as tigers; Golley et al., 1965), and e was
Mautz and Pekins, 1989). An additional energetic requirement that 60% growth efficiency during the 56 day nursing period and 100%
is important to northern mammals is the energetic costs of move- during the post-weaning dependence period (Moen, 1973). We cal-
ment in snow. Unfortunately, there are few studies of the energetic culated age-specific weight gain using Michaelis–Menton growth
costs of travel in snow on large carnivores. Parker et al. (1984) used curves based on wild Amur tiger weights (Slaght et al., 2005).
captive ungulates to develop an empirical model for the additional We estimated daily energy costs after weaning similarly to
costs of moving in snow and found the costs of locomotion in mule adults and added them to the maternal energetic requirements
deer (Odocoileus hemionus) and elk increased curvilinearly in rela- (Oftedal and Gittleman, 1989). Available data from tracking radio-
tion to snow depth and density above a critical snow depth thresh- collared females with cubs suggest that females stay close to the
old. Crête and Larivière (2003) suggested that energetic den site for the first month, and that there is little movement of cubs
requirements of domestic dogs (Canis lupus familiaris) increased through the second month (Miquelle, D.G., unpublished data). From
by 4–6% while traveling in snow compared to travel on hard months 2 to 6, we assumed cubs traveled 2 km during 3 h/day, and
 C.S. Miller et al. / Biological Conservation 170 (2014) 120–129 123

subtracted time spent traveling from time spent resting. From estimated model parameters (weight, movement rates, activity
6 months to dispersal at 19 months (Kerley et al., 2003), we as- patterns) using sensitivity analysis of Eq. (1)to identify the most
sumed cubs traveled with their mother and calculated daily ener- important parameters driving tiger energetics. We iteratively in-
getic costs of cubs based on movements and time budgets of the creased each parameter by 1% to evaluate the influence each
tigress. Finally, we added the total energetic costs of each litter to parameter had in Eq. (1)for males and females during winter and
the total energetic requirements of the tigress over the 19 months summer and then ranked each parameter in order of importance.
from conception to independence of the cubs. We also used independent data from one adult female to vali-
date the energetics model. Pt114 was captured as a solitary adult
3.2. Predicted prey requirements female and gave birth to 3 cubs during our monitoring period.
We compared 4 energetic phases (solitary adult, gestation, lacta-
Our second main research objective was to convert daily tiger tion, and a 4-month post-weaning dependence period for the cubs
energetic estimates into predicted prey requirements in single [after which the collar failed]) to our energetics modeling of aver-
and multi-prey communities. We estimated percent composition age female tigers. No data from Pt114 was used in developing the
of prey species by weight in the diet using kill data from GPS-collar energetics model, making this a valuable out-of-sample test of the
derived cluster searches (Miller et al., 2013). The average weights model.
of prey species in and around SABZ were previously reported for
all sex and age classes (Bromley and Kucherenko, 1983; Danilkin,
4. Results
1999); we used these weight estimates to calculate prey require-
ments based on estimates of caloric demands from Eq. (1)(Ctotal).
4.1. Energetic requirements
The edible portion of whole deer fed to captive cougars was
estimated to be 79% (Ackerman et al., 1986) and whole elk was
From 2009 to 2011, we captured and radiocollared two adult
estimated to be 68% (Wilmers et al., 2003). Following these studies,
males, two adult females, and one sub-adult female tiger with
we considered 79% of each small prey item (using a 40-kg cut point
GPS collars. These tigers were monitored from 36 to 481 days, for
to assign prey species into small and large categories) to be edible
a combined total of 1109 days (Appendix A). Overall, 30.7% of tiger
and only 68% of each large prey item was considered edible. We as-
locations were at known or predicted kill sites, 31.2% were single
sumed 1890 kcal/kg of wet weight for meat and organs of all prey
locations while traveling and searching for prey, and 38.0% were
species based on white-tailed deer values (Odocoileus virginianus;
at clusters formed while resting (Table 1). On average, tigers spent
Davison et al., 1978). The number of ungulates killed per day
7.4 h (95% CI 7.1–7.7) at kill sites, 7.5 h (95% CI 7.2–7.8) traveling,
(PBk) can be estimated using the potential biomass consumed per
9.1 h (95% CI 8.8–9.5) resting, and the remaining 1.44 min were
day, the average weight of the prey species available, the percent
attributed to time spent hunting, or more specifically, attacking
diet composition of each prey species, and the edible portion of
and subduing prey each day (Table 1). Adult male tigers averaged
each carcass. Previous studies estimated that 14% of the energy in-
998 and 1173 m/h and adult females averaged 564 and 828 m/h
gested from deer meat is lost during digestion and assimilation by
during winter and summer, respectively. Additional details from
cougars (Ackerman et al., 1986). As human disturbance in our
the movement rate analyses are located in Appendix B.
study area was limited and tigers that are not disturbed by humans
The energetic costs of resting, traveling, eating, and hunting re-
rarely leave edible portions of a carcass (Kerley et al., 2002), we as-
sulted in an overall daily estimate (Ctotal) of 6858 kcal/day (95% CI
sumed tigers rarely abandoned edible biomass at kill sites. To ac-
6398–7326) for adult male tigers (38.9 kcal/kg) and 5063 kcal/day
count for occasional prey abandonment, scavenging and rapid
(95% CI 4802–5327) for adult non-reproducing females (42.9 kcal/
carcass decay during summer, we followed Yudakov and Nikolaev
kg) during summer (Table C.1). The additional energetic costs of
(1987) and assumed 15% of each kill was lost. Based on these
thermoregulation results in overall daily averages of 7808 kcal/
parameters, we estimated the potential biomass of ungulate prey
day (95% CI 7281–8345) for adult males (44.3 kcal/kg) and
consumed (PBc) following Ackerman et al. (1986) as:
5713 kcal/day (95% CI 5408–6023) for adult non-reproducing fe-
PBc kg=day ¼ C total kcal=ð1890 kcal=kg  0:86  PBu  0:85Þ ð9Þ male tigers (48.5 kcal/kg) during the winter (Table C.2). See Appen-
dix C for detailed predictions from the different components of Eq.
where PBu represents the proportion of tiger diet that are ungulates.
(1); here we only report total energetic costs.
For single-prey systems, we estimated the total number of individ-
The overall energetic costs of a female raising an average-sized
uals killed annually for each prey species. For multi-prey systems,
litter from gestation through dispersal are 10,341 kcal/day (95% CI
we estimated the number of each prey species killed based on the
10,063–10,622), whereas for a female successfully raising a litter of
proportion by weight to the rate found in the diet of GPS collared
four cubs, the average daily energy requirements were 1.5 times
tigers (Miller et al., 2013) with the same equation as Ackerman
greater, averaging 15,679 kcal/day (95% CI 15,402–15,961;
et al. (1986):
Table 2).
PBk ¼ PBc =ðM w  eÞ prey=day ð10Þ
where Mw was the mean live weight of prey in kilograms and e is 4.2. Estimating prey requirements
the edible portion of each prey item.
Using these daily caloric requirements, during summer, an aver-
3.3. Testing predictions of the energetics model age adult male Amur tiger would need to consume a minimum of
5.0 kg of meat/day (95% CI 4.6–5.3) and an average non-reproduc-
We evaluated our model’s ability to predict a tiger’s status tive female would need a minimum of 3.7 kg of meat/day (95% CI
(starving, meeting, or exceeding maintenance or reproduction 3.5–3.9). During winter, these estimates increased to 5.7 kg/day
requirements) by comparing modeled estimates of energy require- for males (95% CI 5.3–6.0) and 4.1 kg/day for non-reproducing fe-
ments to empirical data from tracking GPS radiocollared tigers males (95% CI 3.9–4.4). From gestation to dispersal, a tigress with
from which we estimated consumption rates in the field (Miller an average litter needed to acquire an average of 7.5 kg/day (95%
et al., 2013). We also tested for seasonal differences (summer, win- CI 7.3–7.7) and a female successfully raising 4 cubs to dispersal
ter) between the empirical estimates and the energetics model. We needed to acquire, on average, 11.4 kg/day (95% CI 11.2–11.6)
evaluated the sensitivity of the predicted kill rate of ungulates to during the entire 651 day reproductive period. Converting these
124 C.S. Miller et al. / Biological Conservation 170 (2014) 120–129

Table 1
Number of locations from each Amur tiger (Panthera tigris altaica) in the Russian Far East, from 2009 to 2013, during different activities (at predicted kill sites, resting, and
traveling/searching for prey) used to calculate average daily energy/time budgets and number of missed fix attempts. These energy/time budgets were considered constant
throughout the year.

Tiger ID Total locations At kill site (%) Resting (%) Traveling (%) Missed fixesa (%)
Pt90 477 14.0 42.5 43.4 10.3
Pt97 205 21.4 42.4 36.1 25.5
Pt99b 2988 31.7 35.3 32.9 9.6
Pt100 1529 35.1 41.4 23.4 3.4
Pt114c 6209 45.1 29.4 25.4 8.8
Avg. # 2006 30.1 41.7 28.1 5.2
Avg. $ 3193 31.0 35.8 33.1 10.8
Total 5199 30.7 38.0 31.2 8.7
a
Missed fix attempts were not included in time budgets analyses.
b
Two months of winter location data were removed from this analysis while Pt99 recovered from a poaching attempt.
c
Data from Pt114 were not included in average female or overall average activity budgets.

Table 2
Reproductive costs (kcal/day) for Amur tigers (Panthera tigris altaica) including gestation, lactation, and weaning to dispersal at 19 months, Russian Far East, 2009–2013. Caloric
value for each event is reported for a non-reproductive tigress, a tigress successfully raising a single male or female cub to independence, a tigress with 2–4 cubs (assuming a
50:50 sex ratio), and a tigress with an average litter.

Tiger Gestation Lactation Weaning to 6 months 6 Months to dispersal Total (651 day % of Non-
(103 days) (56 days) (96 days) (396 days) average) reproducing $
Non-repro. $ NA NA NA NA 5314 100
$ + 1 Male cub 430 1248 1826 3552 7919 149.0
$ + 1 Female 430 1546 2009 3420 7892 148.5
cub
$ + 2 Cubs 860 2794 3835 6972 10,497 197.5
$ + 3 Cubs 1290 4190 5753 10,458 13,088 246.3
$ + 4 Cubs 1720 5587 7670 13,944 15,679 295.1
$ + Average 1032 3352 4602 6406 10,341 194.6
littera
a
An average litter assumes a 50:50 sex ratio and is considered to be 2.4 cubs at birth and 1.3 cubs from 1 year until dispersal at 19 months (Kerley et al., 2003).

daily consumption estimates to yearly estimates of biomass results to average 36.8 wild boar/year (95% CI 36.1–37.5), 28.1 red deer/
in an average of 1909 kg/year for an average male Amur tiger, year (95% CI 27.6–28.6), 140.6 roe deer/year (95% CI 138.1–
1404 kg/year for an average lone female tigress, 2732 kg/year for 143.1), 58.3 sika deer/year (95% CI 57.3–59.4), or 434.6 musk
a female raising an average litter, and 4142 kg/year for a female deer/year (95% CI 426.9–442.4; Table 3).
successfully raising a litter of 4 cubs. To meet the average daily energetic estimates under a multiple
To meet these energetic requirements under a single prey sce- prey scenario, based on proportions of prey located at confirmed
nario, an average male tiger would need to kill and consume kill sites, an average male tiger would need to consume 4.8 wild
17.0 wild boar/year (95% CI 15.8–18.1), 13.0 red deer/year (95% boar/year (95% CI 4.5–5.2), 5.3 red deer/year (95% CI 5.0–5.7),
CI 12.1–13.8), 64.8 roe deer/year (95% CI 60.4–69.2), 26.9 sika 4.3 sika deer/year (95% CI 4.0–4.5), and 10.3 other prey/year (95%
deer/year (95% CI 25.1–28.7), or 200.3 musk deer/year (95% CI CI 9.6–11.1), or 24.8 total prey/year (95% CI 23.1–26.5; Table 3).
186.8–214.0; Table 3). In contrast, a reproductive tigress raising A reproductive tigress raising 4 cubs to dependence under a multi-
4 cubs to independence under a single prey scenario would need ple prey scenario would need to consume 10.5 wild boar/year (95%

Table 3
Converting Amur tiger (Panthera tigris altaica) caloric demands into prey requirements under single (S) and multi-prey (M) species scenarios in the Russian Far East, 2009–2013.
For the single prey scenario, we estimated the total number of individuals killed annually for each prey species. For the multi-prey scenario, proportions of each species in the diet
are based on empirical biomass acquisition estimates from field-investigated kill sites (Miller et al., 2013) and are 28.6% wild boar, 41.2% red deer, 15.8% sika deer, and 14.4% all
other prey combined.

Tiger kcal/day kg/day Boar/year Red deer/year Sika deer/year Roe deer/year Musk deer/year Other prey/yearb Total/yearc
S M S M S M S S M M
Avg. # 7225 5.2 17 4.8 13 5.3 26.9 4.3 64.8 200.3 10.3 24.8
Avg. $ 5314 3.9 12.5 3.6 9.5 3.9 19.8 3.1 47.6 147.3 7.6 18.2
$ + 1 Male cub 7919 5.7 18.6 5.3 14.2 5.8 29.5 4.7 71 219.5 11.3 27.2
$ + 1 Female cub 7892 5.7 18.5 5.3 9.5 5.8 29.5 4.6 70.8 218.7 11.3 27.1
$ + 2 Cubs 10,497 7.6 24.6 7 18.8 7.8 39.1 6.2 94.1 290.9 15 36
$ + 3 Cubs 13,088 9.5 30.7 8.8 23.2 9.7 48.7 7.7 117.3 362.8 18.7 44.9
$ + 4 Cubs 15,679 11.4 36.8 10.5 28.1 11.6 58.3 9.2 140.6 434.6 22.5 53.8
$ + Average littera 10,341 7.5 24.3 6.9 18.5 7.6 38.5 6.1 92.7 286.6 14.8 35.5
a
An average litter assumes a 50:50 sex ratio and is considered to be 2.4 cubs at birth and 1.3 cubs from 1 year until dispersal at 19 months (Kerley et al., 2003).
b
Other prey refers to the combined estimates of roe deer (Capreolus pygargus), musk deer (Moschus moschiferus), brown bear (Ursus arctos), Himalayan black bear (U.
thibetanus), badger (Meles leucurus), domestic dog (Canis familiaris), and cattle (Bos taurus).
c
Total prey per year refers to the combined total of all prey required to meet annual caloric demands in the multi-prey scenario.
 C.S. Miller et al. / Biological Conservation 170 (2014) 120–129 125

CI 10.3–10.7), 11.6 red deer/year (95% CI 11.4–11.8), 9.2 sika deer/ obtain 82.2 total prey/year, or 664.6 musk deer/year under the sin-
year (95% CI 9.1–9.4), and 22.5 other prey/year (95% CI 22.1–22.9), gle prey scenario. In SABZ, musk deer exist at an average density of
or 53.8 total prey/year (95% CI 52.8–54.7; Table 3). 1.0/km2 across habitats (Stephens et al., 2006). So, within the
home range of an average adult female, about 390 km2 (Goodrich
4.3. Evaluating predictions et al., 2010), there is no feasible way that the musk deer scenario
would provide sustainable consumption rates for tigers, let alone
We monitored Pt114 for 448 days, which included 208 days as a reproduction. However, with multi-prey densities of larger, pre-
non-reproductive adult, a 103 day gestation period, 56 day wean- ferred prey of 1.0 red deer/km2, 1.5 roe deer/km2, and 0.5 wild -
ing period, and 81 days with 3 dependent young after which the boar/km2 (Stephens et al., 2006), an average tigress home range
collar prematurely failed. Empirical consumption rates obtained would have 390, 585, 195 red, roe deer and wild boar, respectively.
from the field during these periods were 8.2 kg/day, 9.7 kg/, These potential densities and our multiple prey scenario consump-
9.4 kg/day, and 12.3 kg/day, respectively. Energetics-based predic- tion rate of 82 prey/year clearly contrasts with the single prey
tions of consumption rates during the different reproductive scenario, supporting our interpretation that conservation of abun-
phases, although lower, paralleled empirical estimates (Fig. 1). dant populations of larger ungulate prey will be required to en-
Our sensitivity analyses suggested that, for all sexes and sea- hance tiger reproduction. Combining insight from empirical data
sons, tiger weight accounted for 41% of the variation in energetic with modeling, when empirical data are unavailable or impossible
requirements, followed by time spent eating (20%), traveling to collect, can provide valuable insights into the extremely high
(17%), and resting (11%). The last three parameters (travel speed, costs of reproduction in tigers.
distance traveled per day, and time spent hunting) only accounted Prey requirements derived from our model (Table 3) were lower
for 11% of the total variation in total energetic requirements. than field estimates of tiger consumption rates (Appendix A). This
is perhaps expected because our energetics model is built to pre-
dict the energetic needs of a sustenance diet and any prey con-
5. Discussion sumed above these requirements can be used as fuel reserves
during high demand times, such as lactation, or times of prolonged
The strongest scientific and conservation contribution of our hunting failures. Indeed, our empirical estimates of 8.9 kg/day for
energetics modeling approach is the estimate of the energetic costs an average adult non-reproductive tiger (Miller et al., 2013) were
of reproduction that are almost impossible to obtain from wild car- much higher than the energetics estimates of 5.2 kg/day for an
nivores. Based on our energetics model, average energetic costs for adult male and 3.9 kg/day for an adult female (Table 3). In fact,
a tigress raising an average litter from the start of gestation to dis- our energetics-modeled tiger consumption rate estimates are low-
persal age were 95% higher than a non-reproductive tigress. This is er than all previously reported empirical consumption rates
double the only previously published estimate; in Nepal, Sunquist (Appendix D). Similarly, energetics-modeled cougar consumption
et al. (1999) estimated that a single Bengal tigress with two large rate estimates of 1.5–4.3 kg/day (Ackerman et al., 1986; Laundre,
cubs needed 50% more biomass. Furthermore, our energetics mod- 2005) were lower than published empirical estimates of 6.6–
el predicted a tigress with 2 large cubs needs 145% more biomass/ 12.9 kg/day (Knopff et al., 2010). For wolves, Peterson and Ciucci
day during the last few months prior to dispersal than a solitary ti- (2003) developed an energetics model to estimate a consumption
gress. During the 651 day reproductive period, energetic costs of 1 rate of 3.25 kg/day for a 35 kg adult wolf, but empirical estimates
cub are 49% higher than a non-reproductive tigress, for 2 cubs, 98%, from Hebblewhite et al. (2003) averaged 5.4 kg/day and averaging
and for 4 cubs, 195%. Similarly, Laundre (2005) reported over the seasonal results from Metz et al. (2012) produces an estimate of
18-month pregnancy to dispersal period, a female cougar raising 6.8 kg/day. Clearly, comparison of tiger, cougar, or wolf energetic
an average litter (2.6 kittens) needed to obtain an average of needs derived from models are consistently underestimating
160% more biomass/day. Combining our empirical estimates of empirical estimates using GPS collars, regardless of size of species
consumption rates in adult tigers (8.9 kg/day) with this relative being investigated.
adjustment based on average reproduction (95%) suggests that These consistent discrepancies between empirical observations
an adult tigress raising an average litter requires 17.4 kg/day to and energetics modeling raise a series of questions. For example,
meet ‘empirical’ energetic needs. To meet these energetic require- are the models failing to accurately estimate energy requirements
ments under our multiple prey scenario, the tigress would need to or are carnivores killing more than they need to merely meet basal
energetic costs? One factor that might lead to higher empirical kill
rates in comparison to energetic requirements are large carnivore
foraging strategies in the face of high variance in predation rates.
The standard deviation of our predicted consumption rates was
over 1.3 kg/day, suggesting that on many days our field observed
consumption rates were very close to the energetic minimum, or
even below the starvation threshold. Therefore, carnivores likely
strive to exceed starvation thresholds on a daily basis, not merely
meet a basal energetic minimum. Vucetich et al. (2012) found that
high variance in wolf consumption rates often leads wolves to ex-
hibit partial prey consumption to ensure that their consumption
rates are usually higher than a starvation threshold. Therefore,
we conclude that one of the reasons that energetics models under-
predict empirical consumption rates across large carnivore species
is because all energetics models are based on basal metabolic rate,
and not on predicting the consumption rates required to avoid
days of negative energy balance.
Fig. 1. Comparison of empirical consumption rates from Pt114 to our energetics-
Other methodological factors, such as our inability to account
model based estimates for Amur tigers in the Russian Far East at different phases of for additional costs of traveling in snow could lead to underesti-
reproduction. mating energetic requirements. The energetic costs of walking in
126 C.S. Miller et al. / Biological Conservation 170 (2014) 120–129

Table B.1 in snow for large carnivores. A study by Parker et al. (1984) based
Summary of movement rates (meters/hour), corrected for effects of fix-rate, from 5 on wild ungulates found that above a critical snow depth threshold
Amur tigers (Panthera tigris altaica) by season in the Russian Far East, 2009–2013.
Values represent the corrected distance between consecutive locations not associated
determined by chest height, costs increased exponentially. How-
with localized movements at a cluster. ever, this threshold varied between species. The simpler approach
of Crête and Larivière (2003) measured heart rates of domestic
Tiger ID Winter Summer Total
dogs while traveling in snow and found 4–6% higher energetic
Pt90 949.5 1121.6 1075.1 expenditure while traveling in snow. Out of curiosity, we increased
Pt97 636.7 665.0 656.8
Pt99a 558.0 837.3 774.7
the daily cost of travel during winter by 5% (Crête and Larivière,
Pt100 1010.5 1218.6 1100.2 2003) and found average male and female tigers would only need
Pt114b 1172.5 1310.3 1246.5 an additional 13.4 and 10.9 kg/winter, respectively. However, this
Avg. # 998.2 1172.8 1091.5 approach does not account for days when snow depth is above a
Avg. $ 564.0 828.0 767.7
critical threshold (chest height), nor high snowfall events, and
Total 798.5 926.3 886.0
therefore underestimates costs. A further complication is the
a
Two months of winter location data were removed from this analysis while fine-scale habitat selection of large carnivores to avoid deep snow.
Pt99 recovered from a poaching attempt.
b
For example, studying wolves across western Canada, Paquet et al.
Data from Pt114 were not included in calculations for an average female or the
overall averages.
(2012) showed that as average snow-depths increased, gray
wolves showed stronger selectivity for human compacted trails,
reducing the energetic costs of travel in snow. While we acknowl-
edge that our model fails to account for the true energetic costs of
Table B.2 travel in snow during winter, we suggest this is an important area
Summary of total hourly corrected movement distances (meters/hour) from 5 Amur for future research to resolve on large carnivores especially.
tigers (Panthera tigris altaica) by season in the Russian Far East, 2009–2013 and 2 Regardless, high variance in consumption rates and the additional
Bengal tigers (P. t. tigris) in the Bangladesh Sundarbans, 2004 and 2006 (Barlow,
costs of traveling in snow would tend to increase consumption
2009). Values represent the average corrected distance between all consecutive
locations, regardless of activity, and are used in our Sundarbans example.
rates, emphasizing the need for conservative ungulate manage-
ment. Until the discrepancies between field-based observations
Tiger ID Winter Summer Total
and energetics models are resolved, the conservative approach is
Pt90 512.1 575.0 559.2 to rely on the field-based observations for conservation measures
Pt97 249.1 323.8 297.3 focusing on prey requirements for tigers.
Pt99a 187.5 402.4 334.7
Pt100 260.2 576.6 342.3
In addition to the insights into tiger energetic requirements, our
Pt114 414.8 440.0 428.6 estimated tiger movement rates are among the first based on GPS
Amur average 338.6 442.6 396.1 data and support some previously proposed hypotheses about tiger
Sundarbans average 513.6 513.6 spatial ecology that are relevant to understanding energetics. Good-
a
Two months of winter location data were removed from this analysis while rich et al. (2010) showed that male Amur tiger home ranges are lar-
Pt99 recovered from a poaching attempt. gely exclusive and often overlapped 1–5 sympatric females,
whereas female home ranges are correlated with prey biomass (San-
snow are substantial and represent one of the biggest energetic dell, 1989). We found that male tigers traveled an average of 2.3 km
differences between Amur and other populations of tigers. As we further per day than female tigers in winter (6.7 vs. 4.4 km) and
discussed above, however, previous studies did not provide a 1.3 km further in summer (7.9 vs. 6.6 km; Table B.1). The increased
transferable or rigorous platform to estimate costs of movement travel distances we observed in males could, therefore, be a result of

Table C.1
Energetic costs (kcal/day) of each activity during summer (21 April–30 November, 2009–2013) for each Amur tiger (Panthera tigris altaica) in the Russian Far East given tiger-
specific body weight, movement rates, and activity budgets estimated from GPS collars, as well as average male and female costs from averaged weights, movement rates, and
time budgets. Age and weight estimates are based on measurements taken at time of capture.

Tiger ID Sex Age Weight (kg) BMR Traveling Eating Hunting Total kcal/kg
Pt90 # 10 yo Adult 200 1584 3942 1266 674 7467 37.3
Pt97 $ 2 yo Subadult 100 939 1826 1084 351 4201 42.0
Pt99 $ 6–8 yo Adult 130 953 2082 1985 449 5469 42.1
Pt100 # 4 yo Adult 193 1501 2106 3104 652 7363 38.2
Avg # Adult 176.4 1412 2357 2490 599 6858 38.9
Avg $ Adult 117.9 897 1948 1808 410 5063 42.9

Table C.2
Energetic costs (kcal/day) of each activity during winter (1 December–20 April, 2009–2013) for each Amur tiger (Panthera tigris altaica) in the Russian Far East given tiger-specific
body weight, movement rates, and activity budgets estimated from GPS collars, as well as average male and female costs from averaged weights, movement rates, and time
budgets. Age and weight estimates are based on measurements taken at time of capture.

Tiger ID Sex Age Weight (kg) BMR Traveling Eating Hunting Thermoregulation Total kcal/kg
Pt90 # 10 yo Adult 200 1584 3830 1266 674 1103 8458 42.3
Pt97 $ 2 yo Subadult 100 939 1816 1084 351 629 4820 48.2
Pt99 $ 6–8 yo Adult 130 953 1976 1985 449 805 6168 47.4
Pt100 # 4 yo Adult 193 1501 2035 3104 652 1094 8385 43.4
Avg # Adult 176.4 1412 2289 2490 599 1018 7808 44.3
Avg $ Adult 117.9 897 1853 1808 410 745 5713 48.5
 C.S. Miller et al. / Biological Conservation 170 (2014) 120–129 127

reproductive behavior (Goodrich et al., 2010). Also, each of our col- these data to a single prey scenario, similar to above, a tigress in
lared tigers traveled greater distances in the summer than in winter the Sundarbans would need to consume 25.0 spotted deer/year,
(Table B.1). These seasonal differences could reflect concentrations 36.7 wild boar/year, or 294 macaque or adjutant/year. This simple
of prey due to snow conditions (Laundre, 2005), although this has application illustrates the potential utility of our model to
been observed to not occur for at least one primary prey species in understanding tiger energetics, and emphasizes that large bodied
our study area (red deer; Hojnowski et al., 2012). ungulate prey are critically important to tigers (Hayward et al.,
Of course, one of the challenges of any field study on large car- 2012).
nivores such as tigers is sample size. While we developed the ener-
getics model from principles based on the metabolic literature,
small sample sizes limited our calibration of the model to activity 6. Conclusions
data from 5 wild tigers, which we then validated with empirical
consumption rates from 1 wild tigress, whose GPS data was not To meet the Global Tiger Recovery Program’s goals of doubling
used in development of the model. Despite this weakness, the sim- wild tiger numbers by 2022, maintaining or enhancing adequate
ilarity of our empirical consumption rates with previous tiger stud- prey resources will be crucial. Our energetics modeling demon-
ies of consumption rates (Appendix D) provides some reassurance strates that high densities of preferred prey (ungulates of similar
that our results are reasonable. Similarly, the congruence between size to tigers, Hayward et al., 2012) are critical for successful repro-
our conclusions from the energetics model and other carnivore duction, and that even highly abundant small, non-preferred prey
energetics models supports use of our model (Ackerman et al., such as muntjac or musk deer will not ensure successful reproduc-
1986; Laundre, 2005; Peterson and Ciucci, 2003). Other studies of tion because kill rates must be unreasonably high, or because such
tiger ecology and demography (Kerley et al., 2003) have been sim- kill rates will rapidly deplete prey populations. This modeling exer-
ilarly hampered by small sample sizes because of the sheer diffi- cise further shows that a highly reproductive tigress will need 2–3
culty of studying these endangered species. Nonetheless, our times more biomass than a solitary tigress to successfully raise
tiger model provides improvements to our current understanding cubs to independence, emphasizing that enhancing density of prey
of the costs of reproduction that can be used to support tiger con- for tigers will be needed to enhance tiger reproduction. Our obser-
servation policy. We also expect that with growing interest in tiger vations of empirical kill rates provide a baseline for predicting
ecology and conservation, future studies will be able to test and threshold prey requirements for survival and reproduction, and
refine predictions from our tiger energetics model. emphasize the need for conservative management of prey popula-
As an example of the application of our energetics model to tions as a key component of tiger conservation.
other tiger range countries, we assessed prey requirements of Ben-
gal tigers in the mangrove forests of the Bangladesh Sundarbans.
Acknowledgements
Barlow (2009) deployed GPS collars on two adult female tigers
(average weight 76.67 kg) from 2004 to 2006. Combining overall
We would like to thank A. Astafiev, Director, and Y. Pimenova,
daily movement rates estimated from 30 min location data (includ-
Assistant Director of the Sikhote-Alin Zapovednik for their ongoing
ing locations while on a cluster) with our correction factors resulted
support of our research. The Siberian Tiger Project staff, particu-
in daily movement rates of 4.1 km/day, higher than our overall ob-
larly N. Rybin and V. Melnikov, helped collect data in the field.
served Amur tiger movement rates (Table B.2). Applying our time
Comments from J. Stetz and three anonymous reviewers substan-
budgets results in an average travel speed of 513.6 m/h, higher than
tially improved the manuscript. Funding was provided by the
both the 352 m/h for an average Amur tiger (Table B.2) and the
Mohamed bin Zayed Species Conservation Fund, Panthera’s Kaplan
210 m/h reported for Bengal tigers in Nagarahole, India (Karanth
Graduate Award Program, Liz Claiborne and Art Ortenburg Founda-
and Sunquist, 2000). These data result in an estimate of
tion, Save the Tiger Fund, USFWS Tiger Rhino Conservation Fund,
3515 kcal/day, or 2.5 kg/day for a tigress in the Sundarbans. Tigers
University of Montana, and the Wildlife Conservation Society.
in the Sundarbans rely on spotted deer (Axis axis; 80.1% of biomass
consumed, average body mass of 30 kg from Hayward et al. (2012))
and wild boar (10.6%; Khan, 2004). Other, smaller species, such as Appendix A
the rhesus macaque (Macaca mulatta) and lesser adjutant storks
(Leptoptilos javanicus) represent the remaining 9.7% consumed bio- Summary of collared Amur tigers (Panthera tigris altaica) in Rus-
mass. Applying the daily energetic estimates of Bengal tigers in the sian Far East study area, 2009–2013, including length of monitor-
Sundarbans with these consumption estimates results in a mixed- ing period, GPS locations acquired, number of clusters searched,
species prey requirement of 20.0 spotted deer/year, 3.9 wild boar/ and number of kills located. Empirical kill rates (kills/year/tiger;
year, and 24.7 other prey/year, or 48.6 total prey/year. If we apply KR) and consumption rates (kg consumed/day/tiger; CR) were

Tiger Sex Days Good Fix % Clusters Kills Kill rate (kills/ Consumption rate (kg/
ID monitored locations searched confirmed year) day)
Pt90 Male 45 477 89.7 6 1 NA NA
Pt97 Female 36 205 74.5 4 2 NA NA
Pt99a Female 481 3433 90.6 194 50 53.9 8.7
Pt100 Male 99 1529 96.6 48 14 70.1 11.3
Pt114 Female 448 6209 91.2 181 69 58.7 9.4
Total 5 1109 11,853 91.3 433 136 57.8 9.3
Mean 221.8 2370.6 88.5 86.6 27.2 60.9 9.8
SD 223.17 2492.28 8.29 93.88 30.67 8.31 1.35
a
Two months of these data were excluded from analyses while Pt99 recovered from a poaching attempt. However, snow tracking data were used during this time to
generate the 3 h correction factor.
128 C.S. Miller et al. / Biological Conservation 170 (2014) 120–129

calculated based on GPS collared tigers in the field using methods The energetic requirements of thermoregulation (Cth) were
from Miller et al. (2013). added to mean daily energetic estimates during the winter season
to generate overall daily averages of energetic requirements.
Increasing energetic requirements by 15% results in an increase
Appendix B of 1018 kcal/day (95% CI 950–1089) for an average weight adult
male (5.8 kcal/kg) and 745 kcal/day (95% CI 705–785) for an aver-
Estimation of movement rates for Amur tigers in the Russian Far age weight non-reproducing adult female (6.3 kcal/kg; Table C.2).
East from 2009 to 2011.
We snow-tracked Pt100 for a total of 81.1 km between 49 pairs Appendix D
of consecutive locations. The average correction factor for all 49
pairs of locations at a 90 min fix rate was 1.412 (SD = 0.2188). Summary of literature estimating consumption rates (kg/day) of
We snow tracked Pt99 for a total of 52.5 km between 41 pairs of tigers (Panthera tigris) using various methods. All estimates are for
consecutive locations. The average correction factor for all 41 pairs solitary, adult tigers.
of locations at a 3 h fix interval was 1.513 (SD = 0.2866). We were
unable to snow track between any locations at the 2 h fix interval, Author Year Study Methods Consumption
so to calibrate these location data with an actual distance traveled site rates
correction factor we took a weighted average of the 90-min and 3-
h correction factors to estimate a 1.446 correction factor for 2 h fix Novikov 1962 Russian Empirical snow 8 kg/day
intervals. During summer, male tigers averaged 1172.8 m/h (95% CI Far East tracking
1087.6–1258.1; SE 43.3) and female tigers averaged 828.0 m/h Schaller 1967 Kanha, Empirical 5–7 kg/day
(95% CI 780.6–875.4; SE 24.1). Distances traveled during winter NP, India observational
dropped to 998.2 m/h (95% CI 903.5–1092.9; SE 48.1) for male ti- Sunquist 1981 Chitwan, Empirical VHF # – 6–7 kg/
gers (P = 0.0076) and 564.0 m/h (95% CI 490.9–637.1; SE 37.1) for Nepal telemetry day
female tigers (P < 0.001). $ – 5–6 kg/
Combining seasonal movement rates with annual time budgets, day
adult male Amur tigers spent 6.8 h traveling/day (95% CI 6.3–7.2) Pikunov 1988 Russian Empirical snow 5–15 kg/day
and moved 7.9 km/day (95% CI 6.8–9.1) in summer and 6.7 km/ Far East tracking
day (95% CI 5.7–7.9) in winter. Adult female non-reproducing Miller 2013 Russian Empirical GPS 8.9 kg/day
Amur tigers spent 7.9 h traveling/day (95% CI 7.6–8.3) and aver- et al. Far East cluster
aged 6.6 km/day (95% CI 5.9–7.3) in summer and 4.5 km/day sampling
(95% CI 3.7–5.3) in winter. Mean travel speed was estimated to This Russian Energetics # – 5.2 kg/day
be 1.173 km/h (95% CI 1.088–1.258) for male tigers and study Far East modeling
0.828 km/h (95% CI 0.781–0.875) for non-reproducing female ti- $ – 3.9 kg/day
gers during summer and 0.998 km/h (95% CI 0.903–1.093) for
males and 0.564 km/h (95% CI 0.491–0.637) for non-reproducing
female tigers during winter (Table B.1). In every instance, tigers
traveled less during winter than summer.
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