Technical ReportPDF Available

Modeling of tiger and its prey populations in the Balkhash Lake Region as the basis for adaptive management of the Tiger Reintroduction Program in Kazakhstan

Authors:

Abstract and Figures

The southern shore of Lake Balkhash in Kazakhstan as well as the area surrounding and to the east of the Ili River delta has been identified by WWF as a potential site for a tiger restoration program. This area still has vast tugay woodlands and reed thickets that were populated by Caspian tigers (Panthera tigris virgata) up to the middle of 20th century. These riparian ecosystems still cover more than 7,000 km² on the southern shore of Balkhash Lake and may represent suitable habitat for tiger introduction. Currently about 25% of the ecosystems are degraded due to intense annual fires and livestock grazing, but may be restored via fire management, grazing restrictions and support of optimal water regime in Ili River. Tugay woodlands and reed thickets can contain high population densities of wild boar (Sus scrofa) and Bukhara deer (Cervus elaphus bactrianus), main prey species for tiger in Central Asia. Roe deer (Capreolus pygargus) is another traditional prey species for tiger in the area, although its population density is relatively low. Circumstantial historical evidence collected from hunting records and encounters indicate that the Caspian tiger population density in Central Asia was much higher than that of the Amur tiger and was similar to tiger densities in India. Therefore, restoration of the tiger population in the Balkhash region could make a considerable contribution to the Global Tiger Recovery Program. This report provides details of projection modeling of the area of potential tiger habitat – tugay woodlands and reed thickets - as well as population growth models for tiger and its prey species (wild boar, Bukhara deer, and roe deer) on the southern shore of Balkhash Lake and Ili River delta over the next 50 years given three scenarios of possible habitat change and management options for tiger introduction program. The objectives of this modeling exercise are as follows: - To identify possible changes in the distribution and area of potential tiger habitat (tugay woodlands and thicket); - To define time-to-recovery for prey species (wild boar, Bukhara deer, and roe deer) and predict population dynamics given three scenarios of possible habitat changes; - To estimate optimal timing and regimes for tiger introduction given temporal change in the availability of their prey; - To evaluate potential number of tigers that could populate the project area and number of tigers needed for introduction; - To evaluate impact on source populations of tigers and Bukhara deer due to removal of individuals for introduction.
Content may be subject to copyright.
World Wide Fund for Nature (WWF-Russia)
Department of Environmental and Forest Biology, State University of New
York College of Environmental Science and Forestry
1
January 2015
REPORT
Modeling of tiger and its prey populations in the Balkhash Lake Region as the
basis for adaptive management of the Tiger Reintroduction Program in
Kazakhstan
Mikhail Paltsyn,
James P. Gibbs,
Liza Iegorova,
Olga Pereladova
Modeling of tiger and prey in Kazakhstan
2
CONTENTS
INTRODUCTION ....................................................................................................................................... 3
Data and methods ....................................................................................................................................... 4
Habitat and Scenarios ......................................................................................................................... 4
Prey species population growth models .......................................................................................... 13
Tiger and prey models ...................................................................................................................... 15
Model limitations .............................................................................................................................. 17
RESULTS .................................................................................................................................................. 19
Population projections for tiger prey species under different scenarios ...................................... 19
Population projections for tigers under different scenarios and schemes of introduction ......... 29
Population projections for Bukhara deer source population at Karachingil Game Management
Area .................................................................................................................................................... 39
Population projections for source population of Amur tiger in the Russian Far East ............... 41
CONCLUSIONS ....................................................................................................................................... 43
Modeling of tiger and prey in Kazakhstan
3
INTRODUCTION
The southern shore of Lake Balkhash in Kazakhstan as well as the area surrounding and to the
east of the Ili River delta has been identified by WWF as a potential site for a tiger restoration
program
1
. This area still has vast tugay woodlands and reed thickets that were populated by
Caspian tigers (Panthera tigris virgata) up to the middle of 20th century
2
. These riparian
ecosystems still cover more than 7,000 km² on the southern shore of Balkhash Lake and may
represent suitable habitat for tiger introduction. Currently about 25% of the ecosystems are
degraded due to intense annual fires and livestock grazing, but may be restored via fire
management, grazing restrictions and support of optimal water regime in Ili River. Tugay
woodlands and reed thickets can contain high population densities of wild boar (Sus scrofa) and
Bukhara deer (Cervus elaphus bactrianus), main prey species for tiger in Central Asia. Roe deer
(Capreolus pygargus) is another traditional prey species for tiger in the area, although its
population density is relatively low. Circumstantial historical evidence collected from hunting
records and encounters indicate that the Caspian tiger population density in Central Asia was
much higher than that of the Amur tiger and was similar to tiger densities in India
3
. Therefore,
restoration of the tiger population in the Balkhash region could make a considerable contribution
to the Global Tiger Recovery Program.
This report provides details of projection modeling of the area of potential tiger habitat tugay
woodlands and reed thickets - as well as population growth models for tiger and its prey species
(wild boar, Bukhara deer, and roe deer) on the southern shore of Balkhash Lake and Ili River
delta over the next 50 years given three scenarios of possible habitat change and management
options for tiger introduction program. The objectives of this modeling exercise are as follows:
- To identify possible changes in the distribution and area of potential tiger habitat (tugay
woodlands and thicket);
- To define time-to-recovery for prey species (wild boar, Bukhara deer, and roe deer) and
predict population dynamics given three scenarios of possible habitat changes;
- To estimate optimal timing and regimes for tiger introduction given temporal change in
the availability of their prey;
- To evaluate potential number of tigers that could populate the project area and number of
tigers needed for introduction;
- To evaluate impact on source populations of tigers and Bukhara deer due to removal of
individuals for introduction.
1
WWF 2014. Tiger Reintroduction Program in Kazakhstan
2
Sludskiy A.A. 1953. Tiger in the USSR. News of the Academy of Sciences of Kaz. SSR, ser. boil., №8, p.18-73,
(in Russ)
3
WWF 2014. Tiger Reintroduction Program in Kazakhstan
Modeling of tiger and prey in Kazakhstan
4
Data and methods
Habitat and Scenarios
Landsat 5 TM imageries for July-August 1989 and July-August 2010 were used as basis for
habitat mapping in the project area (source: USGS Earth Explorer http://earthexplorer.usgs.gov/).
These imageries were mosaiced and classified into 30 classes using Band 4 (Near-infrared),
Band 1(blue) and Band 2 (green) with the ArcMap 10.2.2. Image Classification function. After
that the imageries were reclassified in four general habitat classes: 1 water; 2 desert and
semi-desert; 3 dry bush (including saxaul forest), meadow and steppe; 4 tugay woodlands
and reed thickets. Map of ecosystems of Ili River delta produced by TERRA Center, Kazakhstan,
was used for reference during habitat classification process.
We assumed that habitat map prepared on the base of Landsat 5 TM 2010 classification
adequately represents current distribution and area of different habitat types at water level in
Balkhash Lake about 343 m above sea level.
4
The habitat map developed on the base of Landsat
5 TM 1989 imagery was assumed to represent general habitat distribution in the conditions of
water scarcity (in 1987-1989 the water level in the lake was 340-341 m above sea level). Also,
we assumed that the habitats can potentially come back to the conditions of 1987-1989 in about
20 years from now if water deficiency has the same effects on habitat as in 1969-1988 (the last
historic period of water deficiency in the Ili River) (Figure 1)
5
.
Figure 1. Dynamic of water level in Balkhash Lake in 1880-2005 (Bragin 2006).
6
4
Bragin E.A. 2006. Review of Balkhash Lake conditions and politics in the field of water resource management in
Ile-Balkhash watershed. Technical Report. WWF-Russia. (in Russ.)
5
Ibid
6
Bragin E.A. 2006. Review of Balkhash Lake conditions and politics in the field of water resource management in
Ile-Balkhash watershed. Technical Report. WWF-Russia. (in Russ.)
Modeling of tiger and prey in Kazakhstan
5
We used MODIS MCD45A1 data (Burned Area Monthly L3 Global 500m of LP DAAS
https://lpdaac.usgs.gov/products/modis_products_table/mcd45a1 ) to map area and intensity of fires in
tugay forest and reed thickets. Data used were from for April-July 2000-2014 - period of wild
fires in the project area. Fires are one of the main causes of degradation of tugay and reed thicket
ecosystems
7
. According to the data of the TERRA Center and the Institute of Geography
8
reed
thickets require about one year to restore after one-time burning and tugay woodlands can restore
themselves in 4-5 years after burning in conditions of periodic flooding (at least once in 3-4
years). Tugay forest and reed thickets represent one type of land cover on our habitat maps,
therefore, for the purpose of modeling we assumed that the areas that were burned once requires
at least 3 years for recovering in average, areas burned twice at least 6 years to recover, areas
burned thrice 9 years, and so on given optimal regime of flooding and current water level in
Balkhash Lake, and absence of intensive grazing. Thus, all areas of tugay forest and reed thicket
burned in 2000-2014 were classified as degraded ecosystems. The map representing current
conditions of tiger and its prey habitat is shown on the Figure 2 A. This map was used as starting
point for three scenarios considered in our modeling exercise.
For the purpose of modeling all areas including potential tiger habitats were divided in three
Tiger Management Units. We assumed that only tugay woodlands and reed thickets represent
optimal habitat for tiger, wild boar and Bukhara deer
9
. Dry bush (including saxaul forest),
meadow and steppe were classified as marginal habitat for these species, but optimal for roe
deer
10
. Burned tugay woods and reed thickets also represent marginal habitats in our modeling,
due to their intensive use as pastures and low quality of cover. Water, deserts and semi-deserts
were designated as non-habitat for all four species listed above. Maximal population densities for
the prey species in different habitats are shown in the Table 1.
Table 1. Estimated maximum population densities of wild boar, Bukhara deer and roe deer
in Central Asia biomes.
Species
Maximal population density (ind./km²)
High quality tugay
woodlands and reed
thickets
Low quality tugay
woodlands and reed
thickets (burned
areas)
Dry bush (including
saxaul forest),
meadow and steppe
Wild boar
6.011
0.5
0.5
Bukhara deer
10.012
0.5
1.013
7
Institute of Geography 2013. Preparation of socio-economic component of long-term Program for tiger
reintroduction in the Ili River delta and southern shore of Balkhash Lake. Research Report. (in Russ.)
8
Ibid
9
WWF 2014. Tiger Reintroduction Program in Kazakhstan
10
Ibid
11
population density of wild boars can achieve up to 5-30 animals/km² in tugay and reed ecosystems of Central Asia
(Geptner et al. 1969; Danilkin 2006; WWF 2014)
12
Pereladova O.B. 2013. Restoration of Bukhara deer (Cervus elaphus bactrianus Lydd.) in Central Asia in 2000-
2011. IUCN Deer Specialist Group News. March 2013. Pp. 19-30.
13
Ibid
Modeling of tiger and prey in Kazakhstan
6
Roe deer
0.5
0.5
1.014
Figure 2. Projected dynamics of tugay woodland and reed thicket ecosystem in the Ili-
Balkhash region under three management scenario: A - No Fire Management, B Fire
Management Years 30-50, and C Water scarcity Year 20. Tiger Management Units
(TMU) planned for introduction: 1 Balkhash, 2 Ili delta, 3 Karatal.
We assumed that habitat quality for tiger and its prey species can be gradually improved through
a process of active fire management and restrictions on livestock grazing, and that increase the
area of high quality tugay forest and reed thickets by 25-27% given flooding regime is constant
and similar to the period of 2010-2013 (Figure 2 B).
Also we considered possibility that water volume in Ili River and Balkhash Lake may drop to the
level of 1987-1989 as a result of increased water consumption in western China in the nearest 20
years. In this situation tugay forest and reed thicket may shrink considerably (Landsat 5 TM
14
WWF 2014. Tiger Reintroduction Program in Kazakhstan
Modeling of tiger and prey in Kazakhstan
7
1989 imagery) in 20 years from now (Figure 2 C) and may continue shrinking further at the same
rate.
Maps shown on Figure 2 A-C were used for developing of three scenarios of future habitat
conditions for tiger and its prey species on the southern shore of Balkhash Lake.
Scenarios
Three scenarios were used for our modeling exercise:
I. No fire management (area and quality of optimal tiger and prey species habitat is
constant in the nearest 50 years and equal to the area of the habitat in 2010);
II. Fire management (area and quality of optimal habitat gradually increase due to fire
control and grazing restrictions);
III. Water scarcity (area and quality of optimal habitats gradually decrease as a result of low
water volume in Ili River).
No fire management scenario
This scenario makes the following assumptions over the next 50 years:
- No fire management is implemented in the project area. Thus, area and frequency of
fires in tugay woodlands and reed thicket remains constant and follows the pattern of
fires in 2000-2014 (Figure 2 A);
- Habitat area and quality is constant and do not change (Figure 2 A); therefore, carrying
capacity for wild boar, Bukhara deer and roe deer is a constant value (Table 2);
- Water volume in Ili River remains constant at the level of 2010-2013;
- Harvesting of prey species is completely stopped in the project area;
- 30 Bukhara deer are brought annually in each TMU during first 10 years of the
program;
- Intensive management is used to achieve maximum population densities of wild boar
and Bukhara deer in the TMUs (see Table 1).
Table 2. Area and carrying capacity of habitat in three TMUs at No Fire Management
Scenario (Figure 2)
TMU
Area,
km²
Carrying capacity, K
Wild boar
Bukhara
Deer
Roe deer
All prey species
Balkhash
909.6
5580
9328
687
15595
27.264
Modeling of tiger and prey in Kazakhstan
8
218.3
Ili River
Delta
3696.3
23887
39599
4484
67970
1565.3
1853.4
Karatal
690.0
4580
7689
1134
13402
182.0
698.0
Estimated carrying capacity of all
TMUs:
34047
56576
6305
96968
Fire Management Scenario
The Fire Management scenario is based on the following assumptions:
- All fires are prevented in tugay woodlands and reed thickets from the first year of the
program;
- Strict restrictions on livestock grazing are implemented starting the first year of the
program (number of livestock gradually decreases in the project area);
- Area of high quality tugay woodlands and reed thicket gradually increases (gradual
move from Figure 2 A to Figure 2 B). Habitat carrying capacity for wild boar and
Bukhara deer gradually increases with increase of the area of restored tugay and reed
ecosystems (Figures 3-5, Table 3);
- Water volume in Ili River remains constant at the level of 2010-2013;
Modeling of tiger and prey in Kazakhstan
9
- Harvesting of prey species is completely stopped in the project area;
- 30 Bukhara deer are brought annually in each TMU during first 10 years of the
program;
- Intensive management is used to generate maximum population densities of wild boar
and Bukhara deer in the TMUs (see Table 1).
Figure 3. Estimated carrying capacity of habitat in Balkhash TMU under Fire
Management Scenario.
Figure 4. Estimated carrying capacity of habitat in Ili River Delta TMU under Fire
Management Scenario.
0
2000
4000
6000
8000
10000
12000
010 20 30 40 50 60
Carrying capacity, K
Years
Wild Boar
Bukhara deer
Roe deer
0
10000
20000
30000
40000
50000
60000
010 20 30 40 50 60
Carrying capacity, K
Years
Wild boar
Bukhara deer
Roe deer
Modeling of tiger and prey in Kazakhstan
10
Figure 5. Estimated carrying capacity of habitat in Karatal TMU under Fire Management
Scenario.
Table 3. Estimated carrying capacity of habitat in three TMUs before and after 30 years of
fire management
TMU
Area of high
quality tugay
woodland and
reed thicket,
km²
Carrying capacity, K
Before
mgmt
After
mgmt
Wild
boar,
before
mgmt
Wild
boar,
after
mgmt
Bukhara
deer,
before
mgmt
Bukhara
deer,
after
mgmt
Roe
deer,
before
mgmt
Roe
deer,
after
mgmt
All prey,
before
mgmt
All prey,
after
mgmt
Balkhash
909.6
936.8
5580
5730
9328
9586
687
687
15595
16003
Ili River Delta
3696.3
5261.6
23887
32496
39599
54470
4484
4484
67970
91450
Karatal
690.0
872.0
4580
5580
7689
9418
1134
1134
13403
16132
All TMUs
5295.9
7070.4
34047
43806
56616
73474
6305
6305
96968
123585
Thus, as a result of fire management carrying capacity of the habitat can be potentially increased
by 27%.
0
1000
2000
3000
4000
5000
6000
7000
8000
9000
10000
010 20 30 40 50 60
Carrying capacity, K
Years
Wild boar
Bukhara deer
Roe deer
Modeling of tiger and prey in Kazakhstan
11
Water Scarcity Scenario
This scenario made the following assumptions over the next 50 years:
- Water volume in Ili River and water level in Balkhash Lake gradually decrease with the
same rate as in 1969-1989 (Figure 1);
- No fire management is implemented in the project area. Thus, area and frequency of
fires in tugay woodlands and reed thicket slowly increases (Figure 2 C);
- Area and quality of tugay woodlands and reed thickets gradually decrease due to water
scarcity with the same rate as in 1969-1989 and come to the conditions shown on Figure
4 (1989) after 20 years from now. The habitat continues to deteriorate and shrink with
the same rate through the nearest 50 years;
- Carrying capacity of the habitat decreases with the same rate as high quality tugay
woodlands and reed thickets deteriorates and shrink (Figure 6-8, Table 4);
- Harvesting of prey species is completely stopped in the project area;
- 30 Bukhara deer are brought annually in each TMU during first 10 years of the
program;
- Intensive management is used to generate maximum population densities of wild boar
and Bukhara deer in the TMUs (see Table 1).
Figure 6. Estimated carrying capacity of habitat in Balkhash TMU under Water Scarcity
Scenario
0
1000
2000
3000
4000
5000
6000
7000
8000
9000
10000
010 20 30 40 50 60
Carrying capacity, K
Year
Wild boar
Bukhara deer
Roe deer
Modeling of tiger and prey in Kazakhstan
12
Figure 7. Estimated carrying capacity of habitat in Ili River Delta TMU under Water
Scarcity Scenario
Figure 8. Estimated carrying capacity of habitat in Karatal TMU under Water Scarcity
Scenario
Table 4. Possible change of habitat carrying capacity in three TMUs under Water Scarcity
Scenario
Habitat/Species
Current
(Figure 2)
Year 10
Year 20
(Figure 4)
Year 30
Year 40
Year 50
Balkhash TMU
Area of high quality tugay woodland and
reed thicket, km²
909.6
752.8
596.0
439.2
282.4
125.6
Wild boar, carrying capacity
5580
4714
3752
2790
1828
866
0
5000
10000
15000
20000
25000
30000
35000
40000
45000
010 20 30 40 50 60
Carrying capacity, K
Years
Wild boar
Bukhara deer
Roe deer
0
1000
2000
3000
4000
5000
6000
7000
8000
9000
010 20 30 40 50 60
Carrying capacity, K
Year
Wild boar
Bukhara deer
Roe deer
Modeling of tiger and prey in Kazakhstan
13
Bukhara deer, carrying capacity
9328
7884
6279
4675
3070
1466
Roe deer, carrying capacity
687
687
687
687
687
687
All prey, carrying capacity
15595
13285
10718
8152
5585
3019
Ili River Delta TMU
Area of high quality tugay woodland and
reed thicket, km²
3696.3
3190.5
2685.0
2179.5
1674
1168.5
Wild boar, carrying capacity
23887
21189
18192
15194
12196
9199
Bukhara deer, carrying capacity
39599
35113
30129
25145
20161
15177
Roe deer, carrying capacity
4484
4484
4484
4484
4484
4484
All prey, carrying capacity
67970
60786
52805
44823
36841
28860
Karatal TMU
Area of high quality tugay woodland and
reed thicket, km²
690.0
595.0
500.0
405
310
215
Wild boar, carrying capacity
4580
4004
3365
2725
2086
1446
Bukhara deer, carrying capacity
7689
6750
5706
4662
3619
2576
Roe deer, carrying capacity
1134
1134
1134
1134
1134
1134
All prey, carrying capacity
13403
11888
10205
8521
6839
5156
All TMUs
All prey, carrying capacity
96968
85959
73728
61496
49265
37035
Under the Water Scarcity Scenario habitat carrying capacity for tiger’s prey species may
decrease by 25% in 20 years and by 62% in 50 years.
Prey species population growth models
Three ungulate species wild boar, Bukhara deer and roe deer were assumed to be main preys
for tigers after introduction in all three TMUs. To describe population growth of the prey species
we implemented a model of density-dependent population (logistic) growth typically used in
wildlife population projection scenarios such as this:
Nt+1 = Nt + rNt(1- Nt/K), where
Nt is abundance of a population in year t ;
Nt+1 is abundance of a population next year (t+1);
r is instantaneous per capita growth rate; and
K is population carrying capacity.
Modeling of tiger and prey in Kazakhstan
14
Population abundance for wild boar, Bukhara deer and roe deer in TMUs in the first year of the
program (N0) as well as r values for these species are shown in the Table 5. Carrying capacity
for prey species at different scenarios are listed in the Tables 2-4.
Table 5. Estimated values of N0 and r =for wild boar, Bukhara deer, and roe deer in three
TMUs
Species
r mean
SD
Population abundance (Year 0)
Balkhash
Ili River
Delta
Karatal
Wild boar
0.34915
0.503
46016
188517
35218
Bukhara
deer
0.14419
0.111
3020
30
30
Roe deer
0.02021
0.143
20022
90023
17024
Environmental stochasticity was added to the deterministic population models via lognormal
distribution of r values
25
. For each scenario a total of 1000 iterations were performed to calculate
mean prey species abundance (and its 95% confidence interval) at years 5, 10, 15, 20, 25, 30, 35,
40, 45, and 50.
At present the Bukhara deer population from Karachingil Game Management Area is targeted
for use as an only source of deer for introduction in the TMUs. Currently this population is at
carrying capacity and has 400-500 animals in the area of 200 km²
26
. We used same population
model as above for Bukhara deer to define how many deer can be harvested in Karachingil for
introduction in the first 5-10 years of the program. The goal of our management for Karachingil
population was to avoid population decline below half of the population carrying capacity
(approximately 225 animals). Newly established populations in one of the TMUs can be used as
an additional source of deer for other two TMUs after 10-15 years of the program.
15
Calculated as average from time-series for wild boar populations in the Russian Far East (Zaumyslova O.Yu.
2005), Belovezhskay Puscha, Crimea, and Europe (Mayer J.J. 2009). No time-series for tugay forest and reed thicket
were found.
16
Our assumption based on Lukarevsky and Baydavletov 2010; WWF 2014
17
Ibid
18
Ibid
19
Calculated as average using time-series from Badai-Tuagai Nature Reserve and Karachingil Game Management
Area in 1999-2011(Pereladova O.B. 2013. Restoration of Bukhara deer (Cervus elaphus bactrianus Lydd.) in
Central Asia in 2000-2011. IUCN Deer Specialist Group News. March 2013. Pp. 19-30.)
20
30 deer will be brought to the selected TMU annually in the first 10 years of the program.
21
Calculated as average from time-series from Russian Far East, Norway, and France ((Ignatova et al., 2004; Nilsen
et al., 2009; Melis et al., 2010). No time-series for tugay forest and reed thicket were found.
22
Our assumption based on Lukarevsky and Baydavletov 2010; WWF 2014
23
Ibid
24
Ibid
25
Hood, G. M. 2010. PopTools version 3.2.5. Available on the internet. URL http://www.poptools.org
26
R. Baidavletov, personal communication
Modeling of tiger and prey in Kazakhstan
15
Tiger and prey models
We used age class matrix for tiger females to build a model of tiger population growth in the
project area. We assumed that density-dependence in our models affects only reproduction, but
not survival rates. For our modeling exercise we used following assumptions:
- tugay forest and reed thickets represent optimal habitat for tiger in the project area
27
;
- dry bush (including saxaul forest), meadow and steppe were assumed to be marginal
habitat where tigers can come occasionally for a short period of time;
- habitat carrying capacity for tigers is limited by prey species population density and
abundance only
28
;
- only sub-adult and adult tigers kill their own prey;
- number of particular prey species killed by a tiger directly depends on the ratio of this
species abundance to all prey species abundance in the project area;
- only sub-adult and adult tiger abundance was used to calculate carrying capacity
depending on prey abundance;
- female/male ratio of tiger population is constant and similar to that of the Amur tiger;
- tiger vital rates in the project area are the same as for Amur tiger. Parameters used for
tiger-prey population models are shown in the Table 6.
Table 6. Vital rates and other parameters used for tiger-prey population models
Age
stages29
Survival
rates30
Fecundity31
Female/male ratio32
Average number of
ungulates killed by 1 tiger
annually
Average prey abundance
necessary to support 1
tiger
Cub (0-1)
0.6
0
1.533
0
-
Juvenile
(1-2)
0.8
0
0
Sub-Adult
(2-3)
0.6
0
6034
45035
Adult (3+)
0.8
0.75
27
Sludskiy A.A. 1953. Tiger in the USSR. News of the Academy of Sciences of Kaz. SSR, ser. boil., №8, p.18 -73,
(in Russ). WWF 2014. Tiger Reintroduction Program in Kazakhstan
28
Karanth K.U., Nichols J.D., Kumar N. S., Link W.A, and J.E. Hines. 2004. Tigers and their prey: Predicting
carnivore densities from prey abundance. PNAS. 101(14). Pp. 48544858.
29
Tian Yu. Et al. 2011. Population viability of Siberian tiger in changing landscape: going, going and gone?
Ecological Modeling. 222. Pp. 3166-3180
30
Ibid
31
Ibid
32
Ibid
33
The actual female/male ratio for adult Amur tigers is between 5/3 and 6/5 based on field surveys (Carroll and
Miquelle, 2006; Miquelle et al., 2006; Tian et al. 2011).
34
According to the Strategy for Amur tiger conservation in Russia (2010) one tiger annually kill 50-70 ungulates.
Karanth et al. 2004 applied the average kill rate of 50 ungulates/tigers per year consistently observed in field studies
of tigers.
35
According to the Strategy for Amur tiger conservation in Russia (2010) one tiger needs population of 400-500
ungulates to support itself with sufficient prey. According to Karanth et al. 2004 average tiger/ungulate ratio should
be about 1/500.
Modeling of tiger and prey in Kazakhstan
16
We used following formula to calculate carrying capacity for tigers in TMUs:
Kt = Nprey t/450, where
Kt is carrying capacity for tigers in a TMU in the year t,
Nprey t is abundance of all prey species (wild boar, Bukhara deer and roe deer) in the TMU,
450 is the mean number of ungulates required to support one tiger.
The effect of tiger predation on a particular ungulate species in the TMUs was calculated using
the following formula:
Nt+1 = Nt + rNt(1- Nt/K) Ns+a t * (Nt / Np t)*R s+a , where
Nt is abundance of an ungulate species population in year t ;
Nt+1 is abundance of an ungulate species population next year (t+1);
r is the instantaneous per capita growth rate of an ungulate species;
K is ungulate species population carrying capacity;
Ns+a t number of sub-adult and adult tigers in year t;
Np t abundance of all ungulate species in year t;
R s+a number of ungulates killed by one tiger annually (constant value equal 60).
Environmental stochasticity was added to the tiger models as changing carrying capacity
determined by variation of prey species numbers. No demographic stochasticity was considered
in our modeling. Tiger population modeling was implemented on the outputs of the ungulate
species population growth models
36
. We explored different schemes of tiger introduction
(bringing different number of tigers in different years to TMUs at different scenarios) to find an
acceptable one. The criterion for acceptability was that the lower level of 95% confidence
interval would not cross quasi-extinction threshold (10 tigers) considerably.
Amur tiger population in the Russian Far East is currently considered as the most probable
source of tigers for introduction in Ili-Balkhash TMUs. According to the data of last tiger counts
in 2005 the population has 428-502 tigers (465 in average)
37
. The population has following sex
and age structure: 25% - adult males, 39% adult females, 22% cubs up to 1.5 years old, 6% -
juveniles and sub-adults (1.5-3 years old)
38
. The same matrix model for females as above was
used to identify number of tigers that can be used for introduction program without considerable
36
Hood, G. M. 2010. PopTools version 3.2.5. Available on the internet. URL http://www.poptools.org
37
Strategy for conservation of the Amur Tiger in the Russian Federation. Approved by the Ministry of Natural
Resources and Environment of the Russian Federation, Decree of July 2, 2010, # 25-r.
38
Ibid
Modeling of tiger and prey in Kazakhstan
17
impact for Russian Far East population (Table 6). For the model we assumed that carrying
capacity for the Amur tiger in the Russian Far East in current conditions is 510-550 (530 in
average)
39
. Following population vector for tiger population in the Russian Far East was used for
the year 1 of modeling:
Age stage
Abundance (females only)
Cub (0-1)
51
Juvenile (1-2)
30
Sub-Adult (2-3)
17
Adult (3+)
181
N total
279
5 tigers (3 females and 2 males) were removed every 3, 4 or five years from the source
population for different number of years according to various schemes of tiger introduction in the
TMUs on the shore of Balkhash Lake. For this model we assumed that the source population will
have same survival and reproduction rates for the nearest 50 years and will not be considerably
affected by poaching, habitat destruction and prey depletion. No environmental and demographic
stochasticity was considered for Amur tiger source population model.
Model limitations
Our models have following limitations that should be recognized and considered for their
implementation as a tool for adaptive management of the tiger introduction program:
Limited data were available wild boar and roe deer current abundance upon which to base
the model estimates of prey population dynamics; field population surveys are necessary
to evaluate current population status and dynamics of these species in the Ili-Balkhash
Basin.
No time-series of population numbers for wild boar and roe deer for tugay forest and reed
thickets in Kazakhstan or other regions of Central Asia were available. Thus, the r mean
values were calculated using time-series for these species populations from other areas
and habitat types. As a result of this approach annual population growth of wild boar and
roe deer have very high variability that effect the precision of the models. The models
could be improved a great deal if time-series for wild boar and roe deer populations in the
tugay woodlands and reed thickets become available. These data may be available from
Nature Chronicles of Central Asian Nature Reserves (Zapovedniks).
Our models are based on the highest population densities for wild boar, roe deer and
Bukhara deer that could be achieved in the tugay woodlands and reed thickets. It may not
be a real option for a 7,000 km² of such habitat in the Ili-Balkhash Basin. Thus, additional
scenarios with average population densities of this species should be considered to make
the models to be an effective tool for adaptive management of the program.
39
Goal 2020 of the Strategy for conservation of the Amur Tiger in the Russian Federation. Approved by the
Ministry of Natural Resources and Environment of the Russian Federation, Decree of July 2, 2010, # 25-r.
Modeling of tiger and prey in Kazakhstan
18
Vital rates of tigers in Ili-Balkhash Basin might be very different from vital rates of tigers
in the Russian Far East used in our tiger population models.
It is extremely difficult to predict actual water balance in the Ili River and Balkhash Lake
over the long term. Thus, our assumptions for the Water Scarcity Scenario based on the
satellite imageries of 1989 and 2010 could be far from perfect.
Our models for wild boar, Bukhara deer, roe deer and tigers do not consider effect of
demographic and genetic stochasticity on the species population growth (although these
are likely to be inconsequential for two reasons: prey population sizes are generally large
and tiger populations can persist even at very low densities given the intrinsic mobility of
tigers).
Model for source population of Amur tiger in the Russian Far East did not consider
demographic, environmental and genetic stochasticities that can make population growth
much slower and less predictable.
No environmental autocorrelation, catastrophes or bonanzas were taken in account in our
models because these phenomena and their effects of prey populations (should they even
occur) are difficult to estimate at this time.
Modeling of tiger and prey in Kazakhstan
19
RESULTS
Population projections for tiger prey species at different scenarios
In all depictions of likely future population scenarios the solid line = mean population abundance
and dashed lines = upper and lower limits of 95% confidence interval.
BALKHASH TMU
No Fire Management Scenario
Wild boar
Bukhara deer
Roe deer
All prey
Years 10-15 of the program is likely to be the earliest year to start tiger introduction in Balkhash
TMU (N mean = 3779-5560, enough prey base for first five tigers (3 females and 2 males)), but
at that time high probability exists that the prey base could be still to low (Lower CL = 902-
0
1000
2000
3000
4000
5000
6000
020 40 60
Number
Year
0
1000
2000
3000
4000
5000
6000
7000
8000
9000
10000
020 40 60
Number
Year
0
100
200
300
400
500
600
700
020 40 60
Number
Year
0
2000
4000
6000
8000
10000
12000
14000
16000
18000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
20
1450). In the worst case introduction can be started at year 20 when there would be minimally 5
tigers.
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 20
Tiger CC Year
20
Prey Year 5040
Tiger CC Year
5041
Nmean
7145
15
14710
32
Upper CL
8907
19
15320
34
Lower CL
2623
5
13463
29
Fire Management Scenario
Wild boar
Bukhara deer
Roe deer
All prey
40
Prey abundance without tiger predation
41
Carrying capacity without effect of tiger predation
-1000
0
1000
2000
3000
4000
5000
6000
7000
020 40 60
Number
Year
0
1000
2000
3000
4000
5000
6000
7000
8000
9000
10000
020 40 60
Number
Year
0
100
200
300
400
500
600
700
020 40 60
Number
Year
0
2000
4000
6000
8000
10000
12000
14000
16000
18000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
21
Fire management will not improve prey species carrying capacity considerably in Balkhash
TMU due to very limited area and frequency of fires in this territory. Still the earliest time to
bring fist 5 tigers (3 females and 2 males) to the TMU will be year 10-15. At Year 20 even in
the worst case scenario it will be appropriate to start introduction.
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 20
Tiger CC Year
20
Prey Year 5042
Tiger CC Year
5043
Nmean
6778
15
15030
33
Upper CL
9033
20
15671
34
Lower CL
2576
5
13667
30
Water Scarcity Scenario
Wild boar
Bukhara deer
Roe deer
All prey
42
Prey abundance without tiger predation
43
Carrying capacity without effect of tiger predation
0
1000
2000
3000
4000
5000
6000
020 40 60
Number
Year
0
1000
2000
3000
4000
5000
020 40 60
Number
Year
0
100
200
300
400
500
600
020 40 60
Number
Year
0
1000
2000
3000
4000
5000
6000
7000
8000
9000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
22
The introduction can be started at the same period as in the two previous scenarios, but since
year 30 carrying capacity for prey species is likely to drop considerably if the water volume
in Ili River will continue to go down. In this case Balkhash TMU will be unlikely to support
even smallest viable tiger population (at least 10 animals) by year 50.
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 20
Tiger CC Year
20
Prey Year 5044
Tiger CC Year
5045
Nmean
5652
12
2869
6
Upper CL
6955
15
4061
9
Lower CL
2940
6
2579
5
ILI RIVER DELTA TMU
No Fire Management Scenario
Wild boar
Bukhara deer
Roe deer
All prey
44
Prey abundance without tiger predation
45
Carrying capacity without effect of tiger predation
0
5000
10000
15000
20000
25000
30000
020 40 60
Number
Year
0
5000
10000
15000
20000
25000
30000
35000
40000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
23
In Ili River Delta TMU introduction can be started at the Year 5-10 of the program as the earliest
in the best case (N mean = 6582-14123), but still some probability exists that the prey base will
be lower than necessary (Lower CL = 1595-2298). But by Year 15 there would likely be enough
prey for the first 5 tigers even in the worst case of prey species population growth. The area has
much larger capacity for tiger population, than Balkhash TMU.
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 15
Tiger CC Year
15
Prey Year 5046
Tiger CC Year
5047
Nmean
20426
45
55643
123
Upper CL
27145
60
63355
140
Lower CL
3430
7
42316
94
46
Prey abundance without tiger predation
47
Carrying capacity without effect of tiger predation
0
500
1000
1500
2000
2500
3000
3500
4000
020 40 60
Number
Year
0
10000
20000
30000
40000
50000
60000
70000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
24
Fire Management Scenario
Wild boar
Bukhara deer
Roe deer
All prey
0
5000
10000
15000
20000
25000
30000
35000
020 40 60
Number
Year
0
10000
20000
30000
40000
50000
60000
020 40 60
Number
Year
0
500
1000
1500
2000
2500
3000
3500
4000
020 40 60
Number
Year
0
10000
20000
30000
40000
50000
60000
70000
80000
90000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
25
Fire management and grazing restrictions can improve tugay forest and red thicket quality and
increase carrying capacity for prey species and tigers by 29-30% in the nearest 30 years. Still the
earliest time for introduction is Year 5-10 and the latest is Year 15.
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 15
Tiger CC Year
15
Prey Year 5048
Tiger CC Year
5049
Nmean
24081
53
71943
159
Upper CL
33993
75
84532
187
Lower CL
3681
8
53960
120
Water Scarcity Scenario
Wild boar
Bukhara deer
Roe deer
All prey
48
Prey abundance without tiger predation
49
Carrying capacity without effect of tiger predation
0
5000
10000
15000
20000
25000
020 40 60
Number
Year
0
5000
10000
15000
20000
020 40 60
Number
Year
0
500
1000
1500
2000
2500
3000
3500
020 40 60
Number
Year
0
10000
20000
30000
40000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
26
Despite severe decline in habitat area and carrying capacity (37-40%) in the conditions of water
deficit Ili River Delta TMU is likely to be able to support a relatively stable tiger population (no
less than 30-50 animals) by the year 50.
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 15
Tiger CC Year
15
Prey Year 5050
Tiger CC Year
5051
Nmean
17821
39
26783
59
Upper CL
23394
51
29840
66
Lower CL
3245
7
22455
50
KARATAL TMU
No Fire Management Scenario
Wild boar
Bukhara deer
Roe deer
All prey
50
Prey abundance without tiger predation
51
Carrying capacity without effect of tiger predation
0
1000
2000
3000
4000
5000
020 40 60
Number
Year
0
2000
4000
6000
8000
10000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
27
Tiger introduction in Karatal TMU is likely to be safely started at the year 20 according to the
model above even at the slowest population growth of prey species. Year 10-15 is the earliest
possible option.
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 20
Tiger CC Year
20
Prey Year 5052
Tiger CC Year
5053
Nmean
6127
13
12251
27
Upper CL
7730
17
12909
28
Lower CL
2811
6
11416
25
Fire Management Scenario
Wild boar
Bukhara deer
52
Prey abundance without tiger predation
53
Carrying capacity without effect of tiger predation
0
200
400
600
800
1000
020 40 60
Number
Year
0
2000
4000
6000
8000
10000
12000
14000
020 40 60
Number
Year
0
1000
2000
3000
4000
5000
6000
020 40 60
Number
Year
0
2000
4000
6000
8000
10000
12000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
28
Roe deer
All prey
Fire management and grazing restrictions can improve carrying capacity of Karatal TMU for
tiger and prey species by 20-23%.
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 20
Tiger CC Year
20
Prey Year 5054
Tiger CC Year
5055
Nmean
7197
15
15186
33
Upper CL
9078
20
15792
35
Lower CL
3170
7
14360
31
Water Scarcity Scenario
Wild boar
Bukhara deer
54
Prey abundance without tiger predation
55
Carrying capacity without effect of tiger predation
0
200
400
600
800
1000
020 40 60
Number
Year
0
5000
10000
15000
20000
020 40 60
Number
Year
0
1000
2000
3000
4000
5000
020 40 60
Number
Year
0
1000
2000
3000
4000
5000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
29
Roe deer
All prey
Prey Abundance/
Tiger carrying
capacity (CC)
Prey Year 20
Tiger CC Year
20
Prey Year 5056
Tiger CC Year
5057
Nmean
6365
14
4846
10
Upper CL
9078
20
5362
12
Lower CL
2643
5
4404
9
In case of progressive water deficit in Ili River for nearest 50 years, Karatal TMU will be
unlikely to support viable tiger population by the year 50.
Population projections for tigers at different scenarios and schemes of introduction
In all graphical depictions of tiger population projections below black solid line = mean
population abundance, black dashed lines = upper and lower limits of 95% confidence interval,
and horizontal red line = quasi-extinction threshold (10 individuals).
BALKHASH TMU
No Fire Management Scenario
56
Prey abundance without tiger predation
57
Carrying capacity without effect of tiger predation
0
100
200
300
400
500
600
700
020 40 60
Number
Year
0
1000
2000
3000
4000
5000
6000
7000
8000
020 40 60
Number
Year
Modeling of tiger and prey in Kazakhstan
30
Scheme 1. 3 tiger females and 2 males relocated to the TMU on the Year 10, 3 more females
and 2 more males added on the Year 15. Nmean that is likely to be achieved by the year 50
using this scheme is 13-14 tigers only. This scheme is not considered as optimal, because
considerable part of 95% confidence interval is located below quasi-extinction threshold, thus
high probability exist the population could go extinct due to rapid prey depletion.
Scheme 2. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 23. This scheme allows to achieve Nmean equal to
17-18 tigers by the year 50, but still part of the confidence interval falls below quasi-extinction
threshold, though probability of extinction is much lower than in the previous case.
0
2
4
6
8
10
12
14
16
18
20
10 15 20 25 30 35 40 45 50 55
Number
Year
0
2
4
6
8
10
12
14
16
18
20
15 20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
31
Scheme 3. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 25. This scheme allows to achieve Nmean equal to
17 tigers by the year 50 and has relatively low probability of the population falling below quasi-
extinction threshold.
Scheme 4. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 28. This scheme allows to achieve Nmean equal to
17-18 tigers by the year 50 and has confidence interval all above the quasi-extinction threshold
since the year 27.
0
2
4
6
8
10
12
14
16
18
20
15 20 25 30 35 40 45 50 55
Number
Year
0
5
10
15
20
20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
32
Scheme 5. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 28, 2 females and 1 male added at the year 33.
This scheme allows to achieve a Nmean equal to 22-23 tigers by the year 50, but population has
high probability to again fall below quasi-extinction threshold after year 45.
Scheme 6. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 28 and 33. This scheme allows reaching Nmean
equal to 23 tigers by the year 50, but population has a probability to fall below quasi-extinction
threshold after year 45 due to prey depletion.
The maximum carrying capacity of Balkhash TMU for tigers is no more than 24-26 individuals
according to the model (includes effect of tiger predation on prey); to achieve this population
some 10-15 tigers (no more than 9 females and 6 males) should be introduced in the TMU in
years 20-33.
Fire Management Scenario
0
5
10
15
20
25
20 25 30 35 40 45 50 55
Number
Year
0
5
10
15
20
25
30
20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
33
For Balkhash TMU this scenario provided very similar results to the previous scenario; therefore
we do not consider its details further.
Water Scarcity Scenario
Scheme 7. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 25. In this case the tiger population can drop
below quasi-extinction threshold by the year 45 and therefore cannot be considered as viable.
Balkhash TMU evidently would not provide safe shelter even for small viable tiger population in
conditions of water deficit in Ili River.
0
2
4
6
8
10
12
14
16
18
15 20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
34
ILI RIVER DELTA TMU
No Fire Management Scenario
Scheme 8. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 25, 30, 35, and 40 (25 tigers introduced in the
TMU in Years 20-40). The total tiger population that is likely to be achieved by year 50 is 43-45
individuals.
Scheme 9. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 24, 28, 32, 36, 40, 44, and 48 (40 tigers
introduced in the TMU in Years 20-48). The total tiger population that is likely to be achieved
by year 50 is 64-69 individuals.
0
10
20
30
40
50
15 20 25 30 35 40 45 50 55
Number
Year
0
10
20
30
40
50
60
70
80
15 20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
35
Scheme 10. 3 tiger females and 2 males are relocated to the TMU on the Year 15, 3 more
females and 2 more males added on the Year 18, 21, 24, 27, 30, 33, 36, 39, 42 (50 tigers
introduced in the TMU in Years 15-42). The total tiger population that is likely to be achieved
by year 50 is 77-88 individuals, but it is likely that the population will drop down to 21-22
individuals as a result of prey depletion.
Maximal population of tigers that could be achieved in Ili River Delta TMU in the nearest 60-80
years at this scenario is 80-84 individuals (actual estimated carrying capacity of the TMU
including effect of tiger predation).
Fire Management Scenario
Scheme 11. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 24, 28, 32, 36, 40, 44, and 48 (40 tigers
introduced in the TMU in Years 20-48). The total tiger population that is likely to be achieved
by year 50 is 66-70 individuals.
0
10
20
30
40
50
60
70
80
90
100
15 20 25 30 35 40 45 50 55
Number
Year
0
10
20
30
40
50
60
70
80
15 20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
36
Scheme 12. 3 tiger females and 2 males are relocated to the TMU on the Year 15, 3 more
females and 2 more males added on the Year 18, 21, 24, 27, 30, 33, 36, 39, 42 and 45 (55
tigers introduced in the TMU in Years 15-45). The total tiger population that is likely to be
achieved by year 50 is 92-98 individuals, but it is likely that the population will drop down to 60
individuals as a result of prey depletion in this case.
Maximal sustainable tiger population that is likely to be achieved at this scenario is 90-94
individuals (actual carrying capacity of the TMU improved by fire management and grazing
restrictions).
Water Scarcity Scenario
Scheme 13. 3 tiger females and 2 males are relocated to the TMU on the Year 15, 3 more
females and 2 more males added on the Year 18, 21, 24, 27, 30, 33, 36, 39, 42 (50 tigers
introduced in the TMU in Years 15-42). In this case the tiger population could drop down to 5-
32 (mean = 18) individuals by Year 50 as result of habitat deterioration and prey depletion.
0
20
40
60
80
100
120
15 20 25 30 35 40 45 50 55
Number
Year
0
10
20
30
40
50
60
70
80
15 20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
37
Scheme 14. 3 tiger females and 2 males are relocated to the TMU on the Year 15, 3 more
females and 2 more males added on the Year 18, 21, 24, 27, 30, and 33 (35 tigers introduced
in the TMU in Years 15-33). In this case the tiger population could drop down to about 37 by
Year 50 as a result of habitat deterioration and prey depletion.
Scheme 15. 3 tiger females and 2 males are relocated to the TMU on the Year 15, 3 more
females and 2 more males added on the Year 18, 21, 24, 27, (25 tigers introduced in the
TMU in Years 15-27). In this case tiger population is likely to survive even the period of
continuous water scarcity in the nearest 50 years.
KARATAL TMU
No Fire Management Scenario
0
10
20
30
40
50
60
70
15 20 25 30 35 40 45 50 55
Number
Year
0
10
20
30
40
50
60
15 20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
38
Scheme 16. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 28 (10 tigers introduced in the TMU in Years
20-28). Population of 17-18 tigers may be achieved using this option by Year 50. Probability that
population will fall below quasi-extinction threshold is low.
Scheme 17. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 25, and 35 (15 tigers introduced in the TMU
in Years 20-35). Tiger population in 21-25 individuals can be achieved by year 50, but
probability exists that population will go below quasi-extinction threshold after year 45 due to
prey depletion. According to the model 21-25 the tiger population is above the TMU carrying
capacity and population will obviously decline after Year 50.
Karatal TMU can support maximal sustainable tiger population of 17-19 individuals for long-
term at this scenario
0
2
4
6
8
10
12
14
16
18
20
15 20 25 30 35 40 45 50 55
Number
Year
0
5
10
15
20
25
30
15 20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
39
Fire Management Scenario
This scenario is similar to the previous one, but carrying capacity of the TMU could increase up
to 23-25 tigers as a result of fire management and grazing restrictions. Thus, three groups of 5
tigers could be brought into the area with interval of 8-10 years.
Water Scarcity Scenario
Scheme 18. 3 tiger females and 2 males are relocated to the TMU on the Year 20, 3 more
females and 2 more males added on the Year 25 (10 tigers introduced in the TMU in Years
20-25). In the conditions of continuous water scarcity tiger population in Karatal TMU is likely
to fall below the quasi-extinction threshold by year 45-50.
Population projections for Bukhara deer source population at Karachingil Game
Management Area
Black solid line = mean population abundance, black dashed lines = upper and lower limits
of 95% confidence interval, horizontal red line = half of the source population carrying
capacity (225 individuals).
0
5
10
15
20
15 20 25 30 35 40 45 50 55
Number
Year
Modeling of tiger and prey in Kazakhstan
40
Scheme 19. 30 deer are removed annually for 10 years for introduction in one of the
TMUs. It is very low probability that the population will fall below half of its carrying
capacity. Thus, this scheme will likely be sustainable.
Scheme 20. 35 deer are removed annually for 9 years for introduction in one of the
TMUs. There is strong probability that the population will fall below half of its carrying
capacity in the area of less sustainable management. This scheme still could be sustainable
but should be implemented with caution.
0
50
100
150
200
250
300
350
400
450
500
010 20 30 40 50 60
Number
Year
0
50
100
150
200
250
300
350
400
450
500
010 20 30 40 50 60
Number
Year
Modeling of tiger and prey in Kazakhstan
41
Scheme 21. 40 deer are removed annually for 8 years for introduction in one of the
TMUs. This scheme has high probability of the source population decline below the half of
its carrying capacity. It is better to avoid this pattern of deer removal for introduction.
Population projections for source population of Amur tiger in the Russian Far East
0
50
100
150
200
250
300
350
400
450
500
010 20 30 40 50 60
Number
Year
460
470
480
490
500
510
520
530
10 20 30 40 50 60
Number
Year
Stable population, no tiger removal
3 F +2M removal Year 20, 28 and 33
3 F +2M removal Year 20, 25 and 35
3 F +2M removal Year 20, 25, 30, 35,
and 40
3 F +2M removal Year 15, 18, 21, 24,
27, 30, 33, 36, 39, 42
3 F +2M removal Year 15, 18, 21, 24,
27, 30, 33, 36, 39, 42 and 45
Modeling of tiger and prey in Kazakhstan
42
Scheme 22. Population projections for Amur tiger population in the Russian Far East at
different schemes of tiger group removal for introduction (3 females and 2 males in each
group). If poaching, habitat degradation and prey depletion do not influence Russian Amur tiger
population it is likely to continue to grow despite any reasonable off-take of individuals for
introduction in the Ili-Balkhash watershed. Effect of tiger removal on the Amur population is
summarized in the table below. First two options (removal of 5 tigers in three groups) are
acceptable if the introduction program starts with Balkhash or Karatal TMU: they have very
minor effect on Amur population (only 2 animal difference with baseline option by Year 50).
The last three options consider tiger introduction if the program starts with Ile River Delta TMU:
25-55 animals are removed in the period of 20-30 years. Last two options have noticeable impact
on the source population and should be carefully considered with different positions including
political context (conservation goals for Amur tiger population). Introduction of 25 tigers in the
Ili River Delta TMU will be enough to achieve population of 45-50 tigers (viable population for
a short-term) in 50-60 years of the program. Of course, poaching and habitat destruction impact
on the Amur population should be carefully considered and monitored at the start of introduction
program before any removal of tigers from Russian Far East.
Options of tiger removal
Amur tiger population abundance
Year 10
Year
20
Year 30
Year 40
Year 50
No removal (baseline)
470
492
508
520
526
3 F+2M removal Year 20, 28 and 33
(Balkhash or Karatal TMU)
470
487
503
515
524
3 F+2M removal Year 20, 25 and 35
(Balkhash or Karatal TMU)
470
487
504
515
524
3 F+2M removal Year 20, 25, 30, 35,
and 40 (Ili Delta TMU)
470
487
499
508
522
3 F+2M removal Year 15, 18, 21, 24,
27, 30, 33, 36, 39, 42 (Ili Delta TMU)
470
485
490
500
514
3 F+2M removal Year 15, 18, 21, 24,
27, 30, 33, 36, 39, 42 and 45 (Ili Delta
TMU)
470
485
490
500
512
Modeling of tiger and prey in Kazakhstan
43
CONCLUSIONS
Beginning of introduction program. Years 10-15 of the program may be considered as
the earliest possible option for tiger introduction in Balkhash and Karatal TMUs (first
group of 3 females and 2 males) in the best circumstances (high population growth of
wild boar first of all). High probability exists that the prey base could be still too low in
the TMUs at that time to initiate tiger introduction. In the worst case, introduction in
these TMUs can be started at years 20-25 when there could be sufficient prey abundance
to support at least 5 tigers.
If the program starts with Ile River Delta TMU first group of 5 tigers may be brought on
Year 5-10 of the program as the earliest possible option. As in the case of Balkhash and
Karatal TMUs it is probably that prey base may be still low at Year 5-10. If true, Year 15
will likely to have sufficient prey base to start introduction.
Number of tigers required for introduction. At least two groups of 5 tigers (3 females
and 2 males) may be introduced in Balkhash and Karatal TMUs with an interval of 5-10
years between releases of the groups (Schemes 3, 4, and 16). Maximum 3 groups of tigers
(15 tigers in total) might be possible to bring to Balkhash TMU (Scheme 6) in years 20-
33 with relatively low probability that the population falls below quasi-extinction
threshold due to prey depletion in the nearest 50 years. Three groups of tigers (15 tigers
Modeling of tiger and prey in Kazakhstan
44
total) will likely to be possible to introduce to Karatal TMU if habitat is improved with
fire management and grazing restrictions.
Ili River Delta TMU has much greater potential for introduction than other TMUs. Up to
40 tigers may be brought in the area in the Years 15-48 without risk of prey base
depletion under No Fire Management Scenario (Scheme 9). Though, introduction of 50
tigers in the TMU at this scenario may be risky, because the population may exceed its
carrying capacity and fall down (Scheme 10). Under Fire Management Scenario
introduction of 50-55 tigers in the TMU is possible, thought brining 55 tigers in the area
may lead to decrease of prey abundance in the year 50 and following decline of tiger
population.
Maximal population of tigers that is likely to be achieved by the Year 50 of the
program. Tiger abundance that is possible to achieve in Balkhash or Karatal TMUs by
the Year 50 is 17-18 individuals if two groups of 5 tigers are introduced in each TMU
(Schemes 3, 4, and 16). Introduction of three groups (15 tigers) in Balkhash TMU may
results in 23-25 tigers by Year 50 if the area has sufficient prey base (Scheme 6). Under
Fire Management Scenario introduction of 3 groups (15 tigers) in Karatal TMU will
likely to result in 22-24 tigers by the same year.
Introduction of 5 groups of tigers (25 tigers total) in Ile River Delta TMU may result in tiger
population of 43-45 individuals by Year 50 (Scheme 8). 40 introduced tigers may provide
maximal population of 64-69 individuals (Scheme 9) under No Fire Management Scenario and
66-70 individuals (Scheme 11) under Fire Management Scenario by Year 50. Release of 55
tigers in Ile River Delta TMU under Fire Management Scenario is likely to achieve 92-98
individuals (population carrying capacity) by the Year 50 (Scheme 12).
Carrying capacity of TMUs. Carrying capacity of Balkhash TMU under No Fire
Management Scenario and Fire Management Scenario does not exceed 24-26 tigers
(adults, sub-adults, juveniles and cubs). Karatal TMU is not able to support tiger
population more than 17-19 tigers under No Fire Management Scenario and 23-25 tigers
under Fire Management Scenario. Ile River Delta TMU has much higher carrying
capacity: 80-84 tigers under No Fire Management Scenario and 90-94 tigers under Fire
Management Scenario.
All three TMUs may have entire tiger population of 121-129 individuals if no fire
management and grazing restrictions are implemented in the area, and 137-145 animals if
habitat are improved under Fire Management Scenario.
Sustainability of tiger populations in the TMU. Tiger populations in Balkhash and
Karatal TMU alone cannot be considered as sustainable in the long-term due to their
small sizes. Water Scarcity Scenario clearly demonstrates that this population may go
extinct in the nearest 50 years if water level in Balkhash Lake drops down more than by
2-3 meters (Schemes 7 and 18). Ili River Delta TMU is likely to be much more
Modeling of tiger and prey in Kazakhstan
45
sustainable under Water Scarcity Scenario and may sustain 18-43 tigers after progressive
water level decline in Ili River for the nearest 50 years (Schemes 13-15). Thus, tiger
introduction in Balkhash and Karatal TMU may be considered as an option only in the
case if Ili River Delta TMU is populated with tigers. Even we recommend to start
introduction program with Ili River Delta TMU first: in this case tiger population in Ili
River Delta may be used as a source population for tiger introduction in Balkhash and
Karatal TMUs.
Optimal number of Bukhara deer removed from Karachingil Game Management
Area (GMA) for introduction in the TMUs. Removal of 30 deer annually for first 10
years of the program from Karachingil GMA for introduction in one of the TMUs may be
considered as the most sustainable option for Karatchingil Bukhara deer population
(Scheme 19). At the same time this option will allow to establish sustainable deer
population (450-800 individuals) in one of the TMU that can be used as a source
population for other TMUs. Removal 35 deer for first 9 years of the program is less
sustainable, because the source population will have high probability to decrease below
the half of the population carrying capacity (225 individuals) (Scheme 20). Removal of
40 deer for 8 years from Karachingil GMA is even less sustainable due to much higher
probability of driving the source population below the half of its carrying capacity
(Scheme 21).
Number of tigers that may be removed from Amur tiger population in the Russian
Far East for introduction. If poaching, habitat degradation and prey depletion do not
influence Russian Amur tiger population in the nearest 50 years it is likely to continue to
grow despite removal of individuals for introduction in the Ili-Balkhash watershed
(Scheme 22). Even removal of 11 groups of 5 tigers (3 females and 2 males each) 55
tigers in total in Year 15-45 of the program will not stop the population from growing
under these ideal conditions of no poaching, prey depletion or habitat loss. Introduction
of 25 tigers from outside into the Ili River Delta TMU will be enough to achieve
population of 45-50 tigers (viable population for a short-term) in 50-60 years of the
program. Of course, poaching and habitat destruction impact on the Amur population
should be carefully considered and monitored at the start of introduction program before
any removal of tigers from the Russian Far East is considered.
Modeling of tiger and prey in Kazakhstan
46
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Summary 1. The Amur or Siberian tiger Panthera tigris altaica forms a relatively small and disjunct population of less than 600 individuals in the Russian Far East. Because tigers in this region require large territories to acquire sufficient prey, current strictly protected areas, comprising 3·4% (10 300 km 2 ) of the region, are unlikely to prevent extirpation of the subspecies in the face of expanding forestry and external demand for tiger parts. 2. We used resource selection function models and spatially explicit population models to analyse the distribution and predict the demographic structure of the population to identify policy options that may enhance population viability. 3. A resource selection function model developed from track distribution data predicted that tigers were most likely to occur in lower altitude valley bottoms with Korean pine forest and low human impacts. 4. The results from the spatially explicit population model suggested that current tiger distribution is highly dependent on de facto refugia with low human impacts but with- out statutory protection, and that small increases in mortality in these areas will result in range fragmentation. Although an expanded reserve network only marginally increases tiger viability under current conditions, it dramatically enhances distribution under potential future scenarios, preventing regional extirpation despite a more hostile landscape matrix. 5. The portion of tiger range most resistant to extirpation connects a large coastal reserve in the central portion of the region with largely unprotected watersheds to the north. A southern block of habitat is also important but more severely threatened with anthropogenic disturbances. The results suggest that preserving source habitat in these two zones and ensuring linkages are retained between blocks of habitat in the north and south will be critical to the survival of the tiger population. 6. Synthesis and applications . Conservation priorities identified in this analysis differ from those suggested by a conservation paradigm focusing only on sustaining and connecting existing protected areas that has been applied to tiger conservation in more developed landscapes with higher prey densities. An alternative paradigm that assesses population viability in a whole-landscape context and develops priorities for both protected area expansion and increasing survival rates in the landscape matrix may be more appropriate in areas where tigers and other large carnivores coexist with low-density human popu- lations. Although landscape connectivity merits increased emphasis in conservation planning, identification of landscape linkages should be tied to broad-scale recommen- dations resulting from spatial viability analyses in order to prevent misdirection of resources towards protecting corridors that add little to population persistence.