Rabbits, Foxes and

Mathematical Modeling
Peter Pang
University Scholars Programme and
Department of Mathematics, NUS

SMS Workshop
July 24, 2004
Mathematical Modeling of Population Dynamics

 How does population grow?

 Denote the population by “x”. Suppose the population x(t) at time t

changes to x + Δx in the time interval [t, t + Δt]. Then the growth rate
is
x
x (t )  t
Time Population Growth Rate 0.1
1 10
2 11
3 12.1
4 13.31
5 14.641
1200
6 16.105
7 17.716
1000
8 19.487
9 21.436
10 23.579 800
11 25.937
12 28.531 600
13 31.384
14 34.523 400
15 37.975
16 41.772 200
17 45.95
18 50.545 0
19 55.599 1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49
20 61.159
21 67.275
22 74.002
23 81.403
24 89.543
25 98.497
26 108.35
27 119.18
28 131.1
 The growth rate depends on many things, such as
 Per capita food supply – call it “s”
 A minimum supply of food, say s0, is needed to sustain life
 Say growth rate is proportional to s – s0

x
 a ( s  s0 )
x(t ) t

x
 a ( s  s0 ) x(t )
t

 Call the constant “a” the growth coefficient

Time Population Food Supply 8
1 10 Min Food Supply 5
2 11 Growth Coeff 0.0333
3 12.1
4 13.31
Population
5 14.64
6 16.104
1200
7 17.715
8 19.486 1000
9 21.434
10 23.578 800
11 25.935
12 28.528 600
13 31.381
14 34.519 400
15 37.97
200
16 41.767
17 45.943 0
18 50.537 1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49
19 55.59
20 61.149
21 67.263
22 73.988
23 81.386
24 89.524
 Is infinite growth realistic?
 Suppose population reaches saturation at x0
 Say the growth coefficient is proportional to x0 – x

x
 b( x0  x(t ))( s  s0 ) x(t )
t
 bx0 ( s  s0 ) x(t )  b( s  s0 ) x(t ) 2

 We can interpret the x2 term as a number proportional to the

average number of encounters between x individuals. Hence it
measures a kind of social friction.
Time Population Food Supply 8
1 10 Min Food Supply 5
2 11 Pop Saturation 1000
3 12.099 Coeff 3.37E-05
4 13.306
5 14.632
6 16.089
7 17.688
8 19.443 600
9 21.369 500
10 23.481
11 25.797 400
12 28.335 300
13 31.117
14 34.162 200
15 37.495
100
16 41.14
17 45.125 0
18 49.477 1 4 7 10 13 16 19 22 25 28 31 34 37 40 43 46 49
19 54.227
20 59.408
21 65.052
22 71.195
23 77.875
24 85.129
Foxes and Rabbits
 Let’s look at the fox population
 Let’s assume that the fox population doesn’t get really huge so that
the issue of “population saturation” can be ignored
 Recall that the model for unlimited growth is
x
 a ( s  s0 ) x(t )
t
 Now, suppose that the only food for foxes is rabbits; then s is
proportional to the population of rabbits
 Denote rabbit population by “y”

x
 a (by (t )  s0 ) x(t )
t
 cx(t ) y (t )  dx(t )
 (cy  d ) x
 As for the rabbit population, let’s again assume we have unlimited
growth while the rabbits are being eaten by the foxes -- we further
assume that the number of rabbits eaten is proportional to the fox
population

y
 f y(t )  g x(t ) y(t )
t
 ( f  gx) y
 The Lotka-Volterra (Predator-Prey) Equations

 Population of foxes – x
 Population of rabbits – y

x
 (cy  d ) x
t
y
 ( f  gx) y
t
 where c, d, f, g are constant parameters
Time Fox Rabbit c 5E-06
1 35000 70000 d 0.3
2 36750 80500 f 0.5
3 40517 91166 g 1E-05
4 46831 99812
5 56153 102975 120000
6 68219 96639
7 80716 79033
8 88397 54757 100000
9 86080 33732
10 74774 21561 80000
11 60403 16220
12 47181 14532
13 36455 14942 60000
14 28242 16966
15 22165 20657 40000
16 17805 26407
17 14814 34909
18 12956 47192 20000
19 12126 64674
20 12410 89169 Fox
0
21 14220 122688 Rabbit
22 18676 166586 1 3 5 7 9 11 13 15 17 19
23 28630 218767
24 51357 265518
 Let’s introduce a saturation for the rabbit population:
Time Fox Rabbit c 0.000005
1 50000 60000 d 0.3
2 50000 60000 f' 0.0000125
3 50000 60000 g 0.00001
4 50000 60000 rab-sat 100000
5 50000 60000
6 50000 60000 62000
7 50000 60000
8 50000 60000 60000
9 50000 60000 58000
10 50000 60000
11 50000 60000 56000
12 50000 60000
54000
13 50000 60000
14 50000 60000 52000
15 50000 60000
50000
16 50000 60000
17 50000 60000 48000
18 50000 60000
19 50000 60000
46000
20 50000 60000 44000
21 50000 60000 Fox
1 3 5 7 9 11 13 15 17 19
22 50000 60000 Rabbit
23 50000 60000
24 50000 60000
 Functional Response

 Lotka-Volterra System (with unlimited growth for prey)

x y
 c yx  d x  f y
gy x
t t p( y)

 The term p(y) measures the number of prey susceptible to each

predator as the prey population changes -- this is known as the
functional response
 p(y) = gy says that the number of prey for each predator is a constant
proportion of the prey population
 What if the prey population really shoots up?
 Michaelis-Menten or Holling type II functional response

my
p( y) 
h y
y p(y) m 10
0 0 h 1
0.5 3.333333
1 5
1.5 6
2 6.666667 10
2.5 7.142857
9
3 7.5
3.5 7.777778 8
4 8
7
4.5 8.181818
5 8.333333 6
5.5 8.461538
6 8.571429
5
6.5 8.666667 4
7 8.75
7.5 8.823529 3
8 8.888889 2
8.5 8.947368
9 9 1
9.5 9.047619 0
10 9.090909
10.5 9.130435
11 9.166667
11.5 9.2
 Sigmoidal functional response

my2
p( y ) 
h  jy  y 2
y p(y) m 10
0 0 h 1
0.5 0.666667 j 5
1 1.428571
1.5 2.093023
2 2.666667
2.5 3.164557 9
3 3.6
3.5 3.98374 8
4 4.324324
7
4.5 4.628571
5 4.901961 6
5.5 5.148936
6 5.373134 5
6.5 5.577558
4
7 5.764706
7.5 5.936675 3
8 6.095238
8.5 6.241901 2
9 6.377953
9.5 6.504505 1
10 6.622517 0
10.5 6.732824
11 6.836158
11.5 6.933159
 Holling type III functional response

my 2
p( y ) 
h  y2
y p(y) m 10
0 0 h 1
0.5 2
1 5
1.5 6.923077
2 8
2.5 8.62069 10
3 9
3.5 9.245283
9
4 9.411765 8
4.5 9.529412
5 9.615385 7
5.5 9.68 6
6 9.72973
6.5 9.768786 5
7 9.8 4
7.5 9.825328
8 9.846154 3
8.5 9.863481 2
9 9.878049
9.5 9.890411 1
10 9.90099 0
10.5 9.910112
11 9.918033
11.5 9.924953
 Holling type IV or Monod-Haldane type functional response
my
p( y ) 
h  jy  y 2
y p(y) m 10
0 0 h 1
0.5 4 j 0
1 5
1.5 4.615385
2 4
2.5 3.448276 10
3 3
3.5 2.641509
9
4 2.352941 8
4.5 2.117647
5 1.923077
7
5.5 1.76 6
6 1.621622
6.5 1.50289 5
7 1.4 4
7.5 1.310044
8 1.230769 3
8.5 1.16041 2
9 1.097561
9.5 1.041096 1
10 0.990099 0
10.5 0.94382
11 0.901639
11.5 0.863039
t x y m 100 g 0.0000001 sat 200000
1 50000 60000 h 1b 0.01
2 49525 60756.667 j 0c 0.3
3 49054.2 61521.149
4 48587.58 62293.351
5 48125.11 63073.174 120000
6 47666.75 63860.515
7 47212.47 64655.266 100000
8 46762.25 65457.32
9 46316.06 66266.561 80000
10 45873.87 67082.873
11 45435.65 67906.135 60000
12 45001.36 68736.224
13 44570.99 69573.012 40000
14 44144.5 70416.368
15 43721.86 71266.158 20000
16 43303.05 72122.245
0 x
17 42888.03 72984.487
18 42476.78 73852.74 1 5 y
9 13 17 21 25 29 33 37 41 45 49
19 42069.26 74726.856
20 41665.46 75606.686
21 41265.34 76492.074
22 40868.87 77382.865
23 40476.03 78278.897
24 40086.78 79180.009
 Ratio-Dependent Theory
 Recall that functional response measures the number of prey
susceptible to each predator as the prey population changes.
 We have seen various types of functional response functions of the
type p(y).
 The ratio-dependent theory asserts that functional response should
be dependent of the ratio of prey to predator (especially if the
predator needs to search for the prey), i.e., instead of being just a
function of y, p should be a function of y/x :

 y
p  p 
 x
t x y m 0.1 g 0.0000008 sat 200000
1 35000 70000 h 1b 0.01
2 35350 74946.667 j 1c 0.3
3 35717.11 80042.496
4 36100.84 85254.179
5 36500.68 90544.076 180000
6 36916.09 95871.161
160000
7 37346.52 101192.22
8 37791.42 106463.19 140000
9 38250.23 111640.68 120000
10 38722.41 116683.35 100000
11 39207.4 121553.29
12 39704.69 126217.13 80000
13 40213.74 130646.91 60000
14 40734.08 134820.62 40000
15 41265.21 138722.39
16 41806.69 142342.41
20000
17 42358.09 145676.56 0 x
18 42918.99 148725.86 1 12 23 34 45 56 67 78 89 y100
19 43489.02 151495.76
20 44067.81 153995.38
21 44655.02 156236.68
22 45250.34 158233.74
23 45853.47 160002.04
24 46464.13 161557.86
 Spatial Dependence

 What can happen to a spatially inhomogeneous population?

 Diffusion
 Cross Diffusion
 Partial differential equations
Two Predators and One Prey

 Defense switching
 Cross diffusion