APAKAH LESLIE MATRIX?

Kaedah yang dinamik untuk mewakilkan usia atau struktur saiz populasi. Gabungan proses populasi (kelahiran dan kematian) untuk membentuk model tunggal. Biasanya digunakan pada populasi kitaran pembiakan tahunan.

APA YG PERLU UTK LESLIE MATRIX?

Konsep VEKTOR POPULASI Kelahiran Kematian

POPULATION VECTOR
N0

N1
N2
s+1 baris dengan 1 colum

N3 . Ns

(s+1) x 1

s= umur maksimum

KELAHIRAN/BIRT H
Note: fecundity here is defined as number of female offspring Also, the term newborns may be flexibly defined (e.g., as eggs, newly hatched fry, fry that survive past yolk sac stage, etc.

BARU LAHIR = (Number of age 1 females) times (Fecundity of age 1 females) plus (Number of age 2 females) times (Fecundity of age 2 females) plus ..

N0 = N1F1 + N2F2 +N3F3 .+FsNs

KEMATIAN/MORTALITY
Number at age in next year = (Number at previous age in prior year) times (Survival from previous age to current age)

Na,t = Na-1,t-1Sa
Another way of putting this is, for age 1 for example:
N1,t = N0,t-1S0-1 + N1,t-1 (0) + N2,t-1 (0) + N3,t-1 (0) +

LESLIE MATRIX
N0 N1 N2

F0 S0

F1 F2 F3 . Fs 0 0 0 . 0

N0 N1 N2

0
0 . 0

S1 0
0 0

0 .

0
0

N3 . Ns
(s+1) x 1

S2 0 . 0 0

N3 . Ns
(s+1) x 1

Ss-1 0

(s+1) x (s+1)

LESLIE MATRIX
N0 N1 N2

F0 S0

F1 F2 F3 . Fs 0 0 0 . 0

N0 N1 N2

0
0 . 0

S1 0
0 0
sxs

0 .

0
0

N3 . Ns
sx1

S2 0 . 0 0

N3 . Ns
sx1

Ss-1 0

LESLIE MATRIX
N0 N1 N2

F0 S0

F1 F2 F3 . Fs 0 0 0 . 0

N0 N1 N2

0
0 . 0

S1 0
0 0

0 .

0
0

N3 . Ns

S2 0 . 0 0

N3 . Ns

Ss-1 0

Nt+1 = A Nt

PROJECTION WITH THE LESLIE MATRIX

Nt+1 Nt+2 Nt+3 Nt+4

= ANt = AANt = AAANt = AAAANt

Nt+n = AnNt

PROPERTIES OF THIS MODEL

Age composition initially has an effect on population growth rate, but this disappears over time (ergodicity) Over time, population generally approaches a stable age distribution Population projection generally shows exponential growth

PROPERTIES OF THIS MODEL

GRAPHICAL ILLUSTRATION

25000 20000 15000

Age 0 Age 1

10000 5000 0 0 5 10 Time 15 20 25

PROPERTIES OF THIS MODEL

GRAPHICAL ILLUSTRATION
6 5

Lambda = Nt+1 / Nt Thus, Nt+1 = Nt

Lambda

4 3 2 1 0 0 5 10 Time 15

20

25

PROPERTIES OF THIS MODEL

GRAPHICAL ILLUSTRATION
25

20

Percent in Age 1

15

10

0 0 5 10 Time 15 20 25

PROJECTION WITH THE LESLIE MATRIX

Given that the population dynamics are ergodic, we really dont even need to worry about the initial starting population vector. We can base our analysis on the matrix A itself

Nt+n = AnNt

PROJECTION WITH THE LESLIE MATRIX

Given the matrix A, we can compute its eigenvalues and eigenvectors, which correspond to population growth rate, stable age distribution, and reproductive value

PROJECTION WITH THE LESLIE MATRIX EIGENVALUES

Whats an eigenvalue? Cant really give you a plain English definition (heaven knows Ive searched for one!) Mathematically, these are the roots of the characteristic equation (there are s+1 eigenvalues for the Leslie matrix), which basically means that these give us a single equation for the population growth over time

PROJECTION WITH THE LESLIE MATRIX CHARACTERISTIC EQUATION

1= F1-1 + P1F2 -2 + P1P2F3 -3 + P1P2P3F4 -4 Note that this is a polynomial, and thus can be solved to get several roots of the equation (some of which may be imaginary, that is have -1 as part of their solution)

The root () that has the largest absolute value is the dominant eigenvalue and will determine population growth in the long run. The other eigenvalues will determine transient dynamics of the population.

PROJECTION WITH THE LESLIE MATRIX EIGENVECTORS

Associated with the dominant eigenvalue is two sets of eigenvectors The right eigenvectors comprise the stable age distribution The left eigenvectors comprise the reproductive value (We wont worry how to compute this stuff in class computing the eigenvalues and eigenvectors can be a bugger!)

PROJECTING VS. FORECASTING OR PREDICTION

So far, Ive used the term projecting what does this mean in technical terms, and how does it differ from a forecast or prediction. Basically, forecasting or prediction focuses on short-term dynamics of the population, and thus on the transient dynamics. Projection refers to determining the long-term dynamics if things remained constant. Thus projection gives us a basis for comparing different matrices without worrying about transient dynamics.

PROJECTING VS. FORECASTING OR PREDICTION

Simple (?) Analogy: The speedometer of a car gives you an instantaneous measure of a cars velocity. You can use to compare the velocity of two cars and indicate which one is going faster, at the moment. To predict where a car will be in one hour, we need more information, such as initial conditions: Where am I starting from? What is the road ahead like? etc. Thus, projections provide a basis for comparison, whereas forecasts are focusing on providing accurate predictions of the systems dynamics.

STAGE-STRUCTURED MODELS LEFKOVITCH MATRIX

Instead of using an age-structured approach, it may be more appropriate to use a stage or size-structured approach. Some organisms (e.g., many insects or plants) go through stages that are discrete. In other organisms, such as fish or trees, the size of the individual is more important than its age.

LEFKOVITCH MATRIX EXAMPLE

N0 N1 N2

F0

F1 F2 F3 . Fs
.. Ts-1 .. Ts-2 .. Ts-3

N0 N1 N2

T0-1 T1-1 T2-1 T3-1

T0-2 T1-2 T2-2 T3-2

T0-3 T1-3 T2-3 T3-3 . T0-s T1-s T2-s T3-s

N3 . Ns

N3 . Ns

.. Ts-s

LEFKOVITCH MATRIX
Note now that each of the matrix elements do not correspond simply to survival and fecundity, but rather to transition rates (probabilities) between stages. These transition rates depend in part on survival rate, but also on growth rates. Note also that there is the possibility for an organism to regress in stages (i.e., go to an earlier stage), whereas in the Leslie matrix, everyone gets older if they survive, and they only advance one age

LEFKOVITCH MATRIX EXAMPLE

N0 N1 N2

F0

F1 F2 F3 . Fs
.. Ts-1 .. Ts-2 .. Ts-3

N0 N1 N2

T0-1 T1-1 T2-1 T3-1

T0-2 T1-2 T2-2 T3-2

T0-3 T1-3 T2-3 T3-3 . T0-s T1-s T2-s T3-s

N3 . Ns

N3 . Ns

.. Ts-s

EXAMPLE APPLICATION: SUSTAINABLE FISHING MORTALITY

An important question in fisheries management is How much fishing pressure or mortality can a population support?

EXAMPLE APPLICATION: SUSTAINABLE FISHING MORTALITY

N0 N1 N2

F0 S0

F1 F2 F3 . Fs 0 0 0 . 0

N0 N1 N2

0
0 . 0

S1 0
0 0

0 .

0
0

N3 . Ns

S2 0 . 0 0

N3 . Ns

Ss-1 0

EXAMPLE APPLICATION: SUSTAINABLE FISHING MORTALITY

S=

-(M+F) e

Knife-edge recruitment, meaning that fish at a given age are either not exposed to fishing mortality or are fully vulnerable

EXAMPLE APPLICATION: SUSTAINABLE FISHING MORTALITY

1.4

Rate of increase (lambda)

1.2 1.0 0.8 0.6 0.4 0.2 0.0 0.5

Age at entry
5 4 Current 3 2 Maintenance level

1.0 1.5 2.0 Instantaneous fishing mortality

2.5

EXAMPLE APPLICATION: SUSTAINABLE FISHING MORTALITY

750 500 250 0 500 250 0 500 250 0

Year 3

Year 4

Year 5

Frequency

500 250 0 500 250 0 500 250 0 500 250 0 500 250 0

Year 6

Year 7

Year 8

Year 9

Year 10

5,000

10,000

15,000

20,000

25,000

Population size

EXAMPLE APPLICATION: SUSTAINABLE FISHING MORTALITY

6,000 4,000 2,000 0 4,000 2,000 0 1,200 800 400 0 800 400 0 800 400 0 800 400 0 800 400 0 800 400 0 800 400 0

Year 1-2 Year 2-3 Year 3-4 Year 4-5 Year 5-6 Year 6-7 Year 7-8 Year 8-9

Frequency

Year 9-10 0.8 1.0 1.2 1.4 1.6 1.8 2

0.6

Population growth rate

EXAMPLE APPLICATION: SUSTAINABLE FISHING MORTALITY

1,200 800 400 1,200 800 400 1,200 800 400

Year 10-20

Year 10-30

Year 10-40

Frequency

1,200 800 400 1,200 800 400 1,200 800 400 1,200 800 400 0

Year 10-50

Year 10-60

Year 10-70

Year 10-150

1.0

1.1

1.2

1.3

1.4

Population growth rate

EXAMPLE APPLICATION: SUSTAINABLE FISHING MORTALITY

1.4

Rate of increase (lambda)

1.2 1.0 0.8 0.6 0.4 0.2 0.0 0.5

Age at entry
5 4 Current 3 2 Maintenance level

1.0 1.5 2.0 Instantaneous fishing mortality

2.5

EXAMPLE APPLICATION: REPRODUCTIVE STRATEGY OF YELLOW PERCH

One of main questions was whether stunting, meaning very slow growth, of yellow perch was caused by reproductive strategy or if reproductive strategy resulted from adaptation to low prey abundance Our goal was to understand what stunted fish should do in terms of age at maturity

REPRODUCTIVE STRATEGY OF YELLOW PERCH

Basic model had a number of assumptions

Energy intake is limited and depends on size of fish

Yellow perch show an ontogenetic shift in diet where indivduals less than 10 grams eat zooplankton, individuals 10 to 30 grams eat benthic invertebrates, and individuals larger than 30 grams eat fish

Net energy intake can only be partitioned to growth or reproduction Reproduction is all or nothing Reproduction may have survival costs (theta)

REPRODUCTIVE STRATEGY OF YELLOW PERCH

REPRODUCTIVE STRATEGY OF YELLOW PERCH

REPRODUCTIVE STRATEGY OF YELLOW PERCH

REPRODUCTIVE STRATEGY OF YELLOW PERCH

REPRODUCTIVE STRATEGY OF YELLOW PERCH

REPRODUCTIVE STRATEGY OF YELLOW PERCH

REPRODUCTIVE STRATEGY OF YELLOW PERCH

REPRODUCTIVE STRATEGY OF YELLOW PERCH

REPRODUCTIVE STRATEGY OF YELLOW PERCH