Ekologi Populasi dan Komuniti BDV3501

Lab : Life Table

INTRODUCTION Within a population of a species, not all individuals are the same. We have already spoken of differences due to sex and genetic makeup. Today, we will examine the differences due to age. The age of an organism can strongly affect both the rate of mortality and natality. Just think of humans. It is very obvious that not all humans have the ability to reproduce as some are too young and some too old. It may be less obvious, but no less true, that mortality is also dependent on age, with the very young and old more likely to die. When considering a population of mixed ages, an ecologist needs to be able to predict the composition of the population in the future based on the population of today. To do this, we must have a means of taking the proportions of the population in various age classes into account. We can do this with a life table: a schedule of the probabilities of death and reproduction that covers the potential life span of an individual organism. The rates (= probabilities) of dying and reproduction in a life table are referred to as age-specific mortality and fecundity. How do we get such a schedule? There are two general ways. One can take a snap-shot of the population at a single time and look at the proportions occurring in each age class. However, this method assumes that the population has reached an equilibrium value, called the Stable Age Distribution. A better way, but more difficult, is to follow a group of individuals born at the same time (called a cohort) until all are dead and record when each female reproduces (and how many offspring are produced). From the life table, we can calculate other ecological parameters such as expected life span (adjusted for age), the eventual stable age distribution for the population if the life table does not change with time, and the growth rate of the population. The techniques used were developed for animal populations and are most highly developed for human populations. For that reason, the science of life tables is called demography (demos is the Greek root for people). Now the term is used for life tables for any organisms, and you often see the phrase "human demography" which is a bit redundant. Recently, plant ecologists have made good use of demographic techniques, which makes good sense if you consider that it should be easier to follow a cohort of plants due to their sessile (fixed in one place) life style.

Ekologi Populasi dan Komuniti BDV3501 LAB OBJECTIVES Through participating in the exercise, students will able to learn: 1. the methods used to establish life table 2. to calculate and build the life table of given population EQUIPMENTS Working indiviually:

lab instruction calculator

INSTRUCTIONS: Life Table

I Life-Table Notation (columns in a life table):

age, measured in years or some other conventional unit. With longer-lived animals and plants this is often 1 year, but for voles it might be 1 week and for some insects 1 day. Often expressed as an interval, e.g., 0-1 years old. the number of individuals surviving at the start of age interval x. Note: n0 = sum(dx) if dx expressed as numbers dying and the survival schedule is complete for all members of the cohort. the proportion (scaled from 0 to 1) of individuals surviving at the start of age interval x. You will also see lx expressed per 1,000, i.e., = (nx / n0)1000 . This parameter is used to plot survivorship curves (see comments below). the number of individuals of a cohort dying during the age interval x to x+1. Note: sometimes calculated as proportion dying. finite rate of mortality during the age interval x to x + 1. Note: this parameter is least affected by bias in the sample and gives the most direct projection of the

x=

nx = nx-1 - dx-1

lx = n x / n 0 =

dx = lx - lx+1 =

qx = dx / lx =

Ekologi Populasi dan Komuniti BDV3501 mortality pattern in a population. It is often used to make comparisons within and between species.

Lx=(lx+lx+1)/2 Tx = Lx + Lx+1

the average number alive in interval x Time units left for all individuals to live from age x onward mean expectation of further life for individuals alive at the start of age interval x. The mean expectation of further life can be used as one way of compressing an entire life table into one number. It has limited application for wildlife studies, but is commonly used in the insurance business.

ex = Tx / lx

II. Extended or Expanded Life Tables = Survival + Fecundity Schedules

fecundity rate (i.e., average number of female offspring produced per female in the population over some period of time, generally a year). The mean number of female offspring produced by females in an age class. This information is used to calculate net reproductive rate (R0) and the instantaneous rate of change ( r ). the net reproductive rate per generation. Or in other words, the mean number of female offspring produced by a female during her lifetime (i.e., the replacement rate). Note: R0 < 1 indicates the members of the population are not replacing themselves (i.e., the population is declining). R0 > 1 denotes an increasing population. R0 = 1 indicates a stationary or "stable" population. the mean length of a generation. Other definitions include (1) the time elapsing between birth of a female and mean time (age) of birth of her offspring and (2) the average age that adult females give birth. the finite rate of population change or the net reproductive rate over some time interval, which is a year in many cases. Lambda >1 indicates an increasing population, lambda=1 indicates a stationary population, and lambda<1 indicates a

mx =

lx mx =

R0 = (lx mx) =

G = (lx mx x) / R0 =

(lambda) = R01/G = er =

Ekologi Populasi dan Komuniti BDV3501 decreasing population. The use of this parameter is geared toward organisms that reproduce during a short breeding season (i.e., discrete growth or birthpulse fertility). Note: your textbook uses "R" to denote lambda.

intrinsic or instantaneous rate of increase (i.e., the

r ~ = ln(R0)/G = ln(lambda) =

change in population size per individual per unit of time). An r>0 indicates an increasing population, r=0 indicates a stationary population, and an r<0 indicates a declining population. Note: the equations listed only give you an approximation of r. You need to solve Euler's equation for a precise estimate of r (see Begon et al. 1996:165).

NOTE: Population ecologists do not all use the same life-table notation. For example, Begon et al. (1996) use different symbols to denote the finite rate of increase and mean generation length. Nevertheless, data and calculations in the respective columns have the same meaning

Many life tables have no age specific birth rate columns and are used only for calculating age specific mortalities and life expectancy. This can be very useful, as we can learn what is the most risky time of life for an organism. Life tables are at the base of the life-insurance business. If the table is to be used to calculate the growth rate, then only females are included in the table, as males do not give birth. Problem Set #1: Life Tables

1. The following data were collected from Gray Squirrel X(year) nx 0-1 530 1-2 134 2-3 56 3-4 39 4-5 23 5-6 12 6-7 5 7-8 2

2. Construct a separate life table for each cohort. Use the headings below: age class - x - nx - dx - qx - lx - Lx - Tx - ex 3. Adding a Fecundity Schedule

Ekologi Populasi dan Komuniti BDV3501 Let use the table above as basis for the construction of the fecundity table. The fecundity table uses the survivorship column, l x from the life table and mx is given below: X(year) mx 0-1 0.0 1-2 1.28 2-3 2.28 3-4 3.42 4-5 3.42 5-6 2.48 6-7 2.28 7-8 2.28

Use the headings : mx-lxmx-xlxmx 4. Calculating the population growth rates a. Calculate the population growth rate R0 and r b. Obtain r exactly: you must proceed by trial and error using Eulers equation. That is, you must make the quantity e-rxlxmx, where r is the unknown, equal exactly 1. c. Is the population increasing, decreasing or stationary? How can you tell? What is the intrinsic rate of increase (r), net reproductive rate per generation (R0), and the finite rate of increase () tell us? What do these population growth parameter sreally mean? d. What is the limitation of life table? SUBMISSION: Discussion and exercise on 21 Nov, Calculator required. Submit by 28 Nov, individually