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Life tables are used to describe age-specific mortality and survival rates for a
population. When this information is combined with fecundity data, life-tables
can be used to estimate rates of population change (e.g., r, lambda, and Ro).


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1.    or 
 or   life tables: data are collected by following a cohort
throughout its life. This is rarely possible with natural populations of animals. Note: a
cohort is a group of individuals all born during the same time interval.
2. x or 
 life tables: age-distribution data are collected from a cross-section
of the population at one particular time or during a short V
  of time, such as through
mortality data. Resulting age-specific data are treated Va cohort was followed through
time (i.e., the number of animals alive in age class must be less than alive in age class -
1). Because of variation caused by small samples, data-smoothing techniques may be
required (see Caughley 1977).
3.     - data are gathered over a number of years and generations using cohort 
time-specific techniques. This method allows the natural variability in rates of survival to
be monitored and assessed (Begon and Mortimer 1986).

x    
2 x  - Individuals that have only a single, distinct period of reproductive output
in their lives, prior to which they have largely ceased to grow, during which they invest
little or nothing in survival to future reproductive events and after which they therefore
die (Begon et al. 1996:147). For annual species, this results in nonoverlapping
generations. Examples other than annual plants and some insects?
2    - Individuals that normally experience several or many such reproductive
events. During each period of reproductive activity the individual continues to invest in
future survival and possibly growth, and beyond each it therefore has a reasonable chance
of surviving to reproduce again (Begon et al. 1996:147). Results in overlapping
generations.

1. ½  - reproductive activity is restricted to a specific breeding season.

Begon et al. (1996) refer to this as "overlapping semelparity."
2. ½ 
  - reproductive events merge into a single extended period.

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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.
x = x-1x-1 the number of individuals surviving at the V  of age interval .
Note: n0 = sum(dx) if dx expressed as numbers dying and the
survival schedule is complete for all members of the cohort.
x = x - x+1 = the number of individuals of a cohort dying  
the age
interval to +1. Note: sometimes calculated as proportion
dying.
x = x / x =      during the age interval to + 1.
Note: this parameter is least affected by bias in the sample and
gives the most direct projection of the mortality pattern in a
population. It is often used to make comparisons within and
between species.
x = (nx-dx)/nx  1 - x =    V  during the age interval to + 1. This
parameter is used in harvest calculations and in population
modeling.   rates cannot be added to get total survival
rate for a longer period of time; however, finite survival rates are
 (i.e., survival from age 0 to 3 = S0 x S1 x S2).
x = x / 0 = the proportion (scaled from 0 to 1) of individuals surviving at the
V  of age interval . You will also see x expressed per 1,000,
i.e., = (x / 0)1000 . This parameter is used to plot survivorship
curves (see comments below).
 = log10 x - log10 x+1 = 
 or a standardized measure of the intensity (rate) of
mortality. Unlike qx-values, kx-values can be added to determine
the mortality rate for a number of age classes. Note: because kx-
values are calculated using log10, you must take the antilog of
the common log to convert back to finite survival (i.e., S = 10-k).


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 x! !"
# x
x = fecundity rate (i.e., average number of female offspring
produced    in the population over some period of time,
generally a year).
xx = 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 ( ).
R0 = [sum (xx)] / 0 = the        
 . Or in other words, the
mean number of female offspring produced by a female during
 her lifetime (i.e., the replacement rate).   $%&' indicates
the members of the population are  replacing themselves
(i.e., the population is declining). $%(' denotes an increasing
population. $% ' indicates a stationary or "stable" population.
G =Sum(xx x) / R0 = 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    
or the net reproductive rate
(lambda) = R01/G = er =
over some time interval, which is a year in many cases.
c(' indicates an increasing population,  '
indicates a stationary population, and &' indicates a
decreasing population. The use of this parameter is geared
toward organisms that reproduce during a short breeding season
(i.e., discrete growth or p V  ).  your
textbook uses "R" to denote lambda.
r ~ = ln(R0)/G = ln(lambda) =  VorV V   V (i.e., the change in
population size per individual per unit of time) An (%
indicates an increasing population,  % indicates a stationary
pop'n, and an &% indicates a declining population.   the
equations listed only give you an approximation of . You need
to solve  V  for a precise estimate of (see Begon
et al. 1996:165).

x= Tx / x = mean expectation of further life for individuals alive at the start

of age interval . The mean expectation of further life can be
 Lx = (x + x+1) / 2 used as one way of compressing an entire life table into one
number. It has limited application for wildlife studies, but is
Tx = (Sum (Lx)) - Lx-1 commonly used in the insurance business.
= stable age distribution (i.e., where the proportion of the
population in each age class remains constant over time). This is
only achieved if the observed survival and fecundity schedules
remain constant over a long period of time.   population
projections from simple deterministic growth models (e.g., the
exponential and logistic growth models) are based on the
assumption that the population has a stable age distribution.
)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


* 
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+
1.  +$ +  the number of individuals dying in successive
intervals of time is recorded for a group of individuals born at the same time. This is the
most precise type of data available because it is based on a single cohort followed
through time. The observed data are the x column of the life table.
2.   x ,$ +  - The number of individuals alive in successive intervals
of time is recorded for a cohort of individuals. These data are similar to those obtained
with Method 1, except that those V 
are tallied, not those dying. These data are
also precise and specific to the cohort studied. Observed data are entered into the x
column of the life table.
3.  +$ 
x!   - Individuals are marked at birth and their
death recorded, as in Method 1, but several cohorts are pooled from different years or
seasons. These data are usually treated as if the individuals were members of one cohort
and the analysis of Method 1 is applied.
4.  x$ +  The number of individuals aged in a population is
compared with the number of these that died before reaching age +1. The number of
deaths in that age interval, divided by the number alive at the start of the age interval,
gives an estimate of x directly.
5.  +$  x +   - $
 
- Often it is possible to find skulls or other remains that give the age at death of an
individual. These data can be tallied into a frequency distribution of deaths and thus give
x directly.
6.  +   $ 
.   x +   
- $
  - In this case, the age distribution is measured directly by
sampling. The number of individuals born is calculated from fertility rates.

  Methods 5 and 6 are based on the  assumption that the

rate of population change is known (or the population is stationary, i.e.,
r=0) and the age distribution is stable. Although methods 5 and 6
appear more realistic in terms of data collection, there are numerous
ways in which time-specific life tables have been calculated
incorrectly, e.g., from hunter kills (see Caughley 1977).


x ,/.  / 
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Caughley (1977) recommended that the cohort consist of at least 150 individuals when basing
survival estimates on the stable-age assumption. However, age determination is difficult for
many species, and if age is not measured carefully the resulting life table may be very inaccurate.



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The main value of a life table lies in what it tells us about the population's strategy for survival,
i.e., life tables help us to understand the dynamics of populations. For example, time-specific life
tables, although often not meeting the assumptions necessary to estimate survival rates, are
valuable to a manager of exploited populations because they show the existence of strong year
classes or help identify weak age classes.

Although we have used age-structured schedules, for some organisms age is not the best life
history variable on which to develop analyses of population change. For example, the stage of
development (egg, larval, pupal, adult stages) of some insects may be a more important life
history variable. If predation on fish is size-dependent, then size rather than age would be the
appropriate stage-variable.


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Opportunities to follow cohorts for long periods of time are rare, which precludes cohort
analysis, and the critical assumption of a stable-age distribution is so difficult to meet that it
makes the use of time-specific life tables (Johnson 1994).

0
#     
Although life tables are rarely used in fisheries work, information on age or size composition of
the population can be used to estimate survival (see Ricker 1975). Catch curves are probably the
most common method for estimating mortality of exploited fish populations (see Ricker 1975,
Krebs 1989).

0
x! !  *  !
2 x! !  ! are usually created by plotting x on the -axis and age on the -
axis. Occasionally you may see nx plotted on the -axis. The axis is usually logarithmic,
i.e., log10 (x), to allow comparisons among different studies and species. In other words,
log transformations standardize the survivorship curve.
2 *  ! are created by plotting qx or kx against age.
2 Compare your survivorship/mortality curves to those described in Begon and Mortimer
(1996:153-154) and Johnson (1992:433).

0
  
0
. x
0
x$

2 ½ /*
/1
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2
$
 
1996. Ecology: Individuals, populations
and communities. 3rd ed. Blackwell Scientific Ltd., Cambridge, Mass. 1068pp.
2 ½ /*
/*
*  
1986. Population ecology: a unified study of animals and
plants, 2nd edition. Blackwell Sci. Publ., Boston, Mass.
 2 /3
1977. Analysis of vertebrate populations. John Wiley & Sons, New York.
234pp.
2 1  /+
1994. Population analysis. Pages 419- 444  T. Bookhout, ed. Research and
management techniques for wildlife and habitats. Fifth ed. The Wildl. Soc., Bethesda,
Md.
2 -/
1
1989. Ecological methodology. Harper & Row, Publ., New York. 654pp.
2 $ 4/5

1975. Computation and interpretation of biological statistics of fish
populations. Fish Res. Board Can. Bull. 191.