Data Analysis Module

ecology

Data Analysis Module
DA  Figure  1  illustrates  stochastic  variation  in  births
and deaths in a population of killer whales (orcas) ( Orci-nus  orca)  that  reside  off  the  coast  of  British  Columbia,
Canada. These data are reproduced in the Excel spread-sheet that accompanies this module as well. You can see
that although the number of reproductive females in the
population  varied  between  16  and  28  individuals,  the
number of young born each year varied between 0 and
8. Annual deaths varied between 0 and 7.
Step 1: Calculate the average rate of population
change and its standard deviation.
Using  the  numbers  of  births  and  deaths  and  the  initial
population  size  of  73  individuals  in  1974,  calculate  the
changes  in  population  size  in  the  Excel  spreadsheet
through 2001.
The average exponential rate of population growth (r)
during year  i is calculated from the number of individuals
at the beginning and at the end of the year as
r
i  ln


N
N
i
i
1


Remember that the exponential growth rate (r) is equal
to the natural logarithm of the geometric growth rate 
(see page 225).
•  Did  the  population  increase  or  decrease  over  this
period? Was the average exponential growth rate positive
or negative?
To  explore  how  stochastic  environmental  variation
affects the size of a population and the probability of its
extinction, we need to develop a model based on random
changes in the factors that determine changes in popula -tion size. Such models often include an upper limit to the
size of the population to incorporate the effect of density
dependence.  However,  in  the  simplest  case,  the  expo-nential rate of increase of a population is independent of
its size (density-independent) and has a mean of  r and a
standard deviation of S. Thus, on average, the population
grows at an exponential rate of  r, but during some peri -ods the growth rate is above this level, and during other
periods it is below this level. If a population experiences
many  periods  of  below-average  growth,  it  runs  the  risk
of  extinction.  The  critical  parameter  that  influences  the
average time to extinction under random environmental
variation is the variance in r.
The variance is the square of the standard deviation,
or S
2
. In the special case in which population size is, on
average, balanced ( r  0; that is, births equal deaths under
Stochastic Extinction with Variable
Population Growth Rates
The random nature of births, the number and sex of off-spring,  and  particularly  deaths  can  lead  to  variation  in
population  size  even  in  a  constant  environment.  Such
stochastic  variation  generally  is  not  a  problem  for  large
populations  because  these  chance  events  average  out
over  many  individuals.  However,  small  populations  can
suffer from random variations in births and deaths, which
can lead to random variation in population size and even
to extinction.
The  kakapo  ( Strigops  habroptilus )  is  a  large,  flightless
parrot  found  only  in  New  Zealand.  Because  it  is  flight-less,  it  is  vulnerable  to  introduced  predators  such  as
cats,  opossums,  and  weasels.  By  1976,  only  14  kakapos
were known to be alive on New Zealand’s South Island;
sadly, all of them were males. If males and females were
hatched  with  equal  frequency  on  average,  what  is  the
probability that 14 individuals would include no females?
If each individual is considered a trial, and being male is
considered a success with probability (p)   0.5, then the
probability that  n trials will all be successes is ( p
n
). If you
think this is unlikely, why might the New Zealand Wildlife
Service have failed to locate any female kakapos? Fortu-nately, additional kakapos were later discovered on Stew-art Island, at the southern tip of South Island, and

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kakapos
were then introduced onto two small islands from which
all  predators  had  been  removed.  (For  more  on  this  fas-cinating  bird,  see  http://en.wikipedia.org/wiki/Kakapo;
http://animaldiversity.ummz.umich.edu/site/accounts/
information/Strigops_habroptila.html.)
When the environment varies, all individuals in a pop -ulation  can  be  affected  in  the  same  way,  and  dramatic
changes, even in large populations, can result. All members
of a population feel a prolonged drought or a cold snap.
Conversely, a period of exceptionally favorable conditions
may  increase  the  fecundity  of  all  individuals  or  increase
their  probability  of  survival.  The  kakapo,  for  example,
breeds primarily in years when the rimu tree, an endemic
conifer of the podocarp group, produces good fruit crops.
Variations  in  environmental  conditions  may  be  random
and essentially unpredictable —what is referred to as sto-chastic  environmental  variation—or  they  may  occur  with
some  regularity.  Understanding  the  connection  between
changes  in  the  environment  and  changes  in  population
size can suggest interventions, such as supplemental feed -ing during critical periods of limited food supply, that can
reduce the chance of population decline and extinction.
Data a nalysis MoD ule
page 2 Data Analysis Module
average conditions), the average time to extinction ( T ) of
a population of size N is
T(N )  
S
2
2
ln(1  S
2
N )  1
Step 2: Estimate the average time until extinction
for a population of killer whales.
Assume that for our killer whale population, the average
growth rate (r) is 0.
•  Starting with the population size in 2001, what is your
estimate of the time to extinction?
•  How does T(N ) change with the size of the initial popu -lation and with the variance in the rate of change in popu -lation size?
•  Fill in the expected times to extinction for the range of
population sizes (N ) and standard deviations of the popu -lation growth rate ( S ) in DA Table 1.
Assuming  that  the  time  units  are  years,  these  values
suggest that small populations in particular have relatively
short life expectancies.
•  What would T(N ) be for the killer whale population at
its largest and smallest sizes? If a population grows just by
chance,  does  this  mean  that  its  prospects  for  long-term
persistence  improve?  Assume  that  the  sample  standard
deviation of r in the spreadsheet accurately estimates the
underlying value of S.
The  implication  of  this  model  for  conservation  is
clearly that populations should be managed to maintain
as large a population size as possible and to prevent strong
depression in the population growth rate during periods of
poor environmental conditions. The latter strategy might
Age and sex structure of population
Births and deaths
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2 000
1974
1975
1976
1977
1978
1979
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2 000
2001
10
15
20
25
30
35
40
45
Deaths
Number
Births
(a)
(b)
KEY
Calves and juveniles
Adult males
Reproductive females Births
Postreproductive females
DA Figure 1  Stochastic variation in
births and deaths of a population of killer
whales. Data from Taylor and Plater, 2001.
Initial population size
S 10 100 1,000 10,000
0.05
0.1
0.2
0.5
Da table 1   Calculating time to extinction
page 3 Data Analysis Module
involve supplemental feeding or predator and pathogen
control programs at critical times.
The model described here lacks density dependence.
Normally, ecologists believe that the growth potential of
populations reduced to low numbers is greatly increased
during  normal  conditions,  which  should  allow  them  to
increase and draw back from the brink of extinction. This
is a fundamental message of the logistic equation and one
of the most basic foundations of ecology.
•  If this were always the case, why should we be worried
about  small  populations?  Under  what  conditions  might
you expect a population not to increase when reduced to
low numbers? This certainly has been the case for many
endangered species that have become extinct or now tee-ter on the brink of extinction. Do some populations simply
not “have what it takes” to maintain healthy numbers?
Let’s  consider  the  effects  of  adding  normal  density
dependence  to  models  incorporating  stochastic  environ-mental variation the intrinsic exponential growth rate, r

.
According to the logistic equation, the average growth rate
of populations reduced below their usual carrying capac-ity always exceeds r   0, and such populations tend to
recover quickly. But a long series of unfavorable periods
might still be enough to drive population size below 1 indi-vidual and cause extinction. The important parameters for
predicting the average time to extinction in models with
density dependence are, first, the product of the average
value of  r

and K, and second, the ratio of the standard
deviation of r

to its mean; in other words,  S/r

. The equa -tions for time to extinction under density dependence are
messy, but as you would expect, the addition of negative
density  dependence  greatly  increases  the  expected  time
to extinction. For example, when N   100,  r

 0.1, and
S    0.22;  T(100)  is  equal  to  nearly  26,000  time  units,
rather than the value of about 81 (see DA Table 1) in the
absence of density dependence.
Literature Cited
Taylor, M., and B. Plater. 2001. Population viability analysis for the
southern  resident  population  of  the  killer  whale  ( Orcinus  orca).
The  Center  for  Biological  Diversity,  Tucson,  Arizona  (http://
www.biologicaldiversity.org/swcbd/species/orca/pva.pdf).

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