butterflies have been observed ina recent study of dispersal in
ametapopulation of asmall falcon,
Dispersal Within a
Metapopulation of Lesser Kestrels
‘The lesser kestrel, Falco naumanni, isa small migratory fal-
con that breeds in colonies of monogamous pairs in Eurasia
and spends its winters in sub-Saharan Africa (fig. 10.12),
Lesser kestrels have suffered a high rate of decline across
their range and are listed as a globally threatened species. In
contrast to its global circumstance, the lesser kestrel popu-
lation of the Ebro River valley of northeast Spain has grown
dramatically in recent years. David Serrano and Jose Tella,
two ecologists who have conducted long-term studies of this
population (Serrano and Tella 2003), documented growth in
this population from 224 pairs distributed among 4 subpop-
ulations in 1993 to 787 pairs living in 14 subpopulations in
2000. Serrano and Tella attribute this regional growth to sus-
tained traditional farming practices in the Ebro River valley.
However, they warn that plans to modernize furming prac-
tices in the Ebro River valley may lead to the population
declines seen elsewhere.
Serrano and Tella used numbered. color leg bands to
‘mark and keep track of individual kestrels in their study pop-
ulation. From 1993 to 1999, they banded 4,901 fledgling
kestrels and 640 adults. Because lesser kestrels breed within
colonies. they are easier to track during the breeding season.
(Once locating a colony, Serrano and Tella would observe the
colony members from a blind, record the numbers of pairs
within the colony and, using a telescope, read the numbers on
the leg band of any banded adult birds in the colony. Serrano
and Tella were able to obtain accurate counts of the entire
‘breeding population of lesser kestrels within the Ebro River
iM &
Figure 102. Tre ese este Flew nommunn reds in
Seatered clos caletivel orm mctapopultion Mos
porultons fh tetes te declined dal ih
Tatlortznion of tuber
Chapter 10 Population Dynamics 239
valley each year. They could also use their observations to
plot the movements of any banded birds they saw. Within
Colonies, the percentage of banded adults of known age
ranged from 60% 10 90%.
‘The data gathered by Serrano and Tella indicate that a
substantial percentage of birds leave the breeding colony
where they hatched to join other subpopulations in their first
year of breeding. However, females in this species are more
likely to move than males. The rate of emigration by first-
breeding females is approximately 30% versus 22% for first-
breeding males. In contrast, less than 4% of older adults
emigrate from a colony on any given year. Though some
lesser kestrels in the study population dispersed more than
100 km, Serrano and Tella found a negative correlation
between distance between colonies and the frequency of dis-
persal between them,
‘The lesser kestrels of the Ebro River valley and the
Rocky Mountain Parnassian butterflies of southern Alberta,
Canada, interact with their environments on greatly ditferent
scales. In addition, the butterfly population appears to be
contracting spatially; the lesser kestrel population is expand-
ing. However, these populations also have a number of fea-
tures in common. First, they both are spatially organized into
metapopulations. Another feature that the two populations
share is the influence of local population size on the ten-
dency to disperse and the direction of dispersal. Like P.
smintheus. lesser kestrels in smaller subpopulations are
‘more likely to emigrate than are individuals in larger sub-
populations. Second, lesser kestrels are more likely to dis-
perse from small colonies to larger colonies.
In the last rwo concept discussions, we have reviewed the
influences of dispersal on populations. Now we compare
patterns of survival in populations. « major contributor to the
‘dynamic nature of populations.
Concept 10.2 Review
1. Figure 10.11 and the upper portion of Figure 21.13.
show the relationship between meadow size and pop
lation size in wo butterfly species. How are hi
tems shown by the two graphs similar? How do |
ier? (note 1 ha = 10.000 m?)
2. The Rocky Mountain Parnassian butrfly tends (dis
pets from small t large meadows. Why is this ise
‘of movement more advantageous than the reverse?
(Hint: See Glanville fritillary studies. chypier 4)
204)
3. Contrast human i
uences on metapopulations of the
Rocky Mountain Parnassian butterfly versus the les
Patterns of Survival
A survivorship curve summarizes the pattern of survival
population. Patterns of survival vary a great deal from
fone species to another and, depending on environmental
circumstances, can vary substantially even within a single240 Section Il Population Ecology
species. Some species produce young by the millions, w'
in turn dic at ahigh rate, ther species produce a few young.
‘and invest heavily in their care. The young of species that
hhave evolved this pattern survive at a high rate, Still other
species show intermediate patterns of reproductive rate
parental care, and juvenile survival. In response to practical
‘challenges of discerning patterns of survival, population biol-
‘ogists have invented bookkeeping devices called life tables
‘that list both the survivorship and the deaths, or mortality in
populations,
Estimating Patterns of Survival
“There are three main ways of estimating patterns of survival
‘within a population, The first and most reliable way isto iden-
y a large number of individuals that are born at about the
same time and keep records on them from birth to death. A
‘group bor af the same time is called a cohort. A life table
‘made from data collected in this way is called a cohort life
table. The cohort studied might be a group of plant seedlings.
that germinated at the same time or all the lambs born into a
population of mountain sheep in « particular yeat.
‘While understanding and interpreting a cohort life table
‘may be relatively easy. obtaining the data upon which
‘cohort life table is based is not. Imagine yourself lying Face
‘down ina meadow painstakingly counting thousands of tiny
seedlings of un annual plant. You most mark their locations
‘and then come back every week for 6 months until the last
‘member of the population dies. Or, if you are studying a
‘moderately long-lived species, such asa barnacle or a peren-
nial herb like a buttercup, imagine checking the cohort
repeatedly over a period of several years. If your study
‘organism is a mobile animal such as a whale or falcon, the
problems multiply. If your species is very long-lived, such as
a giant sequoia, such an approach is impossible within a sin-
tle human lifetime. In such circumstances population biolo-
gists usually resort to other techniques.
‘A second way (0 estimate patterns of survival in wild
populations is vo record the age at death ofa large number of
individuals. This method differs from the cohort approach
because the individuals in your sample are born at different
nes. This method produces a static life table, The table is
called static because the method involves a snapshot of sur-
-vival within a population during a short interval of time, To
produce a static life rable the biologist often needs to estimate
the age at which individuals die. This ean be done by tagging
individuals when they are born and then recovering the tags
after death and recording the age at death. An alternative
procedure is to somehow estimate the age of dead individu-
als, The age of many species can be determined reasonably
well, For instance, mountain sheep can be aged by counting
the growth rings on their horns. There are also growth rings
‘on the carapaces of turtles, in the trunks of trees, and in the
“stems” of soft or hard corals.
A third way of determining patterns of survival is from
the age distribution. An age distribution consists of the
proportion of individuals of different ages within a popula-
tion. You can use an age distribution to estimate survival by
calculating the difference in proportion of individuals in
sueceeding age classes. This method, which also produces 2
static life table, assumes that the difference in numbers of
individuals in one age class and the next isthe result of mor-
tality, What are some other major assumptions underlying the
use of age distributions to estimate patterns of survival? This,
method requires that a population is neither growing nor
declining and that itis not receiving new members from the
‘ouiside or losing members because they migrate away. Since
‘most of these assumptions are often violated in natural pop-
ulations. 2 life table constructed from this type of data tends
to be less accurate than a cohort life table. Static life Lables
are often useful, however, since they may be the only infor-
mation available.
High Survival Among the Young
‘Adolph Murie (1944) studied patterns of survival among Dall
sheep (fig. 10.13) in what is now Mount Danali National
Park. Alaska, Murie estimated survival patterns by collecting
the skulls of 608 sheep that had died from various causes. He
determined the age at which each sheep inhis sample died by
counting the growth rings on their horns and by studying
{tooth wear. The major assumption ofthis study was that the
proportion of skulls in each age class represented the typical
proportion of individuals dying at that age. For example, the
proportion of sheep in the sample that died before the age of,
| year represents the proportion that generally dies during the
first year of life. While this assumption is not Tikely to be
strictly true, the pattern of survival that emerges probably
gives a reasonable picture of survival in the population. par-
ticularly when the sample is as large as Murie’s
Figure 10.14 summarizes the survival patterns for Dall
sheep based on Murie’s sample of skulls. The upper portion
Of the figure shows the stati life table that Murie con-
structed. The first column lists the ages of the sheep, the see:
fond colurnn lists the number surviving in each age class, and
Figure 10.13 Dall sheep, Ovis dill. & mountain seep of Fae
northern North America, was the subject of one ofthe classic
studies of suvivorship.the third column lists the numbers dying in each age class.
Notice that although Murie studied only 608 skulls, the
‘numbers in the table are expressed as numbers per 1,000
individuals. This adjustment is made to ease comparisons
with other populations.
Nuwberot
begining