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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 single 240 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

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