7SCCCTECSIIGISSESSESESCLEE SZ 4 I FAZ4444d44 4494 Unit 4 Analysis of Populations Introduction ‘The population is a basic organizational unit in biology, It is an assemblage of individuals of the same species inhabiting a given area, Unfortunately, a few writers have lapsed carelessly into using the term population to mean aan aggregation of several species in an area, a situation where community or subcommunity would be the appro- priate designation (see U A population has unique structural and functional characteristics not found in the individuals it comprises, In his unit we shall investigate some of these characteris- tics. One, population structure, will be discussed in terms of age distribution (Section 4A) and spatial distri bution (Section 4C). Another, population dynamics, Will be treated in terms of population growth (Section 4B) and survivorship (Section 4), 1374a Age Structure and Survivorship 1. Introduction Populations, whether animal or plant, vary in their pro- portions of young and old individuals. Time units such as ‘weeks, months, or years can describe ages. Or individuals can be assigned to qualitative age classes, such as nest- Jing, juvenile, subadult, and adult, or egg. larva, pupa, and adult, The proportions of individuals belonging to the various age groups are collectively referred to as the age structure or age distribution of the population. ‘Three different procedures may be used for obtain- ing the age structure of a population, The vertical ap- proach follows a particular cohort. A cohort is a group of individuals born within the same time interval. Thus, by knowing the age of cohort members, you ean follow their survival until all have died. The horizontal approach uses data on all ages within a given population at one time; that is, all cohorts in the population are examined at the same time. In the latter method, one assumes a stable age structure and constant birth and death rates. A third ap- proach involves knowing the age at death for members of ‘a population, Such data are commonly obtained for game species. ‘Knowledge of age structure is important, for the age distribution of a population affects its growth and dy~ namics. From a knowledge of age structure, a table of age-specific mortality, survivorship, and life expectancy can be constructed —a life table (Section 4.4). In addi- tion, population growth rates may be estimated from data on births per female in the population. 2, Procedures ‘Age-class data may be collected from a random sampling of a field of laboratory population or from a preserved (but randomly collected) sampling of a population. Methods for aging vertebrates include counting growth 138 rings on horns of mammals or on scales of fishes, exam- ining tooth development and wear, and observing pelage cor plumage changes. Weight or length classes may be tabulated for many species of fishes and invertebrates. Growth rings are also used for aging woody plants as well as certain invertebrates, such as clams and snails. Various environmental variables often affect the measured oF ob- served characters. Therefore, such characters as tooth ‘wear, molt, weight, and length should be standardized to known ages for a given location and time. Insects are often aged by their developmental stage or by molts ‘within stages of metamorphosis. Details on methods of determining age are given in Caughley (1977), Glock (1955), Lagler (1956), Larson and Taylor (1980), Mo- ‘mot (1967), Smith (1980), and Southwood (1978). Record the number of individuals of each age. Gen- erally, the numbers are recorded for age classes, or inter vals, rather than exact ages (e-g., those individuals be- tween three and four years old, those between four and five years old, and so on), The investigator should also record measurements (¢.g., length and/or weight of the body or of some part of the body) other than the charac~ ters used to indicate age of the organism, as these often provide additional information on population growth and vigor. 3. Age Pyramids ‘The number of individuals in each age class may'be plot- ted as a horizontal histogram, forming a “pyramid.” Age is placed on the vertical axis and the number, or the proportion, of individuals in each age class is plotted so that a symmetrical, pyramidal graph results. Often the age-class bars are graphed so that data for males appear fon one side of the pyramid and females on the other (Figure 44,1). Age pyramids such as these usefully compare populations from different sites, or the same population at different times of the year or from year to year. ‘Age structure is dependent on many factors, such as longevity, rate of population increase, mortality, and en~ vironmental influences. In general, a growing population will show an increased proportion of young individuals, 4 stable one will show no increase or decrease in the relative numbers in each age class, and a declining one will show an increase in the proportions of the popula tion in the older age classes and a decrease in member~ ship in the younger age classes. ‘The size of individuals is usually correlated with the age structure of the population. For added information, plot the length, weight, or other size measurement against size class. In an overcrowded population, one may find stunting of individuals (Section 4D.2.2) as well as re~ duced reproduction and a high proportion of older indi~ viduals. 0998) 9979999900009 0000 222727799999 » - &= . “9 So re oe ~3 a ~~ ° ° ° ° . ° ° » ° °e °o ° ° ° ° ° ° ° ° hd e e ee ° 8-9 7-8 6-7 5-6 45 3-4 2-3 bee 0-1 Percent of Pepulaton Figure 44.1. An age pyramid ‘Age Structure and Survivorship 139 Table 4A.1 A Life Table. The data in the x and L, columns were obtained from a population of animals, Then, all other columns of data were derived from them, as described in Section 44.4 Cohort) Number) Number Number, Probability , Probability» Animal-Years (age in | Living | Dying | of Dying” | of Surviving Left to intervay | Cohort, | at Start, during, "uring, | interval, Live, | Expectancy, Agee) | oe L ‘ 4, « be Zs «09 ot 0 3 46 ost 043 6 137 12 \ 16 040 0.60 - 150 23 2 9 0.50 050 17 34 3 4 ost 033 083 4s 4 1 1.00 0.00 050 5-6 5 ° 4, Life Tables In a life table (Table 4.1), various statistics are com- piled for each age class, or cohort (designated x). Data are commonly collected as numbers of individuals in ceach age class. Lis the number of individuals in age class «x Itis assumed that L, is the number alive at the middle of age class x (for example, in Table 4.1, 33 individuals are assumed to be 0.5 year old, even though the true ages of the 33 might range between 0 and 1 year old). ‘We designate /, as the number of individuals alive at the beginning of age class x. Thus, L, may be defined as (e+ bee/2 a (i.e., Li the number alive at the midpoint of age class »), and @