Chapter 6 HUMAN ECOLOGY AND CUCCESSION Objectives: At the end of the chapter, students should be able to: Discuss the scope of human and population ecology. Define the properties of population. Name and discuss the factors influencing population growth. Suggest ways to lessen the impact of rapid population growth. Anglyze an environmental issue and offer solutions im Pie we Human and Population Ecology Defined: Human ecology is interdisciplinary. Neither ordinary biological ecology nor human ecology is a discipline in the usual sense. Both have to draw extensively on other disciplines relevant to understanding the behavior of organisms (in our case, humans). The multidisciplinary commitments of ecology and human ecology arise because the behavior of organisms is so complex. It will often be important to understand how individuals tick, requiring knowledge of human psychology and biology. Human groups are elaborately organized, so the social context is often important. Anthropologists, sociologists, and political scientists specialize in such things. Humans interact intimately with other organisms, so disciplines from agronomy to zoology will sometimes be useful. Some disciplines specialize in theory development, such as economics and evolutionary biology. Others specialize{n certain important empirical tools, such as participant observation in anthropology, laboratory experimentation in psychology, and survey research in sociology and political science, Still other specialized disciplines, such as history, archaeology, and paleontology, are devoted to the long view. Since humans interact with environments, the physical/chemical environmental disciplines such as climatology, hydrology, and soil science will also be important to answer certain kinds of questions. If we want to understand a particular facet cof human behavior, we have to be prepared to go shopping for ideas and methods in a variety of disciplines. Population ecology studies organisms from the point of view of the size and structure of their populations. A population ecologist studies the interaction of organisms with their environments by measuring properties of populations rather than the behavior of individual organisms. Properties of Population Properties of populations include: Population Size Population Density Patterns of Dispersion Demographics Population Growth, Limits of Population Growth All ofthese properties are not those of individual organisms but instead are properties hich exist only fone considers more than one organism at any given time, or over a period of time. “The characteristics of a population are shaped by the interactions between individuals and their environments on both ecological and evolutionary time scales, and natural selection can modify these characteristics in a population” ‘Thus, population ecology also goes beyond consideration of just population parameters and additionally considers how the characteristics of individual organisms impact on population parameters Population. A population in an ecological sense is a group of organisms, of the same species, which roughly occupy the same geographical area at the same time. Individual members of the same population can either interact directly, or may interact with the dispersing progeny of other members of the same population. Population members interact with a similar environment and experience similar environmental limitations ‘A population is a group of individuals of the same species living in the same geograp! area The study of actors thataffect growth, stably, and decline of population is population iynamics. Population size. A populations size depends on how the population is defined. If a population is defined in terms of some degree of reproductive isolation, then that populations size is the size ofits gene pool. Ifa population is defined in terms of some geographical range,then that population’ size is the number of individuals living in the defined area. Ecologists typically are more concerned with the latter means of defining a population since this is both easier to do and is @ more practical measure if one is interested in determining the impact of a given population on a given ecosystem, or vice versa “Although we can determine an average population size for many species, the average is often of less interest than the year to-year or place-to-place trend in numbers. Population density. Given that a population is defined in terms of some natural or arbitrarily defined geographical range, then population density may be defined as simply the number of individual organisms per unit area. Different species, of course, exist at different densities in their environments, and the same species may be able to achieve one density in one environment and another in a different environment. Population densities may additionally be determined in terms of some measure other than population size per unit area such as population mass per unit area Patterns of dispersion. Individual members of populations may be distributed over a ical area in a number of different ways including Clumped distribution (attraction); Uniform distribution (repulsion); and Random distribution (minimal interaction/influence) Clumping may result either from individual organisms being attracted to each other, or individual organisms being attracted more to some patches within a range than they are to other patches; the net effect is that some parts of the range will have a large number of individuals whereas others will contain few or none. Patterns of Dispersion Within a Populations Geographical Range 'A uniform distribution means that approximately the same distance may be round between individual organisms; uniform distributions result from individual organisms actively repelling each other ietributs c individual organisms are found is onl A random distribution means that where individual org " ini fundom ditriteractions with other members ofthe same population, and random distributions are uncommon; “Random spacing occurs in the absence of strong, attractions or repulsions among individuals of a population___ Both clumping and uniform distributions suggest that individual organisms are either interacting with one another (actively seeking each other out or actively avoiding each other), or are all competing with one another for the same limited resources, regardless of the overall population density. Demographics. A population's demographics are its vital statistics, particularly those statistics that can impact on present and future population size. Two statistics that are of particular import are a population’ age structure and a population's gender ratio. Population growth. The simplest case of population growth is that which occurs when there exist no limitations on growth within the environment. All populations undergo three distinct phases of their life cycle: 1. growth 2. stability 3. decline Population growth occurs when available resources exceed the number of individuals able to exploit them, Reproduction is rapid, and death rates are low, producing a net increase in the population size. Population stability is often proceeded by a “crash” since the growing population eventually outstrips its available resources. Stability is usually the longest phase of a population’ life cycle. Decline is the decrease in the number of individuals in a population, and eventually leads to population extinction, Factors Influencing Population Growth Nearly all populations will tend to grow exponentially as long as there are resources available. Most populations have the potential to expand at an exponential rate, since reproduction is generally a multiplicative process, Two of the most basic factors that affect the rate of population growth are the birth rate, and the death rate. The intrinsic rate of increase isthe birth rate minus the death rate. | @)Exponential (un- (b)Logistic (restricted) restricted) growth growth Carrying capacity ‘The growth rate of the population Secolerates Population size Population sie o Time o TimeTwo Modes of Population Growth. ‘The Exponential curve (also known as a J-curve) occurs when there is no limit to population size. The Logistic curve (also known as an S-curve) shows the effect of a limiting factor (in this case the carrying capacity of the environment). Population Growth Potential Is Related to Life History The age within its individual life cycle at which an organism reproduces affects the rate of population increase. Life history refers to the age of sexual maturity, age of death, and other events in that individual’ lifetime that influence reproductive traits. Some organisms grow fast, reproduce quickly, and have abundant offspring each reproductive cycle. Other ‘organisms grow slowly, reproduce at a late age, and have few offspring per cycle. Most organisms are intermediate to these two extremes. Several Basic Controls Govern Population Size The environment is the ultimate cause of population stabilization. Two categories of factors are commonly used: physical environment and biological environment. Three subdivisions of the biological environment are competition, predation, and symbiosis, Physical environment factors include food, shelter, water supply, space availability, and (for plants) soil and light. One of these factors may severely limit population size, even if the others are not as constrained, The Law of the Minimum states that population growth is limited by the resource in the shortest supply. ‘The biological role played by species in the environment is called a niche. Organisms/ populations in competition have aniche overlap ofa scarce resource for which they compete. Competitive exclusion occurs between two species when competition isso intense that one species completely eliminates the second species from an area In nature this is rather rare. ‘While owls and foxes may compete for a common food source, there are alternate sources of food available. Niche overlap is said to be minimal. Population Decline and Extinction Fatinction is the elimination of all individuals ina group. Local extinction is the loss ofall individuals in a population. Species extinction occurs when all members of a species and its component populations go extinct. Scientists estimate that 99% of all species that ever existed are now extinct, The ultimate cause of decline and extinction is environmental change. Changes in one of the physical factors: ‘of the environment may cause the decline and entire tion: likewise the fossil record indicates that some extinctions are caused by migration of a competitor. ‘Dramatic declines in human population happen periodically in response toan infectious disease.Human Impact ‘Human populations have continued to incre ogy that disrupted natural populations. Destabilization of populations leads to possible out , due to use of technology that has ‘+ population growth as previous limits are removed + population decline as new limits are imposed Agriculture and animal domestication are examples of population increase of favored organisms, Pollution Pollutants generally are releases of substances into the air and water. Many lakes often hhave nitrogen and phosphorous as limiting nutrients for aquatic and terrestrial plants. Runoff from agricultural fertilizers increases these nutrients, leading to runaway plant ‘growth, or eutrophication. Increased plant populations eventually lead to increased bacterial populations that reduce oxygen levels in the water, causing fish and other organisms to suffocate. Pesticides and Competition Removal of a competing species can cause the ecological release of a population ‘explosion in that species competitor. Pesticides sprayed on wheat fields often result in a secondary pest outbreak as more-tolerant-to-pesticide species expand once less tolerant competitors are removed. Removal of Predators Predator release is common where humans hunt, trap, or otherwise reduce predator populations, allowing the prey population to increase. Elimination of wolves and panthers have led to increase in their natural prey: deer. Large deer populations often cause over ‘grazing that in turn leads to starvation of the deer. Introduction of New Species Introduction of exotic or alien non-native species into new areas is perhaps the greatest single factor to affect natural populations, More than 1500 exoti than 25 families of alien fish have been introduced; in excess of 3000 plant species have also been introduced. The majority of accidental introductions may fail, however, once an Introduced species becomes established, its population growth is explosive, ic insect species and more Altering Population Growth Humans can remove or alter the constraints on population sizes, with both good and bad consequences, On the negative side, about 17% of the 1500 introduced insect species require the use of pesticides to control them,For example, bees are ex another. These killer bees a honeybee populations. Ona positivenote, human-induced popu for growing human po and sustaining increase ae their population and migrating from one place to ich more aggressive than the natives, and destroy native lation explosions can provide needed resources ulations. Agricul : ; iculture now produces more food per acre, allowin «human population size : : Human action is causin, : fi ; nds of times the natural rate. Eatinction s caused by fst cxntion of species at thou ‘or draining wetlands. Habitat disruption in currently the leading cause of extinction. ‘Changes in the biological environment occur in three ways. 1 Specie troduction: An xt secession no an rs wie i ay ae its population size, or where it can greatly out compete native 2 Overhunting: When a predator population increases or becomes more efficient at Killing the prey, the prey population may decline or go extinct. Examples today include big game hunting, which has in many places reduced the predator (or in this case prey) ‘population. In human prehistory we may have caused the extinction of the mammoths ‘and mastodons due to increased human hunting skill 3. Secondary extinction: Loss of food species can cause migration or extinction of any. species that depends largely or solely on that species as a food source. ‘Overkill is the shooting, trapping, or hunting of a species usually for sport or economic reasons. Unfortunately, this cannot eliminate “pest” species like cockroaches and mice due to their large population sizes and capacity to reproduce more rapidly than we can eliminate them, However, many large animals have been eliminated or had their populations drastically reduced (such as tigers, elephants). “The death of one species or population can cause the decline or elimination of others, a process known as secondary extinction. Destruction of river ecosystem, the place for many fishes, may cause the extinction of the native fish The extinction of the insect has caused ‘some flowering plants to become unable to reproduce new plant. Ecological Succession ‘A directional change in an ecological community, Populations of animals and plants “Through the continual turnover of individuals, a population may ing onthe success of ts members in survival and reproduction. As ‘omposition of communities typically oes not remain static with fluctuations in species abundance related to seasonal changes, ly with time through a recognizable sequence known community may * popalations are replaced by succesive colonists along a more or less predic ma eet a relatively stable community. This process of succession results Pred tat ena between diferent species and between species and their environment which govern the sequence and the rate with which species replace each other. The rate at are in a dynamic state. expand or decline depend a consequence, the species ¢ time, Apart from the regular a community may develop progressivelwhich succession proceeds depends on the time scale of species life histories as well as on the effects species may have on each other and on the environment which supports them. In some cases, seres may take hundreds of years to complete, and direct observation at a given site is not possible. Adjacent sites may be identified as successively older stages of the same cere ft is assumed that conditions were similar when each seral stage was initiated. The course of ecological succession depends on initial environmental conditions Primary succession occurs on novel areas such as volcanic ash, glacial deposits, or bare rock, areas which have not previously supported a community. In such harsh, unstable tnvironments, pioneer colonizing organisms must have wide ranges of ecological tolerance to survive. In contrast, secondary succession is initiated by disturbance such as fre, which removes a previous community from an area, Pioneer species are here constrained not by the physical environment but by their ability to enter and exploit the vacant area rapidly. ‘As succession proceeds, many environmental factors may change through the influence ‘of the community. Especially in primary succession, this leads to more stable, less severe environments, At the same time interactions between species of plant tend to intensify ‘competition for basic resources such as water, light, space, and nutrients. Successional change tesults from the normal complex interactions between organism and environment which lead to changes in overall species composition. Whether succession is promoted by changing ‘environmental factors or competitive interactions, species composition alters in response to svailability of niches. Populations occurring in the community at a point in succession are those able to provide propagules (such as seeds) to invade the area, being sufficiently tolerant of current environmental conditions, and able to withstand competition from members of cther populations present at the same stage. Species lacking these qualities either become locally extinct or are unable to enter and survive in the community. Early stages of succession tend to be relatively rapid, whereas the rates of species turnover and soil changes become slower as the community matures. Eventually an approximation to the steady state is established with a relatively stable community, the nature of which has aroused considerable debate. Earlier, the so-called climax vegetation wvas believed to be determined ultimately by regional climate and, given sufficient time, any community in a region would attain this universal condition, This unified concept of ‘succession, the monoclimax hypothesis, implies the ability of organisms progressively to ‘modify their environment until it can support the climatic climax community. Although plants and animals do sometimes ameliorate environmental conditions, evidence suggests ‘overwhelmingly that succession has a variety of stable end points, This hypothesis, known as the polyclimax hypothesis, suggests that the end point of a succession depends on a complex of environmental factors that characterize the site, such as parent material, topography, local climate, and human influences. ‘Actions of the community on the environment, termed autogenic, provide an important driving force promoting successional change, and are typical of primary succession where initial environments are inhospitable. Alternatively, changes in species composition of a community may result from influences external to the community called allogenic, Whereas intrinsic factors often result in progressive successional changes, that is, changes leading from simple to more complex communities, external (allogenie) forces may induce retrogressive succession, that is, toward a less mature community. For example, ifa grassland is severely overgrazed by cattle, th a caltle, the most palatable species will disappear. As grazing continues, the grass cover is reduced, and in a ‘open te weeds ree itic of initial stages of succession may become established In some instances of succession, the food web is based on photosynthetic organisms, and there is slow accumulation of organic matter, both living and dead. This is termed autotrophic’ succession. In other instances, however, addition of organic matter to an ecosystem initiates a succession of decomposer organisms which invade and degrade it. Such ‘a succession is called heterotrophic. Observed changes in the structure and function of communities result from natural selection of Individuals within their current environment. Three mechanisms by which species may replace each other have been proposed; the relative importance of each apparently depends on the nature ofthe sequential and stage of development. 1. ‘The facilitation hypothesis states that invasion of later species depends on conditions created by earlier colonists, Earlier species modify the environment so as to increase the competitive ability of species which are then able to displace them. Succession thus proceeds because of the effects of species on their environment, 2. The tolerance hypothesis suggests that later successional species tolerate lower levels of resources than earlier occupants and can invade and replace them by reducing resource levels below those tolerated by earlier occupants. Succession proceeds despite the resistance of earlier colonists. 3, The inhibition hypothesis is that all species resist invasion of competitors and ate displaced only by death or by damage from factors other than competition. Succession proceeds toward dominance by longer-lived species. None of these models of succession is solely applicable in all instances; indeed most examples of succession appear to show elements of all three replacement mechanisms Succession has traditionally been regardedas followingan orderly progression of changes toward a predictable end point, the climax community, in equilibrium with the prevailing environment. This essentially deterministic view implies that succession will always follow the same course from a given starting point and will pass through a recognizable series of intermediate states, In contrast, a more recent view of succession is based on adaptations of independent species. Iti argued that succession is disorderly and unpredictable, resulting from probabilistic processes such as invasion of propagules and survival of individuals which make up the community. Sch a stochastic view reflects the inherent variability Tan ct innate and the uncertainty of environmental conditions. In particular, it allows Pannen rei fake alternative pathways and end points dependent on the chance outcome of interactions among species and between species and their environment. of community properties such as energy flow supports the view of is. The rate of gross primary productivity typically becomes earl aepallability of nutrients, now incorporated within the community diy a fist as successive invasions occur, but declines again with by the climax community Consideration succession as an orderly proce: biomass, and deci diversity tends to rise rap! f the elimination of the pioneer species