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The Living World Fifth Edition George B. Johnson Jonathan B. Losos Chapter 38 Populations and Communities Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 38.1 Population Growth • A population is a group of individuals of a species that live together and influence each other’s survival • Populations have several properties that can describe them population size is the number of individuals in the population population density is the population size that occurs in a given area population dispersion is the scatter of individuals within the population’s range Figure 38.1 Population dispersion 38.1 Population Growth • Another characteristic about any population is its capacity to grow • Population growth can be modeled in different ways that identify what factors in nature limit growth 38.1 Population Growth • Biotic potential, symbolized by r, is the rate at which a population of a given species will increase when no limits are placed on its rate of growth • The simplest model of population growth assumes a population growing without limits at its maximal rate 38.1 Population Growth • The exponential growth model is defined by the following formula growth rate = dN/dT = riN N is the population size dN/dT is the rate of change over time ri is the intrinsic rate of natural increase 38.1 Population Growth • The actual rate of population increase, r, is defined as r = (b – d) + (i – e) • b is the birthrate, d is the death rate • e is the amount of emigration out of the area and i is the amount of immigration into the area 38.1 Population Growth • The innate capacity for growth of any population is exponential even when the rate of increase remains constant, the actual increase in the number of individuals accelerates rapidly as the population grows in practice, such patterns prevail for only short periods, usually when an organism reaches a new habitat with abundant resources 38.1 Population Growth • No matter how rapidly populations grow, they eventually reach a limit imposed by shortages of important environmental factors • A population ultimately stabilizes at a certain size, called the carrying capacity the carrying capacity is symbolized by K and is defined as the maximum number of individuals that an area can support 38.1 Population Growth • The growth curve of a population that is approaching its carrying capacity can be approximated by the logistic growth equation dN/dt = rN (1 – N/K) • As N approaches K, the rate of population growth (dN/dt) begins to slow, until it reaches zero at N = K Figure 38.2 Two models of population growth 38.1 Population Growth • The sigmoid growth curve is characteristic of most biological populations • The processes of competition and emigration tend to increase as a population approaches its carrying capacity Figure 38.3 Most natural populations exhibit logistic growth 38.2 The Influence of Population Density • Many factors act to regulate the growth of populations in nature density-independent effects • these effects regulate population growth regardless of population size • for example, weather effects or geological events (i.e., volcanoes) density-dependent effects • the effect that these factors have on population growth depends on population size • these effects grow stronger as the population size increases Figure 38.4 Density-dependent effects 38.2 The Influence of Population Density • In natural systems that are exploited by humans, the aim is to maximize productivity by exploiting the population early in the rising portion of its sigmoid growth curve commercial fisheries, for example, attempt to operate so that they harvest a population near its point of maximal sustainable yield Figure 38.5 Maximal sustainable yield 38.3 Life History Adaptations • Life history describes the complete life cycle of an organism r-selected adaptations • these favor rapid growth in a habitat with unlimited resources or in unpredictable environments • or, in unpredictable environments, organisms have to take advantage of resources when they are available K-selected adaptations • favor reproduction near the carrying capacity of the environment • help survival in an environment in which individuals are competing for limited resources Table 38.1 r-selected and K-selected life history adaptations 38.3 Life History Adaptations • The r/K concept of life histories provides a way to examine more closely related organisms living in different types of habitats populations living in rapidly changing environments tend to exhibit r-selected adaptations populations living in more stable and competitive habitats tend to exhibit more Kselected adaptations Figure 38.6 The consequences of exponential growth 38.4 Population Demography • demography is the statistical study of populations measures characteristics of populations and helps predict how population sizes will change in the future • populations grow if births outnumber deaths and shrink if deaths outnumber births • birth and death rates are dependent on age and sex 38.4 Population Demography • A cohort is a group of individuals of the same age within a population, every cohort has the following characteristics • fecundity, or birthrate, which is defined as the number of offspring produced in a standard time • mortality, or deathrate, which is the number of individuals that die in that period the relative number of individual’s in each cohort defines a populations age structure 38.4 Population Demography • Sex ratio is the proportion of males and fames in a population the number of births is usually directly related to the number of females • Age distribution is the proportion of individuals in different age categories when a population lives in a constant environment for a few environments, its population size remains fairly constant and is said to be a stable population 38.4 Population Demography • A survivorship curve is one way to express the age distribution characteristics of a population survivorship is defined as the percentage of an original population that survives to a given age there are three types of survivorship curves • type I has highest mortality for the youngest individuals • type II has relatively the same mortality risk for all ages • type III has highest mortality for the oldest individuals Figure 38.7 Survivorship curves 38.5 Communities • Community refers to the species that occur at any given locality communities can be characterized by either their constituent species (a list of all species present in the community) or by their properties, such as primary productivity interactions among community members govern many ecological and evolutionary processes • for example, predation, competition, and mutualism affect the population biology of a particular species, as well as the way in which energy and nutrients cycle through the ecosystem Figure 38.8 A Tanzanian savanna community 38.5 Communities • Two views exist on the makeup and functioning of communities individualistic concept • advanced by H. A. Gleason, this concept holds that a community is nothing more than an aggregation of species that happen to co-occur at one place holistic concept • views communities as an integrated unit and this concept was first proposed by F. E. Clements • the community is viewed as a “superorganism” 38.6 The Niche and Competition • The niche an organism occupies is the sum total of all the ways it utilizes the resources of its environment sometimes species are not able to occupy their entire niche because of the presence or absence of other species competition describes the interaction when two organisms attempt to use the same resource when there is not enough of the resource to satisfy both – interspecific competition occurs between individuals of different species – intraspecific competition occurs between individuals of the same species 38.6 The Niche and Competition • Fundamental niche is the entire niche that an organism may theoretically occupy • Realized niche is the actual niche that the organism is able to occupy because of competition Figure 38.9 Competition among two species of barnacles limits niche use 38.6 The Niche and Competition • G.F. Gause demonstrated the principle of competitive exclusion if two species are competing for a resource, the species that uses the resource more efficiently will eventually eliminate the other locally in other words, no two species with the same niche can coexist Figure 38.10 (a) Competitive exclusion among three species of Paramecium Figure 38.10 (b) Competitive exclusion among three species of Paramecium Figure 38.10 (c) Competitive exclusion among three species of Paramecium 38.6 The Niche and Competition • Species in communities act to avoid competition whenever possible when their niches overlap, two outcomes are possible • competitive exclusion (i.e., winner takes all) • resource partitioning, which divides up resources to create two niches thus, persistent competition between two species is rare in natural communities either one species drives the other to extinction or natural selection reduces the competition between the them Figure 38.11 Resource partitioning in warblers 38.6 The Niche and Competition • Resource partitioning can often be seen in similar species that occupy the same geographical area such species are sympatric when a pair of species occupy the same habitat (i.e., when they are sympatric), they tend to exhibit greater differences in morphology and behavior then the same two species do when living in different habitats (i.e., when they are allopatric) • the evident differences are called character displacement and are favored by natural selection to facilitate habitat partitioning and reduce competition 38.7 Coevolution and Symbiosis • Coevolution is the adaptation of a species not only to its physical environment but also to the other organisms that share it examples of coevolution include • plants and animal pollinators • predator-prey interactions • symbiotic relationships Figure 38.13 Pollination by bat 38.7 Coevolution and Symbiosis • In a symbiotic relationship, two or more kinds of organisms live together in often elaborate or more of less permanent relationships there are three major kinds of symbiotic relationships • mutualism • parasitism • commensalism 38.7 Coevolution and Symbiosis • Mutualism is a symbiotic relationship in which both species benefit Figure 38.14 Mutualism: ants and aphids Figure 38.15 Mutualism: ants and acacias 38.7 Coevolution and Symbiosis • Parasitism is a symbiotic relationship in which one species benefits while the other is harmed this interaction is really a form of predator/prey relationship but a parasite usually does not kill its host the parasite is much smaller than the host and remains closely associated with it 38.7 Coevolution and Symbiosis • There are many forms of parasitism in nature external parasites • also known as ectoparasites, these parasites feed on the exterior surface of a host • parasitoids are insects that lay eggs on living hosts internal parasites • also known as endoparasites, these parasites feed internally in their hosts brood parasitism is a form of parasitism in which the parasite does not consume the body of its host • brood parasites are birds, such as cowbirds and cuckoos, that lay their eggs in the nest of other species for the host to raise Figure 38.16 Parasitism 38.7 Coevolution and Symbiosis • Commensalism is a symbiotic relationship that benefits one species but neither hurts nor helps the other there is no clear-cut boundary between commensalism and mutualism Examples of Commensalism Figure 38.17 Commensalism in the sea Figure 38.18 Commensalism between oxpeckers and African cape buffalo 38.8 Predator-Prey Interactions • Predation is the consuming of one organism by another in nature, predators often have large effects on prey populations • population cycles may be, in some situations, stimulated by predators • a classic example is the “10-year cycle” of the snowshoe hare, Lepus americanus, that appears to be under the influence of food plants and predators under laboratory conditions, predators may exhaust their prey species and then starve Figure 38.21 A predator-prey cycle Figure 38.20 Predator-prey in the microscopic world 38.8 Predator-Prey Interactions • Predator-prey interactions are an essential factor in the maintenance of communities that are rich and diverse in species predators prevent or greatly reduce competitive exclusion by reducing the number of individuals of competing species predators that reduce competition and increase community diversity are known as keystone species Figure 38.22 Predation reduces competition 38.9 Plant and Animal Defenses • Pressures from predation have driven the evolution of mechanisms that defend organisms from predation plants produce chemical defenses that make them toxic to herbivores • some herbivores have, as a result, evolved a tolerance to these chemicals and may use them for their own defense many animals have defensive coloration • aposematic coloration is a warning coloration that is characteristic of animals that use poisons • cryptic coloration is color that blends in with surroundings Figure 38.23 Insect herbivores are well suited to their hosts Figure 38.24 A blue jay learns that monarch butterflies taste bad Defensive Coloration Figure 38.25 Cryptic coloration Figure 38.26 Vertebrate chemical defense 38.9 Plant and Animal Defenses • Predation can exert strong selective pressures on prey populations a coevolutionary arms race between predators and prey is likely because • any feature that acts to decrease the probability of capture should thus be strongly favored by selection in prey • natural selection would also favor counteradaptations in predators 38.10 Mimicry • Batesian mimicry is when a palatable species resembles a poisonous one there may also be nonvisual cues, such as olfaction, involved • Müllerian mimicry is when several unrelated, but protected, species come to resemble one another for example, the colors black, yellow, and red are used often in aposematic coloration Figure 38.27 A Batesian mimic Figure 38.28 Müllerian mimics 38.10 Mimicry • Self mimicry is a special case of mimicry in which one animal body part comes to resemble another body part this occurs in both prey and predators • prey might use this form of mimicry to startle a predator or to provide a false target for attack • predators might use this mimicry to simulate bait to lure prey in Figure 38.29 Self mimicry 38.11 Ecological Succession • Succession is the orderly replacement of one community with another primary succession • occurs on bare, lifeless substrates, such as those left behind when a glacier retreats or when a volcanic island emerges • pioneering community is the first to become established secondary succession • occurs after an already established community has been disturbed Figure 38.30 Plant succession produces progressive changes in the soil 38.11 Ecological Succession • Succession happens because species alter the habitat and the resources available in it, often in ways that favor other species • Three dynamic concepts are of critical importance tolerance • early succession stages are characterized by weedy r-selected species that tolerate harsh conditions but do not compete well facilitation • the weedy species introduce local changes in the habitat that favor non-weedy species Inhibition • sometimes the changes in habitat caused by one species may inhibit the growth of the species that caused them Figure 38.31 Primary succession at Alaska’s Glacier Bay Inquiry & Analysis • Are the populations with lower juvenile mortality bigger or smaller than the populations with higher juvenile mortality? • Do the population sizes of these song sparrows appear to exhibit density dependence? Graph of Effects of Population Size on Songbird Success