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Chapter 54 Community Ecology PowerPoint® Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Overview: A Sense of Community • A biological community is an group of populations of various species living close enough for potential interaction Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Concept 54.1: Community interactions are classified by whether they help, harm, or have no effect on the species involved • Ecologists call relationships between species in a community interspecific interactions • Examples are competition, predation, herbivory, and symbiosis (parasitism, mutualism, and commensalism) • Interspecific interactions can affect the survival and reproduction of each species, and the effects can be summarized as positive (+), negative (–), or no effect (0) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Competition • Interspecific competition (–/– interaction) occurs when species compete for a resource in short supply In Alaska, the lynx and the fox must compete with each other for prey such as the snowshoe hare. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Competitive Exclusion • Strong competition can lead to competitive exclusion, local elimination of a competing species • The competitive exclusion principle states that two species competing for the same limiting resources cannot coexist in the same place – One species will be more efficient and thus reproduce more rapidly than the other. This will eventually lead to the local elimination of the inferior competitor. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Ecological Niches • The total of a species’ use of biotic and abiotic resources is called the species’ ecological niche • An ecological niche can also be thought of as an organism’s ecological role (its “job”) • Ecologically similar species can coexist in a community if there are one or more significant differences in their niches Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Resource partitioning is differentiation of ecological niches, enabling similar species to coexist in a community – Ex: Seven species of Anolis Lizards live close together in the Dominican Republic. They all eat the same food. Competition, however, is reduced because each lizard species has a preferred perch, thus occupying a distinct niche. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-2 A. distichus perches on fence posts and other sunny surfaces. A. insolitus usually perches on shady branches. A. ricordii A. insolitus A. aliniger A. distichus A. christophei A. cybotes A. etheridgei • As a result of competition, a species’ fundamental niche may differ from its realized niche – Fundamental niche = niche potentially occupied – Realized niche = part of the niche actually occupied Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-3 EXPERIMENT Chthamalus Two barnacle species have a stratified distribution on coastal rocks. Is this the result of interspecific competition between the two? Balanus Chthamalus realized niche Balanus realized niche To find out, researchers removed the Balanus from the rocks. Ocean Results were that the Chthamalus moved into the area formerly occupied by Balanus. Thus, interspecific competition makes the realized niche of Chthamalus much smaller than its fundamental niche High tide RESULTS Knowing that Balanus cannot survive high on the rocks because it dries out during low tides, how does its realized Ocean niche compare with its fundamental niche? Low tide High tide Chthamalus fundamental niche Low tide Character Displacement • Character displacement is a tendency for characteristics to be more divergent in sympatric populations (geographically overlapping)of two species than in allopatric populations (geographically separate) of the same two species • An example is variation in beak size between populations of two species of Galápagos finches – Where these two species live separately, their beak sizes are about the same---indicating they eat the same seeds. – Where they live together (on Santa Maria and San Cristobal) they must compete for food and overtime have evolved differences that allow for different beak sizes thus, eating different seeds. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-4 G. fuliginosa G. fortis Percentages of individuals in each size class Beak depth 60 Los Hermanos 40 G. fuliginosa, allopatric 20 0 60 Daphne 40 G. fortis, allopatric 20 0 60 Sympatric populations Santa María, San Cristóbal 40 20 0 8 10 12 Beak depth (mm) 14 16 Predation • Predation (+/– interaction) refers to interaction where one species, the predator, kills and eats the other, the prey • Some feeding adaptations of predators are claws, teeth, fangs, stingers, and poison Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Prey display various defensive adaptations Behavioral defenses include hiding, fleeing, forming herds or schools, selfdefense, and alarm calls Animals also have morphological and physiological defense adaptations such as: Canyon Tree Frog Cryptic coloration, or camouflage, makes prey difficult to spot Video: Seahorse Camouflage Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Aposematic coloration bright warning coloration indicating a chemical defense (poison) Poison Dart Frog • Batesian Mimicry —a palatable or harmless species mimics an unpalatable or harmful species Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Hawkmoth larva Green parrot snake • Müllerian mimicry-- two or more unpalatable species resemble each other. Each species gains an advantage because the more unpalatable prey there are, the more quickly predators adapt to avoid prey with that appearance. (This is also why warning colors seem to be universal: yellow, black, and red) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Cuckoo bee Yellow jacket Herbivory • Herbivory (+/– interaction) refers to an interaction in which an herbivore eats parts of a plant or alga • It has led to evolution of plant mechanical and chemical defenses and adaptations by herbivores Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Symbiosis • Symbiosis is a relationship where two or more species live in direct and intimate contact with one another • Symbiotic relationships can be harmful, helpful or neutral Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Parasitism • In parasitism (+/– interaction), one organism, the parasite, derives nourishment from another organism, its host, which is harmed in the process • Parasites that live within the body of their host are called endoparasites (tapeworm); parasites that live on the external surface of a host are ectoparasites (leech) • Parasites may account for 1/3 of all the species on Earth • Many parasites have a complex life cycle involving a number of hosts (blood fluke—infects snails and humans) • Some parasites change the behavior of the host to increase their own fitness Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Mutualism • Mutualism (+/+ interaction), is an interspecific interaction that benefits both species • A mutualism can be Ants get food in the form of nectar – Obligate, where one species cannot survive without the other (termites and digestive microbes) – Facultative, where both species can survive alone (acacia trees and ants) Trees get protection as ants keep the area at the base of the tree free from fungus, herbivores, debris, etc. Video: Clownfish and Anemone Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Commensalism • In commensalism (+/0 interaction), one species benefits and the other is apparently unaffected • Commensal interactions are hard to document in nature because any close association likely affects both species Cattle egrets and water buffalo Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Concept 54.2: Dominant and keystone species exert strong controls on community structure • In general, a few species in a community exert strong control on that community’s structure • Two fundamental features of community structure are species diversity and feeding relationships Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Species Diversity • Species diversity of a community is the variety of organisms that make up the community • It has two components: species richness and relative abundance • Species richness is the total number of different species in the community • Relative abundance is the proportion each species represents of the total individuals in the community Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-9 Which forest is more diverse? Both of these forest communities have the same species richness because they each have four species of trees. A B C D Community 1 A: 25% B: 25% C: 25% D: 25% Community 2 A: 80% B: 5% C: 5% D: 10% Their relative abundance, however is very different. Most people would therefore describe Community 1 as being more diverse. • Diversity can be compared using a diversity index – Shannon diversity index (H): H = –[(pA ln pA) + (pB ln pB) + (pC ln pC) + …] A,B, and C are the species in the community, p is the relative abundance of each species, and ln is the natural logarithm. For the two previous communities: Community 1 H= - 4 x (0.25 ln 0.25) = 1.39 Community 2 H = - ((0.8 ln 0.8) + (0.05 ln 0.05) + (0.05 ln 0.05) + (0.1 ln 0.1) = 0.71 This confirms that Community 1 is more diverse. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Determining the number and abundance of species in a community is difficult, especially for small organisms • Molecular tools can be used to help determine microbial diversity – Read figure 54.10 pg. 1205 Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Trophic Structure • Trophic structure is the feeding relationships between organisms in a community • It is a key factor in community dynamics Video: Shark Eating a Seal Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-11 Food chains link trophic levels from producers to top carnivores Quaternary consumers Carnivore Carnivore Tertiary consumers Carnivore Carnivore Secondary consumers Carnivore Carnivore Primary consumers Herbivore Zooplankton Primary producers Plant Phytoplankton A terrestrial food chain A marine food chain Fig. 54-12 Humans A food web is a branching food chain with complex trophic interactions Smaller toothed whales Baleen whales Crab-eater seals Birds Leopard seals Fishes Sperm whales Elephant seals Squids Carnivorous plankton Euphausids (krill) Copepods Phytoplankton • Species may play a role at more than one trophic level This is true for many omnivores. • Food webs can be simplified by isolating a portion of a community that interacts very little with the rest of the community Sea nettle Fish eggs Juvenile striped bass Zooplankton Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Limits on Food Chain Length • Each food chain in a food web is usually only a few links long (about 5) • Two hypotheses attempt to explain food chain length: the energetic hypothesis and the dynamic stability hypothesis Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • The energetic hypothesis suggests that length is limited by inefficient energy transfer – Only about 10% of the energy stored in one level of a food chain is passed on to the next • The dynamic stability hypothesis proposes that long food chains are less stable than short ones – The longer the food chain, the more slowly top predators can recover from environmental shocks. • Most data support the energetic hypothesis Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Species with a Large Impact • Certain species have a very large impact on community structure • Such species are highly abundant or play a pivotal role in community dynamics Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Dominant Species • Dominant species are those that are most abundant or have the highest biomass – Biomass is the total mass of all individuals in a population – Dominant species exert powerful control over the occurrence and distribution of other species – Ex: Maple trees in Northeastern American Forests impact abiotic factors such as shading and soil. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • How does a species become the dominant species? • One hypothesis suggests that dominant species are most competitive in exploiting resources • Another hypothesis is that they are most successful at avoiding predators – Invasive species lend support to the second hypothesis. They are typically introduced to a new environment by humans and often lack predators or disease, therefore attaining a high biomass. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • The impact of a dominant species on a community can be seen when it is accidentally lost – Ex: American Chestnut: formally the dominant tree of Northeastern American forests. All were killed by a fungus between 1910 and 1950. Few mammals and birds were harmed by the loss, but 7 species of moths and butterflies went extinct. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Keystone Species • Keystone species exert strong control on a community by their ecological roles, or niches • In contrast to dominant species, they are not necessarily abundant in a community – Example: Pacific Sea Otter Observation of sea otter populations and their predation shows how otters affect ocean communities Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-16 Otter number (% max. count) 100 When sea otters are abundant, kelp forests thrive 80 60 40 20 0 (a) Sea otter abundance 300 200 100 0 (b) Sea urchin biomass Number per 0.25 m2 As the otters are hunted by orcas, their numbers decline. Sea urchin populations rise, causing the loss of kelp forests. Grams per 0.25 m2 400 10 8 6 4 2 0 1972 1985 (c) Total kelp density 1989 Year 1993 1997 Food chain Fig. 54-15 EXPERIMENT Field studies of sea stars exhibit their role as a keystone species in intertidal communities. They feed on mussels. Without them, the mussels outcompete all other species and diversity declines. Number of species present RESULTS 20 15 With Pisaster (control) 10 5 Without Pisaster (experimental) 0 1963 ’64 ’65 ’66 ’67 ’68 ’69 ’70 ’71 ’72 ’73 Year Foundation Species (Ecosystem “Engineers”) • Foundation species (ecosystem “engineers”) cause physical changes in the environment that affect community structure • For example, beaver dams can transform landscapes on a very large scale Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-18 Number of plant species Some foundation species act as facilitators that have positive effects on survival and reproduction of some other species in the community 8 6 4 2 0 (a) Salt marsh with Juncus (foreground) (b) With Juncus Juncus prevent salt build up in soil and evaporation thereby increasing diversity Without Juncus Bottom-Up and Top-Down Controls • The bottom-up model of community organization proposes a unidirectional influence from lower to higher trophic levels N V H P • In this case, presence or absence of mineral nutrients (N) determines the amount of vegetation (V) which controls the number of herbivores (H) and the number of predators (P) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • The top-down model, also called the trophic cascade model, proposes that control comes from the trophic level above N V H P • In this case, predators control herbivores, which in turn control the amount of vegetation and the uptake of nutrients • The effects of any manipulation thus move down the trophic structure as alternating +/- effects Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Long-term experimental studies have shown that communities vary in their relative degree of bottom-up to top-down control Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Pollution can affect community dynamics • Biomanipulation can help restore polluted communities • In the example that follows, algal blooms and eutrophication have polluted a pond. If fish are removed, zooplankton levels should raise. Once this happens, algae levels should decline and eutrophication should be slowed. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-UN1 This is an example of biomanipulation using the top-down control. Polluted State Restored State Fish Abundant Rare Zooplankton Rare Abundant Algae Abundant Rare Concept 54.3: Disturbance influences species diversity and composition • Decades ago, most ecologists favored the view that communities are in a state of equilibrium • This view was supported by F. E. Clements who suggested that species in a climax community function as a superorganism • Other ecologists, including A. G. Tansley and H. A. Gleason, challenged whether communities were at equilibrium • Recent evidence of change has led to a nonequilibrium model, which describes communities as constantly changing after being buffeted by disturbances Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Characterizing Disturbance • A disturbance is an event that changes a community, removes organisms from it, and alters resource availability • Fire is a significant disturbance in most terrestrial ecosystems • It is often a necessity in some communities Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • The intermediate disturbance hypothesis suggests that moderate levels of disturbance can foster greater diversity than either high or low levels of disturbance. They do this by opening up habitats for occupation by less competitive species • High levels of disturbance exclude many slow-growing species • Low levels of disturbance allow dominant species to exclude less competitive species Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • The large-scale fire in Yellowstone National Park in 1988 demonstrated that communities can often respond very rapidly to a massive disturbance • The dominant species in Yellowstone is the Lodgepole pine. It releases seeds only after exposure to extreme heat. The fire is actually healthy for a forest like this. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-21 (a) Soon after fire (b) One year after fire Ecological Succession • Ecological succession is the sequence of community and ecosystem changes after a disturbance • Primary succession occurs where no soil exists when succession begins. May take hundreds or thousands of years • Secondary succession begins in an area where soil remains after a disturbance Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Early-arriving species and later-arriving species may be linked in one of three processes: – Early arrivals may facilitate appearance of later species by making the environment favorable – They may inhibit establishment of later species – They may tolerate later species but have no impact on their establishment Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Retreating glaciers provide a valuable fieldresearch opportunity for observing succession • Succession on the moraines in Glacier Bay, Alaska, follows a predictable pattern of change in vegetation and soil characteristics Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Current Iceberg Front Glacier Bay Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-22-1 1941 1907 1 Pioneer stage, with fireweed dominant 0 1860 Glacier Bay Alaska 1760 5 10 15 Kilometers Fig. 54-22-2 1941 1907 2 1 Pioneer stage, with fireweed dominant 0 1860 Glacier Bay Alaska 1760 5 10 15 Kilometers Dryas stage—after 30 years Fig. 54-22-3 1941 1907 2 1 Pioneer stage, with fireweed dominant 0 1860 Dryas stage—after 30 yrs. 5 10 15 Kilometers Glacier Bay Alaska 1760 3 Alder stage—after another few decades Fig. 54-22-4 1941 1907 2 1 Pioneer stage, with fireweed dominant 0 1860 Dryas stage—after 30 yrs. 5 10 15 Kilometers Glacier Bay Alaska 1760 4 Spruce stage—after two centuries 3 Alder stage—after another few decades • Succession is the result of changes induced by the vegetation itself • On the glacial moraines, vegetation lowers the soil pH and increases soil nitrogen content • As this occurs, more and more plant species can develop over time Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-23 60 Soil nitrogen (g/m2) 50 40 30 20 10 0 Pioneer Dryas Alder Successional stage Spruce Human Disturbance • Humans have the greatest impact on biological communities worldwide • Human disturbance to communities usually reduces species diversity • Humans also prevent some naturally occurring disturbances, which can be important to community structure These photos show disturbances on the ocean floor before (top) and after (bottom) deep-sea trawlers have passed. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Concept 54.4: Biogeographic factors affect community biodiversity • Latitude and area are two key factors that affect a community’s species diversity Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Latitudinal Gradients • Species richness generally declines along an equatorial-polar gradient and is especially great in the tropics • Two key factors in equatorial-polar gradients of species richness are probably evolutionary history and climate – The greater age of tropical environments may account for the greater species richness – Climate is likely the primary cause of the latitudinal gradient in biodiversity – Two main climatic factors correlated with biodiversity are solar energy and water availability – They can be considered together by measuring a community’s rate of evapotranspiration – Evapotranspiration is evaporation of water from soil plus transpiration of water from plants Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-25 180 140 Tree species richness Potential evapotranspiration is a measure of potential water loss. 160 120 100 80 60 40 20 0 100 300 500 700 900 Actual evapotranspiration (mm/yr) 1,100 (a) Trees 200 Vertebrate species richness (log scale) Evapotranspiration is higher in hot areas with abundant rainfall 100 50 10 0 500 1,000 1,500 Potential evapotranspiration (mm/yr) (b) Vertebrates 2,000 Area Effects • The species-area curve quantifies the idea that, all other factors being equal, a larger geographic area has more species • A species-area curve of North American breeding birds supports this idea Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-26 Number of species 1,000 100 10 1 0.1 1 10 100 103 104 105 106 107 108 109 1010 Area (hectares) Island Equilibrium Model • Species richness on islands depends on island size, distance from the mainland, immigration, and extinction • The equilibrium model of island biogeography maintains that species richness on an ecological island levels off at a dynamic equilibrium point Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Equilibrium number Number of species on island (a) Immigration and extinction rates Rate of immigration or extinction Rate of immigration or extinction Rate of immigration or extinction Fig. 54-27 Small island Large island Number of species on island (b) Effect of island size Far island Near island Number of species on island (c) Effect of distance from mainland • Studies of species richness on the Galápagos Islands support the prediction that species richness increases with island size Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Number of plant species (log scale) Fig. 54-28 400 200 100 50 25 10 5 10 100 103 104 Area of island (hectares) (log scale) 105 106 Concept 54.5: Community ecology is useful for understanding pathogen life cycles and controlling human disease • Ecological communities are universally affected by pathogens, which include disease-causing microorganisms, viruses, viroids, and prions • Pathogens can alter community structure quickly and extensively Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Pathogens and Community Structure • Pathogens can have dramatic effects on communities • For example, coral reef communities are being decimated by white-band disease As the staghorn coral dies, algal species take its place, and the fish community is dominated by herbivores feeding on the algae. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Human activities are transporting pathogens around the world at unprecedented rates • Community ecology is needed to help study and combat them Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Community Ecology and Zoonotic Diseases • Zoonotic pathogens have been transferred from other animals to humans • The transfer of pathogens can be direct or through an intermediate species called a vector • Many of today’s emerging human diseases are zoonotic Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings • Avian flu is a highly contagious virus of birds • Ecologists are studying the potential spread of the virus from Asia to North America through migrating birds Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 54-UN2 You should now be able to: 1. Distinguish between the following sets of terms: competition, predation, herbivory, symbiosis; fundamental and realized niche; cryptic and aposematic coloration; Batesian mimicry and Müllerian mimicry; parasitism, mutualism, and commensalism; endoparasites and ectoparasites; species richness and relative abundance; food chain and food web; primary and secondary succession Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 2. Define an ecological niche and explain the competitive exclusion principle in terms of the niche concept 3. Explain how dominant and keystone species exert strong control on community structure 4. Distinguish between bottom-up and top-down community organization 5. Describe and explain the intermediate disturbance hypothesis Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 6. Explain why species richness declines along an equatorial-polar gradient 7. Define zoonotic pathogens and explain, with an example, how they may be controlled Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings