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Chapter 53 Community Ecology Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings What Is a Community? • Assemblage of populations of various species living close enough for potential interaction Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Savanna community in southern Africa Figure 53.1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Community’s interactions include competition, predation, herbivory, symbiosis, and disease • Populations are linked by interspecific interactions Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Interspecific interactions Table 53.1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Competition • Species compete for a particular resource that is in short supply Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Competitive Exclusion Principle • Two species competing for the same limiting resources cannot coexist in the same place Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ecological Niches • The total of an organism’s use of the biotic and abiotic resources in its environment Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Ecologically similar species can coexist in a community if there are one or more significant difference in their niches EXPERIMENT Ecologist Joseph Connell studied two barnacle speciesBalanus balanoides and Chthamalus stellatus that have a stratified distribution on rocks along the coast of Scotland. RESULTS When Connell removed Balanus from the lower strata, the Chthamalus population spread into that area. High tide High tide Chthamalus Chthamalus realized niche Balanus Chthamalus fundamental niche Balanus realized niche Ocean Ocean Low tide In nature, Balanus fails to survive high on the rocks because it is unable to resist desiccation (drying out) during low tides. Its realized niche is therefore similar to its fundamental niche. In contrast, Chthamalus is usually concentrated on the upper strata of rocks. To determine the fundamental of niche of Chthamalus, Connell removed Balanus from the lower strata. Figure 53.2 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Low tide The spread of Chthamalus when CONCLUSIONwas removed indicates that Balanus competitive exclusion makes the realized niche of Chthamalus much smaller than its fundamental niche. • As a result of competition – A species’ fundamental niche may be different from its realized niche Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Resource Partitioning • Differentiation of niches that enables similar species to coexist in a community A. insolitus usually perches on shady branches. A. ricordii A. insolitus A. distichus perches on fence posts and other sunny surfaces. A. alinigar A. christophei A. distichus A. cybotes A. etheridgei Figure 53.3 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Character Displacement • There is a tendency for characteristics to be more divergent in sympatric populations of two species than in allopatric populations of the same two species G. fortis G. fuliginosa Beak depth Percentages of individuals in each size class Santa María, San Cristóbal 40 Sympatric populations 20 0 Los Hermanos 40 G. fuliginosa, allopatric 20 0 Daphne 40 20 G. fortis, allopatric 0 8 10 12 Beak depth (mm) Figure 53.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 14 16 Predation • One species, the predator, kills and eats the other, the prey Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Feeding adaptations of predators include – Claws, teeth, fangs, stingers, and poison • Animals also display – A great variety of defensive adaptations Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Cryptic coloration, or camouflage – Makes prey difficult to spot Figure 53.5 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Aposematic coloration – Warns predators to stay away from prey Figure 53.6 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Batesian mimicry – A palatable or harmless species mimics an unpalatable or harmful model (b) Green parrot snake Figure 53.7a, b (a) Hawkmoth larva Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Müllerian mimicry – Two or more unpalatable species resemble each other (a) Cuckoo bee Figure 53.8a, b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (b) Yellow jacket Herbivory • Herbivore eats parts of a plant – evolution of plant mechanical and chemical defenses and consequent adaptations by herbivores Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Parasitism • One organism, the parasite, derives its nourishment from another organism, its host, which is harmed in the process Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Disease • Pathogens, disease-causing agents – Are typically bacteria, viruses, or protists Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mutualism (mutualistic symbiosis) • Interspecific interaction that benefits both species Figure 53.9 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Commensalism • One species benefits and the other is not affected Figure 53.10 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Dominant and keystone species exert strong controls on community structure • In general, a small number of species in a community – Exert strong control on that community’s structure Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Species Diversity • Variety of different kinds of organisms that make up the community – 2 components Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Species richness – total number of different species in the community • Relative abundance – proportion each species represents of the total individuals in the community Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 2 different communities same species richness, but a different relative abundance A B C D Community 1 A: 25% B: 25% C: 25% D: 25% Community 2 Figure 53.11 A: 80% Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings B: 5% C: 5% D: 10% • A community with an even species abundance – Is more diverse than one in which one or two species are abundant and the remainder rare Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Trophic Structure • Feeding relationships between organisms in a community – key factor in community dynamics Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Food chains – Link the trophic levels from producers to top carnivores Quaternary consumers Carnivore Carnivore Tertiary consumers Carnivore Carnivore Secondary consumers Carnivore Carnivore Primary consumers Zooplankton Herbivore Primary producers Plant Figure 53.12 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings A terrestrial food chain Phytoplankton A marine food chain Food Webs • branching food chain, more complex Humans Smaller toothed whales Baleen whales Crab-eater seals Birds Sperm whales Elephant seals Leopard seals Fishes Squids Carnivorous plankton Copepods Euphausids (krill) Phytoplankton Figure 53.13 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Food webs can be simplified – By isolating a portion of a community that interacts very little with the rest of the community Juvenile striped bass Sea nettle Fish larvae Figure 53.14 Fish eggs Zooplankton Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Dominant Species • Species in a community that are most abundant or have the highest biomass • Exert powerful control over the occurrence and distribution of other species Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • One hypothesis suggests that dominant species – Are most competitive in exploiting limited resources • Another hypothesis for dominant species success – Is that they are most successful at avoiding predators Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Keystone Species • Are not necessarily abundant in a community • Exert strong control on a community by their ecological roles, or niches Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings With Pisaster (control) 20 present Number of species • Sea stars, keystone species in intertidal communities 15 Without Pisaster (experimental) 10 5 0 1963 ´64 (a) The sea star Pisaster ochraceous feeds preferentially on mussels but will consume other invertebrates. Figure 53.16a,b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings ´65 ´66 ´67 ´68 (b) When ´69 ´70 ´71 ´72 Pisaster was removed from an intertidal zone, mussels eventually took over the rock face and eliminated most other invertebrates and algae. In a control area from which Pisaster was not removed, there was little change in species diversity. ´73 • Observation of sea otter populations and their predation Otter number (% max. count) 80 60 40 20 0 (a) Sea otter abundance 400 Grams per 0.25 m2 – Shows the effect the otters have on ocean communities 100 300 200 100 0 Number per 0.25 m2 (b) Sea urchin biomass 10 8 6 4 2 0 1972 1985 1989 Year Figure 53.17 Food chain before killer whale involvement in chain Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (c) Total kelp density 1993 1997 after killer whales started preying on otters Food chain Ecosystem “Engineers” (Foundation Species) • Beaver dams – Can transform landscapes on a very large scale Figure 53.18 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Some foundation species act as facilitators – That have positive effects on the survival and reproduction of some of the other species in the community Number of plant species 8 6 4 2 0 Figure 53.19 Salt marsh with Juncus (foreground) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings With Juncus Without Juncus Conditions • Pollution – Can affect community dynamics • But through biomanipulation – Polluted communities can be restored Polluted State Fish Restored State Abundant Rare Zooplankton Rare Abundant Algae Abundant Rare Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings What Is Disturbance? • Changes a community • Removes organisms from community • Alters resource availability Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Fire – significant disturbance in most terrestrial ecosystems – often a necessity in some communities Figure 53.21a–c (a) Before a controlled burn. A prairie that has not burned for several years has a high proportion of detritus (dead grass). (b) During the burn. The detritus serves as fuel for fires. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (c) After the burn. Approximately one month after the controlled burn, virtually all of the biomass in this prairie is living. • The large-scale fire in Yellowstone National Park in 1988 – Demonstrated that communities can often respond very rapidly to a massive disturbance (a) Soon after fire. As this photo taken soon after the fire shows, ( b) One year after fire. This photo of the same general area taken the burn left a patchy landscape. Note the unburned trees in the distance. Figure 53.22a, b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings the following year indicates how rapidly the community began to recover. A variety of herbaceous plants, different from those in the former forest, cover the ground. Human Disturbance • most widespread agents of disturbance Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ecological Succession • Sequence of community and ecosystem changes after a disturbance Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Primary succession – no soil exists when succession begins • Secondary succession – where soil remains after a disturbance Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Early-arriving species 1. May facilitate the appearance of later species by making the environment more favorable 2. May inhibit establishment of later species 3. May tolerate later species but have no impact on their establishment Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Retreating glaciersfield-research opportunity on succession Canada Grand Pacific Gl. 1940 Alaska 0 1912 1948 1879 1935 Miles 1941 1 899 1907 5 1879 1948 1931 1911 1900 1892 1879 1913 1949 1860 Reid Gl. Johns Hopkins Gl. 1879 Glacier Bay 1830 1780 1760 Pleasant Is. Figure 53.23 McBride glacier retreating Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 10 • Succession on the moraines in Glacier Bay, Alaska – Follows a predictable pattern of change in vegetation and soil characteristics (a) Pioneer stage, with fireweed dominant (b) Dryas 60 Soil nitrogen (g/m2) 50 40 30 20 10 0 Figure 53.24a–d Pioneer Dryas Alder Spruce Successional stage (d) Nitrogen fixation by Dryas and alder increases the soil nitrogen content. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (c) Spruce stage stage • Biogeographic factors affect community diversity • 2 key factors correlated with a community’s species diversity – Are its geographic location and its size Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Species richness generally declines along an equatorial-polar gradient • The greater age of tropical environments greater species richness Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Climate – Is likely the primary cause of the latitudinal gradient in biodiversity Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 2 main climatic factors correlated with biodiversity – solar energy input and water availability 180 200 140 Vertebrate species richness (log scale) Tree species richness 160 100 120 100 50 80 60 40 20 0 100 (a) Trees 10 1 300 Actual 900 1,100 evapotranspiration (mm/yr) 500 700 Figure 53.25a, b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (b) Vertebrates 500 1,000 1,500 2,000 Potential evapotranspiration (mm/yr) Area Effects • All other factors being equal, the larger the geographic area of a community, the greater the number of species Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • A species-area curve of North American breeding birds Number of species (log scale) 1,000 100 10 1 1 10 100 Figure 53.26 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 103 104 105 Area (acres) 106 107 108 109 1010 Island Equilibrium Model • Species richness on islands – Depends on island size, distance from the mainland, immigration, and extinction Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Studies of species richness on the Galápagos Islands – Support the prediction that species richness increases with island size FIELD STUDY Ecologists Robert MacArthur and E. O. Wilson studied the number of plant species on the Galápagos Islands, which vary greatly in size, in relation to the area of each island. RESULTS 400 Number of plant species (log scale) 200 100 50 25 10 5 0 0.1 10 1 100 1,000 Area of island (mi2) (log scale) CONCLUSION Figure 53.28 The results of the study showed that plant species richness increased with island size, supporting the species-area theory. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings