5/13/13 Community = An assemblage of populations (species) in a particular area or habitat. Here is part of a community in the grassland of the Serengetti. Trophic downgrading of planet Earth: What escapes the eye is a much more insidious kind of extinction: the extinction of ecological interactions Daniel Janzen Wolves in Yellowstone Wolves-the unexpected impacts Elk were eating riparian (river) trees and shrubs Birds and steelhead had declined ecosystem change Wolves drove Elk up into higher ground Riparian systems are recovering, ecosystem is shifting back Top down Ecosystem change: an example Ecosystem change: an example Killer whales and their appetite for sea otters Reduction of coral = increase in Algae. This shifts functional Groups of species, and affects Primary productivity. Bottom up 1
Dead zones in the oceans Eutrophication Stratification and oxygen depletion on the bottom Can affect all trophic levels, but it takes time Some important concepts What is a niche? - The role of an organism Its use of abiotic and biotic resources, and its reaction to them (eg. Tolerance plays a role here) Competitive Exclusion Principle No two species can occupy the same niche - it will always result in competition Generalists Specialists Vultures: multiple niches or multiple species per niche? Table 53.1 A simplistic summary of the kinds of species interactions. 2
Species Interactions: interspecific -/- (competition) What leads to competition? What is the result? Leave Adapt: resource partitioning Competitive Exclusion Principal Competition Realized Niche vrs. Fundamental Niche Who exists in which? Figure 53.2 Testing a competitive exclusion hypothesis in the field. Removal of Balanus results in Chthamalus occupying a larger part of the rock. Thus, Balanus is competitively excluding Chthamalus from parts of the rock. Figure 53.2 Testing a competitive exclusion hypothesis in the field. Removal of Balanus results in Chthamalus occupying a larger part of the rock. Thus, Balanus is competitively excluding Chthamalus from parts of the rock. Figure 53.2 Testing a competitive exclusion hypothesis in the field. Removal of Balanus results in Chthamalus occupying a larger part of the rock. Thus, Balanus is competitively excluding Chthamalus from parts of the rock. Figure 53.4 Coevolutionary impact of competition: Character displacement. Two species have different beak sizes when occurring together and competing for seeds, compared to when they live apart. 3
Competition Competitive dominance (balanus/thamalus) Resource partitioning Character displacement - resource partitioning Effects of competition One species dominates (within a functional group): low species diversity. What is the role of lesser species within the same functional group (eg diversity)? Resource partitioning leads to variation (phenotypic) in sympatric species Competition can lead to genotypic differences - speciation Competition is an important factor in maintaining ecosystem function. When one species is removed, the structure of competition is changed too. Introduced species? Ecosystem change: an example Reduction of coral = increase in Algae. This shifts functional Groups of species, and affects Primary productivity Species with large impact Keystone species - is there such a thing? A species whose role or niche has a major impact on the structure of an ecosystem Indicator Species Sensitive to change Dominant species: most biomass Ecosystem engineers: foundation species/ facilitators Sea Otters and Sea Urchins: a kelp forest paradigm Figure 53.17 Kelp forests can be radically restructured depending on who the top predator is. Sea Urchins eat kelp, especially new recruits If kept in check, they eat drift kelp If populations expand, they will eat established kelp Sea Otters eat urchins, especially exposed ones They will keep sea urchin populations in check The Aleutian Island studies Sea Otters as a keystone predator 4
Community Ecology What is species diversity: both Species richness Relative abundance What leads to high diversity? Habitat heterogeneity High levels of competition Other interspecific interactions, functional group diversity, abiotic factors? Some important concepts Functional groups Several related niches, eg. scavengers, grazers, decomposers etc. Table 53.1 A simplistic summary of the kinds of species interactions. Species Interactions: interspecific Positive Negative Neutral (can it be?) The reality of species interactions 1. There is a continuum of possibilities between the major four types. For example, an interaction between a plant and a mycorrhizal fungus can range from mutualism to commensalism to parasitism as soil nutrients range from scarce to abundant. 2. Most species interactions are asymmetric, i.e. Species A has a much greater impact on Species B than Species B has on Species A. Species Interactions: interspecific +/- (predation and parasitism) Plant defenses against herbivory Animal defenses against predation Parasites: endo and ecto Co-Evolution 5
Predation Predation includes a variety of interactions in which one species consumes all or part of another organism: predators, herbivores, and parasites all engage in predation Examples: Lynx eating the snow shoe hare Caterpillar eating plant leaves Parasitic nematode living in the gut of a dog Species Interactions: interspecific +/o, +/+ (mutualism & commensalism) Does commensalism exist? Mutualism requires evolution of adaptations in both species: a true partnership. Nitrogen fixation on plant roots Digestion track bacteria Zooxanthellae in corals Anemones and fish Cleaning stations on reefs Mutualism Mutualisms include mutually beneficial interactions between a wide variety of species. Examples: Plants and their pollinators, seed dispersers, mycorrhizal fungi, and Rhizobia Lichens: fungi and their symbiotic algae or cyanobacteria Cleaning symbiosis: mutualism Figure 53.9 Mutualism between Acacia trees and ants. Trees provide food for the ants in the form of nectar and fatty Beltian bodies (yellow objects at leaf margins), and also provide shelter in their thorns. Ants remove and/or kill other insects, fungal spores, and clip off the leaves of any vegetation that grows too close to the Acacia. In a commensalism, one species benefits, and the other is not affected. Shown here is a commensalism between a bird and mammal, whereby the bird picks small parasites off the mammal. The bird benefits, but the mammal probably does not. 6
Zooxanthellae: the key to coral reef productivity mutualism Temporal Community Development (succession) Facilitation model Inhibition model Sere: successional stage defined by a community of living organisms of various trophic levels Climax communities? Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Primary forest succession Figure 3.27 The role of ecosystem disturbance Equilibrium and stability: over what time scale do we measure this? Natural and anthropogenic disturbance Short term and long-term disturbance Succession and disturbance: climax communities? What does disturbance adapted mean? 3-14 7
Figure 53.19 Patchiness and recovery following a large-scale disturbance: Mount St. Helens Figure 53.18 Patchiness and recovery following a large-scale disturbance by fire. After a disturbance, the process of succession begins: a habitat is colonized by a suite of ruderal or weedy species, which are gradually replaced by a series of other species. Figure 53.24 The Pattern of Succession following retreat of glaciers at Glacier Bay, Alaska. Succession and diversity Non-equilibrium model: succession can take a variety of pathways Depends on the size, frequency and severity of the disturbance By destabilizing community structure, disturbances initiate change, and affect its course Vacated niches Changes in competition for resources Habitat changes Disturbance and diversity Nonequilibrial model: diversity is maintained by environmental patchyness eg. Caused by frequent abiotic disturbance Dynamic equilibrium hypothesis: species diversity depends on the effect of the disturbance on competitive interactions. Diversity is enhanced when competitive exclusion is prevented Intermediate disturbance hypothesis: species diversity is greatest where disturbance is moderate in frequency and severity. Key concepts in community ecology Trophic forcing: bottom up, top down Niche Effects of competition Competitive exclusion Niche partitioning Fundamental vrs. Realized niche Character displacement Species with large impact Species diversity Predation Symbiosis Community succession and the role of disturbance 8
Biogeography Regions defined by species assemblages Dispersal (range/tolerance/adaptations), and survivability account for geographical ranges of species Geology and geographical barriers play a key role Range explanations A species never dispersed beyond its present boundaries Pioneers failed to survive Range has been reduced over evolutionary time Pioneers became isolated/separated: phylogeography Phylogeography: ecological genetics Study of populations (not species), their distributions, and the degree of genetic separation between populations Investigates the cause of population structure Island Biogeography Depth of the water, or degree of difficulty to cross a barrier Distance away from source Size of the island Species diversity is highest on large near islands: immigration and extinction (competitive exclusion and small islands) Figure 53.20 Biogeographical realms Figure 53.21 The hypothesis of island biogeography 9
Figure 53.22 Species richness and island size Key Points Species diversity (richness and abundance) is determined in part by habitat heterogeneity and species interactions. Niches and the CEP - resource partitioning or exclusion Keystone species and functional groups Interspecific interactions are a driving force in evolution: examples? Ecosystem disturbance affects community structure, composition, and diversity Species assemblages can be explained in part through historical ranges and the establishment of barriers. 10