Ecological Communities 45. Concept 45.1 Communities Contain Species That Colonize and Persist...

Ecological Communities

45

Concept 45.1 Communities Contain Species That Colonize and Persist

Community—a group of species that coexist and interact with one another within a defined area

Biologists may designate community boundaries based on natural boundaries (e.g., the edge of a pond) or arbitrarily.

They may restrict study to certain groups (e.g., the bird community).

Concept 45.1 Communities Contain Species That Colonize and Persist

Communities are characterized by species composition; that is, which species they contain and the relative abundances of those species.

A species can occur in a location only if it is able to colonize and persist there.

A community contains those species that have colonized minus those that have gone extinct locally.

Concept 45.1 Communities Contain Species That Colonize and Persist

Local extinctions can occur for many reasons:

• Species unable to tolerate local conditions

• A resource may be lacking

• Exclusion by competitors, predators, or pathogens

• Population size too small; no reproduction

Concept 45.1 Communities Contain Species That Colonize and Persist

In 1883 the volcano on Krakatau erupted, killing everything on the island.

Scientists studied the return of living organisms. Within 3 years, seeds of 24 plant species had reached the island.

Later, as trees grew up, some pioneering plant species that require high light levels disappeared from the island’s now-shady interior.

Species composition continues to change as new species colonize and others go extinct.

Figure 45.1 Vegetation Recolonized Krakatau (Part 1)

Figure 45.1 Vegetation Recolonized Krakatau (Part 2)

Concept 45.2 Communities Change over Space and Time

Ecologists have documented recurring patterns of species compositional change.

Species composition varies along environmental gradients, after disturbances, and with changing climate.

Concept 45.2 Communities Change over Space and Time

Species composition changes along environmental gradients, such as elevation or soil types.

A transect is a straight line used for ecological surveys.

A transect along an environmental gradient will show the species turnover through space.

Figure 45.2 Species Turnover along an Environmental Gradient

Concept 45.2 Communities Change over Space and Time

Many animal species are associated with particular plant communities:

The plants they eat may be there; or because plants modify physical conditions and contribute to habitat structure.

Habitat structure determines the animal’s ability to get food or avoid predators.

Figure 45.3 Many Animals Associate with Habitats of a Particular Structure (Part 1)

Figure 45.3 Many Animals Associate with Habitats of a Particular Structure (Part 2)

Concept 45.2 Communities Change over Space and Time

In any community there is ongoing colonization and extinction, and thus a steady turnover in species composition.

Species turnover can result from disturbance events: volcanic eruptions, wildfires, hurricanes, landslides, human activities.

Some or all the species are wiped out, and environmental conditions are changed.

Concept 45.2 Communities Change over Space and Time

Species often replace one another in a predictable sequence called succession.

Example: A patch of elephant dung is colonized by a sequence of dung beetle species.

Figure 45.4 Dung Beetle Species Composition Changes over Time

Concept 45.2 Communities Change over Space and Time

Some species are better at colonizing than others.

Early-arriving dung beetles tend to be strong fliers with a good sense of smell, or “hitchhikers” that ride on the dung-producers.

On Krakatau, the first plants were species that have seeds that are easily dispersed by sea or wind.

Concept 45.2 Communities Change over Space and Time

After a disturbance, environmental conditions change with time.

Examples:

Dung starts out wet and dries over time.

As trees grow, the forest canopy closes and light conditions change.

Concept 45.2 Communities Change over Space and Time

After a disturbance, succession often leads to a community that resembles the original one.

Example: On Krakatau, tropical forests eventually came back.

Concept 45.2 Communities Change over Space and Time

If the original community is not reestablished, there is an ecological transition to a different community.

Example: Conversion of grasslands to shrublands in the U.S.–Mexico Borderlands after intensive cattle grazing.

Concept 45.2 Communities Change over Space and Time

Climate change can also cause temporal variation in communities.

As physical conditions change, the geographic ranges of species necessarily change with them.

One way to reconstruct such change is analysis of fossilized plant remains in packrat middens. Biologists can show how plant communities of the Borderlands changed over the last 14,000 years as the climate became drier.

Figure 45.5 Species Composition Changes with Climate Change

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Each species in a community has a unique niche.

This concept refers to the environmental tolerances of a species, which define where it can live.

Also refers to the ways a species obtains energy and materials and to patterns of interaction with other species in the community.

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Consumer–resource, or trophic interactions cause energy and materials to flow through a community.

Trophic levels—feeding positions

Primary producers, or autotrophs, convert solar energy into a form that can be used by the rest of the community.

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Heterotrophs get energy by breaking apart organic compounds that were assembled by other organisms.

Primary consumers (herbivores) eat primary producers.

Secondary consumers (carnivores) eat herbivores.

Tertiary consumers eat secondary consumers.

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Omnivores feed from multiple trophic levels.

Decomposers, or detritivores, feed on waste products or dead bodies of organisms.

Decomposers are responsible for recycling of materials; they break down organic matter into inorganic components that primary producers can absorb.

Table 45.1 The Major Trophic Levels

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Trophic interactions are shown in diagrams called food webs.

Arrows indicate the flow of energy and materials —who eats whom.

Figure 45.6 A Food Web in the Yellowstone Grasslands

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Gross primary productivity (GPP)—total amount of energy that primary producers convert to chemical energy.

Net primary productivity (NPP)—energy contained in tissues of primary producers and is available for consumption.

Change in biomass of primary producers (dry mass) per unit of time is an approximation for NPP.

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Ecological efficiency is about 10%:

Only about 10% of the energy in biomass at one trophic level is incorporated into the biomass of the next trophic level.

This loss of available energy at successive levels limits the number trophic levels in a community.

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Ecological efficiency is low because:

• Not all the biomass at one trophic level is ingested by the next one.

• Some ingested matter is indigestible and is excreted as waste.

• Organisms use much of the energy they assimilate to fuel their own metabolism.

Figure 45.7 Energy Flow through Ecological Communities

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

The per capita growth rate of a species is related to the sum of positive and negative contributions of species with which it interacts.

Succession can be driven by such interactions.

Examples: Late-colonizing dung beetles inhibit early colonizers by competing for nutrients.

Late-arriving plant species shade out pioneering species.

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Consumer–resource interactions can have ripple effects across trophic levels, resulting in a trophic cascade.

In Yellowstone National Park, wolves were extirpated by hunting by 1926.

Elk were culled each year to prevent them from exceeding carrying capacity, until 1968. Elk population then rapidly increased.

The elk browsed aspen trees so heavily that no young aspens could get a start.

Concept 45.3 Trophic Interactions Determine HowEnergy and Materials Move through Communities

Elk also browsed streamside willows to the point that beavers (who depend on willows for food) were nearly exterminated.

Wolves were reintroduced in 1995 and preyed primarily on elk.

Aspen and willows grew again, and the beaver population increased.

Figure 45.8 Removing Wolves Initiated a Trophic Cascade (Part 1)

Figure 45.8 Removing Wolves Initiated a Trophic Cascade (Part 2)

Figure 45.8 Removing Wolves Initiated a Trophic Cascade (Part 3)

Concept 45.4 Species Diversity Affects Community Function

Species diversity has two components:

Species richness—the number of species in the community.

Species evenness—the distribution of species’ abundances

Figure 45.9 Species Richness and Species Evenness Contribute to Diversity

Concept 45.4 Species Diversity Affects Community Function

Both aspects of diversity affect community function.

A species’ influence in a community depends on its interactions, and also its abundance.

Communities with a few very abundant species are largely defined by them, rather than by the many rare ones.

Concept 45.4 Species Diversity Affects Community Function

Communities can be thought of as systems with inputs and outputs.

Important measures of community function are the total flow of energy into the community (GPP), and net energy available for consumption by heterotrophs (NPP).

Concept 45.4 Species Diversity Affects Community Function

Community outputs vary with species diversity.

Within a community type, NPP is generally greater and more stable as species diversity increases.

A long-term study of prairie plant communities found that above-ground biomass increased as species diversity increased.

Figure 45.10 Species and Functional Group Diversity Affect Grassland Productivity (Part 1)

Concept 45.4 Species Diversity Affects Community Function

Possible reasons that species diversity affects community function:

• Sampling: communities with more species are more likely to have some with a strong influence on community output.

• Niche complementarity: communities with more species may be better able to use all available resources.

Concept 45.4 Species Diversity Affects Community Function

In the prairie plant experiment, the most species-rich plots also had the most plant functional groups:

Plant groups differing in traits such as ability to grow in warm versus cool seasons, associations with N-fixing bacteria, allocation to growth versus reproduction, etc.

Figure 45.10 Species and Functional Group Diversity Affect Grassland Productivity (Part 2)

Concept 45.5 Diversity Patterns Provide Clues to Determinants of Diversity

Geographic patterns of species richness suggest factors that affect diversity.

Early explorer–naturalists noticed that species richness varies with latitude.

Greatest diversity of many plant and animal groups occurs in the tropics.

Figure 45.11 Species Richness Increases toward the Equator

Concept 45.5 Diversity Patterns Provide Clues to Determinants of Diversity

Why the tropics support more species is an unresolved question.

Possibilities:

• Climatic conditions have been stable and not affected by the glacial cycles that caused massive shifts in geographic ranges in temperate regions.

Concept 45.5 Diversity Patterns Provide Clues to Determinants of Diversity

• Tropics have abundant solar energy and high productivity. Greater energy flow through communities could facilitate coexistence of more species with narrow, specialized niches.

• Environmental heterogeneity: in general, diversity is higher in more structurally complex habitats.

Figure 45.12 Structurally Complex Habitats Support Greater Diversity

Concept 45.5 Diversity Patterns Provide Clues to Determinants of Diversity

Oceanic islands have fewer species than areas of comparable size on nearby mainlands.

Small islands contain fewer species than large islands, and isolated islands contain fewer species than comparable-size islands closer to a mainland.

These patterns could not be explained by productivity, habitat heterogeneity, or disturbance rate.

Figure 45.13 Area and Isolation Influence Species Richness on Islands (Part 1)

Figure 45.13 Area and Isolation Influence Species Richness on Islands (Part 2)

Concept 45.5 Diversity Patterns Provide Clues to Determinants of Diversity

In 1963, MacArthur and Wilson formulated the theory of island biogeography:

Equilibrium species richness on islands depends on relative rates of colonization and extinction.

Small islands—small populations sizes, greater likelihood of extinctions

More isolated islands (farther from a colonizing source)—less likely that colonizers will reach it

Figure 45.14 MacArthur and Wilson’s Theory of Island Biogeography

Concept 45.5 Diversity Patterns Provide Clues to Determinants of Diversity

The theory of island biogeography has been tested in many natural communities and has been one of the most successful explanatory theories in ecology.

Figure 45.15 The Theory of Island Biogeography Can Be Tested (Part 1)

Figure 45.15 The Theory of Island Biogeography Can Be Tested (Part 2)

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

Ecological communities provide humans with critical goods and services, which depend on community diversity.

These ecosystem services include food, clean water, clean air, fiber, building materials, fuel, flood control, soil stabilization, pollination, and climate regulation.

Table 45.2 Some Major Ecosystem Goods and Services

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

The role of properly functioning communities in providing ecosystem services is often taken for granted.

Example: European settlers in Australia brought cattle. Native dung beetles were adapted to dry, fibrous dung of marsupials and ignored the wet dung produced by cattle.

Cattle dung piled up, pastures lost productivity (no recycling of nutrients) and populations of flies that lay eggs in dung exploded.

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

Ecosystem services have economic value.

Example: In the U.S. wild native pollinating insects contribute $3 billion annually to crop production.

Some services, such as greenhouse gas regulation, are more difficult to place a value on, but are very valuable.

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

Faced with the need to improve drinking water supplies, New York City considered a new water treatment facility that would cost $6–$8 billion to build and $300 million annually to run.

Instead, they invested $1.5 billion in land protection and better sewage treatment in the Catskills where the water reservoirs are located.

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

Humans are rapidly converting natural communities into less diverse, human-managed communities such as croplands, pastures, and urban areas.

Large areas of habitat are being fragmented. The fragments can be seen as habitat “islands” surrounded by “seas” of human-modified habitat.

Figure 45.16 Habitat Fragmentation in Tropical Forests

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

Habitat fragmentation causes loss of species:

• Total amount of habitat decreases, average patch size decreases, and patches become more isolated from one another.

• Populations become smaller and more prone to extinction.

• The human-modified habitat may be a barrier to dispersal, reducing colonization.

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

The theory of island biogeography suggests ways to minimize effects of fragmentation.

Enhance colonization: cluster habitat fragments together, connect fragments with dispersal corridors

Reduce extinctions: retain large patches of original habitat, maintain ability of the fragments to support healthy populations.

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

In a large-scale experiment in Brazil, land owners agreed to preserve forest patches laid out by biologists.

The patches were surveyed before and after forest cutting.

Species began to disappear: monkeys that travel over large areas of forest; army ants and the birds that follow them.

Small, isolated patches lost species most rapidly.

Figure 45.17 A Large-Scale Study of Habitat Fragmentation

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

Conservation efforts often target species that have important roles in community structure and function.

Example: the wolves in Yellowstone National Park are critical in maintaining healthy aspen forests and watersheds via trophic cascades.

The Yellowstone to Yukon Conservation Initiative aims to maintain a continuous corridor of wolf habitat between Yellowstone and similar areas to the north.

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

Restoration ecology focuses on restoring function to degraded ecosystems.

One goal is to restore original species diversity, drawing on our knowledge of the factors that shape diversity.

Figure 45.18 Species Richness Can Enhance Wetland Restoration (Part 1)

Figure 45.18 Species Richness Can Enhance Wetland Restoration (Part 2)

Concept 45.6 Community Ecology Suggests Strategiesfor Conserving Community Function

Disturbance sometimes results in an ecological transition to a very different community.

It may be very difficult to reverse the transition and restore the original community.

Answer to Opening Question

Coffee cultivation can be improved by using principles of community ecology:

• Increase diversity by growing crops together in functional groups

• Attract wild bee pollinators by planting coffee close to intact forest patches and leave flowering weeds in place

Figure 45.19 Traditional Coffee Cultivation and Community Diversity

Answer to Opening Question

Traditional low-intensity cultivation may yield less per acre, but are profitable because they avoid costs of chemicals and labor.

It also avoids pollution and helps maintain natural communities and their species.