Focuses on the variety of species within a community.
The trophic structure of a community is a hierarchy of trophic levels.
These levels are defined by the feeding relationships among species.
First trophic level (primary producers or autotrophs):
Plants and photosynthetic organisms capture sunlight and convert it to chemical energy.
Animals are consumers (heterotrophs):
They acquire energy and nutrients by consuming other organisms or their remains.
Second trophic level (primary consumers):
Herbivores that eat plants.
Third trophic level (secondary consumers):
Carnivores that eat herbivores.
Fourth trophic level (tertiary consumers):
Carnivores that eat other carnivores.
Omnivores:
Organisms like humans and some bears feed at multiple trophic levels simultaneously.
Scavengers (detritivores):
Animals that ingest dead organisms, digestive wastes, and cast-off body parts (e.g., earthworms and vultures).
Decomposers:
Small organisms like bacteria and fungi that feed on dead or dying organic material.
Ecological function:
Detritivores and decomposers reduce organic material to inorganic molecules, which producers can assimilate.
Food chain:
The trophic structure where one organism eats another.
Each link points from the food to the consumer.
Straight-line food chains:
Rare in nature.
Food web:
A set of interconnected food chains with multiple links, portraying complex relationships.
Links between trophic levels contribute to community stability when environmental disturbances eliminate some species.
In species-rich communities, losing one or two species has minor effects on overall community stability.
The proportions of species at high, middle, and low trophic levels are reasonably constant across communities.
Regardless of species richness, a community includes two to three prey species for every predator species.
Interspecific Competition:
Can cause local extinction or prevent new species from establishing, decreasing species richness.
More studies focus on competition in K-selected species than in r-selected species, which may overestimate the importance of competition.
In communities where resource partitioning and character displacement occur due to past competition, its current importance may be underestimated.
Ecologists' Views:
Some ecologists are undecided about the influence of interspecific competition on species composition and community structure.
Plant and vertebrate ecologists believe competition profoundly affects species distributions and resource use.
Insect and marine ecologists suggest predation, parasitism, and physical disturbance govern community structure.
Predators can increase species richness by stabilizing competitive interactions among prey.
Example: Sea Stars and Mussels:
Predatory sea stars preferentially eat mussels, reducing their numbers.
This allows other species to grow.
When sea stars (keystone species) were removed, mussels outcompeted other species, reducing diversity from 18 to 2 or 3.
Periwinkle Snails (Keystone Species):
Periwinkle snails in Massachusetts graze on the green alga Enteromorpha.
In tidepools:
Enteromorpha outcompetes other algae.
Moderate herbivory allows less competitive species to grow, increasing species richness.
On high rocks:
The red alga Chondrus is dominant.
Periwinkles feeding on Enteromorpha reduce algal species richness.
Great Barrier Reef Study:
From 1963 to 1992, researchers tracked the effects of five major cyclones.
Changes resulted from external disturbances removing coral colonies and internal processes (growth and recruitment).
The community never reaches equilibrium due to slow growth and recruitment, and frequent disturbances.
Species richness is greatest in communities with fairly frequent disturbances of moderate intensity.
This allows K-selected species to survive while creating openings for r-selected species to colonize.
Severe and frequent disturbances:
Communities include only r-selected species with fast life cycles.
Mild and rare disturbances:
Communities are dominated by long-lived K-selected species.
Ecological succession:
A series of changes in species composition in response to disturbance.
Primary succession:
Begins when organisms colonize terrestrial habitats without soil (e.g., after volcanic eruptions or glacier retreat).
Lichens are usually the first visible colonizers.
They erode rock, initiate soil development, and produce organic material.
Soil Accumulation:
As soil accumulates, r-selected plants colonize the site.
Their decaying remains enrich the soil (facilitated by detritivores and decomposers).
Development:
As soil gets deeper and richer, bushes and eventually trees are supported.
Climax Community:
A relatively stable, late successional stage (climax community) is often dominated by long-lived K-selected species.
It persists until an environmental disturbance eliminates it, allowing other species to invade.
Glaciers retreat, leaching minerals (especially nitrogen) from the newly exposed substrate.
Lichens and mosses establish.
Mountain avens (Dryas) grow on the nutrient-poor soil, benefiting from nitrogen-fixing bacteria.
Shrubby willows (Salix), cottonwoods (Populus), and alders (Alnus) take hold in drainage channels, also symbiotic with nitrogen-fixing microorganisms.
Young conifers (hemlocks - Tsuga, and spruce - Picea) join the community.
After 80 to 100 years, dense forests of Sitka spruce (Picea sichensis) and western hemlock (Tsuga heterophylla) crowd out other species.
Occurs after existing vegetation is destroyed or disrupted (e.g., by fire, storm, or human activity).
Early stages proceed rapidly because the soil is ready for colonization and may contain numerous seeds.
Later stages parallel those of primary succession.
Similar climax communities sometimes arise from alternative successional sequences.
Example: Pond to Hardwood Forest (Aquatic Succession):
Debris from rivers and runoff accumulates, transforming a pond into a swamp.
Transpiration by larger plants dries the soil, allowing other species to colonize.
Eventually, the swamp may become a meadow or forest with moist, low-lying ground.
Facilitation Hypothesis:
Species modify the local environment to make it less suitable for themselves but more suitable for the next successional stage.
Succession is orderly and predictable.
Inhibition Hypothesis:
New species are prevented from occupying a community by existing species.
Succession is neither orderly nor predictable.
Tolerance Hypothesis:
Succession proceeds because competitively superior species replace competitively inferior ones.
Early-stage species neither facilitate nor inhibit later-stage species.
Competition eliminates species that cannot harvest scarce resources.
Succession results from a combination of facilitation, inhibition, and tolerance with differences in dispersal, growth, and maturation rates.
Disturbance and density-independent factors play important roles, sometimes speeding up successional change.
In other cases, disturbance inhibits successional change, establishing a disturbance climax (disclimax community).
Species richness follows a latitudinal gradient for many plant and animal groups, with the most species in the tropics and a decline toward the poles.
Hypotheses for high tropical species richness:
High reproductive rates.
Low migration.
Few environmental disturbances.
Hypotheses for maintaining high tropical species richness:
High availability of energy and other resources.
The equilibrium theory of island biogeography addresses variations in species richness on islands of different sizes and levels of isolation.
The number of species on an island is determined by the equilibrium between immigration and extinction.
The mainland forms a species pool from which species immigrate to offshore islands as new colonizers.
As the number of species on an island increases, the extinction rate rises due to competition and predator-prey interactions.
At equilibrium, large islands have more species than small islands.
Reasons:
Large islands have higher immigration rates (larger target).
Large islands have lower extinction rates (support larger populations, greater range of habitats and resources).
At equilibrium, islands closer to a mainland source have more species than more distant islands.
Reasons:
Islands near the mainland have higher immigration rates (easier to locate).
Distance does not affect extinction rates.