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Chapter 18, 19, 20, 21, 22, 23, 24, Exam Review
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Ecological Succession
Process of change in the species composition of an ecological community over time
What happens following a disturbance or the initial colonisation of a new habitat
Henry Chandler Cowles
Studied vegetation development on sand dunes on the shores of Lake Michigan (Indiana Dunes)
Recognised that vegetation on dunes of different ages might be interpreted as different stages of a general trend of vegetation development on dunes
Frederic Clements
Was raised in the prairies
Had a large-scale view of communities, leading to the idea of an organismal concept of communities
This concept dominated community ecology until the mid-20th century
Organismal Community Concept
Clements regarded a community as a closely integrated holistic entity of mutually interdependent organisms and relatively discrete borders
Succession has a predictable end-point, a climax community
Is a stable condition or equilibrium
Henry Gleason
Was raised at the prairie-forest border, which cannot be fully described by a single climax community (non-equilibrium)
Developed the individualistic (continuum) concept of communities
This concept dominated community ecology during the second half of the 20th century
Clement’s Predicted
Discrete community types with sharp ecotones
Gleason’s Suggested
There is continuous variation in communities
Robert Whittaker Findings
Species show individualistic distribution patterns along gradients of environmental factors, resulting in a continuous change of community composition
This supports Gleason’s view
Modern Synthesis (of Theory)
Gleason’s view is more widespread among scientists but Clements’ aspects of a community are frequently accepted
E.g. dominant tree species influence a lot of other organisms
Discrete community units are pragmatic for applied disciplines such as forestry
Effects of Disturbance
Leads to changes in community structure and composition over time
Examples: volcanoes, earthquakes, storms, fires, and climate change
Intermediate Disturbance Hypothesis
Species richness is the highest at intermediate levels of disturbance
At High Levels of Disturbance
Many species fail to get established (non-equilibrium)
At Low Disturbance Levels
Competitively superior species suppress the others species
Premises Base for The Intermediate Disturbance Hypothesis
Ecological disturbances have major effects on species richness within the area of disturbance
Interspecific competition results from one species driving a competitor to extinction and becoming dominant in the ecosystem
Moderate ecological scale disturbances prevent interspecific competition
Intermediate Disturbance Hypothesis Example
Rabbit grazing →
Mean number of plants per plot in sand dunes is the highest at intermediate levels of rabbit grazing
Communities in Early Succession
Are dominated by fast-growing, well-dispersed species (opportunist, fugitive, or r-selected life-histories)
As succession proceeds, these species will be replaced by more competitive (k-selected) species
Patterns of Diversity - Succession
Formerly seen as having a stable end-stage called the climax, shaped primarily by the local climate
This idea has been largely abandoned by modern ecologists in favour of non-equilibrium ideas of ecosystems dynamics
Secondary Succession Example
Bare rock
Mosses/grasses
Grasses/perennials
Woody pioneers
Fast growing trees
Climax forest
Primary Succession Characteristics
Begins on rock formations
Lichen, algae, fungi develop soil
These are replaced by plants, like grasses and shrubs
Water and nutrient levels increase over time
Nitrogen-producing species play a large role
N is limited
Secondary Succession Characteristics
Follows disturbance (fire, flood) or removal of an existing community (logging)
Strongly influenced by pre-disturbance conditions (e.g. soil, and seed banks)
Can be relatively rapid
Species that dominate are usually present from the start
Community Composition
Some species are naturally rare/abundant
Some communities are dominated by one or a few species, some are species-rich
How Biodiversity is Measured
Alpha diversity (α-diversity)
Beta diversity (β-diversity)
Gamma diversity (γ-diversity)
Alpha Diversity (α-Diversity)
Mean species diversity in a site at a local scale
Number of species
Beta Diversity (β-diversity)
Ratio between regional and local species diversity (gamma diversity divided by alpha diversity)
Rate at which species composition changes across a region
Overemphasises the role of rare species
Example: high diversity in matrices of recently disturbed land
Gamma Diversity (γ-Diversity)
Total species diversity in a landscape
Scale is important to distinguish between alpha and gamma diversity
Autogenic Succession
Brought about by changes in the soil caused by the organisms there
Includes accumulation of organic matter in litter or humic layer, alteration of soil nutrients, or change in the pH of soil due to the plants growing there
Structure of the plants can also alter the community
Autogenic Succession Example
When larger species like trees mature, they produce shade on the forest floor that excludes light-requiring species → as a result, shade-tolerant species will invade the area
Allogenic Succession
Caused by external environmental influences and not by the vegetation
Animals play an important role as they are pollinators, seed dispersers, and herbivores
Allogenic Succession Example
Soil changes due to erosion, leaching, or the deposition of silt and clays. This can alter the nutrient content and water relationships in the ecosystem
(Main) Models of Successional Change
Facilitation
Tolerance
Inhibition
Facilitation Model
Presence of an initial species aids and increases the probability of growth of a second species
Facilitation Model Example
Presence of Alder plants aids the growth of Willow and Poplar seedlings in an Alaskan floodplain by increasing the availability of nitrogen
Alder roots contain nitrogen-fixing bacteria (which greatly increase inorganic nitrogen in soils)
Tolerance Model
The climax community is composed of the most “tolerant” species that can co-exist with other species in a more densely populated area
Eventually, dominant species replace or reduce pioneer species abundance through competition
Tolerance Model Example
Forest succession with survival in conditions of shade
Species better adapted to shady conditions will become dominant
Inhibition Model
Earlier successional species inhibit growth of later successional species and reduce growth of colonising species already present
The environment is thus less hospitable to other potential colonising species
Inhibition Model Example
Lantana (Lantana camara) sprawling shrubs exclude and inhibit the growth of certain tree species
Succession in Consumers
Initially, fauna is sparse (few mites, ants, and spiders living in cracks and crevices)
The animal population increases and diversifies with the development of the forest climax community (nematodes, insect larvae, ants, etc.)
Vertebrates enter the scene, such as squirrels, foxes, mice, etc.
Alternative Stable States (Questions)
Does succession under given environment always lead to the same climax community?
Does the final stage depend on the identity of colonising species?
Are there different stable states for a community?
Alternative Stable States - Normal Successional Trajectory
There is a predictable way a “ball” will fall
Alternative Stable States - Disturbance
The input of disturbance can push the “ball” back up the hill after it’s fallen
Alternative Stable States - Redirection
The redirection of energy can lead to an alternative stable state
Succession in Mixed-Hardwood Forest
When moose browsing is added to the equation, we would expect one of two possible outcomes:
If moose browse all species equally and they are sufficiently abundant, they may actually delay succession, keeping the forest in a more-open, shorter state
If moose brows preferentially, avoiding species they dislike (such as Spruce), they may redirect the trajectory of the forest. Mature forest which eventually takes hold will be dominated by these browse-resistant tree species
Energy
Is required by most complex metabolic pathways (often in the form of adenosine triphosphate, ATP)
Especially those responsible for building large molecules from smaller compounds, and life itself is an energy-driven process
Living organisms would not be able to assemble macromolecules from their monomeric subunits without a constant energy input
Food Chains
Linear representation of feeding interactions
Only a part of the energy content of the food can be absorbed and utilised
Do not accurately describe most ecosystems
Food Chain Example
Aphid eats plants
Ladybugs eats aphid
Food Web(s)
Representation of all feeding interactions
It expresses connections among the food chains in an ecosystem
Three basic ways in which organisms get food are:
Producers (Autotrophs)
Consumers (Heterotrophs)
Decomposers (Detritivores)
Producers (Photoautorophs)
Are responsible for almost all productivity of the biosphere
Use sunlight as an energy source
Chlorophyll and other pigments capture photons
Photoautotrophs
Plants, algae, and photosynthetic bacteria
Serve as the energy source for a majority of the world’s ecosystems
Producers (Chemoautotrophs)
Specialised bacteria that get their energy from inorganic chemicals (e.g., by oxidising sulphide minerals)
Energy released by these exothermic reactions is used to drive biosynthesis of CO2 and H2O to form glucose
Hydrothermal Vent Ecosystem
Supported by chemoautotrophic bacteria and organic material that sinks from the ocean’s surface
Deep-Sea Vents
Species in deep-sea ecosystems have adapted to interact with each other in many ways - symbiosis between many species and chemosynthetic bacteria
Whale Falls
Are an important influx of nutrients to the sun-starved deep ocean
Consumers (Heterotrophs)
Consume rather than produce biomass energy as they metabolise, grow, and add to levels of secondary production
There are different kinds of feeding relations: herbivory, carnivory, scavenging, and parasitism
The Ghost Plant (Monotropa uniflora)
NOT an autotroph, is a parasitic mycoheterotroph and gets its energy from the surrounding trees, via a mycorrhizal fungus
Lacks chlorophyll
Occur in dark forests, where there isn’t enough light for photosynthesis
Decomposers - Fungi
Heterotrophs that break down dead or decaying organisms
Carry out decomposition, a process possible by only certain kingdoms, such as fungi
Fungi
Are the primary decomposers in most environments
Decomposers - Detritivores
Heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces)
Ingest and digest dead matter internally
Decomposers - General
Directly absorb nutrients through external chemical and biological processes
Decomposers - Scavengers
Heterotrophs that consume dead organisms that have died from causes other than predation or have been killed by other predators
Scavenging
Generally, refers to carnivores feeding on carrion, but is also a herbivorous feeding behaviour
Positions in a Food Web
Producers
Primary consumers
Secondary consumers
Tertiary consumers
Apex predators
Trophic Levels - Producers
Plants and algae thats make their own food
Level 1
Trophic Levels - Primary Consumers
Herbivores that eat plants
Level 2
Trophic Levels - Secondary Consumers
Carnivores that eat herbivores
Level 3
Trophic Levels - Tertiary Consumers
Carnivores that eat other carnivores
Level 4
Trophic Levels - Apex Predators
Have no predators and are at the top of their food web
Level 4
Second Trophic Level Example
Rabbits eat plants at the first trophic level, so they are primary consumers
Third Trophic Level Example
Foxes eat rabbits at the second trophic level, so they are secondary consumers
Fourth Trophic Level Example
Golden eagles eat foxes at the third trophic level, so they are tertiary consumers
Decomposer Example
The fungi on trees feed on dead matter, converting it back to nutrients that primary producers can use
Ecological Pyramids
Are created due to inefficiency of energy transfers, the productivity declines with increasing trophic level
If producers are productive but short-lived, their biomass may be less than that of the primary consumers
Are not necessarily upright
Kinds of Ecological Trophic Pyramids
Pyramid of numbers
Pyramid of biomass
Pyramid of energy
Ecological Pyramid Example - Aquatic Ecosystem
Sea lion (Top)
Herring
Zooplankton
Phytoplankton (Bottom)
Ecological Pyramid Example - Terrestrial Ecosystem
Snakes (Top)
Mice
Grasshoppers
Grasses (Bottom)
Transfer Efficiency
Proportion of energy transferred from one trophic level to the next
Lindeman’s 10% Law of Transfer of Energy
Was not originally called a “law”, and was also cited ranging from 0.1% to 37.5%
10% of the transferred energy is stored as flesh
Remaining is lost during transfer (broken down in respiration, or lost to incomplete digestion by higher trophic level)
Food Chain Length
Way of describing food webs as a measure of the number of species encountered as energy or nutrients move from the plants to top predators
Mean Chain Length
Entire web is the arithmetic average of the lengths of all chains in a food web
Food Chain Length Example
Simple predator-prey example
Deer is one step removed from the plants it eats (chain length = 1)
Wolf that eats the deer is 2 steps removed from the plants (chain length = 2)
Connectance
Fraction of all possible links that are realised in a network
C = L / [S(S - 1)/2]
L is the number of trophic links
S(S - 1)/2 is the maximum number of binary connections among S species
Mathematical Models on Networks
Are used to determine structure, stability, and laws of food web behaviours relative to observable outcomes
Trophic Cascades
Powerful indirect interactions that can control entire ecosystems
Occur when a trophic level in a food web is suppressed
Next lower trophic level is released from predation (or herbivory if the intermediate trophic level is a herbivore)
Trophic Cascade Example
A top-down cascade will occur if predators are effective enough in predation to reduce the abundance, or alter the behaviour of their prey
Top-Down Cascade Example
Otters eat sea urchins but have declined due to overhunting and disease
Sea urchins eat kelp holdfasts, leading to a reduction in this ecosystem engineer
Low sea otters → lead to sea urchins eating more kelp
Pacific Kelp Forest Trophic Environment
Sea otters feed on sea urchins
Sea otters have been hunted to extinction →
Sea urchins increase in abundance →
Kelp populations are then reduced
Yellowstone Park Trophic Environment
Reintroduction of wolves (Canis lupus)
Reduced the number, and changed the behaviour of elk (Cervus canadensis)
Freed several plant species from grazing pressure
Led to the transformation of riparian ecosystems
Nutrients
Substances necessary for the healthy physiology of organisms
Must be sufficiently available
The most limiting element that determines growth
Macronutrients
Are required in larger quantities
Carbon (C), Oxygen (O), and Hydrogen (H); form the backbone of most organic molecules
Photosynthesis
How plants get Carbon, Oxygen, and Hydrogen
Mineral Nutrients
Elements from the Earth and foods that organisms need to develop and function normally
Examples: Nitrogen (N), Phosphorus (P), Calcium (Ca), Magnesium (Mg), Potassium (K), and Sulphur (S)
General Characteristics of Cycles
Energy flows directionally through ecosystems. Entering as sunlight (or inorganic compounds) and leaving as heat during the many transfers between trophic levels
Matter that makes up living organisms is conserved and recycled
Most Common Elements Associated with Organic Molecules
Carbon (C), Nitrogen (N), Hydrogen (H), Oxygen (O), Phosphorus (P), and Sulphur (S)
Can take various chemical forms, and may exist for long periods in the atmosphere, on land, in water, or beneath the Earth’s surface
Biogeochemical Cycles
The recycling of inorganic matter between living organisms and their environment
Anthropogenic
Are human activities that alter all major ecosystems and the biogeochemical cycles they drive
Hydrosphere
Where water movement and storage occurs
Important for leaching certain components of organic matter into rivers, lakes, and oceans, and is a reservoir for carbon
Reservoir
Place where a certain kind of material is stored, or resides, for some period of time
Place where material moves into and out of
Exchange Pool/Pool
Holds an element or water for a short period of time
Exchange Pool Examples
Atmosphere is an exchange pool for water - holds water (water vapour) for just a few days
Glaciers; the soil layer; the aggregate of bodies of fresh water on the continents (rivers and lakes)
Flux
Rate at which a given material moves between reservoirs
Steady State
If the flux of material into and out of a given reservoir is the same for some period of time
Commonly, flux in and the flux out are not equal