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Lectures 5 - 8: Decomposers & Nutrient Cycling, Niche & Competition, Biodiversity, Climate & Soil
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Scavengers
consume carrion
animals that eat animals that they don’t kill
usually; some scavengers “finish off” animals that are close to death
Carrion
easily digestible biomatter of dead animals
Detritovores
consume detritus
animals that eat what other things (scavengers) have left behind
Detritus
organic debris
Decomposers
consume what’s left
microscopic and very hard to digest organic material
bacteria and fungi breaking down organic matter into inorganic matter and harvesting last iota of energy
Decomposers, detritovores, and scavengers
consumers
harvest last remnants of energy
allow matter to be recycled
prey and hosts
occasionally predator and parasite
Where is the line?
scale
scavengers — consume large pieces of animal
ingestion
detritivores — consume medium-ish pieces of animal
ingestion
decomposers — consume organic molecules
secrete enzymes externally
does it matter?
not usually
they perform the same functions
many species fit multiple categories
Stoichiometry
Efficiency and residence time are affected by nutrient balance
Nutrient balance varies by species and life stage
Diatoms need lots of Si
Vertebrates need lots of Ca and P
Faster growing organisms usually need more P
If don’t obtain correct balance
Consume more -> reduces efficiency
Consume different things
Grow slower
Matter Cycling
Cosmic inputs of matter are negligible … now
Loss of matter is negligible
Matter on Earth can be treated as if in closed cycles
Each substance has own cycle
Elements of life: CHNOPS
Important parts of most cycles
Photosynthesis
Respiration
Decomposition
Water Cycling
water evaporates
falls as precipitation
flows downhill
Anthropogenic impacts on hydrologic cycle
impervious surfaces → more runoff, less filtration
remove vegetation → more runoff
removal of groundwater → reduced terrestrial water, more water cycling
climate change → increased evaporation → increased precipitation
Carbon cycling
photosynthesis removes CO2 from the atmosphere and incorporates it into organic matter
respiration releases C into the atmosphere as CO2
non-respired C can be buried
buried C is released through weathering and volcanism
atmospheric CO2 diffuses into water
CO2 + H2O ←→ H2CO3 ←→H+ + HCO3- ←→ 2H+ + CO32-
CO32- combines with Ca and is buried
dissolved CO2 diffuses into the atmosphere
Anthropogenic impacts on the carbon cycle
extraction and combustion of buried C (fossil fuels)
extraction and processing of buried CaCO3 (concrete)
Nitrogen Cyling
nitrogen fixation = conversion of N2 into forms primary producers can use
requires a lot of energy
NH3 → NH4+ — via bacteria
NO3 — via lightning and wildfire
nitrification = NH4+ → NO2- → NO3-
releases energy
assimilation = NO3- incorporated into organic matter
mineralization = organic matter → inorganic compounds
releases N from organic matter
N can then be assimilated
denitrification = biologically usable N converted into N2
NO3- → NO2- → NO
NO → N2O → N2
Anthropogenic impacts on the Nitrogen cycle
NO produced from combustion of fossil fuels
NO + O2 → NO3-
Planting crops that promote nitrogen fixing
crop rotation
direct production of nitrogen-containing fertilizers
Phosphorous cycling
PO43- containing rocks are weathered
assimilation of PO43- by primary producers
mineralization of organic matter
unbound PO43- carried by water
unbound PO43- combines with Ca or Fe
bound P precipitates and becomes part of sediment
is buried
Anthropogenic impacts on phosphorous cycle
mine PO43- containing rocks to use as fertilizer
PO43- detergents
use is now banned in many countries
Terrestrial nutrient cycling
nutrient cycling mostly occurs in soil (also removed in soil by primary producers)
decomposition of plant material
water leaches minerals and small organic compounds
detritivores consume organic matter
bacteria and fungi break down organic molecules
nutrients are leached from soil
nitrogen fixing and weathering replenishes leached nutrients
Terrestrial decomposition
aerobic
occurs near primary producers
affected by:
temperature
water
nutrient balance
varies
Aquatic nutrient regeneration
nutrient recycling mostly occurs in sediment (and removed all throughout the water column — especially at the surface)
decomposition of plant material
same process as in terrestrial systems
different detritivores
bacteria and fungi are still decomposers
decomposition of phytoplankton
much faster than plant decomposition (plants are hard to break down per primary consumers lectures)
mostly in sediments after dead cells have sunk
some occur in water column
Aquatic decomposition
anaerobic
slow
far from primary consumers
occurs in sediment
deeper water systems are less productive
affected by
temperature
nutrient balance
varies
phytoplankton decomposition
so fast it can deplete dissolved oxygen in water column
Eutrophication
excess production of organic carbon
too much primary production
nutrient pollution = contamination of excess nutrient inputs
eutrophication can occur naturally
anthropogenic/cultural eutrophication is widespread
decreased water quality
hypoxia (low dissolved oxygen) / anoxia (no dissolved oxygen)
Hypoxia/anoxia
hypoxia = reduced O2
anoxia = no O2
occurs naturally in most aquatic sediments
rare in water column
“jubilee” in Alabama
“dead zones”
mass mortality or exodus of aquatic fauna due to low dissolved oxygen
Stable isotopes in ecological research
≥ 80 elements have multiple stable isotopes
Tin (Sn) with 10 and xenon with 9 have the most
≥ 19 elements have just 1 stable isotope
CHNOS multiple stable isotopes
12C, 13C, 1H, 2H, 14N, 15N, 16O, 17O, 18O, 32S, 33S, 34S, 36S
heavier isotopes react slower and form more stable covalent bonds
light isotopes concentrate in products and gases
heavier isotopes concentrate in reactants and solids/liquids
all of this is generally speaking, specific reactions and conditions vary
Research uses of stable isotopes
photosynthesis
enriched heavy C in C4 photosynthesis than C3
enriched heavy H in CAM compared to C3 & C4
source tracking
15N becomes enriched as you move to higher trophic levels
sources:
marine has more heavy N
terrestrial has more light N
anthropogenic sources are rich in heavy N
different geographical regions have distinct isotopic baselines
Radioisotopes in ecological research
all elements have radioactive isotopes
spike a source with radioisotopes and track where decay is occurring
examples
14C to measure photosynthesis
204Hg to measure biomagnification
32P to track nutrient uptake and phosphorous cycling
35S to track sulfate reduction
The niche concept pre-1800
religion, philosophy, naturalism
each organism has a place within a harmonious system
diet, habitat, behavior, distribution
“natural beings are complementary and tend to a common purpose” - A. Pocherville
The niche concept 1800-1900
interdependency of environment and organisms
biotic relationships
Darwin: organisms have a place in nature to which they are adapted by natural selection
Niche as a place
“a recess in a wall especially for a statue” - Merriam-Webster
Roswell Johnson 1910 and Joseph Grinnell 1913
a place occupied by a species in the environment
everything that conditions the existence of a species to a location
abiotic
biotic
vacant niches can exist
Niche as a job
Charles Elton 1927
“the ‘niche’ of an animal means its place in the biotic environment, its relations to food and enemies
based on location within food chains that combine to make a food cycle
location on trophic pyramid
different organisms in different systems can occupy similar niches
Niche as a hypervolume
G. Evelyn Hutchinson 1957
niche is a space of environmental variables representing the limits of species viability
multidimensional space (hypervolume) that determines “where” a species can exist
niche is an attribute of the species
niche can be quantified
fundamental niche = hypervolume niche that permits the species to exist
realized niche = region of the fundamental niche from which the species is not excluded by predators
Hypervolumes
regions in more than 3 dimensions
line is 1D
square is 2D
cube is 3D
hypervolume is 4D+
hypervolume coordinate systems
each axis is a separate variable
temperature
pH
moisture
PO43-
NO3-
Niche now
based on resources, abiotic conditions, and biotic interactions
fundamental niche = range of abiotic conditions under which a species can persist
biotic interactions and limiting factors are not considered
facilitation?
realized niche = actual conditions a species lives under
all environmental and biotic factors are considered
Facilitation
positive, non-harmful interaction in which one species improves the fitness of the other
mutualism = all species benefit
commensalism = at least one of the involved species is not impacted
can work through
direct interaction
indirect interaction
abiotic
biotic
Facilitation and niche
facilitation may counteract negative biotic interactions
refuge from predation
removal of parasites
facilitation may allow persistence outside of non-facilitated niche space
expansion of fundamental niche
OR extension of realized niche beyond fundamental niche
examples:
nurse plants
macroalgae in rocky intertidal habitats
Range of tolerance
“better” conditions are needed for reproduction than growth
“better” conditions are needed for growth than survival
range of reproduction heavily affects occurrence of species
individuals can disperse to areas with limited reproductive potential
Environmental gradient
populations will not be found where the species cannot survive
population size will be limited in regions of physiological stress
population size will be highest in regions where conditions are optimum
Example of gradient on a small scale
grass species:
too close to a lake/pond → too much water for species to survive
medium distance from the water → species can survive, grow, and reproduce
too far from the lake/pond → not enough water for species to survive
Interspecific competition
species A sequesters a resource before species B
species A and species B are both negatively affected
but species B is more negatively affected
species A may
be better able to sequester that resource
need less of the resource
take resource from species B
competitive exclusion = a species is excluded from a habitat due to competition with another species
Niche and competition
competition occurs where niches overlap
competition prevents utilization of full niche hypervolume
Coexistence of competitors: resolving the plankton paradox
environment is neither isotropic nor unstructured
fluid motion
chaotic
wind-driven
light attenuation
changing nutrient concentrations
daily, seasonal, annual variation
grazing/predation
selective by shifting zooplankton assemblages
viruses
environment never equilibrium
natural selection favors reduced competition
Environment and competition
abiotic factors influence competitive outcomes
adaptations to abiotic factors determine “winner”
niche partitioning
Natural selection and competition
natural selection leads to minimization of competition
competitive exclusion
niche partitioning
specialization
character displacement
Natural selection
direction selection: selection favors individuals showing an extreme phenotype (underdominance)
disruptive selection: selection favors individuals at both extremes of a phenotype (underdominance)
stabilizing selection: selection favors individuals with an intermediate phenotype (overdominance)
Competitive separation
allopatric = geographic separation
sympatric = without geographic separation
character displacement
Ecological guilds and functional groups
species with similar niches
through using similar resources = ecological guild
through feeding in similar manners = functional group
same guilds
spiders and bats
same functional group
mosquito and aphids
same guild and same functional group
barnacles and mussels
Ecological guilds
defined by resource use
foraging/feeding guilds
habitat based guilds
pollinator guilds
often competitors
functional redundancy
Functional groups
defined by method of resource acquisition or ecosystem function
Biodiversity
aka diversity, biological diversity, organismal diversity, ecological diversity
variation within and between organisms and ecosystems
Categories of diversity
genetic
allele ratios
taxomonic — shared ancestry
taxa that share a common ancestor
usually species
community/ecosystem — shared geography
place and time
habitat and abiotic conditions
guild — shared resources
exploitation of shared resources
ex. detritivores
ensemble — shared ancestry, geography, and resources
taxa that share a common ancestor, inhabit the same habitat, and utilize the same resource(s)
ex. herbivorous reef fish
Scales of diversity
inventory diversity — measured diversity
point diversity, α-diversity, γ-diversity, & ε-diversity
differential diversity — comparisons of measured diversity
pattern diversity β-diversity & δ-diversity
alpha (α) — local diversity
within a habitat or ecosystem
beta (β) — comparison between two habitats or ecosystems
comparison of 2 α diversities
gamma (γ) — regional diversity
large scale diversity covering multiple habitats
Point and Pattern Diversity
within/between sample diversity
smallest scale
point diversity = diversity from a single sample
pattern diversity = variation in diversities from samples taken within a single habitat
useful for examining diversity within a habitat
multiple of these samples combined to determine α-diversity
α-diversity
Diversity of a defined assemblage, habitat, or system
Geographic scale determined by researcher
Diversity from a set of samples
From the same habitat
Expressed as
Species richness
Species evenness
Diversity index
β-diversity
Comparison of α-diversity between 2 habitats or systems
Or between 2 time periods
Species change or replacement
Species turnover
Habitats/systems being compared must be measured in the same scale
Scale must be appropriate for the species
Expressed as
Ratios
β = γ / α
α species richness / mean α species richness
β species richness = number of unique species
γ-diversity
Diversity of a large region encompassing several habitats and systems
Landscape
Geographic scale determined by researcher
Combination of α-diversities
Same measurement methods as α-diversity
Affected by α & β
Habitats with similar α-diversity but different species will yield high γ-diversity
δ-diversity
Delta diversity
Comparison of γ -diversity between 2 regions
Species change or replacement
Scales must be similar and appropriate
Same methods as β-diversity
ε-diversity
Epsilon diversity
Larger scale than γ-diversity
Continental or large geographic province
Same measurement methods as α & γ –diversities
If you can measure abundances on that scale
In practice usually species richness
Very common to look at Family (instead of species) diversity
Hyperdominant / oligarchs
Occurs when a small subset of species represent most individuals
“Most” = >~50%
γ & ε –diversity only
Does not apply to α-diversity
Demonstrated in tropical forests
More research needed for other habitats and organisms
Community vs assemblage
Community – group of populations living in the same area and interacting with one another
Assemblage – group of species that are taxonomically related and live in the same area
Component of the community
Most biodiversity research focuses on assemblages
Habitat/community boundaries
Zonation – distribution of organisms across an environmental gradient
Arrangement of species in a habitat / across habitats
Species composition along the landscape
Boundaries may be clearly defined or shade from one zone into another
Lake vs forest
Elevation zones as one ascends a mountain
Habitat edges
Changes in population and community structure at the boundary between habitats
Zone that extends into both habitats
Ecotone – sharp change in environmental conditions over a short distance
Large change in species composition
Promotes interaction among different community types
Reduces suitable habitat because part
of the habitat is habitat edge
Species distributions
Why are species found where they are?
Niche
Environmental conditions
Resource availability / primary productivity
Keystone species
Dispersal / immigration
Disturbance
Climate change
Continental drift / plate tectonics
Species-area relationship
Number of species increases area examined increases
Increase area by 10 times, double number of species
Darlington’s rule
S = cAz
S = number of species, A = area
c and z are fitted constants
log(S) = log(c) + z log(A)
log(c) = y-intercept, z = slope
Species and Area
More habitat diversity as area examined increases
Different species adapted to different habitats -> higher β/δ –diversity
Larger areas can fit more and larger species
Equilibrium model of island biogeography
Equilibrium model of island biogeography
Island biogeography + extinction rate
Number of species present is dependent on immigration of new species & local extinction of existing species
Ŝ = IP / (I + E)
Ŝ = equilibrium number of species
I = immigration rate, E = local extinction rate
P = number of species in source pool = maximum number of species
Turnover rate = replacement rate of species
IE / (I + E)
Biodiversity and latitude
More species near equator, fewer near poles
Ecological heterogeneity
Land – solar energy and precipitation
Ocean – water temperature
Ecosystem functions and services
Ecosystem functions – natural processes and interactions occurring within an ecosystem
Soil formation, nutrient cycling, pollination
Ecosystem services – benefit of ecosystem functions derived by humans
Air purification, flood control, seafood
Much but not complete overlap
Biodiversity and ecosystem function
more diversity → more and better ecosystem function
ecosystem functions do not exist without organisms
How does increased diversity lead to better function?
Must have an organism to perform function
Niche partitioning
Functional redundancy
Increased resilience
Resilience = speed at which a system returns to its original state after a perturbation
Intrinsic value of biodiversity
inherent worth
species and ecosystems have value by existing
independent of human use
aesthetic value
right to exist
moral obligation to ensure species continue to exist
Instrumental value of biodiversity
Benefit to humans
Economic value
Supporting services – allow ecosystems to exist
Regulating services – control abiotic environment
Provisioning services – harvestable resources
Ecosystem goods
Cultural services – aesthetic, spiritual, and recreational value
Valuation of ecosystem services (1997)
Constanza et al. 1997
$33 trillion in 1994 $
1.8 times global net production
NOT TO BE TAKEN AT FACE VALUE
Extinction
Complete eradication of a taxon
Extinct in the wild = taxon is only present in captivity
Functionally extinct = taxon can no longer perform its role in the ecosystem
Background extinction rate / normal extinction rate
Varies by group
Mass extinctions – event in which > 75% of known existing species become extinct
“Event” = < 2,000,000 years
5 mass extinctions in last 500 million years
Current extinction rate
6th mass extinction?
Threats to biodiversity
Habitat loss – destruction or degradation of habitat
Overharvesting – removing organisms at unstainable rates
Pollution – addition of a harmful substance or energy to an environment
Climate change – long-term shift in temperature and weather patterns
Introduced species – transportation of an organism to a location it is not native to
Habitat loss
Destruction or degradation of habitat
Most impactful
Land use changes and over harvesting
Habitat fragmentation
Overharvesting
Removing organisms at unstainable rates
Harvesting methods become more effective
Many regulations allow harvesting at unsustainable rates
Subsidies to otherwise non-profitable harvests
Pollution
Addition of a harmful substance or energy to an environment
Chronic vs acute
Point source vs nonpoint source
Single species impacts
Multispecies impacts
Biomagnification – increase in the concentration of a substance as it moves to higher trophic levels
Habitat loss
Climate change
Long-term shift in temperature and weather patterns
Alters the location of species fundamental niche
Populations move if possible
Alters community interactions
Habitat loss
Introduced species
Transportation of an organism to a location it is not native to
Competitors, predators, pathogens to native species
Habitat loss
Not always harmful
… but leads to increase in diversity?
Biotic homogenization
everything becomes the same in the end, so is actually an overall loss of diversity
Conservation of Biodiversity
Ecological conservation = the practice of protecting and restoring species, habitats, and ecosystems
Includes governance of human usage
Mitigate the threats to biodiversity
Common thread of all threats: human population and resource acquisition
Habitat protection and restoration
Biggest threat to biodiversity is habitat loss
Challenges
Protected area must be large enough
Some populations’ requirements change
Some populations migrate
Societal needs
Most successful
Cover large areas
Low human density
Support of local community
Enforcement
Restoration – restoring habitat to natural conditions
Remove invasive species
Remove pollutants
Remove artificial structures
Restore water flow
Reintroduce native species
Reduced harvesting
seems simple
impacts livelihoods
usually successful
Species reintroductions
captive rearing of individuals to be released
very expensive
mixed success
Weather and climate
Weather = short-term local atmospheric conditions
Day-to-day temperature, precipitation, etc.
Climate = long-term “average” atmospheric conditions
“Average” and variability of weather
• “Climate is what you expect, weather is what you get.”
Factors affecting climate
Latitude – proximity to the equator
Atmospheric circulation – large-scale air movement
Topography – mountains and valleys
Continentality – landmasses as well as distance to and size of water bodies
Oceanic circulation – large-scale water movement
Latitude
Solar energy reaching Earth’s surface decreases as latitude increases
Angle – low angle spreads the radiation over a larger area
Atmosphere – absorbs, scatters, and reflects radiation
Global net radiation
unequal heating of the Earth’s surface
the major cause of climate, and weather, is the uneven heating of the Earth’s surface
Movement of heat
heat moves from warmer/hotter areas to cooler/colder areas
Earth’s rotation
causes the Coriolis effect:
in the northern hemisphere, increasing deflection of moving water parcel to the right
equator has no deflection
in the southern hemisphere, increasing deflection of moving water parcel to the left
Atmospheric circulation
6 convection cells
Persistent high- and low-pressure areas
Predictable winds
Hot deserts
mostly found at 30º N and 30º S due to warm, dry air circulating in from Hadley cells
Topography
Physical features of a landscape
Mountains
Mountains force air up
As air rises, it cools
Cool air can hold less moisture
Precipitation forms and falls
Air descending the mountain is dry
high elevations are cooler
Continentality and jet streams
Coastal areas are cooler and wetter with small temperature ranges
Center of continents have large temperature ranges
Often drier
Air masses = large body of air with mostly uniform temperature and humidity
Defined by source
More time over source = more likely will acquire properties of source
Jet streams = narrow bands of strong wind in upper atmosphere
West to east
Persistent but strength varies
Location shifts N or S
Oceanic circulation
Energy from wind + Coriolis effect
5 major ocean gyres
North Atlantic
South Atlantic
North Pacific
South Pacific
Indian
Transfers energy from tropics to high latitudes
European climate
climate in Europe is generally much more mild than similar latitude in North America and Central and East Asia
this is because of jet streams that pick up warm air heated by the warm water of the Gulf Stream current that bring warm water into the Atlantic from the Gulf of Mexico
Climate change
long term shifts in temperatures and weather patterns
Some ecological consquences
Increased mortality
20-30% of evaluated species are at risk of extinction if temperatures reach projected 2100 levels
Timing of seasonal cycles
Habitat loss
Range shifts
Parasites included
Food web disruptions
Synonyms for soil
Earth, ground, dirt, loam, turf, humas, marl, clay, dust, mud, topsoil, muck, mold, silt, clod, sand, subsoil, kaolin, alluvium, loess, sediment, colluvium, gley, substrate, detritus, gault, blackland, shingle, substratum, guck, duff
Soil
Loose surface material that covers most land
Unconsolidated mixture of mineral particles, organic matter, water, and air that covers the Earth's surface and supports plant life
Soil formation: weathering
Physical weathering breaks parent material into smaller pieces
Frost weathering
Thermal stress
Biomechanical weathering
Chemical weathering and transformation alters the minerals present
Dissolution
Oxidation
Hydration
Biochemical weathering