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What is primary production, GPP, and NPP?
Primary production: overall process of carbon fixation
GPP: total rate of carbon fixation by producers
NPP: available fixed carbon after accounting for carbon used in respiration
fixed carbon: organic carbon
fixed nitrogen: inorganic
What controls productivity in terrestrial systems?
Temperature + moisture
Actual evapotranspiration (AET): amount of moisture evaporated in a system over time
nutrient availability only matters locally
**for map questions:
flatlands > mtns (if all else same)
closer to equator with less rain > temperature zone w/ more rain
What controls productivity in aquatic systems?
Nutrients!!!!!
How are nutrients brought up from the aphotic zone?
Coastal upwelling: winds blow water away from shore; forces cooler, nutrient-rich up from the aphotic zone.
Equatorial swelling: winds blow away from equator
Seasonal overturn: when surface water reaches 4C (max density) and sinks to the bottom
Wind-blown nutrients: soil blown into water
What is stratification?
clear layering of water driven by differences in density (primarily driven by temp. differences)
commonly near equator → never reaches 4C → low NPP
What are the Tropics?
btwn 23.5 N + S; highest latitudes that receive perpendicular sunlight once per year
Why is it rainy in Tropics/Equator? +what is coriolis effect?
hot air rises more → as it rises, it cools → condenses into clouds → hot, dry air drops back down at 30
Coriolis effect makes winds (and other moving objects) appear to curve or move at an angle (deflected) rather than straight north or south because of Earth's rotation, causing rightward deflection in the Northern Hemisphere and leftward in the Southern Hemisphere, creating curved global wind patterns and spinning storms.
What are the three cells?
Hadley cell (0-30) - air rises at 0, hits tropopause
Ferrell cell (30-60) - influenced by other cells, air rises at 60, drops at 30
Polar cell (60-90) - air rises at 60, drops at 90
El Nino/La Nina
El Nino: easterlies are weaker than usual; warmer water “sloshes” back eastward; coastal/equatorial upwelling stops
La Nina: stronger trade winds; cooler SST + more nutrients
Describe the N-cycle.
Nitrogen gas (N₂) in the atmosphere is fixed into ammonium (NH₄⁺) by nitrogen-fixing bacteria. ✅
Ammonium is converted into nitrite (NO₂⁻), and nitrite is converted into nitrate (NO₃⁻) by nitrifying bacteria. ✅
Nitrate is converted back into nitrogen gas (N₂) by denitrifying bacteria. ✅
Mineralized nitrogen (NH₄⁺ and NO₃⁻) is readily available for uptake by plants and microbes. ✅
Mineralization is the conversion of organic nitrogen into inorganic nitrogen (NH₄⁺) by microbes. ✅
Immobilization is the conversion of inorganic nitrogen (NH₄⁺ or NO₃⁻) into organic nitrogen in microbial biomass.
Describe the P-cycle.
Phosphate (PO₄³⁻) is released from rocks and minerals by weathering.
Plants and microbes take up phosphate and incorporate it into organic molecules.
Phosphorus moves through the food web as animals consume plants and other organisms.
Microbial decomposition mineralizes organic phosphorus back to inorganic phosphate.
Some phosphate is immobilized in microbial biomass temporarily.
Phosphate can be lost from soil to water or sediments, eventually returning to rocks over long time scales.
Describe Walker-Syers Model.
Young soils:
Most P is still in primary minerals (like apatite) in the rock.
Only a small fraction is non-occluded, so available P is limited despite high total P.
Intermediate soils:
Weathering of primary minerals releases P.
Enough time has passed for P to become non-occluded and available, but not so long that it has been heavily leached.
Result: available P peaks at this stage.
Old soils:
Most primary minerals are gone.
P is largely occluded in Fe/Al minerals or lost to leaching.
Available P declines, even though total P may still be present.
Evolution + Natural Selection (def, diff)
Evolution: changes in gene frequency in a population
Natural Selection: differential reproductive success based on phenotypic variation
Correct this misconception: Natural selection promotes the good of the species
Natural selection acts on individuals, not species. Traits are favored if they increase an individual’s reproductive success, even if they harm others.
Correct this misconception: Differences in reproductive success are the outcome of natural selection
Differences in relative reproductive success are the cause of natural selection, not the outcome. Natural selection occurs because individuals differ in how many offspring they produce, even when population size is stable.
What evidence needs to be present for ongoing evolution to be occurring?
phenotypic variation
phenotype matches selection pressure
phenotype is heritable
differential reproductive success
change in gene frequency
What is the comparative method, and how are phylogenetically independent contrasts used?
Comparative method: Compares traits across species while accounting for shared ancestry; effective sample size = number of independent evolutionary events.
Phylogenetically independent contrasts (PICs): Differences in trait values calculated at each independent evolutionary event + average all to find contrasts
more evolutionary events = more independent appearances of the trait with the same environment
How does exstrinsic mortality affect trade-offs?
High extrinsic mortality favors early reproduction and greater allocation to current reproduction, often at the cost of growth, survival, or offspring size. Low extrinsic mortality allows investment in survival and delayed reproduction.
r-selection vs k-selection
r-selected: early reproduction, many small offspring, little care, unstable environment.
K-selected: late reproduction, few large offspring, high care, stable environment.
How does population density influence life-history trade-offs?
High density favors fewer, larger offspring with more investment per offspring due to competition, while low density favors many, smaller offspring with less investment.
Applied vs Basic research questions?
Applied: addresses specific, current real-world problems for practical solutions
Basic: curiosity; to expand general knowledge
Mark-recapture (for estimating population size)?
capture, mark, and release Nm individuals
return, capture Nc2 individuals; note # of individuals who were already marked (=Nr)
Pm = proportion of 2nd capture that were marked (Nr/Nc2)
Nt= Nm/Pm
mark-recapture (to estimate survival rate)
capture + mark
at least two recaptures
Assumptions/limitations: individual could still be alive just not spotted; mark doesnt affect survival; closed population; marking doesnt affect behavior
Life-tables (variables, Ro, use, assumptions)
Ro = sum(lxmx) = net. reproductive rate = # of female offspring per female per generation
if Ro = 1; pop = stable
lx = proportion of surviving to age X
mx = avg # of female offpsring per female
Assumptions
all individuals are identical
closed population
Age-specific survival and fecundity are constant over time (constant environment)
do not account for overlapping generations!!
Generation time (formula, why its important)
T = sum(xlxmx)/Ro
avg. amount of time between birth of female and her daughter
more indicative of population trends in unstable environments; if T is short, then there is more chance for adaptation
Life-stage transition models + population matrices
Life-stage transition models:
boxes w/ size/stages + arrows between stages (growth, survival, reproduction)
better when demographic characteristics like survival and reproduction are more dependent on an individual's biological stage, size, or condition than on its chronological age
allows individuals to stay in same stage
Population matrices:
predict how populations change over time (lambda)
column → row (arrow from = column; arrow to = row)
*Assumptions: individuals in each stage are identical, closed population, no density-dependence
***remember that survival rate at each stage = both the arrow moving from it + the arrow repeating that stage
How do you determine which stage transition has the greatest effect on lambda?
Sensitvity: vary transitions by fixed increments
Elasticity: vary transitions by fixed proportions
neither of these take into account whether the tested increment is biologically possible
Life-simulation analysis: vary transitions to the extent of variability observed in the population
-create many random projection matrices w/ observed values; then plot them w/ the trait on the x-axis and lambda on y-axis; look for trends
Exponential population growth.
r = per-capita rate of increase
rmax = rate of increase w/ no limitations (potential)
dn/dt = rmaxN or Nt=Noe^rt
**DO NOT ACCOUNT FOR DENSITY!!!!!!
Logistic population growth
dn/dt = rmaxN(K-N/K)
As N→ K, numerator gets smaller, making fraction smaller; lower dn/dt
As N decreases, numerator gets larger, making fraction larger; higher dn/dt
Not always accurate: if N is too low, there are not enough mates for population to be growing
Assumptions: closed population, constant environment, resources are limiting, individuals have identical effects on resources
Exploitation/harvesting of wild populations (MSY, graphical representation, assumptions)
Max. sustainable yield: max # of individuals that can be harvested continuously without reducing the long-term population size
Graphical representation:
N on x-axis, dn/dt on y-axis. Mark K on x-axis, mark H on y-axis, find the dn/dt where H + current N are.
N will move towards K unless H is greater than current dN/dt.
Risky to harvest at Hmsy: population size and growth estimates may be inaccurate, environmental changes can reduce growth, and MSY occurs at intermediate N, leaving little buffer against overharvesting
Define the different types of interactions (herbivory, predation, parasitism, mututalism, competition)
Herbivory: consume plants (primary consumers)
Predation: one organims hunts, kills, and eats another
Parasitism: one organims benefits at the expense of another by utilizing its resources
Mutualism: both organisms benefit
Competition: one organism limits another access to resources
Resource (exploitation) competition
one organism uses a resource before another one can
Interference competition
direct competition; one organism actively prevents another from accessing a resource
Symmetric vs Asymmetric competition
Symmetric: both organisms have an equal effect on each other, proportionate to each of their body sizes
Asymmetric: both organisms are not equally effected by competition, effect is disproportionate to body size
Conclusions from meta-analysis about competition. Why is meta-analysis useful?
competition is present at varying strengths and in different forms across species
meta-analysis increases the overall sample size, which enhances the statistical power to detect effects and provides more precise estimates of the effect magnitude
looking at mutliple species increases ability to generalize _ can help identify similar factors that lead to similar competition
Lotka-Volterra model + competition coefficient
calculates change in pop. size when accountinf for intraspecific + interspecific competition
competition coefficient (𝛼12) quantifies the impact of one individual of species 2 on the growth of species 1, expressed in terms of how many individuals of species 1 it's equivalent to
*if asked to find α12 —-find N2 and N1 in terms of K1; set equal to 0
ASSUMPTIONS: no interspecific competition outside of species 2, all individuals of species 2 are identical (all individuals have the same effect on resources)
Competitive exclusion principle
if two species occupy the same niches, on must be excluded
Define niche segregation.
competing species in a community adapt to use different resources or habitats
Species sorting + community assembly?
Species sorting: community composition is based on which organisms can disperse to an area, persist in the abiotic conditions, and not be outcompeted.
Community assembly: overall how communities form (dont overthink this one!!)
Why do trade-offs allow multiple plant species to coexist?
Trade-offs prevent any one species from dominating because no plant can be good at all traits. Different species are optimized for different resource conditions (light, water, nutrients), so each becomes the best competitor in some environments but not others. This niche differentiation allows coexistence even though all plants require similar resources.
What is disturbance, succession, faciliation and inhibition?
Disturbance: an event that removes biomass quicker than it can replace itself
Succession: predictable replacement/turnover of species over time
Facilitation: species from earlier stages of succession support success of future species
Inhibition: species from earlier stages of succession make it more difficult for later species to establish
What is neutral theory?
Communities are made from random dispersal events. (not niche/exclusion). variation observed in communities is due to random chance alone .
What are null models? How to use for neutral theory vs competitive exclusion principle?
Used to test if observed patterns in a community are due to random chance.
Null hypothesis: species differences arise randomly (neutral theory).
Method: measure a trait, calculate observed differences, randomize trait values many times, and compare observed to random distribution.
Interpretation:
Observed > random → pattern likely deterministic (niche, competition)
Observed ≈ random → pattern could arise by chance (neutral processes)
How do humans impact nutrient cycles?
Eutrophication: nutrient enrichment; excess nutrients (N+P) in water bodies cause excessive algal growth; as algae dies, O2 is depleted (decomposition); causes dead zones (fish need O2 for cell. resp.)
Decreased nutrient retnetion
increased runoff + leaching from agriculture, deforestation, urbanization
What are the most limiting nutrients in different systems (ocean, land, freshwater)?
N: ocean + land
P: freshwater
What is decomposition?
Decomposition: breakdown of organic material that releases CO2 + nutrients
Decomposers + detritivores
Detritivores (like earthworms) ingest dead matter and break it down internally and mechanically, while Decomposers (like fungi and bacteria) secrete enzymes externally to chemically digest and absorb nutrients from dead organic matter
What controls the rate of decomposition?
AET - postively correlated
microbes needs H2O; heat speeds up rate of reactions (but not TOO much H2O, or their is lack of O2)
quality of carbon
Fastest: Glc → Cellulose → Lignin: slowest
Nutrient content (C:N)
have separate card for this :)
Litter quality (C:N + C quality)
determined by nutrient availability and longevity of leaves
high nutrient + low longevity = faster decomposition!
How does C:N content affect rate of decomposition + [N] in soil/leaves?
low N limits decomposition
but immobilzation of mineralized N is quicker
high N increases decompositon rate
not limiting so less is immoblized immediately; can be leached or taken up by plants
Why dead leaves have relatively low N?
as leaves died, N moves away (to shoots/stems + roots)
Why organic Carbon accumulates in some soil?
slow decomposition rate
high altitudes: low O2, low temp
moisture (too high or too low)
What is rarefaction?
Standardizes species richness across communities with unequal sampling effort.
Steps:
Choose the smallest total sample size among communities.
Randomly take many subsamples:
For a single standardized richness comparison: only subsamples of that size.
For a rarefaction curve: subsamples of all sizes ≤ the standardized size.
Calculate the average number of species in each subsample.
Plot a rarefaction curve (X = sample size, Y = average species richness).
Compare richness at the same standardized effort to determine true differences.
α- and β-Diversity & Body Size
1. α-Diversity (local richness)
Larger-bodied species have larger home ranges.
To capture the true local richness (α-diversity), the sampling area must be larger.
Smaller species → small home ranges → small sampling area is sufficient.
2. β-Diversity (turnover)
Larger-bodied species can disperse farther.
Nearby communities are more likely to share the same species, so turnover is lower at small spatial scales.
To observe meaningful β-diversity patterns, you must sample over larger spatial scales.
Smaller-bodied species → limited dispersal → higher turnover at finer scales.
Top-Down Regulation & Trophic Cascade
Top-Down Regulation: predators control herbivore population size rather than resources.
Trophic Cascade: effects of predators propagate through lower trophic levels.
Example: fewer herbivores → less plant damage → plants can invest more in growth and competition rather than defense.
Bottom-up regulation
Definition: Population sizes are limited by resources at the base of the food web, usually plants or primary producers.
Mechanism:
Scarce or low-quality plants
Plants with stronger defense mechanisms
→ fewer herbivores → fewer predators.
Specialists: more affected because they rely on a narrow set of resources; plant defenses or scarcity strongly impact their populations.
Trophic Level, Body Size, Population Density, & Extinction Risk
Higher trophic levels → greater energetic limitation → larger home ranges, lower population densities, higher extinction risk
Body size is not equivalent to trophic level; herbivores and carnivores span wide size ranges
Home range size often increases with body size and energetic demands
Climate Change & Ecological Communities – Species Responses
1. Extinction
Species fail to survive under new climate conditions.
Evidence: fossil record (past climate-driven extinctions), contemporary declines, model forecasts.
High risk: specialists, low-dispersal species, narrow thermal tolerance.
2. Geographic Range Shifts
Species move to track suitable climates.
Upslope: higher elevations (limited by mountain height).
Poleward: higher latitudes (cooler areas).
Evidence: contemporary (plants, butterflies, birds), fossil record (glacial-interglacial cycles).
3. Evolutionary Adaptation
Populations genetically adapt to new conditions if heritable variation and strong selection exist.
Evidence: phenology shifts, thermal tolerance changes in some plants, insects, birds.
Limitation: slow adaptation in long-lived species or low genetic diversity.
What is ecosystem ecology?study of the cycling of nutrients and the flow of energy through
systems
cycling of nutrients and flow of energy through systems
What is biodiversity, richness, and evenness?
Biodiversity: variety of life
Richness: # of species
Evenness: relative abundance of species
What is Equilibrium Theory of Island Biogeography? + what does it say about isolation + area
species richness is a dynamic equilibrium btwn colonization + extinction rate
as island area increases: pop. size increases, species richness increases, extinction decreases
as island isolation increases: colonization rate decreases, species richness decreases, endemic species increase
What is the ETIB graphical model? Assumptions?
x-axis: species richness; y-axis: colonization + extinction rate (colonization decreases w/ inc. species richness; extinction rate inc. w/ inc. species richness
Assumptions:
distance from mainland determines isolation
no speciation, new species only through colonization
island area predicts extinction
constant environment
Why is isolation effect in ETIB less reliable than area effect?
dispersal ability varies among species and environments
Ex: mountains have high richness despite isolation (high spatial heterogeneity)
How does island age impact richness?
Young islands: low richness
low colonization rate due to fewer resources/niches
Intermediate age: highest richness
high colonization; lots of resources + niches available
Old: low richness
high extinction rate as island erodes + resources become more limited
What dictates species richness on continents?
Spatial heterogeneity: more niches
Climate stability: less extinction, more time to adapt and diversify
Energy (NPP)
species -energy theory: more energy means more species
**Not entirely supported: doesnt explain all patterns, likely not only factor
Why is diversity hard to quantify?
unequal sampling effort!
species-accumulation curve (x: # of individuals sampled; y: # of species identified)
increases then plateaus
proportions is not a good way to correct for this: relationship btwn effort + species identified is NOT linear!!
RAREFACTION!!!!!!
Rank-abundance curve
x: rank (most abundant = 1; least abundant = higher ##)
y: relative species abundance
more flat slope = more evenness
longer slope/line = more richness
What is alpha, beta, and gamma diversity?
Alpha: local, homogenous community
Beta: across mult. communities (species turnover/diff. of species btwn them)
Gamma: across large regions
What is NEE? What does it mean when it is negative/positive?
NEE: Net ecosystem exchange (consumer resp. - NPP)
indicates direction + magnitude of CO2 exchange btwn land + atmosphere
negative NEE = more stored carbon than lost (stored in lignin + soil)
What are the direct biological effects of increased CO2 on terrestrial systems?
Increased NPP: higher CO₂ can boost photosynthesis and plant growth, but only if nutrients (N, P) are not limiting.
C3 vs C4 plants:
C3 plants (~90% of species) are better adapted to elevated CO₂ because their photosynthesis is normally limited by CO₂; at low CO₂, O₂ competes with CO₂ for Rubisco.
C4 plants: have a CO₂-concentrating mechanism in which CO₂ is first fixed into a 4-carbon compound in mesophyll cells and then delivered to bundle sheath cells for photosynthesis. This allows them to thrive in low CO₂, high temperature, and high light conditions. Because of this mechanism, C4 plants are less responsive to elevated CO₂.
Implication: C3 species may increase in growth and potentially richness relative to C4 species under elevated CO₂.
C3 Plants under Elevated CO₂ (+ Rubisco)
Rubisco adjustment: C3 plants rely on Rubisco for CO₂ fixation and are normally limited by CO₂ availability.
Elevated CO₂ allows Rubisco to operate efficiently → plants decrease Rubisco production.
Nitrogen demand: Rubisco contains a lot of nitrogen. Less Rubisco → lower nitrogen requirements.
Carbon fixation: More CO₂ available → more carbon is fixed overall, increasing biomass.
C:N ratio: Because carbon accumulation outpaces nitrogen, C:N ratio increases.
Key takeaway for C3 plants: They are more responsive to elevated CO₂ than C4 plants because they can increase carbon gain while reducing nitrogen investment in Rubisco.
more efficient + fix more carbon
This is an adaptive response that is potentially beneficial for the individual plant's resource optimization but is detrimental to the consumers of those plants
How increased CO2 affects oceans
ocean acidification:
co2 reacts w/ carbonate → bicarbonate
inhibits formation of calcium based shells in organisms
Why are no-analog communities sometimes cited as evidence that species sorting and niche segregation are not important in determining community composition?
If species sorting and niche segregation were important, it would be expected that communities would remain similar over time, as coexisting species have adaptations to specific niches that allow them to coexist with each other but not with other species outside of their community. Since no-analog communities exist, this indicates that species can respond to changing conditions differently than species they previously coexisted with. These species disperse to and persist in communities that match their abiotic niche requirements regardless of what species from their previous community do, suggesting that under rapidly changing environmental conditions, species’ niches are influenced more by abiotic factors than by the species they previously coexisted with.