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Population density
Measuring relative vs. absolute density, density dependance, and distribution and abundance
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Population
- A group of individuals of the same species occupying a particular area at the same time
- They are natural biological units: inidividuals are portentially interfertile and share the same resources (interact)
- These are the basic unit of evolution: natural selection operates on organisms, but the population evolves
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Occupying a particular area
- Boundaries of populations are sometimes clear, sometimes vague (sufficient size that reproduction and survival maintains the population for many generations)
- May exist in heterogeneous landscapes, but only suitable habitats are occupied within the particle area
- This is dynamic (disperal immigration and emmigration among local populations)
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Population boundaries in practice
Often set arbitrarily: an elk in Elk Island National Park
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Population Properties Structure
- Static, snapshot description of the population
- How many? spatial distribution? Ages? Sizes? Sex ratio? Survival patterns?
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Population Properties Dynamics
- Changes through time (rates)
- Birth rate, death rate, immigration rate, emmigration rate...?
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Population "size"
- Abundance (how many individuals? how much biomass?)
- Density- numbers (biomass) per unit area or volume (absolute (crude) density: density without regard to habitable area or variation in habitat quality. Absolute density: population density in appropriate habitat only)
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Problems with population density
- What is an individual? (clonal organisms, seedling vs. adult, larvae vs. adult)
- Biomass density, or cover for plants
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Measuring density
- Relative estimate (indices presumed to be related to the abundance of target organisms: used for comparison)
- Absolute estimates (direct counts, sampling)
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Trends in density
- # of individuals/area declines with increasing body size
- Many ecological processes are desity dependent
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Density dependence
- An effect whose magnitude is increased (positively or negatively) with population density (denser = more effect)
- Examples are predation, competition, disease, schools, allee effects
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Density independence
An effect whose magnitude is unaffected by population density (denser/less dense=no matter)
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Random distribution
- Reference pattern (what you would expect if the species where completely indifferent), position of individuals is independent of other individuals
- Relatively rare: homogeneous environment, random dispersal, no social structure, or at least neurtal interactions
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Uniform
- Inidividuals are MORE EVENLY spaced than expected by chance
- Relatively common: intra-specific competition, territoriality in a relatively uniform environment
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Clumped
- neighbors are CLOSER than expected by chance
- Common: patchy habitat/resources, reproduction and dispersal, social structure
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Distribution pattern can:
Reveal underlying spatial patterns in the environment, reflect past and/or future interactions, influence the ecological density of the population, influence accuracy/precision of population estimates
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Distribution pattern can be due to:
- Intrinsic characteristics of the species
- Extrinsic factors
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Population distribution has many potential causes:
- Geography/climate
- Predation
- Competition
- Disease
- Social structure
- Etc
- It is dependant on the niche (where conditions suit a species' niche, the species should thrive
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Metapopulations
- A population made up of sub-populations
- Sub-populations joined by the movement of individuals
- Any subpopulation can go entinct; but, on average, sources produce more emigrants than they recieve immigrants, and sinks do the opposite
- Small sub-populations = greater risk of extinction
- Density-dependent and density independent effects also contribute
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Species range and tolerance
- Geographic range (broad or restricted)
- Habitat tolerance (broad or narrow)
- Local population (large or small)
- The fewer characteristics of RARITY, and the more of ABUNDANCE, that an organism has, the better its chances of survival in the face of environmental change
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Dispersal
Can increase or decrease local density, can be density independent (passive: wind, gravity, etc) or density dependent (competition, habitat quality or size)
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Sources
Consistently produce more emigrants than immigrants while maintaining their existing population
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Sinks
Consistently recieve more then they produce
A patch of insuficient size or quality may become a temporary source while going extinct
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Population structure and variation
Variation among individuals within populations can (potentially strongly) influence dynamics such as birth rates, death rates, immigration, emigration, etc
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Age frequency distributions
- Dominant age classes may be visible, likely reflects non-random recruitment
- environment (climate? fire?)
- intra-specific/inter-age-class inhibition?
- (class make ingerences about the species history)
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Even-aged stands
- Referrs to forrest that most times the trees are the smae age, instead of patchy mixed bag of ages like in the past
- Natural fire cycles
- Human activity
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Cohort - age frequency distributions
A cohort of larger fish preys upon smaller fish, until they "grow out of it"
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Age pyramids
Snapshots of age (and sex) structure
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3 "ecological" age classes
- Juvenile phase dominated by growth
- Reproductive phase
- Post-reproductive phase
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Age distribution
- Reflects a population's history = clues for the ecologist
- survival
- reproduction
- potential for future growth
- May be complex in a highly variable environment
- (allows us to predict "forward" what may happen)
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Sex ratio = the result of natural selection
- 1:1 sex ratio is an evolutionarily stable strategy
- more females than males
- males would have more reproductive chances-> fitness
- the same advantage (=selection pressure) declines near 1:1
- same logic works if more males than females
- --> frequency dependent selection
- May be exceptions (reed frog which begins as female)
- Humans can effect this as well, with herbicides
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Simple life tables
- A useful tool for summarizing survivorship
- Data can be used to create a survivorship curve, which will reveal different life histories
- x= age
- nx= # surviving to age "x" (cohort size)
- lx=proportion suriving to age "x" (age-specific survivorship
- mx= (summation) Gross reproductive rate (age-specific repro)
- lxmx=(summation) net reproductive rate
- xlxmx= (summation / R0) equals T (generation time)
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Type 1 survival
Juvenile survival is high and most mortality occurs amoung older individuals
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Type 2 survival
Individuals die at equal rates regardless of age
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Type 3 survival
Individuals die at high rates as juveniles an then at much lower rates later in life.
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Population growth (basics)
- BIDE
- B- births
- I - immigration
- D - deaths
- E - emigration
For comparisions amoung populations, rates of change can be more useful (signified by small case letters)
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Rate
An amount of change (in something) over time
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Discrete rate
Average rate of change during delta time
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Instantaneous rate
Rate of change over a very short time (t-->0)
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Crude rate
Total change in numbers over time (delta N/delta t)
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Specific rate
Rate scaled per organism (delta N/N0 delta t)
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Reproductive rate (natility)
Specific reproductive rate = # of offspring per individual per unit time
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Reproductive rate and age
- Reproductive rate often varies with age, every age group will have its own R0
- Age specific reproductive rate (mx)
- # of offspring per female of age "X" per unit of time
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Gross Reproductive rate (GRR)
# of offspring produced during a female's life IF she lives to age omega (sum of mx)
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Age-specific survivorship (lx)
- Need to know what % of population survives or dies at each age "X"
- lx = % of cohrot alive at start of age "X"
- lx = nx/n0
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Geometric rate of increase
- # of daughters born in generation t+1/# of daughters born in gerenation t = Nt+1/Nt = lambda
- lambda > 1, pop'n growing
- lambda = 1, pop'n stable
- lambda < 1, pop'n shrinking
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Net reproductive rate (R0)
- If R0 > 0 pop'n growing
- R0 = 0 pop'n remains constant
- R0 < 0 pop'n is declining
- sum of lxmxlittle r is per capita rate of increase (otherwise same concept)
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Generation time (T)
- The average age at which a female gives birth
- Typically the larger you are, the longer it takes to become reproductively active
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Nx=N0R0x
The rate a population wuld increase if R0 remains constanct and >1 and there are no limits on population growth
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Geometric growth
- Discrete, non-overlapping generations
- Nt=N0lambdatNt= # of individuals at time t
- N0=# of inidividuals at start
- Lambda = geometric rate of increase
- t = # of time intervals or generations
- Or if you only care about the next generation:
- Nt+1=Ntlambda
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Exponential growth
- Continuous population growth can be modelled exponentially
- dN/dt = rmaxN
- dN/dt = change in the # of individuals per unit time
- rmax= intrinsic rate of increase
- ->max possible under ideal conditions
- Size of population at any time can be calculated as
- Nt=N0ermaxtNt=# of individuals at time t
- N0=intial # of individuals
- e = base of natural logarithms
- rmax =intrinsic rate of increase
- t = # of time intervals
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Exponential growth under stable conditions
- r -> rmax
- But.... rmax is balanced by extrinsic factors
- As population density increases:
- per capita resources decline
- Growth decreases
- age at maturity increases, fecundity decreases
- Higher densities attract predators, so mortality increases
- Social strife within populations and disease increase
- Environmental resistance increased (birth rate decrease, or death rate increases, or both)
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Upper equilibrium level
carrying capacity (k)
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Populations below K
- below inflection point, growth is accelerating
- above inflection point, growth > 0 but decelerating
- Populations above K decrease
- Populations at K remain approx. constant
K is a property of the environment; the max. sustainable population size
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Logistic population growth
- dN/dt=rmaxN((K-N)/K)
- rmax=intrinsic rate of increase
- K=carrying capacity
- When N nears K, the right side of the equation nears zero, (change in pop'n size approaches 0)
- When N is low, the right side of the equation is near 1 (change in pop'n size is exponential)
- Size of pop'n at any time can be calculated as
- Nt= K/(1+(K/N0-1)e-rmaxt)
- Nt = # of inidividuals at time t
- N0 = intial # of individuals
- e = base of natural logarithms
- rmax=intrinsic rate of increase
- t = # of time intervals
- K = carrying capacity
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Density (in)dependence
- Density independence - effects independent of N, e.g. severe weather
- Density dependence - effects depent on N, e.g. "environmental resistance" (two examples; song sparrows in BC, at high density, more floater males, lower fecundity, lower juvenile survival, and plants, at high density decreased growth rate, decreased survival, decreased seed production)
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Why different cycles?
Identifying and understanding the processes that affect plant and animal structure is a major challenge in ecology
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K-strategists
- slower development, delayed reproduction, longer lifespan, larger size, competitive
- growth regulated by intrinsic factors
- Many exhibit logistic growth and populations are stable near the carrying capacity (k)
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r-strategists
- Faster development, faster reproduction, shorter lifespan, smaller size, not competitive
- Growth regulated by external factors
- Many exhibit irruptive population dynamics in response to temporary environmental change
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Irruptive dynamics
- Due to seasonal variation in K
- Npeak depends on rmax and length of season -> weather dependent
- Density factors can vary in actual importance
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Cyclic dynamics (observations)
- 1. A species may not be cyclic everywhere
- 2. Not all related species in a region are cyclic
- 3. The cyclic species in a region may or may not be in phase
- 4. Density ratios (max vs. min) can be really high (in the hundreds)
- 5. Period of cycle more regular than amplitude
- 6. During early increase phase: high reproductive rates and high dispersal. During high but declining phase: low reproduction and dispersal
- 7. Aggressiveness varies throughout cycle
- 8. Cycle not symmetrical: rapid explosion, rapid crash, several low density year
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Cyclic dynamics: Proximate mechanisms
- Populations gain "momentum"... high birth rates at low densities
- Populations "overcompensate"... low survival and birth rates at high dens.
- Population response is time delayed
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Cyclic dynamics: untimate hypothesis
- Abiotic (weather, sunspots)
- Biotic - intrinsic factors (stress, territoriality, dispersal)
- Biotic - extrinsic factors (food, predation, disease)
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Cyclic dynamics: summary
- It's complicated
- Population regulation is complex
- Even components (e.g.K) can be complex
- r is affected by both extrinsic (abiotic) and intrinsic (biotic) factors
- Negative feedback between r and N - ie., density dependence - is necessary for population regulation, but can vary in relative importance
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Expirpation
Extinction from a specific global region
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Background extinction
- Species lifespan = 1-10 million years
- Causes are highly variable; local disturbance, Pseudo-extinction, or co-extinstion
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Mass extinction
Sudden change in intensity, such as a volcano or meteroites
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Anthropogenic extinction
Extinction caused by humans, caused by climate change, or fragmentation by roads, exploiting at a large rate
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Extinction risk
- Larger populations are likely to persist for longer
- Age and spatial structure of the population also affects extinction risk
- Population resillience - long "return time" = low resillience, short "return time' = high resillience
- Body size, longevity, and population size interact to affect extinction risk (Large organisms should have the advantage in small populations, and small organisms in large populations)
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Conservation
- The preservation of ecological systems in a natural or near-natural condition
- (requires knowledge to decide waht is and isn't harmful)
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Resoration ecology
- The branch of ecology concerned with restoring ecological sytems to their natural condition
- (requires it to decide what is and isn't vital to restore)
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