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FW364 Ecological Problem Solving. Class 11: Population Regulation. October 13, 2013. Outline for Today. Continue to make population growth models more realistic by adding in density dependence Objectives from Last Monday : Introduce density dependence (modeling style)
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FW364 Ecological Problem Solving Class 11: Population Regulation October 13, 2013
Outline for Today Continue to make population growth models more realistic by adding in density dependence Objectives from Last Monday: Introduce density dependence (modeling style) Discuss scramble and contest competition Objectives for Today: Continue discussion of scramble and contest competition Introduce Allee effects (inverse density dependence) Text (optional reading): Chapter 3 Note: Midterm 2 is going to be moved to Nov. 6th, Revised syllabus available online.
Recap from Last Week How does exponential growth come to an end? • Two broadly defined limits to population growth: • Density independent factors • Do not regulate populations • Density dependent factors • Regulate populations Contest Scramble versus Exploitative, free-for-all competition Interference competition Resources shared equally Resources shared unequally Density dependence affects some individuals lightly (territory owners) other individuals strongly (non-owners) Density dependence affects all individuals equally
Contest and Scramble Comparison y-intercept: 2.00 lnλmax Contest growth rate declines faster 1.50 1.00 0.50 steady state 0.00 lnλ(discrete) r (continuous) -0.50 Carrying capacity (K) contest -1.00 Scramble growth rate does not slow down -1.50 -2.00 scramble -2.50 0 200 400 600 800 1000 1200 N • Let’s look at λ versus population size…
Contest and Scramble Comparison 6.0 y-intercept: • Contest: Population growth rate declines faster at first, then slows more λmax 5.0 4.0 scramble λ(discrete) 3.0 2.0 contest steady state 1.0 Carrying capacity (K) 0.0 0 200 400 600 800 1000 1200 N
Distinguishing Density Dependence Types • How can we tell which type of density dependence best • describes the population we are interested in? • Make detailed behavioral observations • OR • Do a model fitting exercise • i.e., Collect data on population density over time for many generations • Fit the data to models of scramble and contest competition • Determine which model (i.e., scramble vs. contest) fits best scramble λ contest N
Distinguishing Density Dependence Types • How can we tell which type of density dependence best • describes the population we are interested in? • Make detailed behavioral observations • OR • Do a model fitting exercise scramble λ • Keep in mind: • Are working with ideal categories… fit will not be perfect… • …but categorizing is still helpful for modeling and making predictions contest N
Population Dynamics • Why do the different density dependence types matter? • Scramble vs. contest has big difference on population stability • i.e., change in population size through time • Scramble (extreme effect) • Contest • Large fluctuations • Increased extinction risk • Difficult to predict • Stable population • Reduced extinction risk • Easier to predict
Population Dynamics • Why do the different density dependence types matter? • Scramble vs. contest has big difference on population stability • i.e., change in population size through time • Scramble (extreme effect) • Contest • Large fluctuations • Increased extinction risk • Difficult to predict • Stable population • Reduced extinction risk • Easier to predict • Let’s investigate population dynamics more closely… • … need to look at equations
General Approach to Equations • Goal: Want equations for how population growth rate (λ, r) changes with density lnλmax steady state lnλ(discrete) r (continuous) Carrying capacity (K) contest scramble N • Want equations to describe these functions for contest and scramble
General Approach to Equations • Goal: Want equations for how population growth rate (λ, r) changes with density • Start with our exponential growth equations: • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • We want to “build” equations for λ and r… • Need to define new parameters: • rmaxand K • λmaxand K • λmaxand rmaxare maximum population growth rates • Occur when population size is very small • (as density approaches 0, population growth rates approach maximum) • Kis carrying capacity • Maximum sustainable population size
Scramble Equation • Goal: Want equations for how population growth rate (λ, r) changes with density • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • (1 – Nt/K) • (1 – N/K) • λ =( λmax ) • r =rmax • Logistic growth equations • Identical to Ricker stock-recruitment equation • Scramble
Scramble Equation • Goal: Want equations for how population growth rate (λ, r) changes with density • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • (1 – Nt/K) • (1 – N/K) • λ =( λmax ) • r =rmax • Challenge: What happens to λ when: • Nt is very small? • Ntequals carrying capacity (K)? • Nt is larger than carrying capacity (K)? • Scramble
Scramble Equation • Goal: Want equations for how population growth rate (λ, r) changes with density • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • (1 – Nt/K) • (1 – N/K) • λ =( λmax ) • r =rmax • Challenge: What happens to λ when: • Nt is very small? λ ≈ λmax(exponent ≈ 1) • Nt equals carrying capacity (K)? λ = 1 (exponent = 0) • Nt is larger than carrying capacity (K)? λ < 1 (exponent < 0) • Scramble
Scramble Equation • Goal: Want equations for how population growth rate (λ, r) changes with density • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • (1 – Nt/K) • (1 – N/K) • λ =( λmax ) • r =rmax • Challenge: What happens to r when: • N is very small? • N equals carrying capacity (K)? • Nis larger than carrying capacity (K)? • Scramble
Scramble Equation • Goal: Want equations for how population growth rate (λ, r) changes with density • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • (1 – Nt/K) • (1 – N/K) • λ =( λmax ) • r =rmax • Challenge: What happens to r when: • N is very small? r ≈ rmax • N equals carrying capacity (K)? r = 0 • Nis larger than carrying capacity (K)? r < 0 • Scramble
Contest Equation • Goal: Want equations for how population growth rate (λ, r) changes with density • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • No continuous analog • λmax K • λ = • Ntλmax – Nt + K • Beverton-Holt equation • Contest
Contest Equation • Goal: Want equations for how population growth rate (λ, r) changes with density • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • No continuous analog • λmax K • λ = • Ntλmax – Nt + K • Challenge: What happens to λ when: • Nt is very small? • Ntequals carrying capacity (K)? • Nt is larger than carrying capacity (K)? • Contest
Contest Equation • Goal: Want equations for how population growth rate (λ, r) changes with density • Discrete growth: • Continuous growth: • dN/dt = r N • Nt+1 = Ntλ • No continuous analog • λmax K • λ = • Ntλmax – Nt + K • Challenge: What happens to λ when: • Nt is very small? λ ≈ λmax(denominator ≈K) • Nt equals carrying capacity (K)? λ = 1 (denominator ≈λmax K) • Nt is larger than carrying capacity (K)? λ < 1 (denominator > numerator) • Contest
Predicting Population Size • We now have equations for population growth with density dependence! • Just considering discrete growth: • λmax K • (1 – Nt/K) • Nt+1 = Nt • Nt+1 =Nt ( λmax ) Contest Scramble • Ntλmax – Nt + K • We can use these equations to calculate population size between any two consecutive years • We’ll do this in a graphically interesting way using replacement curves in Lab tomorrow
Replacement Curves • Replacement Curves: • Plot of next year’s population (Nt+1) size against current population size (Nt) • (for right now, think about points having the ability to fall anywhere in this space) • Helpful for seeing how different types of • density dependence influence actual population dynamics Nt+1>Nt λ > 1 1 : 1 line Steady state line Nt+1 Nt+1 = Ntλ = 1 • Time does not “move” from left to right in these figures!!! • Example in Lab tomorrow Nt+1 < Nt λ < 1 Nt
Scramble vs. Contest Comparison • Comparison of scramble and contest density dependence • Similarity: • Both scramble and contest density dependence (competition) result in population regulation • Regulation slows population growth at high density… • … which draws the population toward an equilibrium (carrying capacity) • Differences: • Scramble: Additional individuals always reduce the population growth rate • linear relationship between lnλ (r) and N • Population will overshootK • Contest: Effect of each additional individual on population growth decreases as more and more individuals are added • Population will not overshootK
Scramble vs. Contest Comparison • Comparison of scramble and contest density dependence 6.0 • Differences Con’t: • For two populations with same λmaxand K 5.0 4.0 • Population decreases faster (λ) for scramble when N > K • Population increases faster (λ) for scramble when N < K 3.0 2.0 scramble λ 1.0 • Combination drives population fluctuations for scramble (Lab 5) contest steady state 0.0 K 0 200 400 600 800 1000 1200 N
Another Type of Density Dependence • So far, density dependence has meant: • “Increased density leads to lower per capita population growth (lower λ)” • The reverse can also be true in some situations! • i.e., Increased density leads to higher per capita population growth (higher λ) • Inverse density dependence: Allee effects • Only occurs at low population sizes • (at high densities, near K, regular density dependence always applies)
How Alee Effects Happen • Per capita reproduction may decrease at low population sizes • Mate limitation: density gets so low that mates cannot find each other • Inbreeding: density so low that inbreeding occurs • For plants, low density may mean fewer pollinators attracted • Mate limitation • Inbreeding • Pollinator attraction • Per capita mortality may decrease at higher population sizes (i.e., increase at low size) • “Safety in numbers” situation, e.g. colony nesting • For plants, greater density may mean greater soil stability (less erosion) • Colony nesting • Soil erosion
How Alee Effects Happen • Constant harvest amount also functions like an Allee effect • Lab 3: Nt+1 = Nt (1 + b’ – d’) - H • When a constant number is harvested, • the proportion of the population harvested changes as population size changes • For H = 100 • When N = 1000, the proportion harvested is 0.10 ( = 100 / 1000) • When N = 400, the proportion harvested is 0.25 ( = 100 / 400) • When N = 200, the proportion harvested is 0.50 ( = 100 / 200) • Blue whale harvest • Proportional harvest mortality increases • as population decreases • (like a death rate increasing at low population sizes) • … inverse density dependence • Constant harvests are a risky practice
Alee Effects Dynamics • Allee effects involve positive feedback at low population sizes • i.e., population decline leads to further decline • Result: Dip in the population growth rate versus density curve • λ > 1 • scramble without Allee effect • scramble with Allee effect • Allee effect gets stronger from top to bottom curve • λ < 1 • Based on Fig. 3.13
Alee Effects Dynamics • Allee effects involve positive feedback at low population sizes • i.e., population decline leads to further decline • Result: Dip in the population growth rate versus density curve • λ > 1 • scramble without Allee effect • has one equilibrium = K • scramble with Allee effect • has two equilibria • λ < 1 • Based on Fig. 3.13
Allee Effects – Population Forecast Allee Replacement Curve Stable equilibrium Unstable equilibrium • N0 above first equilibrium • Carrying capacity • N0 below first equilibrium • Population crash
Allee Effects – Population Forecast Allee Replacement Curve Stable equilibrium Unstable equilibrium • Fate of population depends on starting conditions (starting size)
Allee Effects – Population Forecast Allee Replacement Curve λ > 1 • Increase • to K λ < 1 • Population fate is extinction if it falls below unstable equilibrium • Decline to extinction • Decrease • to K
Allee Effects – Wrap Up • Allee effects have major implications for management of endangered species and fisheries • It is crucial that Alleeeffects be identified and quantified • Must manage to keep population far above “the well” • (unstable equilibrium) • Possible that Allee effects are the cause of • collapses of some overharvested fisheries • Bluefin tuna • Even if population does not decline at low densities, a slowed rate of growth will keep the population low for longer periods (slow to recover) • More vulnerable to random extinction
Lab Tomorrow • We will be working with some data in excel and doing some practice problems similar to what you will be seeing on Midterm 2 (now scheduled for Nov. 6th) • We will look at scramble and contest competition • We will look at Allee effects • You won’t need to bring the RAMAS software, as we will be working with excel and mainly analyzing data.