Demographic PVAs Simulating Demographic Stochasticity and Density Dependence Demographic stochasticity • Simulated by performing so-called Monte Carlo simulations: the fate of each individual in a certain class and a certain year is decided by a set of independent random choices, all of which are based on the same set of mean vital rates • However this can greatly slow a program Variability caused by demographic stocahsticity in binomial vital rates 0.14 0.12 Variance in outcome 0.1 0.08 0.06 0.04 0.02 0 0 5 10 15 20 Number of individuals 25 30 35 Techniques to how to perform Monte Carlo simulations • For vital rates that are probabilities: Pick a uniform random number and compare its value to the probabilities of different fates an individual might experience How to use a uniform random number to decide between fates in a Monte Carlo simulation 0 Survive and shrink to class 3 a34 Survive and stay to class 4 a34+a44 Survive and grow to class 5 a34+a44+a54 Survive and grow to class 6 a34+a44+a54+a64 Die 1 Adding demographic stochasticity to reproduction • Determine if the individual lives • Use a Poisson or another discrete distributions to obtain the individual fertility 40 Observed 35 Predicted Frequency 30 Number of flowers 25 20 Avg. # Flowers 15 10 5 0 0 1 2 3 5 Number of flowers Population 1; 2000 40 Observed 35 Predicted Frequency 30 25 20 15 10 5 0 0 1 2 3 5 pop1 pop2 pop3 pop4 pop5 pop6 2000 1.45 1.39 3.41 3.22 5.71 2.69 2001 0.75 0 1.89 3.62 1.73 0.69 2002 1.26 1.63 4.10 5.00 4.86 1.65 Number of flowers Population 2; 2000 P (y=r) = (e –μ μ r) / r! Sampling from Poisson distributions to estimate flower production Avg. # Seeds Number of seeds per flower Population 1; 2000 8 6 4 2 Std. Dev = 28.97 Mean = 63.2 N = 30.00 0 12.0 40.0 26.0 68.0 54.0 96.0 82.0 124.0 110.0 number of seeds per flower 138.0 2000 2001 2002 pop1 63 63 53 pop2 56 62 42 pop3 54 43 43 pop4 55 49 38 pop5 63 54 41 pop6 68 50 33 Sampling from normal distributions to estimate seed production Including density dependence • Two factors make it more difficult to account for density dependence in demographic PVAs 1. We will rarely have as many years of vital rate estimates from a demographic study 2. There are many more variable that are potentially density dependent Three questions we must consider: • Which vital rates are density-dependent? • How do those rates change with density? • Which classes contribute to the density that each vital rate “feels”? Two more limited approaches to building density-dependent projection matrix models 1. Assume that there is a maximum number or density of individuals in one or more classes, or of the population as a whole , that the available resources can support, and construct a simulation model that prevents the population vector from exceeding this limit. Two more limited approaches to building density-dependent projection matrix models 2. Choose one or at most a few vital rates that, on the basis some evidence are suspected to be strongly density dependent Placing a limit on the size of one or more classes Caps or ceilings on population density are most often used to introduce density dependence when the focal species is territorial The Iberian Lynx Gaona et al. (1998) lynx population in Doñana National Park post breeding census birth-pulse population Cubs Juveniles Floaters Breeders Cubs 0 0 0 b x c x p x s4 Juveniles s1 0 0 0 Floaters 0 s2 (1-g) s3 (1-g) 0 Breeders 0 s2 (g) s3 (g) s4 b = probability that a territory-holding female will breed in a given year c = number of cubs produced by females that do breed p = proportion of cubs that are female Gaona et al. (1998) lynx population in Doñana National Park post breeding census birth-pulse population • Density dependence acts on g, which represent the probability that a surviving juvenile or floater will acquire a territory next year • Just before the birth pulse that precedes census t+1, there will be s4n4(t) breeding females still in possession of a territory and K- s4n4(t) vacant territories available. Gaona et al. (1998) lynx population in Doñana National Park post breeding census birth-pulse population • g= [K-s4n4(t)]/[s2n2(t)+ s3n3(t)] Density dependent functions s0 E t s0 0exp E t The Ricker function Density dependent functions s0 0 s0 E t 1 E t Beverton-Holt model Density dependent functions Et bi fi ni (t ) I II III IV V I 0 0 b3f3s0E(t) b4f4s0E(t) b5f5s0E(t) II s1 0 0 0 0 III 0 s2 0 0 0 IV 0 0 s2 (1-b3) 0 0 V 0 0 0 s3 (1-b4) 0