Count Based PVA: Density-Independent Models Count Data • Of the entire population • Of a subset of the population (“as long as the segment of the population that is observed is a relatively constant fraction of the whole”) • Censused over multiple (not necessarily consecutive years Population dynamics in a random environment • The theoretical results that underlie the simplest count-based methods in PVA. The model for discrete geometric population growth in a randomly varying environment Nt+1=λtNt Assumes that population growth is density independent (i.e. is not affected by population size, Nt) Nt+1=λtNt • If there is no variation in the environment from year to year, then the population growth rate λ will be constant, and only three qualitative types of population growth are possible λ>1 Geometric increase Stasis λ=1 Geometric decline λ<1 By causing survival and reproduction to vary from year to year, environmental variability will cause the population growth rate, to vary as well • A stochastic process Three fundamental features of stochastic population growth • The realizations diverge over time • The realizations do not follow very well the predicted trajectory based upon the arithmetic mean population growth rate • The end points of the realizations are highly skewed t=10 t=20 t=40 t=50 The best predictor of whether Nt will increase or decrease over the long term is λG • Since Nt+1=(λt λt-1 λt-2 …λ1 λ0) No λG is defined as (λG)t =λt λt-1 λt-2 …λ1 λ0 ;or λG =(λt λt-1 λt-2 …λ1 λ0)(1/t) Converting this formula for λG to the log scale μ= lnλG =lnλt+ln λt-1+ …lnλ1 +ln λ0 t The correct measure of stochastic population growth on a log scale, μ, is equal to the lnλG or equivalently, to the arithmetic mean of the ln λt values. μ>0, then λ>1 the most populations will grow μ<0, then λ<1 the most populations will decline t=30 6 6 5 5 4 4 3 3 2 2 1 1 0 -200 t=15 6 5 5 4 4 3 3 2 2 1 1 0 200 400 600 N 800 1000 200 400 600 800 1000 1200 1400 1600 0 1 N 6 0 -200 0 1200 1400 0 1 1600 2 3 4 5 6 7 8 Ln(N) 8 7 2 3 4 5 ln(N) 6 7 8 6 ln(N) 5 4 3 0 10 20 30 t 40 50 To fully characterize the changing normal distribution of log population size we need two parameters: • μ: the mean of the log population growth rate • σ2 : the variance in the log population growth rate 8 6 4 2 0 -2 -4 -6 0 2 4 6 8 10 12 14 16 18 20 8 6 4 2 0 -2 -4 -6 0 2 4 6 8 10 12 14 16 18 20 The inverse Gaussian distribution • g(t μ,σ2,d)= (d/√2π σ2t3)exp[-(d+ μt)2/2σ2t] • Where d= logNc-Nx • Nc = current population size • Nx =extinction threshold The Cumulative distribution function for the time to quasi-extinction To calculate the probability that the threshold is reached at any time between the present (t=0) and a future time of interest (t=T), we integrate • G(T d,μ,σ2)= Φ(-d-μT/√σ2T)+ • exp[-2μd/ σ2) Φ(-d-μT/√σ2T) • Where Φ(z) (phi) is the standard normal cumulative distribution function The probability of ultimate extinction Calculated by taking the integral of the inverse Gaussian distribution from t=0 to t =inf • G(T d,μ,σ2)=1 when μ< 0 • exp(-2μd/ σ2) when μ>0 Three key assumptions • Environmental perturbations affecting the population growth rate are small to moderate (catastrophes and bonanzas do not occur) • Changes in population size are independent between one time interval and the next • Values of μ and σ2 do not change over time Estimating μ,σ2 • Lets assume that we have conducted a total of q+1 annual censuses of a population at times t0, t1, …tq, having obtained the census counts N0, N1, …Nq+1 • Over the time interval of length (ti+1 – ti) Years between censuses i and i+1 the logs of the counts change by the amount log(Ni+1 – Ni)= log(Ni+1/Ni)=logλi where λi=Ni+1/Ni Estimating μ,σ2 • μ as the arithmetic mean • σ2 as the sample variance • Of the log(Ni+1/Ni) Female Grizzly bears in the Greater Yellowstone Adult Females . 120 100 80 60 40 20 0 1950 1970 Year 1990 Estimating μ,σ2 • μ =0.02134; σ2 =0.01305 Model R Adjusted R Square R Square 1 0.186005 Std. Error of the Estimate 0.034598 0.008506 Durbin-Watson 0.114241 2.570113 ANOVA Model Sum of Squares 1 df Mean Square Regression 0.017305 1 0.017305 Residual 0.482884 37 0.013051 Total 0.500189 38 F Sig. 1.325996 0.256906 Coefficients Model 1 INTERVAL Unstandardized Coefficients Standardized Coefficients B Beta Std. Error 0.02134 0.018532 0.186005 t Sig. 1.151519 0.256906