The suggested citation for this analytic report is:
Thorley, J.L., Andrusak, G.F. and Dalgarno, S.I.J. (2018) Kaslo Bull Trout Productivity 2017. A Poisson Consulting Analysis Report. URL: https://www.poissonconsulting.ca/f/443864882.
The Kaslo River and Keen Creek, which is a tributary of the Kaslo River, are important Bull Trout spawning and rearing tributaries on Kootenay Lake. From 2012 to 2017, field crews have night-snorkeled both systems in the fall and recorded all juvenile Bull Trout between 30 mm and 350 mm in length. Snorkel and electrofishing marking crews have also captured and tagged juvenile Bull Trout for the snorkel crews to resight. Redd counts have been conducted in both systems since 2006.
The primary goal of the current analyses is to answer the following questions:
What is the observer efficiency when night-snorkeling for juvenile Bull Trout in the Kaslo River and Keen Creek?
What are the densities of age-1 Bull Trout in the Kaslo River and Keen Creek?
What is the relationship between the number of redds and the resultant densities of age-1 Bull Trout two years later?
The data were cleaned, tidied and databased using R version 3.4.4 (R Core Team 2015).
Table 1. Bull Trout length cutoffs by age-class.
| Age Class | Length (mm) |
|---|---|
| Age-0 | 30 - 90 |
| Age-1 | 91 - 150 |
| Age-2+ | 151 - 350 |
Model parameters were estimated using Maximum-Likelihood (ML) and Bayesian methods. The Maximum-Likelihood Estimates (MLE) were obtained using TMB (Kristensen et al. 2016) while the Bayesian estimates were produced using JAGS (Plummer 2015). For additional information on ML and Bayesian estimation the reader is referred to Millar (2011) and McElreath (2016), respectively.
Unless indicated otherwise, the Bayesian analyses used uninformative normal prior distributions (Kery and Schaub 2011, 36). The posterior distributions were estimated from 1500 Markov Chain Monte Carlo (MCMC) samples thinned from the second halves of 3 chains (Kery and Schaub 2011, 38–40). Model convergence was confirmed by ensuring that \(\hat{R} \leq 1.1\) (Kery and Schaub 2011, 40) and \(\textrm{ESS} \geq 150\) for each of the monitored parameters (Kery and Schaub 2011, 61). Where \(\hat{R}\) is the potential scale reduction factor and \(\textrm{ESS}\) is the effective sample size.
The parameters are summarised in terms of the point estimate, standard deviation (sd), the z-score, lower and upper 95% confidence/credible limits (CLs) and the p-value (Kery and Schaub 2011, 37, 42). For ML models, the point estimate is the MLE, the standard deviation is the standard error, the z-score is \(\mathrm{sd}/\mathrm{MLE}\) and the 95% CLs are the \(\mathrm{MLE} \pm 1.96 \cdot \mathrm{sd}\). For Bayesian models, the estimate is the median (50th percentile) of the MCMC samples, the z-score is \(\mathrm{mean}/\mathrm{sd}\) and the 95% CLs are the 2.5th and 97.5th percentiles. A p-value of 0.05 indicates that the lower or upper 95% CL is 0.
Model averaging and selection was implemented using an information theoretic approach (Burnham and Anderson 2002). ML models were compared using the marginal Akaike’s Information Criterion corrected for small sample size (AICc, Burnham and Anderson 2002). The support for fixed terms in models with identical random (Kery and Schaub 2011, 75) effects was assessed using the Akaike’s weights (\(w_i\)) which were also used for model averaging (Burnham and Anderson 2002).
Where relevant, model adequacy was confirmed by examination of residual plots for the full model(s).
The results are displayed graphically by plotting the modeled relationships between particular variables and the response(s) with the remaining variables held constant. In general, continuous and discrete fixed variables are held constant at their mean and first level values, respectively, while random variables are held constant at their typical values (expected values of the underlying hyperdistributions) (Kery and Schaub 2011, 77–82). When informative the influence of particular variables is expressed in terms of the effect size (i.e., percent change in the response variable) with 95% confidence/credible intervals (CIs, Bradford et al. 2005).
The analyses were implemented using R version 3.4.4
(R Core Team 2015) and the
mbr family of packages.
All resighted fish with a tag were allocated to the closest unallocated marked fish (with the same colour tag) by fork length and distance. The marked fish were analysed using a logistic regression model. Key assumptions of the full Maximum Likelihood logistic regression include:
Preliminary analysis indicated that river kilometre received similar support to watershed area.
The final Bayesian logistic regression assumed that:
Both systems were broken into 100 m sites by bank. The lineal density at each site was estimated using an over-dispersed Poisson Generalized Linear Mixed Model. Key assumptions of the full Maximum Likelihood GLMM include:
Preliminary analysis indicated that river kilometre was better supported as a predictor of lineal density than watershed area.
The final Bayesian GLMM dropped sinuosity and gradient and took into account the uncertainty in the observer efficiencies as estimated by the observer efficiency model.
The stock-recruitment relationship was estimated using a Bayesian Beverton-Holt stock-recruitment curve. Key assumptions of the final BH SR model include:
#include <TMB.hpp>
template<class Type>
Type objective_function<Type>::operator() () {
DATA_VECTOR(Observed);
DATA_FACTOR(System);
DATA_VECTOR(Tagged);
DATA_VECTOR(Length);
DATA_VECTOR(Gradient);
DATA_VECTOR(Sinuosity);
DATA_VECTOR(Rkm);
PARAMETER(bIntercept);
PARAMETER(bSystem);
PARAMETER(bLength);
PARAMETER(bLength2);
PARAMETER(bGradient);
PARAMETER(bSinuosity);
PARAMETER(bRkm);
vector<Type> eObserved = Observed;
int n = Observed.size();
Type nll = 0.0;
for(int i = 0; i < n; i++){
eObserved(i) = invlogit(bIntercept + bSystem * System(i) + bLength * Length(i) + bLength2 * pow(Length(i), 2) + bGradient * Gradient(i) + bSinuosity * Sinuosity(i) + bRkm * Rkm(i));
nll -= dbinom(Observed(i), Tagged(i), eObserved(i), true);
return nll;
Template 1. Full model.
model{
bIntercept ~ dnorm(0, 5^-2)
bLength ~ dnorm(0, 5^-2)
for(i in 1:length(Observed)){
logit(eObserved[i]) <- bIntercept + bLength * Length[i]
Observed[i] ~ dbern(eObserved[i])
}
Template 2. Final model.
#include <TMB.hpp>
template<class Type>
Type objective_function<Type>::operator() () {
DATA_VECTOR(Count);
DATA_VECTOR(Length);
DATA_VECTOR(Coverage);
DATA_VECTOR(Efficiency);
DATA_VECTOR(Dispersion);
DATA_VECTOR(Gradient);
DATA_VECTOR(Sinuosity);
DATA_VECTOR(Rkm);
DATA_FACTOR(System);
DATA_FACTOR(Site);
DATA_INTEGER(nSite);
DATA_FACTOR(Year);
PARAMETER_MATRIX(bSystemYear);
PARAMETER_VECTOR(bSite);
PARAMETER(bGradient);
PARAMETER(bSinuosity);
PARAMETER(bRkm);
PARAMETER_VECTOR(bDispersion);
DATA_INTEGER(nDispersion);
PARAMETER(log_sDispersion);
PARAMETER(log_sSite);
Type sSite = exp(log_sSite);
Type sDispersion = exp(log_sDispersion);
ADREPORT(sSite);
ADREPORT(sDispersion);
vector<Type> eDensity = Count;
vector<Type> eCount = Count;
vector<Type> eNorm = pnorm(bDispersion, Type(0), Type(1));
vector<Type> eDispersion = qgamma(eNorm, pow(sDispersion, -2), pow(sDispersion, 2));
Type nll = 0.0;
for(int i = 0; i < nSite; i++){
nll -= dnorm(bSite(i), Type(0), sSite, true);
for(int i = 0; i < nDispersion; i++){
nll -= dnorm(bDispersion(i), Type(0), Type(1), true);
eDensity(i) = exp(bSystemYear(System(i), Year(i)) + bGradient * Gradient(i) + bSinuosity * Sinuosity(i) + bRkm * Rkm(i) + bSite(Site(i)));
eCount(i) = eDensity(i) * Length(i) * Coverage(i);
nll -= dpois(Count(i), eCount(i) * Efficiency(i) * eDispersion(i), true);
return nll;
Template 3. The full model.
model{
bEfficiency ~ dnorm(Efficiency[1], EfficiencySD[1]^-2) T(0, )
for(i in 1:nSystem) {
for(j in 1:nYear) {
bSystemYear[i,j] ~ dnorm(0, 5^-2)
}
}
log_sSite ~ dnorm(0, 5^-2)
log(sSite) <- log_sSite
for(i in 1:nSite) {
bSite[i] ~ dnorm(0, sSite^-2)
}
bRkm ~ dnorm(0, 5^-2)
log_sDispersion ~ dnorm(0, 5^-2)
log(sDispersion) <- log_sDispersion
for(i in 1:length(Count)) {
log(eDensity[i]) <- bSystemYear[System[i],Year[i]] + bRkm * Rkm[i] + bSite[Site[i]]
eDispersion[i] ~ dgamma(sDispersion^-2, sDispersion^-2)
Count[i] ~ dpois(eDensity[i] * Length[i] * Coverage[i] * bEfficiency * eDispersion[i])
}
Template 4. The final model.
model {
log_bAlpha ~ dnorm(5, 5^-2)
log(bAlpha) <- log_bAlpha
for(i in 1:nSystem) {
log_bBeta[i] ~ dnorm(0, 5^-2)
log(bBeta[i]) <- log_bBeta[i]
}
log_sRecruits ~ dnorm(0, 5^-2)
log(sRecruits) <- log_sRecruits
for(i in 1:length(Recruits)){
eRecruits[i] <- bAlpha * Stock[i] / (1 + bBeta[System[i]] * Stock[i])
Recruits[i] ~ dlnorm(log(eRecruits[i]), sRecruits^-2)
}
bCarryingCapacity <- bAlpha / bBeta
Template 5. Final model.