Horse Lake Exploitation Analysis 2019

The suggested citation for this analytic report is:

Dalgarno, S. and Thorley, J.L. (2019) Horse Lake Exploitation Analysis 2019. A Poisson Consulting Analysis Appendix. URL: https://www.poissonconsulting.ca/f/248424047.

Background

Horse Lake supports a recreational fishery for Lake Trout (Salvelinus namaycush). To estimate their natural and fishing mortality, individuals were caught by angling and tagged with acoustic transmitters and/or external reward tags. Acoustically tagged fish were detected by a series of seven receivers distributed through the lake as well as annual mobile receiver surveys. The external reward tags included a phone number for anglers to report their recaptures. In 2019, eight Rainbow Trout with acoustic transmitters and external reward tags were transplanted from Greenlee Lake.

All the data were provided by the Ministry of Forests, Lands and Natural Resource Operations (MFLNRO) Cariboo Region.

Methods

Data Preparation

The data were prepared for analysis using R version 3.6.1 (R Core Team 2019). Receivers were assumed to have a detection range of 500 m. Detections were aggregated daily, where for each transmitter the receiver with the most detections was chosen. In the case of a tie, the receiver with the greatest coverage area was chosen.

The Seasons were Winter (December - February), Spring (March - May), Summer (June - August) and Autumn (September - November).

Data Analysis

Hierarchical Bayesian models were fitted to the data using R version 3.6.1 (R Core Team 2019) and JAGS 4.2.0 (Plummer 2015) which interfaced with each other via the jmbr package. For additional information on hierarchical Bayesian modelling in the BUGS language, of which JAGS uses a dialect, the reader is referred to Kery and Schaub (2011, 41–44).

Unless indicated otherwise, the Bayesian analyses used normal and half-normal prior distributions that were vague in the sense that they did not constrain the posteriors (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 \(\hat{R} \leq 1.05\) (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 (Brooks et al. 2011).

The posterior distributions of the fixed (Kery and Schaub 2011, 75) 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). The estimate is the median (50th percentile) of the MCMC samples, the z-score is \(\mathrm{sd}/\mathrm{mean}\) 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.

The results are displayed graphically by plotting the modeled relationships between particular variables and the response 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). Where informative the influence of particular variables is expressed in terms of the effect size (i.e., percent change in the response variable) with 95% CIs (Bradford, Korman, and Higgins 2005).

Model Descriptions

Condition

The expected weight of a fish of a given length was estimated from the data using a allometric mass-length model (He et al. 2008)

\[ L = \alpha W^\beta\]

Key assumptions of the condition model include:

  • The prior distribution on the power term (\(\beta\)) is a normal distribution with a mean of 3.18 and a standard deviation of 0.19 (McDermid, Shuter, and Lester 2010).
  • The residual variation in weight is log-normally distributed.

Growth

Growth was estimated from length and scale age data for Horse Lake using a Von Bertalanffy growth curve model (von Bertalanffy 1938).

\[ L = L_\infty \cdot (1 - \exp(-k \cdot (A - t_0) ))\]

where \(A\) is the scale age in years and \(t_0\) is the extrapolated age at zero length.

Key assumptions of the growth model include:

  • The residual variation in length is normally distributed.

1 fish with a fork length \(\geq\) 500 mm was excluded from the analysis.

Survival

The natural mortality and capture probability for individuals \(\geq\) 300 mm was estimated using a Bayesian individual state-space survival model (Thorley and Andrusak 2017) with monthly intervals. The survival model incorporated natural and handling mortality, acoustic detection, movement, T-bar tag loss, recapture and reporting. Key assumptions of the survival model include:

  • The tagged individuals are a random sample from the population.
  • The only effects of tagging are change in the natural mortality for three months following acoustic tagging.
  • The prior probability on the annual interval individual external tag loss rate is a uniform distribution between 1 and 30% (Fabrizio et al. 1996).
  • All individuals are correctly identified.
  • There is no emigration out of the lake.
  • There are no unmodelled individual differences in the probability of recapture in each time interval.
  • There are no unmodelled individual differences in the probability of survival in each time interval.
  • There are no unmodelled individual differences in the probability of a living individual with an active acoustic transmitter being detected moving among sections in each time period.
  • The fate of each individual is independent of the fate of any other individual.
  • The sampling periods are instantaneous and all individuals are immediately released.

The survival model treats all recaptured fish as if they had been retained. Only individuals with a fork length (FL) \(\geq\) 300 mm that were tagged with an acoustic tag and/or $100 and $10 reward tags were included in the survival analysis. Preliminary analysis indicated that the difference between the natural mortality and fishing mortality rates for individuals \(\geq\) 500 mm were not sigificant.

Yield-Per-Recruit

The optimal capture rate (to maximize number of individuals harvested assuming 100% retention) was calculated using yield-per-recruit analysis (Walters and Martell 2004). A separate yield-per-recruit analysis was performed for the two different Lake Trout ecotypes in Horse Lake.

Results

Templates

Condition

.model {
  bWeight ~ dnorm(0, 1^-2) 
  bWeightLength ~ dnorm(3.18, 0.19^-2)

  sWeight ~ dnorm(0, 1^-2) T(0,) 
  for(i in 1:length(LogLength)) {
    eWeight[i] <- bWeight + bWeightLength * LogLength[i]
    Weight[i] ~ dlnorm(eWeight[i], exp(sWeight)^-2)
  }
..

Block 1. Condition model description.

Growth

.model {
  bLInf ~ dnorm(500, 100^-2) T(0, )
  bK ~ dnorm(0, 1^-2) T(0, )
  bT0 ~ dnorm(0, 1^-1)

  sLength ~ dnorm(0, 100^-2) T(0, )
  for (i in 1:length(Length)) {
    eLength[i] <- bLInf * (1 - exp(-bK * (Age[i] - bT0))) 
    Length[i] ~ dnorm(eLength[i], sLength^-2) T(0, )
  }
..

Block 2. Growth model description.

Survival

.model{
  bMortality ~ dnorm(-3, 3^-2)
  bMortalityHandling ~ dnorm(0, 2^-2)
  bTagLoss ~ dunif(1 - (1 - 0.01)^(1/12), 1 - (1 - 0.30)^(1/12))
  bDetected ~ dunif(0, 1)
  bMoved ~ dunif(0, 1)
  bRecaptured ~ dunif(0, 1 - (1 - 0.50)^(1/12))
  bReported ~ dunif(0.9, 1.00)

  for (i in 1:nCapture){
    logit(eMortality[i,PeriodCapture[i]]) <- bMortality + bMortalityHandling
    eTagLoss[i,PeriodCapture[i]] <- bTagLoss
    eDetected[i,PeriodCapture[i]] <- bDetected
    eMoved[i,PeriodCapture[i]] <- bMoved
    eRecaptured[i,PeriodCapture[i]] <- bRecaptured
    eReported[i,PeriodCapture[i]] <- bReported
    InLake[i,PeriodCapture[i]] <- 1
    Alive[i,PeriodCapture[i]] ~ dbern(1-eMortality[i,PeriodCapture[i]])
    TBarTag100[i,PeriodCapture[i]] ~ dbern(1-eTagLoss[i,PeriodCapture[i]])
    TBarTag10[i,PeriodCapture[i]] ~ dbern(1-eTagLoss[i,PeriodCapture[i]])
    Detected[i,PeriodCapture[i]] ~ dbern(Monitored[i,PeriodCapture[i]] * eDetected[i,PeriodCapture[i]])
    Moved[i,PeriodCapture[i]] ~ dbern(Alive[i,PeriodCapture[i]] * Monitored[i,PeriodCapture[i]] * eMoved[i,PeriodCapture[i]])
    Recaptured[i,PeriodCapture[i]] ~ dbern(Alive[i,PeriodCapture[i]] * eRecaptured[i,PeriodCapture[i]])
    Reported[i,PeriodCapture[i]] ~ dbern(Recaptured[i,PeriodCapture[i]] * eReported[i,PeriodCapture[i]] * step(TBarTag100[i,PeriodCapture[i]] + TBarTag10[i,PeriodCapture[i]] - 1))
  for(j in (PeriodCapture[i]+1):nPeriod) {
    logit(eMortality[i,j]) <- bMortality + bMortalityHandling * step(PeriodCapture[i] - j + 2)
    eTagLoss[i,j] <- bTagLoss
    eDetected[i,j] <- bDetected
    eMoved[i,j] <- bMoved
    eRecaptured[i,j] <- bRecaptured
    eReported[i,j] <- bReported
    InLake[i,j] ~ dbern(InLake[i,j-1] * (1-Recaptured[i,j-1]))
    Alive[i,j] ~ dbern(Alive[i,j-1] * (1-Recaptured[i,j-1]) * (1-eMortality[i,j-1]))
    TBarTag100[i,j] ~ dbern(TBarTag100[i,j-1] * (1-Recaptured[i,j-1]) * (1-eTagLoss[i,j]))
    TBarTag10[i,j] ~ dbern(TBarTag10[i,j-1] * (1-Recaptured[i,j-1]) * (1-eTagLoss[i,j]))
    Detected[i,j] ~ dbern(InLake[i,j] * Monitored[i,j] * eDetected[i,j])
    Moved[i,j] ~ dbern(Alive[i,j] * Monitored[i,j] * eMoved[i,j])
    Recaptured[i,j] ~ dbern(Alive[i,j] * eRecaptured[i,j])
    Reported[i,j] ~ dbern(Recaptured[i,j] * eReported[i,j] * step(TBarTag100[i,j] + TBarTag10[i,j] - 1))}}
  }

Block 3. Survival model description.

Tables

Captures

Table 1. Number of fish captured by year, species and size class. Eight Rainbow Trout were transplanted in 2019.

Year Species < 300 mm 300 - 500 mm >= 500 mm Total
2017 Kokanee 3 20 0 23
2017 Lake Trout 1 153 7 161
2017 Rainbow Trout 0 3 0 3
2018 Kokanee 0 2 0 2
2018 Lake Trout 4 176 19 199
2019 Lake Trout 3 154 7 164
2019 Rainbow Trout 0 8 0 8
Total 11 516 33 560

Table 2. Number of Lake Trout captured by size class, and tag class. Any fish > 500mm is classified as ‘large’.

Year Size Acoustic and T-Bar T-Bar Only No Tag Total
2017 < 300 mm 0 1 0 1
2017 300 - 500 mm 29 113 11 153
2017 >= 500 mm 3 2 2 7
2018 < 300 mm 0 2 2 4
2018 300 - 500 mm 43 103 30 176
2018 >= 500 mm 11 4 4 19
2019 < 300 mm 0 3 0 3
2019 300 - 500 mm 56 98 0 154
2019 >= 500 mm 4 3 0 7
Total 146 329 49 524

Recaptures

Table 3. All Lake Trout recaptures to date. Recaptures with natural mortality were tags found in the stomach of another fish!.

Date Capture Fork Length (mm) Date Recapture External Tag Number 1 External Tag Number 2 Harvested Natural Mortality
2017-06-06 445 [mm] 20 17-07-07 278 544 Ye s No
2017-05-20 405 [mm] 20 17-09-08 86 337 Ye s No
2017-06-01 441 [mm] 20 18-01-29 289 532 Ye s No
2017-05-31 385 [mm] 20 18-02-02 441 NA Ye s No
2017-06-07 439 [mm] 20 18-02-08 413 258 Ye s No
2018-05-25 400 [mm] 20 18-05-28 1390 2315 Ye s No
2017-05-18 360 [mm] 20 18-05-29 326 76 No No
2018-05-29 379 [mm] 20 18-05-29 151 445 No No
2017-05-31 357 [mm] 20 18-05-29 151 445 No No
2018-05-25 436 [mm] 20 18-06-28 1391 2316 Ye s No
2018-05-30 341 [mm] 20 18-07-03 2370 NA Ye s Ye s
2018-06-11 406 [mm] 20 18-07-03 136 552 Ye s Ye s
2017-05-21 370 [mm] 20 18-07-05 340 89 Ye s No
2017-06-07 416 [mm] 20 18-07-16 404 251 Ye s No
2018-05-25 334 [mm] 20 18-07-18 2320 1395 Ye s No
2018-05-30 416 [mm] 20 18-07-24 2368 1335 No No
2017-06-02 360 [mm] 20 18-09-10 538 NA No No
2018-05-31 411 [mm] 20 19-01-15 2386 362 Ye s No
2018-06-08 382 [mm] 20 19-01-25 2340 377 Ye s No
2017-05-18 360 [mm] 20 19-03-09 76 326 Ye s No
2018-06-07 325 [mm] 20 19-03-09 2327 390 Ye s No
2017-05-30 402 [mm] 20 19-03-31 56 512 No No
2018-06-11 411 [mm] 20 19-05-07 556 140 Ye s No
2019-05-23 421 [mm] 20 19-05-23 129 462 No No
2018-05-29 356 [mm] 20 19-06-06 1347 2354 Ye s No
2017-06-01 421 [mm] 20 19-06-06 291 530 Ye s No
2017-05-29 429 [mm] 20 19-06-06 70 523 Ye s No
2017-06-06 405 [mm] 20 19-06-07 547 275 Ye s No
2017-06-06 343 [mm] 20 19-06-11 270 424 Ye s No
2018-05-28 421 [mm] 20 19-06-11 221 1321 No No
2019-06-11 474 [mm] 20 19-06-11 221 1321 No No
2018-06-07 386 [mm] 20 19-06-12 2326 391 No No
2019-06-12 389 [mm] 20 19-06-12 2326 391 No No
2018-06-11 410 [mm] 20 19-07-01 137 553 Ye s No
2017-05-29 425 [mm] 20 19-07-01 65 519 Ye s No
2017-05-30 480 [mm] 20 19-07-04 167 429 Ye s No
2019-05-23 377 [mm] 20 19-07-06 46 596 Ye s No
2018-06-11 407 [mm] 20 19-07-07 142 558 Ye s No
2018-06-08 394 [mm] 20 19-07-15 149 565 Ye s No
2018-05-25 395 [mm] 20 19-07-22 2321 1396 Ye s No
2018-10-24 592 [mm] 20 19-07-28 134 466 Ye s No
2019-06-06 349 [mm] 20 19-08-13 186 1490 Ye s No
2019-06-06 500 [mm] 20 19-09-04 187 1491 Ye s No
2017-05-31 450 [mm] 20 19-09-05 500 201 Ye s No
2018-05-29 415 [mm] 20 19-09-05 210 1310 Ye s No

Condition

Table 4. Parameter descriptions.

Parameter Description
bWeight Intercept of eWeight
bWeightLength Effect of Length on bWeight
eWeight Log expected Weight
LogLength Log-transformed and centered fork length (mm)
sWeight Standard deviation of residual variation in eWeight
Weight Mass (kg)

Table 5. Model coefficients.

term estimate sd zscore lower upper pvalue
bWeight -0.0036990 0.1670595 -0.036993 -0.3419239 0.3149792 0.9773
bWeightLength 3.1710022 0.1832047 17.325243 2.8035274 3.5389078 0.0007
sWeight 0.0193256 0.0294564 0.984291 0.0008586 0.1062319 0.0007

Table 6. Model summary.

n K nchains niters nthin ess rhat converged
38 3 3 500 10 1304 1 TRUE

Growth

Table 7. Parameter descriptions.

Parameter Description
Age[i] Scale age at capture (yr)
bK Growth coefficient (mm/yr)
bLinf Mean maximum length (mm)
bT0 Age at zero length (yr)
Length[i] Fork length at capture (mm)
sLength SD of residual variation in Length

Table 8. Model coefficients.

term estimate sd zscore lower upper pvalue
bK 0.2235899 0.0414649 5.4716353 0.1561642 0.3193449 0.0007
bLInf 449.7879798 14.0211403 32.1310130 425.7101733 479.2432565 0.0007
bT0 -0.2442682 0.7104428 -0.3837309 -1.7881108 1.0088049 0.7267
sLength 22.3656309 3.1870066 7.1616761 17.6684189 30.1849697 0.0007

Table 9. Model summary.

n K nchains niters nthin ess rhat converged
32 4 3 500 100 1270 1.001 TRUE

Survival

Table 10. Parameter descriptions.

Parameter Description
bDetected Monthly probability of detection if inlake
bMortality Log odds monthly probability of dying of natural causes
bMortalityHandling Effect of capture and handling on bMortality
bMoved Monthly probability of being detected moving between sections if alive
bRecaptured Monthly probability of being recaptured if alive
bReported Monthly probability of being reported if recaptured with one or more T-bar tags
bTagLoss Monthly probability of loss for a single T-bar tag

Table 11. Model coefficients.

term estimate sd zscore lower upper pvalue
bDetected 0.8966457 0.0066512 134.794960 0.8826788 0.9096324 0.0007
bMortality -5.1998574 0.3843645 -13.618153 -6.0669245 -4.5710811 0.0007
bMortalityHandling 0.5746340 0.5599843 1.018155 -0.5624161 1.6418533 0.3200
bMoved 0.8539965 0.0081915 104.235398 0.8374648 0.8696397 0.0007
bRecaptured 0.0061447 0.0009421 6.590763 0.0045532 0.0081882 0.0007
bReported 0.9844942 0.0200206 48.858289 0.9216218 0.9993276 0.0007
bTagLoss 0.0028539 0.0015934 1.981170 0.0010039 0.0071287 0.0007

Table 12. Model summary.

n K nchains niters nthin ess rhat converged
14130 7 3 500 300 135 1.027 FALSE

Yield-per-Recruit

Table 13. Summary of parameters in yield-per-recruit model.

Parameter Ecotype Estimate Description Source
tmax Small 3.00e+01 The maximum age (yr). Current Study
k Small 2.24e-01 The VB growth coefficient (yr-1). Current Study
Linf Large 7.50e+01 The VB mean maximum length (cm). Professional Judgement
Linf Small 4.50e+01 The VB mean maximum length (cm). Current Study
t0 Small -2.44e-01 The (theoretical) age at zero length (yr). Current Study
k2 Small 1.50e-01 The VB growth coefficient after length L2 (yr-1). Default
Linf2 Small 1.00e+02 The VB mean maximum length after length L2 (cm). Default
L2 Small 1.00e+03 The length (or age if negative) at which growth switches from the first to second phase (cm or yr). Default
Wb Small 3.17e+00 The weight (as a function of length) scaling exponent. Current Study
Ls Large 5.00e+01 The length (or age if negative) at which 50 % mature (cm or yr). Professional Judgement
Ls Small 3.75e+01 The length (or age if negative) at which 50 % mature (cm or yr). Spin Gillnetting
Sp Small 1.00e+02 The maturity (as a function of length) power. Default
es Small 1.00e+00 The annual probability of a mature fish spawning. Default
Sm Small 0.00e+00 The spawning mortality probability. Default
fb Small 1.00e+00 The fecundity (as a function of weight) scaling exponent. Default
tR Small 1.00e+00 The age from which survival is density-independent (yr). Default
BH Small 1.00e+00 Recruitment follows a Beverton-Holt (1) or Ricker (0) relationship. Default
Rk Small 2.50e+01 The lifetime spawners per spawner at low density. Myers et al. 1999
n Small 1.50e-01 The annual interval natural mortality rate from age tR. Default
nL Small 6.39e-02 The annual interval natural mortality rate from length Ln. Current Study
Ln Small 5.00e+01 The length (or age if negative) at which the natural mortality rate switches from n to nA (cm or yr). Default
Lv Small 3.50e+01 The length (or age if negative) at which 50 % vulnerable to harvest (cm or yr). Professional Judgement
Vp Small 2.50e+01 The vulnerability to harvest (as a function of length) power. Professional Judgement
Llo Small 0.00e+00 The lower harvest slot length (cm). Default
Lup Small 1.00e+02 The upper harvest slot length (cm). Default
Nc Small 0.00e+00 The slot limits non-compliance probability. Default
pi Small 7.13e-02 The annual capture probability. Current Study
rho Small 0.00e+00 The release probability. Default
Hm Small 0.00e+00 The hooking mortality probability. Default
Rmax Small 1.00e+00 The number of recruits at the carrying capacity (ind). Default
Wa Small 5.64e-03 The (extrapolated) weight of a 1 cm individual (g). Current Study
fa Small 1.00e+00 The (theoretical) fecundity of a 1 g female (eggs). Default
q Small 1.00e-01 The catchability (annual probability of capture) for a unit of effort. Default

Table 14. Lake Trout large ecotype yield-per-recruit calculations including the capture probability (pi), exploitation rate (u) and average age, length and weight of fish harvested.

Type pi u Yield Age Length Weight Effort
actual 0.0712952 0.0712952 0.2996335 8.921062 58.59212 2608.401 0.7020118
optimal 0.1595423 0.1595423 0.3731851 6.514245 53.53010 1984.422 1.6496570

Table 15. Lake Trout small ecotype yield-per-recruit calculations including the capture probability (pi), exploitation rate (u) and average age, length and weight of fish harvested.

Type pi u Yield Age Length Weight Effort
actual 0.0712952 0.0712952 0.1185490 10.578893 39.61731 673.1236 0.7020118
optimal 0.3541295 0.3541295 0.2344587 8.064988 37.37973 557.0976 4.1491465

Figures

Captures

figures/capture/CaptureHistogram.png
Figure 1. Lake Trout captures by fork length, year and tag type.
figures/capture/CaptureMap.png
Figure 2. Lake Trout capture location by tagging.
figures/capture/CaptureRate.png
Figure 3. Capture rate by species and date for fish with fork length > 300 mm.

Coverage

figures/receiver/ReceiverMap.png
Figure 4. Location of sites with deployed receivers and estimated coverage (500m radius). Although receivers move slightly over time and after being redeployed, the centroid of all known locations for each receiver is shown.

Detections

figures/detection/DetectionOverview.png
Figure 5. Date of detections by species for each acoustically tagged fish. Grey segments indicate estimated tag life from capture date. Detections have been aggregated daily.
figures/detection/DetectionSeason.png
Figure 6. Percent of Lake Trout detections by receiver group and season.

Condition

figures/condition/condition.png
Figure 7. Estimated weight by length (with 95% CIs).

Growth

figures/growth/growth.png
Figure 8. Estimated length by age (with 95% CIs).

Survival

figures/survival/annual.png
Figure 9. The estimated annual interval probabilities (with 95% CIs).

Yield-per-Recruit

figures/yield/AgeLength.png
Figure 10. Lake Trout assumed age by length and ecotype.
figures/yield/AgeWeight.png
Figure 11. Lake Trout assumed length by weight and ecotype.
figures/yield/LengthSpawning.png
Figure 12. Lake Trout assumed length by spawning and ecotype.
figures/yield/LengthVulnerability.png
Figure 13. Lake Trout assumed length by vulnerability to capture and ecotype.
figures/yield/WeightFecundity.png
Figure 14. Lake Trout assumed weight by fecundity and ecotype.
figures/yield/Yield.png
Figure 15. Lake Trout calculated yield as percent of total unfished population by annual interval fishing mortality rate and ecotype.

Recommendations

Recommendations include:

  • Continue annual mobile receiver surveys.

Acknowledgements

The organisations and individuals whose contributions have made this analysis report possible include:

  • MFLNRO - Cariboo Region
    • Russ Bobrowski
    • Scott Horley

References

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