Is interbreeding of wild and artificially propagated animals prevented by divergent life history?

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Is interbreeding of wild and artificially propagated animals prevented by divergent life history? Todd R. Seamons, Lorenz Hauser, Kerry Naish and Tom Quinn School of Aquatic and Fishery Sciences University of Washington, Seattle, WA Evolutionary Applications doi:10.1111/j.1752-4571.2012.00247.x

Problems with hatchery supplementation Disease Overharvest of wild fish Behavioral and ecological Genetic Naish et al. 2007 Advances in Marine Biology

Methods to minimize or eliminate risks Use one of two approaches to broodstock management: Wild Wild Wild fish in broodstock Hatchery fish on spawning ground < 5% Hatchery Hatchery fish are integrated with wild fish Hatchery Hatchery fish are segregated from wild fish After Mobrand et al. 2005

Methods to minimize or eliminate risks Use artificial selection to increase separation of the populations Wild Hatchery < 5% Lorenzen et al. 2010 Hatchery fish are segregated from wild fish After Mobrand et al. 2005

Main aims of the study Determine whether segregation based on life history is effective Examine temporal trends in relative proportions of wild fish Estimate proportions of hatchery, wild and hybrid fish Evaluate explanatory variables/isolating mechanism

New hatchery steelhead propagation program starting 1996 Sampled fish since 1996 Weir to facilitate sampling hatcheryproduced and wild fish

Genetic and statistical methods Two methods of genetic assignment Mixture proportions via individual assignment (GENECLASS2) Baseline collections: Clipped and unclipped adults from 1996, 1997, and 1998 Hatchery F ST = 0.02 Wild Admixture proportions (STRUCTURE) Piry et al. 2004. J. Heredity Falush et al. 2003. Genetics Hauser et al. 2006. Mol. Ecol. Unclipped, natural origin of unknown ancestry Unknowns

Proportion of collections genetically identified as wild ancestry declines over time

Individual assignment criteria From GENECLASS2 with confident assignment Decision criteria Prob. of pure H or W > 0.95 Results in Wild Hatchery unassigned

Significant overlap in adult migration timing

Individual assignment criteria From GENECLASS2 with confident assignment Decision criteria Prob. of pure H or W > 0.95 Results in Wild Hatchery unassigned Who are these unassigned individuals? Are they hatchery/wild hybrids? Just assignment errors?

Estimate assignment test error rates From baseline collections simulated100,000 offspring genotypes each of Pure wild Pure hatchery F1 hybrid Run them through GENECLASS2 Using HYBRIDLAB, Nielsen et al. 2006. Molecular Ecology Notes.

Individual assignment error rates True ancestry (simulations) Estimated ancestry (GENECLASS2) Hatchery F1 hybrid Wild Hatchery 0.942 0.372 0.007 Unassigned 0.056 0.411 0.126 Wild 0.001 0.216 0.868 From these error rates and given some assumptions we can calculate the estimated proportions of hatchery, wild and hybrid individuals in our collections

Solve for corrected proportions Obs(hatchery) = 0.942x + 0.007y + 0.372z Obs(wild) = 0.001x + 0.868y + 0.216z Assignment test error rates Obs(unassigned) = 0.056x + 0.126y + 0.411z Obs(hatchery) Obs(wild) Obs(unassigned) Observed proportions Unknowns x = proportion hatchery y = proportion wild z = proportion hybrid

Solve for corrected proportions Obs(hatchery) = 0.942x + 0.007y + 0.372z Obs(wild) = 0.001x + 0.868y + 0.216z Assignment test error rates Obs(unassigned) = 0.056x + 0.126y + 0.411z Obs(hatchery) = 0.198 Obs(wild) = 0.475 Obs(unassigned) = 0.327 Example observed data: 1998 smolt collection Solve in R using matrix math for each yearly collection, adult and smolt

Critical assumption only hatchery, wild, or hybrid individuals sampled No strays Shouldn t be very many of these since steelhead are good homers No rainbow trout No evidence of a rainbow trout population No cutthroat Genetically identified and booted out No unclipped hatchery-produced fish (cheaters) Genetically identified (parentage) and booted out

Estimates of wild, hatchery, and hybrid proportions smolt collections

Estimates of wild, hatchery, and hybrid proportions smolt collections

Estimates of wild, hatchery, and hybrid proportions smolt collections

Evaluate possible explanatory variables/isolating mechanisms River discharge during hatchery spawning (Dec Feb) Possible isolating mechanism? USGS Willapa River stream gauge data Number of hatchery-produced adults on the spawning grounds Clipped fish counts at the hatchery Number of wild fish on the spawning grounds WDFW SaSI data for entire Willapa River

Willapa River discharge Willapa River mean daily discharge (m 3 s -1 )

Escapement estimates # hatchery fish at hatchery # adult steelhead WDFW redd count based estimate

Evaluate covariates methods Forward model selection analysis Add covariate to statistical model (also includes a temporal variable) Evaluate using information theoretic criteria (AIC c )

Best fit models explaining hatchery, wild, and hybrid proportions -sample year -discharge -discharge -discharge -sample year -proportion hatchery

Summary Wild fish declining over time Decline may be caused by Hybridization likely due to A constant influx of hatchery-produced adults and Variable environmental conditions Discharge likely negatively affects early spawner reproductive success But not an effective segregating mechanism Divergent migration/spawning timing not an effective segregating mechanism

Acknowledgements Faculty Lab and Field Paul Bentzen, Dalhousie U Lyndsay Newton Duy Mai Will Atlas Former Students Many others... Greg Mackey Jennifer McLean Michael Dauer Other contributions Ian Stewart, George Pess, Ian Taylor Martin Liermann, Thomas Beuhrens Hatchery Crew George Britter, WDFW Rob Allan, WDFW Dave Shores, WDFW Kevin Flowers, WDFW Jenny Allan Merle Hash Larry Sienko Funding Weyerhaeuser H. Mason Keeler Endowment Hatchery Science Reform Group National Science Foundation (DEB-9903914) Bonneville Power Administration (2003-050-00)