How small can isolated brook trout populations become and still respond to environmental change?

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Dylan J. Fraser Department of Biology, Concordia University Research Collaborators: Dr. Jacquelyn Wood Dr. Paul Debes Matthew Yates (PhD candidate) Thais Bernos (MSc candidate) Zachery Wells (MSc candidate) How small can isolated brook trout populations become and still respond to environmental change?

Habitat loss and fragmentation, climate change Small population size (N) Environmental stochasticity Demographic stochasticity

Small population size (N e ) Genetic stochasticity Genetic drift, inbreeding Inbreeding depression

Brook trout Waterton Lakes N.P. Newfoundland Algonquin Park Northern Quebec

Habitat fragmentation, population size and adaptive potential: brook trout, Cape Race NL Common ancestor, pristine Cripple Cove Wood et al. 2014 Ecol Appl; Wood & Fraser 2015 Ecology

Brook trout, Cape Race NL Population demarcation and isolation Comprehensive sampling Belmar et al. 2012 Eco/Evo, Wood et al. 2014 Ecol Appl, Fraser et al. 2014 Proc R Soc Lond Biol Sci

Brook trout, Cape Race NL Population differentiation N = adult census size Nb = effective number of breeders 2Nb to 3Nb ~ Ne for brook trout 100 fold difference in N 15 fold difference in Nb 10 fold..allelic richness (microsatellites) 5 to 12 fold..heterozygosity (microsatellites/snps) 4 fold..mean adult size 6 fold..mean egg size Mean age at maturity: 2.1 3.3yrs Body shape Colouration Hermitage Pond Watern Cove River Behaviour N Nb = 2 5 N == 50 60, 6000 10000, Nb = 180 400

1) Brook trout population assessments: links between habitat size, N and Nb N = adult census size Nb = effective number of breeders 2Nb to 3Nb ~ Ne for brook trout Wood et al. (2014) Ecological Applications; Bernos & Fraser in prep

2) Summer thermal refuge + groundwater seeps = brook trout Typically acidic bogs Dried-up stream bed Seep Upstream of seep Spawning aggregate on seep

3) Many habitats are associated with small population size Habitat 14-19 populations Transects (n=875): ph Dissolved oxygen Conductivity Temperature Channel width Mean depth Mean undercut depth Mean velocity % riparian cover % in-stream vegetation Species of vegetation % large boulder % small boulder % cobble % course gravel % fine gravel % silt Population size N: Schnabel or Petersen Tagging in summer Recapture a week later Nb: >6000 trout @ 12 microsatellites Cohorts (age 0+) for 2009-2014 Population designation (Structure, FST) Wood, et al. 2014, Ecological Applications; Bernos & Fraser 2015, in prep.

Greater variation in (i) mean habitat variables and (ii) around habitat means, across small than large populations Statistically more spread at 8 of 17 habitat variables 2010 2011 Wood, et al. 2014, Ecological Applications

4) Small populations exhibit considerable adaptive genetic differentiation Small popn pairs Large popn pairs Percentage of SNPs under selection 14 populations Nb 11-249, N = 73-8416 Fraser et al. 2014 Proc R Soc

5) Small populations retain genetic variation at fitnessrelated quantitative traits Additive genetic variance (VA) (heritable genetic variation that selection acts on) 9 popn common garden experiment (30-75 half-sib families per popn; median 52) Different aspects of phenotype 3 behavioural traits 5 early life history traits 7 morphological traits Wood et al. 2015 Evolution

Population size and VA Wood et al. 2015 Evolution

6) Can small brook trout populations respond ally to climate warming? Disperse (To better habitat) Evolve (Genetic change) Be (No genetic change) Often not possible Requires Genetic diversity Trait expression Temp 1 Temp 2

6) Small and large populations have similar responses to elevated temperature Magnitude of ity 8 populations, common garden (6-30 full-sib families per popn; median 18) Three incubation temps, six traits 5 C ( normal ); 7 C, 9 C (climate warming) normal climate warming Small effect sizes, low upper CI Wood & Fraser Ecology

General conclusions: there is some hope for small brook trout populations Populations with 100 adults can be as genetically robust as populations with 1000-10000 adults Environmental stochasticity >> genetic stochasticity: habitat quality is key Adaptive population differentiation at fine scales and at small population sizes

Twitter: @thefraserlab dylan.fraser@concordia.ca Thanks! Questions? Funding, logistical support: Mistaken Point Ecological Reserve

Magnitude of ity Reduced? Less Less Less Less Increased? More More More More Trait expression Family A Family B Temp 1 Temp 2 Absolute values of the slopes More variable responses? Less Less Magnitude and directionality (+/-) of within-population family slopes Residual variance of the slopes Variability in ity More Similarly Popn size

Replicated Recapture transplants survivors from to new ponds environments in Fall

Responding to environmental change: driving factors in nature 9 popns, 5 years, ~30 (total) translocations/year Contemporary population size (5 year time series)? Standing genetic variation (genome-wide)? Population history of environmental variability? Extent of environmental change (between home and new environment)?

Responding to environmental change: driving factors in nature Variables affecting survival Z-value Change in ph 4.4*** Change in veg. - 2.8** Change in temp -3.3*** Extent of environmental change matters most Genetic factors: little if any role? Initial body size 2.5* *<0.01, **<0.005, ***<0.001 Best fit models do not include genetic factors (or interaction terms with genetic factors)