Detecting historical population structure among highly impacted White Sturgeon populations of the Upper Columbia River

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Detecting historical population structure among highly impacted White Sturgeon populations of the Upper Columbia River Dr. R. John Nelson University of Victoria Victoria, British Columbia, Canada

Acispenserformidae Sturgeon and paddlefish 27 Species in the Northern Hemisphere

White Sturgeon Largest fish in North American freshwater To 6 m in length - 880 kg - over 100 years old

White Sturgeon Life history Slow growing In upper Columbia River females mature at about 30 years and males at about 20 years. Spawns every 4 to 5 years. Females lay 100,000 to 1,000,000 eggs. Eggs sink to bottom and hatch in about 1 week After hatching larvae drift downstream until they reach suitable habitat. Feed on invertebrates and fishes.

White Sturgeon - Range Alaska to California and is found in lakes and rivers as well as brackish. and marine waters.

In Canada only in British Columbia Currently six recognized populations

Range in Western British Columbia

Range in Eastern British Columbia

White Sturgeon - Human Use Commercial, Native and recreational fisheries. Commercial catch statistics Fraser River, 1880 to 1993. (Figure from Echols 1995).

White Sturgeon other impacts

White Sturgeon All harvest in British Columbia banned since 1994. Catch and release fishery allowed in Lower Fraser River Some sport fishing activities involve research - impact of catch and release not understood.

White Sturgeon Population Status All populations have declined from historical levels Lower Fraser - 47,000 Mid-Fraser 3,700 Upper Fraser 850 Nechako 571 Kootenay 760 Upper Columbia 1500 Endangered Endangered Endangered Endangered Endangered = facing imminent extirpation or extinction (data from Golder 2006, Irvine et al. 2007)

Upper Columbia River Historic geographic context Historically fish had access to over 600 km of river/lake habitat Now river impounded by 4 major dams

Keenlyside dam completed in 1968

Keenlyside dam - HLK 185 megawatts

Waneta Dam completed in 1954

Waneta Dam 450 megawatts

Brilliant Dam completed in 1944

Brilliant Dam 145 megawatts

Grand Coulee Dam completed in 1942

Grand Coulee Dam 6800 megawatts

After completion of Keenlyside dam in 1968 widespread recruitment failure..

BC Ministry of Environment Fisheries Management Data 100 80 Unders Midsize Oversiz 65% 60 6 Percent 40 33 24% 20 11% 0 3% 1981-199 Changes in WS size composition = recruitment failure beginning ~ 1970

Spawning occurs but juveniles are not being recruited into the population - cause uncertain

White sturgeon recovery initiative started 2001 First recovery plan produced in 2002 collaboration between govt, communities, USA and Canada.

UCR currently considered as one population

White sturgeon UCR Five principle habitat areas within the Upper Columbia Recovery Area. AR HLK BRL WAN - ROOS (Hildebrand et al., 1999)

Population Structuring? Knowing population units important for determining cause of recruitment failure and for management and recovery planning. How can we gain insight into population structure, historical and contemporary? Genetics can provide insight.

Defining the Question. In the approximately 10,000 years since colonization, did white sturgeon in this region segregate into reproductively isolated populations?

Defining the Question. If there was genetic structure could we detect it given change to water flows and potential blockage of migration routes and behaviour?

White Sturgeon genome Octoploid nuclear genome 250 chromosomes Nuclear genetic markers such as microsatellites - usually highly variable (require large samples) can be difficult to score and analyse in complex genomes.

Mitochondrial genome haploid, easy to analyse. PCR amplification Avoiding complex Repeated region. First we sequenced region and did Restriction Fragment Length Polymorphism analysis in silico. Chose three restriction enzymes. SfcI, Hsp92II, MseI

Restriction Fragment Length Polymorphism old technique for ancient fish DNA extraction > PCR > enzyme digestion > gel electrophoresis SfcI Hsp92II MseI + + = 9 haplotypes

First genetic results.. Kootenay Lake population was almost monomorphic for haplotype 2 and this distinguished this group from the others.

Genetic structure suggested between other sites but Work was done with samples of mixed age catch location was not well defined How was age of the fish affecting the results?? Could we use catch location data to more rigorously define the samples? Roosevelt fish?

Setting up the analysis. Age of fish. Recruitment failure began around 1970. Pre- and post- recruitment failure fish are present in the system. Habits of older fish more likely to reflect historical life history i.e. population structure. Approach: remove post dam fish from genetic analysis

Setting up the analysis. Age of fish. Define pre-recritment failure fish as those born prior to 1968. Ageing done by fin ray analysis as per Brennan and Cailliet 1989. If fin ray information was lacking: Age = 0.0006 x FL 2 + 0:0862 x FL + 3.9914 Application of this criteria resulted in the removal of 102 fish from the analysis.

Setting up the analysis. Use Tagging and recapture data to define sample location evaluated for HLK, BRL, WAN.

Tagging and recapture

Tagging and recapture

Recapture and read.

Tagging and recapture - stats Database maintained by UCWSRI From 1993 to 2006. 607 fish - 802 captures. 270 fish captured more than once. 249 fish were caught exclusively within one zone. Equal captures in multiple locations occurred for 22 fish.

Tagging and recapture assignment to high use area evaluated for HKD, BRL, WAN. Fish assigned to area most frequently captured Fish with equal number of detections pooled and randomly split Fish with detections that could not be evenly split were removed.

Results of Capture data HLK (110) BRL (15) WAN (145) 84 % of subsequent detections in same locations (p<0.001)

Tagging and recapture 84 % of subsequent detections in same location (p<0.001) 92% HLK 64.5 % BRL (lower fidelity may be due to low water in summer) 93.4% WAN

Sonic telemetry evaluated for HKD, BRL, WAN, ROOS.

Sonic telemetry

Sonic telemetry

Sonic telemetry.

Sonic telemetry- stats 73 fish were monitored for 1.5 4 years between 2002 and 2007 Array of 26 sonic detectors fixed locations (see Golder Associates 2006, Howell & McLellan 2006). 40, 278 location x day detections

Sonic telemetry assignment to high use area evaluated for HKD, ROOS, WAN, BRL..

Results of telemetry data HLK (18) BRL (5) WAN (7) 90 % of subsequent detections in same locations (P<0.001)

Results of telemetry data ROOS(43) 90 % of subsequent detections in same locations (P<0.001)

Sonic telemetry 90 % of subsequent detections in same locations (P<0.001) 94% - HLK 98% - BRL 95% - WAN 95% - ROOS

Haplotype frequencies N 1 2 3 4 5 6 7 8 9 AR 24 0.04 0.33 0.50 0.04 0.00 0.00 0.00 0.04 0.04 HLK 56 0.09 0.29 0.43 0.09 0.02 0.02 0.00 0.07 0.00 WAN 21 0.00 0.43 0.38 0.14 0.00 0.00 0.00 0.05 0.00 BRL 28 0.04 0.71 0.14 0.04 0.00 0.00 0.00 0.04 0.04 KL 28 0.00 0.96 0.04 0.00 0.00 0.00 0.00 0.00 0.00 ROOS 50 0.06 0.52 0.16 0.26 0.00 0.00 0.03 0.00 0.00 Most of the signal coming from haplotypes 2, 3 and 4. AR and HLK have more 3 than 2 KL has less diversity than all others

Haplotype frequencies

Pairwise differentiation test KL ( = 0.05). Can we reject the null hypothesis of genetic homogeneity? H o samples are the same AR HLK WAN BRL KL ROOS AR no no no + + + HLK no no no + + + WAN no no no no + no BRL + + no no 0.06 no KL + + + 0.06 no + ROOS + + no no + no KL differentiated from all others. F ST range from 0.1 to 0.46 P < 0.05

Bonnington Falls Historical obstacle to fish passage.

Pairwise differentiation test KL = 0.05 Can we reject the null hypothesis of genetic homogeneity? H o samples are the same AR HLK WAN BRL KL ROOS AR no no no + + + HLK no no no + + + WAN no no no no + no BRL + + no no 0.06 no KL + + + 0.06 no + ROOS + + no no + no F ST KL-BRL = 0.1 P <0.05 BRL differentiated from KL

Brillant dam

BRL and KL linked by downward migration.

Pairwise differentiation AR and HLK Can we reject the null hypothesis of genetic homogeneity? AR HLK WAN BRL KL ROOS AR no no no + + + HLK no no no + + + WAN no no no no + no BRL + + no no 0.06 no KL + + + 0.06 no + ROOS + + no no + no F ST = 0.0 AR and HLK samples not different.

AR and HLK samples not different. HLK dam broke up a population that spawned up river. Spawing observed in HLK.

Pairwise differentiation AR and HLK Can we reject the null hypothesis of genetic homogeneity? AR HLK WAN BRL KL ROOS AR no no no + + + HLK no no no + + + WAN no no no no + no BRL + + no no 0.06 no KL + + + 0.06 no + ROOS + + no no + no AR and HLK unique population (sample size of WAN may be limiting test) F ST 0.09 to 0.46 P<0.05

ROOS.

Pairwise differentiation ROOS Can we reject the null hypothesis of genetic homogeneity? AR HLK WAN BRL KL ROOS AR no no no + + + HLK no no no + + + WAN no no no no + no BRL + + no no 0.06 + KL + + + 0.06 no + ROOS + + no + + no ROOS different than all but WAN. F ST range from 0.02 to 0.23

ROOS, WAN Unique population? Contraction of habitat may have caused mixing or introgression of mitochondrial DNA haplotypes prior to recruitment failure but old habits retained.

Haplotype frequencies to calculate genetic distance

Overall genetic structure CSE distance

Overall genetic structure

Overall genetic structure

Overall genetic structure Due to habitat contraction?.

Interpretations: Genetics when combined with other approaches is very powerful. When fish dispersed to different areas within the region after initial colonization, genetic divergence occurred. Before the dams were built, AR and HLK fish could were one population that spawned above the HLK dam.

Interpretations. ROOS and WAN -Contraction of habitat may have caused mixing or introgression of mitochondrial DNA haplotypes prior to recruitment failure but old habits retained. BRL linked to KL by unidirectional migration.

Overall Interpretation Even in severely disrupted habitats with declining populations fishes may retain genetic structure and habits. Genetics here used to detect homogenization, disruption and unidirectional migration. Such information will be useful as a basis for future study and design of recovery plans.

Acknowledgements BC Hydro and BC Ministry of Environment for funding. Jason McLellan for samples and telemetry data. Glenn Cooper, Kathryn Clark, Chantal Rajotte for lab expertise.

Thank you.questions?