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Population and Evolutionary Genetic Analyses Using Multiple Nuclear and Mitochondrial Loci in Steller Sea Lions. John W. Bickham, Ryan M. Huebinger, Thomas R. Gelatt, Greg O’Corry-Crowe, and Joseph I. Hoffman ACKNOWLEDGEMENTS Project began in 1992 I thank the following for providing samples: T. Loughlin, T. Gelatt, V. Burkanov and D. Calkins Southeastern Alaska and PWSADF&G Gulf of Alaska and Aleutian Is.-NOAA sponsored cruises ACKNOWLEDGEMENTS Russia: Vladimir Burkanov, Nikolay Pavlov, Vladimir Vertynkin, Evgeny Mamaev. The Alaska SeaLife Center, Don Calkins For help in the laboratory and for data analysis I thank Alyson Baker, Robert Trujillo, John Patton, Cole Matson and Dmitri Politov. 200 Eastern Stock East of 144°W Population Estimates in Thousands 180 160 Western Stock West of 144°W 140 120 100 80 60 40 20 0 1960s 1970s 1980s 1990s 1994 Population Estimates for Steller Sea Lions (1960-1994) Population Trends The total SSL population numbered between 240,000 and 300,000 during the late 1950’s and early 1960’s. Only 116,000 were estimated range-wide in 1989. -1969- -1979- -1989- RUSSIA ALASKA PWS 60° BER OKH KAM CGA SEA COM WGA EAL WAL 45° CAL BRC KUR Sampled Rookeries ORE NCA JAPAN 30° 135°E 180° 135°W Previous Genetic Studies Bickham et al. (1996): Discovered high genetic differentiation between populations on either side of 144˚W; subsequently recognized an eastern stock (threatened) and western stock (endangered) Also found high diversity of “mtDNA haplotypes and a clear pattern of macrogeographic variation” The Western and Eastern Stocks RUSSIA ALASKA PWS Cape Suckling 60° BER OKH KAM CGA SEA COM WGA EAL WAL 45° CAL BRC KUR Sampled Rookeries ORE 144°W NCA JAPAN WESTERN STOCK 30° 135°E 180° EASTERN STOCK 135°W Previous Genetic Studies (cont) Bickham et al. (1998) Found evidence for three possible groups: Asian and central groups within the western stock and an eastern stock Could have arisen due to two, possibly three, glacial refugia that separated the populations during the Pleistocene era A Third Stock? Previous research suggests that animals sampled from Russian rookeries are genetically distinct from other western stock sea lions. RUSSIA ALASKA PWS Cape Suckling 60° BER OKH KAM CGA SEA COM WGA EAL WAL 45° CAL BRC KUR Sampled Rookeries ORE 144°W NCA JAPAN 30° 135°E ASIAN STOCK WESTERN STOCK 180° EASTERN STOCK 135°W Previous Genetic Studies (cont) Baker et al. (2005) Subdivided the Western Stock into Asian Stock (OKH, KUR, KAM) and Western Stock (COM east through PWS) In this presentation we will: Summarize a 14-year range-wide populationgenetic analysis based upon the mtDNA control region of Steller sea lions. Present preliminary results from an ongoing complementary study of cytochrome b sequences. Compare phylogeographic and population genetic patterns for control region vs. cyt b sequences. Present data from nuclear microsatellite loci that corroborate some of the findings of stock structure in Steller sea lions. Control Region Sample Sizes Stock Region EASTERN STOCK NCA (1 ROOKERY) 51 ORE (1 ROOKERY) 85 BRC (2 ROOKERIES) 23 SEA (3 ROOKERIES) 76 WESTERN STOCK TOTAL 235 PWS (2 ROOKERIES) 153 BER (1 ROOKERY) 42 CGA (6 ROOKERIES) 197 WGA (5 ROOKERIES) 108 EAL (6 ROOKERIES) 275 CAL (10 ROOKERIES) 247 WAL (3 ROOKERIES) 98 COM (1 ROOKERY) TOTAL ASIAN STOCK Sample Size 126 1246 KAM (2 ROOKERIES) 71 KUR (5 ROOKERIES) 207 OKH (2 ROOKERIES) 237 TOTAL 515 GRAND TOTAL 1,710 Cytochrome b Sample Sizes Stock Region EASTERN STOCK NCA (1 ROOKERY) 27 ORE (1 ROOKERY) 32 BRC (2 ROOKERIES) 15 SEA (3 ROOKERIES) 341 TOTAL 415 WESTERN STOCK PWS (2 ROOKERIES) 27 BER (1 ROOKERY) 21 CGA (7 ROOKERIES) 114 WGA (5 ROOKERIES) 32 EAL (6 ROOKERIES) 84 CAL (9 ROOKERIES) 73 WAL (3 ROOKERIES) 51 COM (1 ROOKERY) 46 TOTAL ASIAN STOCK Sample Size 448 KAM (2 ROOKERIES) 63 KUR (5 ROOKERIES) 74 OKH (2 ROOKERIES) 133 TOTAL 165 GRAND TOTAL 878 Microsatellite Sample Sizes Stock Region EASTERN STOCK NCA (1 ROOKERY) 25 ORE (1 ROOKERY) 25 BRC (2 ROOKERIES) 18 SEA (3 ROOKERIES) 45 TOTAL WESTERN STOCK 113 PWS (2 ROOKERIES) 75 BER (1 ROOKERY) 13 CGA (3 ROOKERIES) 43 WGA (3 ROOKERIES) 30 EAL (4 ROOKERIES) 172 CAL (3 ROOKERIES) 70 WAL (1 ROOKERY) 12 COM (1 ROOKERY) 25 TOTAL ASIAN STOCK Sample Size 440 KAM (1 ROOKERY) 25 KUR (4 ROOKERIES) 40 OKH (2 ROOKERIES) 50 TOTAL 115 GRAND TOTAL 668 L-shaped distribution of control-region haplotypes No evidence of a significant bottleneck or reduction in variation Individual Haplotype Totals N= 1,433 Individuals N= 132 Haplotypes 50 40 30 20 10 Number of Individuals per Haplotype 10 5 12 8 18 2 25 2 86 31 35 36 51 16 17 21 22 6 7 8 10 11 12 13 3 4 5 0 1 2 Number of Haplotypes 60 Control Region Haplotype Distributions Haplotype LLL Haplotype Frequency Haplotype Frequency Haplotype BB 0.50 0.40 0.30 0.20 0.10 0.00 0.10 0.08 0.06 0.04 0.02 0.00 0.12 0.10 0.08 0.06 0.04 0.02 0.00 Haplotype H Haplotype Frequency Haplotype Frequency Haplotype E 0.50 0.40 0.30 0.20 0.10 0.00 Cytochrome b Haplotype Distributions Haplotype 4 1.00 0.90 0.90 Haplotype Frequency Haplotype Frequency Haplotype 1 1.00 0.80 0.70 0.60 0.50 0.40 0.30 0.20 0.10 0.00 0.80 0.70 0.60 0.50 0.40 0.30 0.20 0.10 0.00 Haplotype 10 1.00 0.90 0.90 0.80 0.70 0.60 0.50 0.40 0.30 0.20 0.10 0.00 Haplotype Frequency Haplotype Frequency Haplotype 2 1.00 0.80 0.70 0.60 0.50 0.40 0.30 0.20 0.10 0.00 NJ Tree for Rookeries Based on Fst Estimates Control Region Legend Asian Stock Western Stock Eastern Stock Adak Island CAL Akun Island EAL Seal Rocks PWS Clubbing Rocks WGA Yunaska Island CAL Ugamak Island EAL Chirikof Island CGA Amak Island EAL Bogoslof Island EAL Pinnacle Rock WGA Chowiet Island CGA Marmot Island CGA Adugak Island EAL Attu Island WAL Amchitka Island CAL Seguam Island CAL Medney Island COM Kasatoshi Island CAL Sugarloaf Island CGA Akutan Island EAL Agattu Island WAL Kozlova Cape RUS Walrus Island BER Fish Island PWS Kiska Island Lief Cove CAL Kiska Island Cape St. Stephen CAL Buldir Island WAL Gramp Rock CAL Atkins Island WGA Tag Island CAL Ogchul Island EAL Antsiferova Island KUR Zheleznaya Bay KAM Raykoke Island KUR Iony Island OKH Tuleniy Island OKH Srednego Island KUR Yamsky Island OKH Lovushki Island KUR Chernyye Bratya KUR Forrester Island SEA North Danger Rocks BRC St. George Reef NCA Triangle Island BRC Hazy Islands SEA Rogue Reef ORE White Sisters Islands SEA 0.005 changes NJ Tree Regions OKH Based on Fst Estimates Control Region KUR KAM COM CAL EAL CGA WGA PWS BER WAL Legend Eastern Stock Western Stock Asian Stock SEA BRC ORE NCA 0.005 changes NJ Zalophus A XXXX UUUUU IIII EEEEE NNN TTTTT E NJ tree of control region haplotypes shows two distinct eastern stock haplotype lineages. F XX VV AAAAA B R FFFF XXXXX ZZZ FFFFF RRRR UU P SSSS III C YYYYY H Q D GGGG BBBBB SS YY LL M DDDDD KKK JJJJ SSS HHHH MMMMM OOOOO Y NNNNN QQQ OOOO KKKK PPPPP LLLL KKKKK LLLLL W PPP X HH II JJJJJ NN U SSSSS 0.001 substitutions/site DDDD EEEE QQQQ ZZZZZ XXX WWW MMMM TTTT MM KK HHHHH DD HHH NNNN AAAAAA T EE UUU AAA FF V BBB GG TT JJ PPPP OOO VVV CC G QQ YYY RRRRR GGG YYYY LLL TTT BBBB MMM RRR VVVVV AAAA RR S O WWWW JJJ FFF DDD QQQQQ PP BB EEE CCCC WWWWW IIIII GGGGG L VVVV K AA J ZZ BBBBBB CCC N UUUU OO WW Z CCCCC I ZZZZ 16 29 5 Minimum Spanning Network of Steller Sea Lion Cytochrome b haplotypes n = 878, 1040 bp 34 33 28 18 20 22 1 ? 24 7 19 9 29 Variable Positions Ti/Tv = 26/3 13 Silent, 13 Nonsynonymous Transitions 1 Silent, 2 Nonsynonymous Transversions 1st Position: 10 (9 AAS) 2nd Position: 6 (6 AAS) 3rd Position: 13 (0 AAS) 14 Total Variable Amino Acids 25 17 21 23 8 27 11 12 3 4 14 13 32 200+ 15 25-50 26 100-200 WEST 2 31 EAST 10-25 50-100 30 ? 10 ASIAN 6 White Line: Silent Mutation Red Line: Amino Acid Shift 2-10 ALL 1 Regional Cytochrome b Slatkin’s Linearized FST Midpoint Rooted NJ Phylogram OKH KUR KAM COM CAL CGA WAL WGA BER EAL PWS Legend Eastern Stock Western Stock Asian Stock BRC OR E SEA NCA 0.05 changes NJ Phylogram of Rookeries Based on Microsatellite Data Yamsky Island -OKH Srednego Island -KUR Raykoke Island -KUR Kozlova Cape -KAM Iony Island -OKH Lovushki Island -KUR Kiska Island -CAL Seguam Island -CAL Sugarloaf Island -CGA Yunaska Island -CAL Akutan Island -EAL Chirikof Island -CGA Ogchul Island -EAL Ugamak Island -EAL Pinnacle Rocks -WGA Medny Island -COM Adugak Island -EAL Antsiferova Island -KUR Clubbing Rocks -WGA Walrus Island -BER Seal Rocks -PWS Buldir Island -WAL Chowiet Island -CGA Fish Island -PWS Atkins Island -WGA Rogue Reef -ORE St. George Island -NCA North Danger Rocks -BRC Triangle Island -BRC Forrester Island -SEA White Sisters Island -SEA Hazy Island -SEA 0.01 Asian Stock Asian Stock Western Stock Eastern Stock Legend Eastern Stock Western Stock Asian Stock NJ Phylogram of Regions Based on Microsatellite Data. OKH Asian Stock KAM KUR WAL CAL COM EAL Western Stock BER WGA CGA Legend Eastern Stock Western Stock Asian Stock PWS ORE NCA BRC 0.01 SEA Eastern Stock Conclusions Control Region Phylogeography Despite the generally high sample sizes in this study, resolution of population substructure increased when rookeries were grouped into their respective regions. Control region sequences clearly support the recognition of three stocks. Cytochrome b Phylogeography • Clearly differentiates the eastern and western stocks but does not resolve the Asian stock. Microsatellite Analysis • Nuclear microsatellites show a pattern nearly the same as for the cytochrome b gene. • Clearly differentiates the eastern and western stocks but does not resolve the Asian stock. The Asian Stock The recognition of three management units may be appropriate for Steller sea lion conservation. RUSSIA ALASKA PWS Cape Suckling 60° BER OKH KAM CGA SEA COM WGA EAL WAL 45° CAL BRC KUR Sampled Rookeries ORE 144°W NCA JAPAN 30° 135°E ASIAN STOCK WESTERN STOCK 180° EASTERN STOCK 135°W