The History and Future of Whales Dr. Stephen R. Palumbi Hot Science - Cool Talks Volume 45 Produced by and for the Hot Science – Cool Talks Outreach Lecture Series of the Environmental Science Institute. We request that the use of any of these materials include an acknowledgement of Dr. Stephen R. Palumbi and the Hot Science – Cool Talks Outreach Lecture Series of the Environmental Science Institute of the University of Texas at Austin. We hope you find these materials educational and enjoyable. The History and Future of Whales Stephen R. Palumbi Stanford University's Hopkins Marine Station Ocean neighborhoods have changed The Sea Around Us Project: D. Pauly and coworkers at UBC The Sea Around Us Project: D. Pauly and coworkers at UBC The Sea Around Us Project: D. Pauly and coworkers at UBC Results of effort-based management The Sea Around Us Project: D. Pauly and coworkers at UBC Decline in state of the World’s fisheries Vitousek et al. 1997 What fish is that on my plate? Microdocs.org - Cooking with DNA - NPR’s Splendid Table Package of whale meat what species is it? What whale meat is legal? • International Whaling Commission (IWC) Sets catch limits and manages population recovery • Convention on International Trade of Endangered Species (CITES) Requires permits for international shipment CITES Convention for the International Trade of Endangered Species • Regulates imports and exports of endangered species and derivatives of endangered species • Distinguishes endangered and commercially threatened species • Member nations follow strict import/export regulations Scientific whaling • IWC members issue themselves permits for scientific whaling. • Review is necessary but approval is not. • Commercial use of products is encouraged after research use. Forensic monitoring of resource use NOAA Forensics and the future DNA testing of retail whale products examines relationship between policy and practice Scott Baker on first Tokyo meat mission (1993) Scott Baker tries for a GQ cover Matt Hare on 1998 Tokyo mission PCR copied whale genes are separated from native whale DNA in the field so we can strictly follow CITES regulations Simplified phylogeny of whales used for assigning species names to meat samples Molecular identification of baleen whale products N. minke S. minke Brydes Pygmy Brydes Sci Humpback Fin Blue Grey 339 369 10 2 9 7 49 2 7 Data from Palumbi, Baker labs 1993-2000 Species other than baleen whales in retail markets Sperm Pygmy Sperm Other Beaked Whales Baird’s Cuvier’s Porpoise Killer Whale Dolphin Non-Whale 5 1 5 56 6 19 2 104 7 Data from Palumbi, Baker labs 1993-2000 Conclusion • Many species of whales and dolphins are for sale in retail markets. • Does this failure to protect put any whales at risk of extinction? Consequences of selling the wrong whales Threatened Sea of Japan minke meat masquerading as legal take from the North Pacific North Pacific genotypes Sea of Japan genotypes Genotypes from market Surveys show 1:2 mix of Sea of Japan and North Pacific animals The consequence of scientific whaling Predicted extinction of Sea of Japan minke whales (Baker et al. 2000) Dolphins and porpoises make up a major fraction of the whale meat market in Japan. We’ve known for a while that unsafe toxin loads (heavy metals, PCBs, Dioxins) are found most commonly in dolphin and porpoise products. So, is dolphin meat labeled as whale meat unsafe? Yes, dolphin meat is often toxic How bad? Dolphins > Pacific minke whales> southern minkes 20 18 Southern minke whale 16 14 Northern minke whale 12 Odontocetes 10 8 6 4 2 0 .1 .2 .3 .4 .5 Mercury content .6 .7 (micrograms/gram) .8 .9 1 Mercury in Whale Meat (samples from 1999) 50 One Stenella dolphin sample @ 200 µg/g 45 40 35 Dolphins and porpoise meat has high mercury loads Count 30 25 20 15 Unsafe levels 10 5 0 1 2 3 4 5 Mercury content 6 7 (micrograms/gram) 8 9 10 The history of whales revealed by genetics How many whales were there before hunting? Catch records for Antarctic whales are well known. From Hilborn et al. 2004 Where History Comes From • Written records • Oral records • Fossils • Tree rings • Ice cores • Reef cores Byrd Polar Research Center 2005 Climate History Tree rings and ice cores record climate Arnold Paul Family history My Great Aunts What do we know about the histories of whale populations? “It is impossible to describe either the number of whales or their familiarity,” with breath that “caused a most annoying stench” Jean Francoise de La Perouse, 1786 Trouble with conventional wisdom about whale history (e.g. Atlantic fin whales) Current estimate: 47,300 Fin Whales North Atlantic 1969-89 Mean: 47,300 Range: 27,700 - 82,000 www.iwcoffice.org/conservation/estimate.htm Past estimate: 30-50,000 (Sargeant 1977) www.calacademy.org Genetic estimates of population size Inbreeding decreases genetic diversity Mutation increases genetic diversity Genetic estimates of population size Large populations have low inbreeding High diversity Genetic estimates of population size Small populations have strong inbreeding Low diversity For stable, single populations and neutral mtDNA variation: = 2Ne(f) : genetic diversity : substitutions/generation Atlantic collaborator Joe Roman at Harvard University World-wide sample of humpback whales from biopsies Data from Palumbi, Baker, Palsbol, Rosenbaum Phylogenetic Trees Branch Length and Genetic Diversity Shared recent ancestors (similarity of DNA signified by shorter branch length) Shared ancient ancestors (greater differences in DNA signified by longer branch length) High global mtDNA diversity for humpback whales Genetic diversity predicted for 115,000 whales wa2 003 wa1 a1 905 04 w 9 wa0 a0 707 06 w gi22 08 2 gi1 22 00 2 9 gi 1 gi 12 1 ea0 81 11 ea0 8 12 gia0 03 218 6 5 w g1 020 19 tt g1 0 wa0 a0 514 13 w 5 wa0 515 10GL91 12G L91 04 6 5 4 12G 6 12G 6 12GL91 L91 64 4 12G L91 6 4 12GL91 L91 64 4 12G 6 t g0 21 gi 38 2 8 M A2 N VAJ O 900 9002 7 MV OLLE T RY H C M A N f0 3 gi10 2L91 4 65 11G M CS03 N f 8 0 wa0 823 9029 5 900 9041 8 I ce 0 I ce 0 M I2 B LB M R C ANO E MT CIYD S LL AT M R N E M W RAP N 2 Nff 0 5 0 Nff 1 7 0 N N F1 2 NR F1 4 D 0 4 15W 17G G0 L90 0 87 7 01 17G L90 17GL90 L90 07 7 17G 0 17G L90 0 7 17G L90 0 17GL90 07 7 17GL90 L90 07 7 17G 0 17G L90 0 7 17GL90 L90 07 7 17G 0 17GL90 L90 07 7 17G 0 17GL90 L90 07 7 17G 0 17G L90 0 7 17G 0 17GL90 L90 07 7 17GL90 L90 07 7 17G 0 17G L90 0 7 17GL90 L90 07 7 17G 0 17GL90 L90 07 7 17G 0 17GL90 07 17GL90 L90 07 7 17G 0 18I L 911 5 9009 3 902 NR f0 1 D 5 1 MT M ASN LI KE K M A LO 14W G 814 13G 16G L91 M 2 0 9 56 16GM 9 07 56 9014 8 900 04G M 0 9 04GM M9 015 9 15 04G 0 15 04GM M9 015 9 15 04G 0 04GL91 M0 03 9 15 08G 08G L91 03 08G L91 0 3 08GL91 03 08GL91 L91 03 3 08G 0 08G L91 0 3 08G 03 D R1 8L91 06GL91 L91 13 3 06G 1 09IL910 3 05G L90 19 9010 N F1 6 Nff 0 7 1 N 6 DRB 1901 02S 9 02S B901 901 9 02S B 9 02S B 901 9 02S B 9 02S B901 901 9 02S B901 901 9 02S B 9 02S B 9 02S B901 901 9 02S B 901 9 02S B901 901 9 02S B 9 02S B901 9 D 8 3 N fR1 8 M P S E A NF0 9 R 01GL91 L91 00 0 01G 0 01G L91 0 0 03G 22 D R1 3L91 DR R3 9 1 D DR1 2 07S B903 903 9 07S B 9 902 0 wa2 a2 117 16 w 1 ea1 227 26 ea0 2 ea0 229 28 ea0 2 g0 931 30 ttt g0 9 g0 932 I ce 0 9033 2 D 0 DR R2 7 3 M LI PP C ER MH OLIM C ET M E X M W ORD S N f1 F1 0 5 23G L90 30 23G 3 23GL90 L90 30 0 23GL90 L90 30 0 23G 3 23G L90 3 0 23GL90 L90 30 0 23G 3 23GL90 L90 30 0 23G 3 23GL90 30 23GL90 L90 30 0 23G 3 23G 3 23GL90 L90 30 0 23GL90 L90 30 0 23G 3 23G L90 3 0 23GL90 L90 30 0 23G 3 27GL90 L91 09 2 28G 2 28GL90 29 28GL90 L90 29 9 28G 2 28G L90 2 28GL90 L90 29 9 28G 2 9 28GL91 L90 20 9 29G 3 29G L91 30 9005 1 900 M E B LTA N EET MS I LHOU 25GL90 L90 14 9 26G 0 26G L90 0 4 26GL90 L90 04 4 26G 0 26GL90 L90 04 4 26G 0 26GL90 L90 04 4 26G 0 M O LA P RI S 24G M 0 9 wa1 833 58 nz0 135 34 nz0 1 nz0 1 36 138 37 tnz0 g1 5 gi1 14 30 6 9 gi gi16 41 6 gi1 16 43 6 2 gi 4 gi 1 4 gi2 16 44 6 5 gi 4 5 6 t g0 2 ea1 356 50 a2 851 twg1 4 w a1 552 53 ea0 5 gi17 654 0 gi1 17 62 7 1 gi 6 gi 1 6 7 3 t g1 gi 41 655 2 4 gi2 34 59 0 7 gi 3 0 5 8 ap0 266 65 ap0 2 ap0 2 67 C a0 8 B Ca1 1. AP Ca1 a1 3. AP C 4 M 15. B x ar t p 268 x 3. A PB tM g0 5 gi0 01 74 1 3 gi 7 gi01 75 gi0 01 77 1 6 gi 7 gi 0 7 1 8 gi0 01 70 1 9 gi 8 wa0 170 69 ea0 6 ea0 672 71 ap0 1 gi3 37 48 6 7 gi 4 gi07 49 gi11 8 G I11 1 C gia2 02 83 5 2 gi 0 8 5 G I0 I05b 5a G Ca0 a0 1.. A AP B P C 6 B C a0 7 . AP B C 6 B Ca1 a2 a0 3.. A 5 AP B P M 2.7.AP x PB A C a0 B H i0 .4C 2 SB Hi0 i03 .C CS SB B H .1 Hi0 i05 .C 4 CS SB H . Hi06 . CSB B HE i0A .C 7 BB S 0 1.SA P S E 0 A 2. P SEA 03. A PB A B SE EA 07. A 6. P PB A B S 0 M 08. A x PB A Mx 27. x 2. A PB A B M 0 P Mx 25. A PB 0.001 substitutions/site 4% 3% 2% 1% Branch lengths (percent substitution) 0 Genetic estimates of world-wide population Genetic diversity predicted for 115,000 whales wa2 003 wa1 a1 905 04 w 9 wa0 a0 707 06 w gi22 08 2 gi1 22 00 2 9 gi 1 gi 12 1 ea0 81 11 ea0 8 12 gia0 03 218 6 5 w g1 020 19 tt g1 0 wa0 a0 514 13 w 5 wa0 515 10GL91 12G L91 04 6 5 4 12G 6 12G 6 12GL91 L91 64 4 12G L91 6 4 12GL91 L91 64 4 12G 6 t g0 21 gi 38 2 8 M A2 N VAJ O 900 9002 7 MV OLLE T RY H C M A N f0 3 gi10 2L91 4 65 11G M CS03 N f 8 0 wa0 823 9029 5 900 9041 8 I ce 0 I ce 0 M I2 B LB M R C ANO E MT CIYD S LL AT M R N E M W RAP N 2 Nff 0 5 0 Nff 1 7 0 N N F1 2 NR F1 4 D 0 4 15W 17G G0 L90 0 87 7 01 17G L90 17GL90 L90 07 7 17G 0 17G L90 0 7 17G L90 0 17GL90 07 7 17GL90 L90 07 7 17G 0 17G L90 0 7 17GL90 L90 07 7 17G 0 17GL90 L90 07 7 17G 0 17GL90 L90 07 7 17G 0 17G L90 0 7 17G 0 17GL90 L90 07 7 17GL90 L90 07 7 17G 0 17G L90 0 7 17GL90 L90 07 7 17G 0 17GL90 L90 07 7 17G 0 17GL90 07 17GL90 L90 07 7 17G 0 18I L 911 5 9009 3 902 NR f0 1 D 5 1 MT M ASN LI KE K M A LO 14W G 814 13G 16G L91 M 2 0 9 56 16GM 9 07 56 9014 8 900 04G M 0 9 04GM M9 015 9 15 04G 0 15 04GM M9 015 9 15 04G 0 04GL91 M0 03 9 15 08G 08G L91 03 08G L91 0 3 08GL91 03 08GL91 L91 03 3 08G 0 08G L91 0 3 08G 03 D R1 8L91 06GL91 L91 13 3 06G 1 09IL910 3 05G L90 19 9010 N F1 6 Nff 0 7 1 N 6 DRB 1901 02S 9 02S B901 901 9 02S B 9 02S B 901 9 02S B 9 02S B901 901 9 02S B901 901 9 02S B 9 02S B 9 02S B901 901 9 02S B 901 9 02S B901 901 9 02S B 9 02S B901 9 D 8 3 N fR1 8 M P S E A NF0 9 R 01GL91 L91 00 0 01G 0 01G L91 0 0 03G 22 D R1 3L91 DR R3 9 1 D DR1 2 07S B903 903 9 07S B 9 902 0 wa2 a2 117 16 w 1 ea1 227 26 ea0 2 ea0 229 28 ea0 2 g0 931 30 ttt g0 9 g0 932 I ce 0 9033 2 D 0 DR R2 7 3 M LI PP C ER MH OLIM C ET M E X M W ORD S N f1 F1 0 5 23G L90 30 23G 3 23GL90 L90 30 0 23GL90 L90 30 0 23G 3 23G L90 3 0 23GL90 L90 30 0 23G 3 23GL90 L90 30 0 23G 3 23GL90 30 23GL90 L90 30 0 23G 3 23G 3 23GL90 L90 30 0 23GL90 L90 30 0 23G 3 23G L90 3 0 23GL90 L90 30 0 23G 3 27GL90 L91 09 2 28G 2 28GL90 29 28GL90 L90 29 9 28G 2 28G L90 2 28GL90 L90 29 9 28G 2 9 28GL91 L90 20 9 29G 3 29G L91 30 9005 1 900 M E B LTA N EET MS I LHOU 25GL90 L90 14 9 26G 0 26G L90 0 4 26GL90 L90 04 4 26G 0 26GL90 L90 04 4 26G 0 26GL90 L90 04 4 26G 0 M O LA P RI S 24G M 0 9 wa1 833 58 nz0 135 34 nz0 1 nz0 1 36 138 37 tnz0 g1 5 gi1 14 30 6 9 gi gi16 41 6 gi1 16 43 6 2 gi 4 gi 1 4 gi2 16 44 6 5 gi 4 5 6 t g0 2 ea1 356 50 a2 851 twg1 4 w a1 552 53 ea0 5 gi17 654 0 gi1 17 62 7 1 gi 6 gi 1 6 7 3 t g1 gi 41 655 2 4 gi2 34 59 0 7 gi 3 0 5 8 ap0 266 65 ap0 2 ap0 2 67 C a0 8 B Ca1 1. AP Ca1 a1 3. AP C 4 M 15. B x ar t p 268 x 3. A PB tM g0 5 gi0 01 74 1 3 gi 7 gi01 75 gi0 01 77 1 6 gi 7 gi 0 7 1 8 gi0 01 70 1 9 gi 8 wa0 170 69 ea0 6 ea0 672 71 ap0 1 gi3 37 48 6 7 gi 4 gi07 49 gi11 8 G I11 1 C gia2 02 83 5 2 gi 0 8 5 G I0 I05b 5a G Ca0 a0 1.. A AP B P C 6 B C a0 7 . AP B C 6 B Ca1 a2 a0 3.. A 5 AP B P M 2.7.AP x PB A C a0 B H i0 .4C 2 SB Hi0 i03 .C CS SB B H .1 Hi0 i05 .C 4 CS SB H . Hi06 . CSB B HE i0A .C 7 BB S 0 1.SA P S E 0 A 2. P SEA 03. A PB A B SE EA 07. A 6. P PB A B S 0 M 08. A x PB A Mx 27. x 2. A PB A B M 0 P Mx 25. A PB >1 million humpbacks 0.001 substitutions/site 4% 3% 2% 1% Branch lengths (percent substitution) 0 North Atlantic humpback mtDNA diversity = 2.2% (n = 188) = 2*(# breeding females) * (mutation/generation) ==> female size = 68,000 ==> population size = 240,000 Current view of the history of humpback whales Original population - 115,000 North Atlantic original - 30,000 North Atlantic current - 10,000 MSY North Atlantic - 16,000 Nearly ready for harvest Genetic view of the history of humpback whales Original population - 1,500,000 North Atlantic original - 314,000 North Atlantic current - 10,000 MSY North Atlantic - 170,000 Ready for harvest in 150 years Antarctic whales existed in huge numbers What does DNA tell us about their past? Frank goes back to Japan for Antarctic Title samples = 2Ne (µ/year)( (years/generation) = 0.02 for cyt b µ/year = 0.34% permillion years (cyt b) Generation time = 15 yr /2 * 5.1% per million generations => 195,000 females => 1,150,000 population size = 2 Ne (µ/year)(years/generation) = 0.09 for entire HVI of control region µ/year = 2.0% per million years (CR) Generation time = 15 yr /2 * 30% per million generations => 150,000 females => 900,000 population size B A 20 C D No evidence that minke whales prevent recovery of cousins in the Antarctic Blue Whale E 0 Fin Whale Number of Whales (unit: 10,000) 40 E 20 0 20 Sei Whale E 0 Genetic estimate of historic population of minke whales (900,000) 80 Minke Whale 60 40 20 E 0 1900 20 40 60 80 2000 Year Data: Dr. Seiji Ohsumi Genetic surveys to date that show large historic population sizes 5 species 8 populations 3 ocean basins 6 genes 2004 IWC recommendation for continuing genetic estimates • • • • • • • Verify substitution rate Analyze multiple loci Measure population dynamics Measure variance Evaluate ghost populations Get better measures for log records Estimate food levels needed Summary • The past history of whales is a very important part of our view of what the future should be. • Estimates based on whaling ship log books appear to underestimate populations. The genetics of whale populations suggest that they were much more numerous than we believed. • Genetic techniques are useful tools for understanding population levels, species diversity, and human exploitation of whale resources. • Knowledge of the history of whale populations and resource use are essential components of whale conservation efforts. Dr. Stephen R. Palumbi Professor of Biological Sciences, Stanford University's Hopkin Marine Station Stephen R. Palumbi is a professor of Biological Sciences at Stanford University's Hopkin Marine Station. He and other research scientists in the Palumbi Lab study genetics, evolution, conservation, population biology, and systematics of a diverse array of marine organisms. He uses molecular genetic techniques in conservation-related research, including the identification of whale and dolphin products available in commercial markets and the genetics of marine reserves designed for conservation and fisheries enhancement. His 2003 publication in the journal Science on Whales before Whaling in the North Atlantic suggests that whale populations were 10 times larger than historical records indicate, which has critical implications for the future of whaling and whale conservation. Dr. Palumbi received his Ph.D. from University of Washington in marine ecology in 1984. In 1996, he received a Pew Fellowship for Marine Conservation Research. He has published on the genetics and evolution of butterflyfishes, bryozoans, cone snails, corals, sea urchins, sharks, spiders, shrimps, and whales. His recent books include The Evolution Explosion: How humans cause rapid evolutionary change and Marine Reserves: An Ecosystem Tool for Marine Management and Conservation.