Madden, K. M., Fuiman, L. A., Williams, T. M., & Davis, R. W. (2015). Weddell seal foraging dives: comparison of free-ranging and isolated-hole paradigms. Antarctic Science, 27(1), 57–68. (pdf)

Kanatous, S. B., Hawke, T. J., Trumble, S. J., Pearson, L. E., Watson, R. R., Garry, D. J., Williams, T.M., Davis, R. W. (2008). The ontogeny of aerobic and diving capacity in the skeletal muscles of Weddell seals. The Journal of Experimental Biology, 211, 2559–65. (pdf)

Davis, R.W., Polasek, L., Watson, R., Fuson, A., Williams, T.M., and Kanatous, S.B. (2004)  The diving paradox: new insights into the role of the dive response in air-breathing vertebrates.  Comparative Biochemistry and Physiology Part A 138:263-268. (pdf)

Williams, T.M., Fuiman, L.A., Horning, M., and Davis, R.W. (2004)  The cost of foraging by a marine predator, the Weddell seal Leptonychotes weddellii: pricing by the stroke.  Journal of Experimental Biology 207:973-982. (pdf)

Davis, R.W., Fuiman, L.A., Williams, T.M., and Horning, M. (2003)  Classification of Weddell seal dives based on three-dimensional movements and video recorded observations.  Marine Ecology Progress Series 264:109-112. (pdf)

Williams, T.M. (2003) Sunbathing seals of Antarctica. Natural History 112: 50-55.

Fuiman, L.A., Davis, R.W., and Williams, T.M. (2002)  Behavior of midwater fishes under the Antarctic fast ice: observations by a predator.  Marine Biology 140:815-822. (pdf)

Kanatous, S.B., Davis, R.W., Watson, R., Polasek, L., Williams, T.M., and Mathieu-Costello, O. (2002)  Aerobic capacities in the skeletal muscles of Weddell seals: key to longer dive durations?  Journal of Experimental Biology 205:3601-3608. (pdf)

Davis, R.W., Fuiman, L.A., Williams, T.M., Collier, S.O., Hagey, W.P., Kanatous, S.B., Kohin, S., and Horning, M. (1999)  Hunting behavior of a marine mammal beneath the Antarctic fast ice.  Science 283:993-996. (pdf)


Helm, R.C., Costa, D.P., DeBruyn, T.D., O’Shea, T.J., Wells, R.S., and Williams, T.M. (2015) Overview of effects of oil on marine mammals. In Handbook of Oil Spill Science and Technology (Fingas, M., eds.). John Wiley & Sons, Inc.: Hoboken, NJ. Pp. 455-476.

Williams, T.M. & J.L. Maresh. (2015). Exercise energetics in Marine Mammal Physiology: Requisites for Ocean Living (Castellini & Mellish, eds.). SPi Global: Madison, WI. Pp. 47-68.

Williams, T. M., Fuiman, L.A., & Davis, R.W. (2015). Locomotion and the cost of hunting in large, stealthy marine carnivores. Integrative and Comparative Biology 55:673-682. (pdf)

Davis, R.W. and Williams, T.M. (2012) The marine mammal dive response is exercise modulated to maximize aerobic dive duration. Journal of Comparative Physiology A 198:583-591. (pdf)

Williams, T. M., Bengtson, P., Steller, D. L., Croll, D. A., & Davis, R. W. (2015). The healthy heart: Lessons from nature’s elite athletes. Physiology30(7), 349–357. (pdf)

Davis, R.W. and Williams, T.M. (2012) The marine mammal dive response is exercise modulated to maximize aerobic dive duration. Journal of Comparative Physiology A 198:583-591. (pdf)

Hooker, S.K., Fahlman, A., Moore, M.J., Aguilar de Soto, N., Bernaldo de Quiros, Y., Brubakk, A.O., Costa, D.P., Costidis, A.M., Dennison, S., Falke, K.J., Fernandez, A., Ferrigno, M., Fitz-Clarke, J.R., Garner, M.M., Houser, D.S., Jepson, P.D., Ketten, D.R., Kvadsheim, P.H., Madsen, P.T., Pollock, N.W., Rotstein, D.S., Rowles, T.K., Simmons, S.E., Van Bonn, W., Weathersby, P.K., Weise, M.J., Williams, T.M., and Tyack, P.L. (2012) Deadly diving? Physiological and behavioural management of decompression stress in diving mammals. Proceedings of the Royal Society B 279:1041-1050. (pdf)

Williams, T.M., M. Zavanelli, M.A. Miller, R.A. Goldbeck, M. Morledge, D. Casper, D.A. Pabst, W. McLellan, L.P. Cantin, and D.S. Kliger. (2008) Running, swimming and diving modifies neuroprotecting globins in the mammalian brain. Proceedings of the Royal Society B 275:751-758. (pdf)

Burns, J.M., Williams, T.M., Secor, S.M., Owen-Smith, N., Bargmann, N.A., and Castellini, M.A. (2006) New insights into the physiology of natural foraging. Physiological and Biochemical Zoology 79(2):242-249. (pdf)

Estes, J.A., Danner, E.M., Doak, D.F., Konar, B., Springer, A.M., Steinberg, P.D., Tinker, M.T., and Williams, T.M. (2004) Complex trophic interactions in kelp forest ecosystems. Bulletin of Marine Science 74(3):621-638. (pdf)

Williams, T.M., Estes, J.A., Doak, D.F., and Springer, A.M. (2004) Killer appetites: Assessing the role of predators in ecological communities. Ecology 85(12):3373-3384. (pdf)

Springer, A.M., Estes, J.A., van Vliet, G.B., Williams, T.M., Doak, D.F., Danner, E.M., Forney, K.A., and Pfister, B. (2003)  Sequential megafaunal collapse in the North Pacific Ocean: An ongoing legacy of industrial whaling? Proceedings of the National Academy of Sciences 100(21):12223-12228. (pdf)

Noren, S.R., Williams, T.M., Pabst, D.A., McLellan, W.A., and Dearolf, J.L. (2001)  The development of diving in marine endotherms: preparing the skeletal muscles of dolphins, penguins, and seals for activity during submergence.  Journal of Comparative Physiology – B, Biochemical, Systemic, & Environmental Physiology 171:127-134. (pdf)

Williams, T.M. (2001)  Intermittent swimming by mammals: a strategy for increasing energetic efficiency during diving. American Zoologist 41:166-176. (pdf)

Williams, T.M., Haun, J., Davis, R.W., Fuiman, L.A., and Kohin, S. (2001)  A killer appetite: metabolic consequences of carnivory in marine mammals.  Comparative Biochemistry and Physiology Part A 129:785-796. (pdf)

Williams, T.M., Davis, R.W., Fuiman, L.A., Francis, J., LeBoeuf, B.J., Horning, M., Calambokidis, J., and Croll, D.A. (2000)  Sink or swim: Strategies for cost-efficient diving by marine mammals.  Science 288:133-136. (pdf)

Williams, T.M. (1999)  The evolution of cost-efficient swimming in marine mammals: Limits to energetic optimization. Phil. Trans. R. Soc. Lond. 354:193-201. (pdf)


Williams, T.M., Blackwell, S.B., Richter, B., Sinding, M-H. S., and Heide-Jørgensen, M.P. (2017). Paradoxical escape responses by narwhals (Monodon monoceros). Science 358:1328-1331. (pdf)

Williams, T.M., Kendall, T.L., Richter, B.P., Ribeiro-French, C.R., John, J.S., Odell, K.L., Losch, B.A., Feuerbach, D.A., Stamper, M.A. (2017). Swimming and diving energetics in dolphins: a stroke-by-stroke analysis for predicting the cost of flight responses in wild odontocetes. Journal of Experimental Biology 220:1135-1145. (pdf)

Fish, F.E., Legac, P., Williams, T.M, and Wei, T. (2014) Measurement of hydrodynamic force generation by swimming dolphins using bubble DPIV. Journal of Experimental Biology 217:252-260.

Noren, D.P., Holt, M.M., Dunkin, R.C., and Williams, T.M. (2013). The metabolic cost of communicative sound production in bottlenose dolphins (Tursiops truncatus). Journal of Experimental Biology 216:1624-1629. (pdf)

Noren, S.R., Williams, T.M., Ramirez, K., Boehm, J., Glenn, M., and Cornell, L. (2012). Changes in partial pressures of respiratory gases during submerged voluntary breath hold across odontocetes: is body mass important? Journal of Comparative Physiology B 182:299-309. (pdf)

Noren, S.R., Kendall, T., Cuccurullo, V., and Williams, T.M. (2012) The dive response redefined: underwater behavior influences cardiac variability in freely diving dolphins. Journal of Experimental Biology 215:2735-2741. (pdf)

Williams, T.M., Noren, S.R., and Glenn, M. (2010) Extreme physiological adaptations as predictors of climate-change sensitivity in the narwhal, Monodon monocerosMarine Mammal Science 27(2):334-349. (pdf)

Estes, J. A., Doak, D. F., Springer, A. M., Williams, T. M., & Van Vliet, G. B. (2009). Trend data do support the sequential nature of pinniped and sea otter declines in the North Pacific Ocean, but does it really matter? Marine Mammal Science, 25, 748–754. (pdf)

Estes, J. a, Doak, D. F., Springer,  a M., & Williams, T. M. (2009). Causes and consequences of marine mammal population declines in southwest Alaska: a food-web perspective. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 364(1524), 1647–1658. (pdf)

Springer, A. M., Estes, J. A., Van Vliet, G. B., Williams, T. M., Doak, D. F., Danner, E. M., & Pfister, B. (2008). Mammal-eating killer whales, industrial whaling, and the sequential megafaunal collapse in the North Pacific Ocean: A reply to critics of Springer et al. 2003. Marine Mammal Science, 24(2), 414–442. (pdf)

Noren, S.R., Cuccurullo, V., and Williams, T.M. (2004) The development of diving bradycardia in bottlenose dolphins (Tursiops truncatus). Journal of Comparative Physiology B 174:139-147. (pdf)

Noren, S.R., Lacave, G., Wells, R.S., and Williams, T.M. (2002)  The development of blood oxygen store in bottlenose dolphins (Tursiops truncatus): implications for diving capacity.  Journal of Zoology, London 258:105-113. (pdf)

Noren, S.R. and Williams, T.M. (2000)  Body size and skeletal muscle myoglobin of cetaceans: adaptations for maximizing dive duration.  Comparative Biochemistry and Physiology Part A 126:181-191. ()

Noren, D.P., Williams, T.M., Berry, P., and Butler, E. (1999)  Thermoregulation during swimming and diving in bottlenose dolphins, Tursiops truncatus.  Journal of Comparative Physiology B 169:93-99. (pdf)

Skrovan, R.C., Williams, T.M., Berry, P.S., Moore, P.W., and Davis, R.W. (1999)  The diving physiology of bottlenose dolphins (Tursiops truncatus), II. Biomechanics and changes in buoyancy at depth.  Journal of Experimental Biology 202:2749-2761. (pdf)

Williams, T.M., Haun, J.E., and Friedl, W.A. (1999)  The diving physiology of bottlenose dolphins (Tursiops truncatus), I. Balancing the demands of exercise for energy conservation at depth.  Journal of Experimental Biology 202:2739-2748. (pdf)

Williams, T.M., Noren, D., Berry, P., Estes, J.A., Allison, C., and Kirtland, J. (1999)  The diving physiology of bottlenose dolphins (Tursiops truncatus), III. Thermoregulation at depth.  Journal of Experimental Biology 202:2763-2769. (pdf)

Shaffer, S.A., Costa, D.P., Williams, T.M., and Ridgway, S.H. (1997)  Diving and swimming performance of white whales, Delphinapterus leucas: An assessment of plasma lactate and blood gas levels and respiratory rates.  Journal of Experimental Biology 200:3091-3099. (pdf)

Pabst, D.A., Rommel, A., McLellan, W.A., Williams, T.M., and Rowles, T.K. (1995) Thermoregulation of the intra-abdominal testes of the bottlenose dolphin (Tursiops truncatus) during exercise. Journal of Experimental Biology 198:221-226. (pdf)

Rommel, S.A., Pabst, D.A., McLellan, W.A., Williams, T.M., and Friedl, W.A. (1994) Temperature regulation of the testes of the bottlenose dolphin (Tursiops truncatus): evidence from colonic temperatures. Journal of Comparative Physiology B 164:130-134. (pdf)

Williams, T.M., Friedl, W.A., and Haun, J.E. (1993)  The physiology of bottlenose dolphins (Tursiops truncatus): Heart rate, metabolic rate and plasma lactate concentration during exercise.  Journal of Experimental Biology 179:31-46. (pdf)

Williams, T.M., Friedl, W.A., Fong, M.L., Yamada, R., Sedivy, P., and Haun, J.E. (1992) Travel at low energetic cost by swimming and wave-riding bottlenose dolphins. Nature 355:821-823. (pdf)


Liwanag, H. E. M., Oraze, J., Costa, D. P., & Williams, T. M. (2014). Thermal benefits of aggregation in a large marine endotherm: Huddling in California sea lions. Journal of Zoology, 293(3), 152–159. (pdf)

Maresh, J.L., Simmons, S.E., Crocker, D.E., McDonald, B.I., Williams, T.M., and Costa, D.P. (2014) Free-swimming northern elephant seals have low field metabolic rates that are sensitive to an increased cost of transport. Journal of Experimental Biology 217:1485-1495. (pdf)

Davis, R.W., Fuiman, L.A., Madden, K.M., and Williams, T.M. (2013) Classification and behavior of free-ranging Weddell seal dives based on three-dimensional movements and video-recorded observations. Deep-Sea Research II 88-89:65-77. (pdf)

Noren, D.P., Budge, S.M., Iverson, S.J., Goebel, M.E., Costa, D.P., and Williams, T.M. (2013) Characterization of blubber fatty acid signatures in northern elephant seals (Mirounga angustirostris) over the postweaning fast. Journal of Comparative Physiology B doi 10.1007/s00360-013-0773-0. (pdf)

Liwanag, H.E.M., Berta, A., Costa, D.P., Abney, M., and Williams, T.M. (2012) Morphological and thermal properties of mammalian insulation: the evolution of fur for aquatic living. Biological Journal of the Linnean Society 106:926-939. (pdf)

Liwanag, H.E.M., Berta, A., Costa, D.P., Budge, S.M., and Williams, T.M. (2012) Morphological and thermal properties of mammalian insulation: the evolutionary transition to blubber in pinnipeds. Biological Journal of the Linnean Society 107:774-787. (pdf)

Williams, T.M., Richter, B., Kendall, T., and Dunkin R. (2011) Metabolic demands of a tropical marine carnivore, the Hawaiian monk seal (Monachus schauinslandi): Implications for fisheries competition. Aquatic Mammals 37(3):372-376. (pdf)

Liwanag, H.E.M., Williams, T.M., Costa, D.P., Kanatous, S.B., Davis, R.W., and Boyd, I.L. (2009) The effects of water temperature on the energetic costs of juvenile and adult California sea lions (Zalophus californianus): the importance of skeletal muscle thermogenesis for thermal balance. Journal of Experimental Biology 212:3977-3984. (pdf)

Williams, T.M., Rutishauser, M., Long, B., Fink, T., Gafney, J., Mostman-Liwanag, H., and Casper, D. (2007) Seasonal variability in otariid energetics: Implications for the effects of predators on localized prey resources. Physiological and Biochemical Zoology 80(4):433-443. (pdf)

Rutishauser, M.R., Costa, D.P., Goebel, M.E., and Williams, T.M. (2004) Ecological implications of body composition and thermal capabilities in young Antarctic fur seals (Arctocephalus gazella). Physiological and Biochemical Zoology 77(4):669-681. (pdf)

Noren, D.P., Crocker, D.E., Williams, T.M., and Costa, D.P. (2003)  Energy reserve utilization in northern elephant seal (Mirounga angustirostris) pups during the postweaning fast: size does matter.  Journal of Comparative Physiology – B, Biochemical, Systemic, & Environmental Physiology 173:443-454. (pdf)

Davis, R.W., Fuiman, L.A., Williams, T.M., and Le Boeuf, B.J. (2001)  Three-dimensional movements and swimming activity of a northern elephant seal.  Comparative Biochemistry and Physiology A – Molecular and Integrative Physiology 129:759-770. (pdf)

Kohin, S., Williams, T.M., and Ortiz, C.L. (1999)  Effects of hypoxia and hypercapnia on aerobic metabolic processes in northern elephant seals.  Respiration Physiology 117:59-72. (pdf)

Boyd, I.L., Woakes, A.J., Butler, P.J., Davis, R.W., and Williams, T.M. (1995) Validation of heart rate and doubly labelled water as measures of metabolic rate during swimming in California sea lions. Functional Ecology 9(2):151-160. (pdf)

Castellini, M.A., Davis, R.W., Loughlin, T.R., and Williams, T.M. (1993) Blood chemistries and body condition of Steller sea lion pups at Marmot Island, Alaska. Marine Mammal Science 9(2):202-208. (pdf)

Davis, R.W., Castellini, M.A., Williams, T.M., and Kooyman, G.L. (1991) Fuel homeostasis in the harbor seal during submerged swimming. Journal of Comparative Physiology B 160:627-635. (pdf)

Williams, T.M., Kooyman, G.L., and Croll, D.A. (1991)  The effect of submergence on heart rate and oxygen consumption of swimming seals and sea lions.  Journal of Comparative Physiology  – B, Biochemical, Systemic, & Environmental Physiology 160(6):637-644. (pdf)

Davis, R.W., Williams, T.M., and Kooyman, G.L. (1985) Swimming metabolism of yearling and adult harbor seals Phoca vitulinaPhysiological Zoology 58(5):590-596. (pdf)

Williams, T.M. and Kooyman, G.L. (1985)  Swimming performance and hydrodynamic characteristics of harbor seals Phoca vitulina.  Physiological Zoology 58(5):576-589. (pdf)


Thometz N.M., Kendall T., Richter B., Williams T.M. (2016) The high cost of reproduction in sea otters necessitates unique physiological adaptations. The Journal of Experimental Biology 219(15): 2260-2264. (pdf)

Estes, J.A., Tinker, M.T., and Williams, T.M. (in press) Recent advances in the physiology, behavior and ecology of sea otters. In Biology and Conservation of the Musteloids (badgers, otters, skunks, raccoons and their kin). Oxford University Press, Oxford.

Kitchener, A., Melero, C., and Williams, T.M. (in press) Mustelid form and function. In Biology and Conservation of the Musteloids (badgers, otters, skunks, raccoons and their kin). Oxford University Press, Oxford.

Thometz, N.M., Tinker, M.T., Staedler, M.M., Mayer, K.A., and Williams, T.M. (2014) Energetic demands of immature sea otters from birth to weaning: implications for maternal costs, reproductive behavior, and population-level trends.  Journal of Experimental Biology 217(12): 2053-2061. (pdf)

Yeates, L.C., Williams, T.M., and Fink, T.L. (2007) Diving and foraging energetics of the smallest marine mammal, the sea otter (Enhydra lutris). Journal of Experimental Biology 210: 1960-1970. (pdf)

Williams, T.M., Ben-David, M., Noren, S., Rutishauser, M., McDonald, K., and Heyward, W. (2002)  Running energetics of the North American river otter: do short legs necessarily reduce efficiency on land?  Comparative Biochemistry and Physiology Part A 133: 203-212. (pdf)

Ben-David, M., Williams, T.M., and Ormseth, O.A. (2000)  Effects of oiling on exercise physiology and diving behavior of river otters: a captive study.  Canadian Journal of Zoology 78: 1380-1390. (pdf)

Estes, J.A., Tinker, M.T., Williams, T.M., and Doak, D.F. (1998)  Killer whale predation on sea otters linking oceanic and nearshore ecosystems.  Science 282: 473-476. (pdf)

Williams, T.M. (1989) Swimming by sea otters: adaptations for low energetic cost locomotion. Journal of Comparative Physiology A 164: 815-824. (pdf)

Davis, R.W., Williams, T.M., Thomas, J.A., Kastelein, R.A., and Cronell, L.H. (1988) The effects of oil contamination and cleaning on sea otters (Enhydra lutris). 2. Metabolism, thermoregulation, and behavior. Canadian Journal of Zoology 66(12): 2782-2790. (pdf)

Williams, T.M., Kastelein, R.A., Davis, R.W., and Thomas, J.A. (1988)  The effects of oil contamination and cleaning in sea otters (Enhydra lutris). 1. Thermoregulatory implications based on pelt studies.  Canadian Journal of Zoology 66(12): 2776-2781. (pdf)

Williams, T.M. (1986)  Thermoregulation of the North American mink during rest and activity in the aquatic environment. Physiological Zoology 59(3): 293-305. (pdf)

Williams, T.M. (1983)  Locomotion in the North American mink, a semi-aquatic mammal. I. Swimming energetics and body drag.  Journal of Experimental Biology 103: 155-168. (pdf)

Williams, T.M. (1983)  Locomotion in the North American mink, a semi-aquatic mammal. II. The effect of an elongate body on running energetics and gait patterns.  Journal of Experimental Biology 105: 283-295. (pdf)


Pagano, A.M., & Williams, T.M. (2019) Estimating the energy expenditure of free‐ranging polar bears using tri‐axial accelerometers: A validation with doubly labeled water. Ecology and Evolution. (pdf)

Suraci, J.P., Frank, L.G., Oriol‐Cotterill, A., Ekwanga, S., Williams, T.M. , & Wilmers, C.C. (2019) Behavior‐specific habitat selection by African lions may promote their persistence in a human‐dominated landscape. Ecology. (pdf)

Pagano, A.M., Cutting, A., Nicassio-Hiskey, N., Hash, A., & Williams, T.M. (2019) Energetic costs of aquatic locomotion in a subadult polar bear. Marine Mammal Science, 35,649-659. (pdf)

Pagano, A.M., Carnahan, A.M., Robbins, C.T., Owen, M.A., Batson, T., Wagner, N., Cutting, A., Nicassio-Hiskey, N., Hash, A., & Williams, T.M. (2018) Energetic costs of locomotion in bears: Is plantigrade locomotion energetically economical? Journal of Experimental Biology, 221. (pdf)

Pagano, A.M., Durner, G.M., Rode, K.D., Atwood, T.C., Atkinson, S.N., Peacock, E., Costa, D.P., Owen, M.A., & Williams, T.M. (2018) High energy-high fat lifestyle challenges an Arctic apex predator, the polar bear. Science, 359,568-572. (pdf)

Bryce, C.M., Williams, T.M., & Wilmers, C.C. (2017) Energetics and evasion dynamics of large predators and prey: pumas vs. hounds. PeerJ, 5, e3701, 1-23. (pdf)

Wilmers, C.C., Isbell, L.A., Suraci, J.P., & Williams, T.M. (2017) Energetics-informed behavioral states reveal the drive to kill in African leopards. Ecosphere, 8, e01850. (pdf)

Bryce, C.M., & Williams, T.M. (2017) Comparative locomotor costs of domestic dogs reveal energetic economy of wolf-like breeds. Journal of Experimental Biology, 220, 312–321. (pdf)

Pagano, A.M., Rode, K.D., Cutting, A., Owen, M.A., Jensen, S., Ware, J. V, Robbins, C.T., Durner, G.M., Atwood, T.C., Obbard, M.E., Middel, K.R., Thiemann, G.W., & Williams, T.M. (2017) Using tri-axial accelerometers to identify wild polar bear behaviors. Endangered Species Research, 32, 19–33. (pdf)

Williams, T.M., Wolfe, L., Davis, T., Kendall, T., Richter, B., Wang, Y., Bryce, C., Elkaim, G.H., and Wilmers, C.C.(2014) Instantaneous energetics of puma kills reveal advantage of felid sneak attacks. Science 346(6205): 81-85. (pdf)

Dunkin, R.C., Wilson, D., Way, N., Johnson, K., and Williams, T.M. (2013) Climate influences thermal balance and water use in African and Asian elephants: physiology can predict drivers of elephant distribution. Journal of Experimental Biology 216: 2939-2952. (pdf)

Wilmers, C.C., Wang, Y., Nickel, B., Houghtaling, P., Shakeri, Y., Allen, M.L., Kermish-Wells, J., Yovovich, V., and Williams, T.M. (2013) Scale dependent behavioral responses to human development by a large predator, the puma. PLOS ONE 8(4): e60590. doi:10.1371/journal.pone.0060590. (pdf)

Williams, T.M., Dobson, G.P., Mathieu-Costello, O., Morsbach, D., Worley, M.B., and Phillips, J.A. (1997)  Skeletal muscle histology and biochemistry of an elite sprinter, the African cheetah.  Journal of Comparative Physiology B 167: 527-535. (pdf)

Williams, T.M. (1990)  Heat transfer in elephants: Thermal partitioning based on skin temparature profiles. Journal of Zoology, London 222: 234-245. (pdf)

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