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"Nothing in evolutionary biology makes sense except in the light of ecology"

- Peter & Rosemary Grant -

Publications (peer-reviewed articles)

Please contact me  at (pwestley at for reprint requests.

*Paper featured on journal cover

‡ Shared lead authorship

†Student at time of research


M. Vignon, M. Zhou, A.R. McIntosh, C. Correa, P.A.H. Westley, L. Jacquin, J. Labonne, A.P. Hendry. Trait variation in a successful global invader: a large-scale analysis of morphological variance and integration in the brown trout. Biological Invasions, DOI: Link



Walther, E. J., D.E. Arthur, A. Cyr, K.M. Fraley, T. Cubbage, E. Hinkle, J. McMahon, P.A.H. Westley. Ecotoxicology of mercury in burbot (Lota lota) from interior Alaska and insights towards human health. Chemosphere, DOI: Link

Walther, E.J., M.S. Zimmerman, J.A. Falke, P.A.H. Westley. Species distributions and the recognition of risk in restoration planning: A case study of salmonid fishes.

Ecological Applications, DOI: Link

Von Biela, V.R. , C.J. Sergeant, M.P. Carey, Z. Liller, C. Russell, S. Quinn‐Davidson, P.S. Rand, P.A.H. Westley, C.E. Zimmerman. Premature mortality observations among Alaska’s Pacific Salmon during record heat and drought in 2019. Fisheries, DOI: Link


Ulaski, M.E., H. Finkle, A.H. Beaudreau, P.A.H. Westley. Climate and conspecific density inform phenotypic forecasting of juvenile Pacific salmon body size.

Freshwater Biology, DOI: Link



Westley, P.A.H., J. Black, C. Carothers, D. Ringer. State of Alaska's salmon and people: introduction to a special feature. Ecology and Society, DOI: Link

Stratton, M.E., H. Finkle, J.A. Falke, and P.A.H. Westley. 2021. Assessing potential stock structure of adult coho salmon in a small Alaska watershed: quantifying run timing, spawning locations, and holding areas with radiotelemetry. North American Journal of Fisheries Management, DOI: Link

Walther, E.J.M.S. Zimmerman, and P.A.H. Westley. 2021. Landscape attributes explain salmonid ecological neighbourhoods across a complex river network. Freshwater Biology, DOI: Link

Jalbert, C.S., J.A. Falke, J.A. López, K.J. Dunker, A.J. Sepulveda, and P.A.H. Westley. 2021. Vulnerability of Pacific salmon to invasion of northern pike (Esox lucius) in Southcentral Alaska. Plos One, DOI:10.1371/journal.pone.0254097. Link

Manishin, K.A., C.J. Cunningham, P.A.H. Westley, and A.C. Seitz. 2021. Can late stage marine mortality explain observed shifts in age structure of Chinook salmon? Plos One, DOI:10.1371/journal.pone.0247370. Link

O’Toole, C., K.P. Phillips, C. Bradley, J. Coughlan, E. Dillane, I.A. Fleming, T.E. Reed, P.A.H. Westley, T.F. Cross, P. McGinnity, P.A. Prodöhl. Population genetics reveal patterns of natural colonisation of an ecologically and commercially important invasive fish. Canadian Journal of Fisheries and Aquatic Sciences, DOI: Link


Okasaki, C., M.L. Keefer, P.A.H. Westley, and A.H. Berdahl. 2020. Collective navigation can facilitate passage through human-made barriers by homeward migrating Pacific salmon. Proceedings of the Royal Society B, DOI:10.1098/rspb.2020.2137. Link

Oke, K.B., C.J. Cunningham, P.A.H. Westley, M.L. Baskett, S.M. Carlson, J. Clark, A.P. Hendry, V.A. Karatayev, N.W. Kendall, J. Kibele, H.K. Kindsvater, K.M. Kobayashi, B. Lewis, S. Munch, J.D. Reynolds, G.K.Vick, and E.P. Palkovacs. 2020. Recent declines in salmon body size impact ecosystems and fisheries. Nature Communications, DOI: 10.1038/s41467-020-17726-z. Link

Westley, P.A.H. 2020. Documentation of en route mortality of summer chum salmon in the Koyukuk River, Alaska and its potential linkage to the heatwave of 2019. Ecology and Evolution, DOI:10:10296-10304. Link

Ulaski, M.E.†, H. Finkle, and P.A.H. Westley. 2020. Direction and magnitude of natural selection on body size differs among age classes of seaward migrating Pacific salmon. Evolutionary Applications, DOI: 10.1111/eva.12957. Link

Sawyer A.C., J.W. Moore, D.E. Schindler, P.A.H. WestleyConnecting Salmon Science in an Era of Global Change. Fisheries, DOI: Link


C. Carothers, T.L. Sformo, S. Cotton, J.C. George, P.A.H. Westley. Pacific Salmon in the rapidly changing Arctic. Arctic, DOI: Link

McConnell, C.J†, S. Atkinson, D. Oxman, and P.A.H. Westley. Is blood cortisol or vateritic otolith composition associated with natal dispersal or reproductive performance on the spawning grounds of straying and homing hatchery-produced chum salmon (Oncorhynchus keta) in Southeast Alaska?. Biology Open, DOI: Link

Cathcart, C.N., K. J. Dunker, T. P. Quinn, A.J. Sepulveda, F. A. von Hippel, A. Wizik, and D. B. Young, and P.A.H. Westley 2019. Trophic plasticity and the invasion of a renowned piscivore: A diet synthesis of northern pike (Esox lucius) from the native and introduced range of Alaska, U.S.A. Biological Invasions 21: 1379-1392.

Westley, P.A.H, 2019. Ocean Ecology of Pacific Salmon and Trout. Edited by Richard Beamish. Invited Book Review, Quarterly Review of Biology 94: 296

Manishin, K.A., K.J. Goldman, M. Short, C.J. Cunningham, P.A.H. Westley, A.C. Seitz. Prey consumption estimates for salmon sharks. Marine and Freshwater Research, DOI: Link

Berghaus, K.I., J.R. Spencer, P.A.H. Westley. Contemporary phenotypic divergence of an introduced predatory freshwater fish, the northern pike (Esox lucius). Evolutionary Ecology Research. Link

Carothers, C.C., T.L. Sformo, S. Cotton, J.C. George, and P.A.H. Westley. Pacific salmon in the rapidly changing Arctic: Exploring local knowledge and emerging fisheries in Utquiġvik and Nuiqsut, Alaska. Arctic 72: 273-288.


Manishin, K. A.†, K. J. Goldman, M. Short, C. J. Cunningham, P.A.H. Westley, and A. C. Seitz. Exploring the potential role of an apex marine predator: Prey consumption estimates of salmon sharks in the North Pacific Ocean. Marine and Freshwater Research

Langdon, S., P.A.H. Westley, D. Rinella. Deep Time—Salmon and People in Alaska Before Contact SASAP: State of Alaska Salmon and People. Link

Sparks, M.M. †, J.A. Falke, T.P. Quinn, M.D. Adkison, D.E. Schindler, K. Bartz, D. Young, and P.A.H. Westley. In press. Influences of spawning timing, water temperature, and climatic warming on early life history phenology in western Alaska sockeye salmon. Canadian Journal of Aquatic and Fisheries Sciences 123-135.


Cunningham, C.J., P.A.H. Westley, and M.D. Adkison. Signals of climate drivers, hatchery enhancement, and marine factors in Yukon River Chinook salmon survival revealed with a Bayesian life history model. Global Change Biology 24: 4399-4416.

Courtney, M., E. Schoen, A. Wizik, and P.A.H. Westley. Quantifying the net benefits of suppression: truncated size structure and consumption of native salmonids by invasive Northern Pike in an Alaska lake. North American Journal of Fisheries Management 38: 1306-1315.  


Berdahl, A.M., A. B. Kao, A. Flack, P.A.H. Westley, E.A. Codling, I.D. Couzin, A.I. Dell, and D. Biro. Collective animal navigation and migratory culture: from theoretical models to empirical evidence. 2018. Philosophical Transactions B 373: 20170009.


Yeakel, J.D., J.P. Gilbert, T. Gross, P.A.H Westley, and J.W. Moore. Eco-evolutionary dynamics, density-dependent dispersal and collective behaviour: implications for salmon metapopulation robustness. Philosophical Transactions B 373: 20170018.


Westley, P.A.H, A.M. Berdahl, C.J. Torney, and D. Biro. Collective movement in ecology: from emerging technologies to conservation and management. Philosophical Transactions B 373: 20170004.


McConnell, C.J†., P.A.H. Westley, and Megan V. McPhee. Differences in fitness-associated traits between hatchery and wild chum salmon despite long-term immigration by strays. Aquaculture Environment Interactions 10: 99-113.


Sparks, M. M.†, P.A.H. Westley, J. A. Falke, and T. P. Quinn. Thermal adaptation and phenotypic plasticity in a warming world: Insights from common garden experiments on Alaskan sockeye salmon*. Global Change Biology 23: 5203-5217.


Kovach, R. P., C. C. Muhlfeld, R. Al-Chokhachy, S. J. Amish, J. L. Kershner, R. F. Leary, W. H. Lowe, G. Luikart, P. Matson, D. A. Schmetterling, B. B. Shepard, P. A. H. Westley, D. Whited, A. Whiteley, and F. W. Allendorf. No evidence for ecological segregation protecting native trout from invasive hybridization. Global Change Biology 23: e11-e12 (response to editor)


Muhlfeld, C. C., R. P. Kovach, R. Al-Chokhachy, S. J. Amish, J. L. Kershner, R. F. Leary, W. H. Lowe, G. Luikart, P. Matson, D. A. Schmetterling, B. B. Shepard, P. A. H. Westley, D. Whited, A. Whiteley, and F. W. Allendorf. Legacy introductions and climatic variation explain spatiotemporal patterns of invasive hybridization in a native trout. Global Change Biology 23: 4663-4674.


Berdahl, A., P.A.H. Westley, and T. P. Quinn. Social interactions shape the timing of spawning migrations in an anadromous fish. Animal Behaviour 126:221-229.


Hauser, W.J., P.A.H. Westley‡, C. Kerkvliet, and N. Dudiak. Homing of Pacific Salmon to a Marine Release Site: A Case Study of the Homer Spit Fishing Hole, Alaska. Northwest Science 91:314-323.


Bond, M.H, P.A.H. Westley, A. H. Dittman, D. Holecek, T. Marsh, and T. P. Quinn. Combined effects of rearing location, barge transportation, and river environments on the migration rate and homing of Snake River Chinook salmon. Transactions of the American Fisheries Society 146: 60-73.


Berdahl, A., P.A.H. Westley, S. Levin, I. Couzin, and T.P. Quinn. A collective navigation hypothesis for homeward migration in anadromous salmonids. Fish and Fisheries, 17: 525-542.


Westley, P.A.H, A.H. Dittman, E.J. Ward, and T.P. Quinn. Signals of climate, conspecific density, and watershed features in patterns of homing and dispersal by Pacific salmon. Ecology, 96: 2823-2833.


Carlson, S.M, C. Cunningham, and P.A.H. Westley‡. Evolutionary rescue in a changing world*. Trends in Ecology and Evolution, 29: 521-530.

Veinott, G., P.A.H Westley, C.F. Purchase, and L.Warner. Experimental evidence simultaneously confirms and contests assumptions implicit to otolith microchemistry research. CJFAS, 71: 356-365


Westley, P.A.H, R. Stanley, and I.A. Fleming. Experimental tests for heritable morphological color plasticity in non-native brown trout. PlosONE, DOI: 10.1371/journal.pone.0080401


Oke, K†., P.A.H. Westley, D.T.R. Moreau, and I.A. Fleming. Hybridization between genetically-modified Atlantic salmon and wild brown trout reveals novel ecological interactions. Proceedings of the Royal Society B 280: 20131047 (Faculty 1000 recommended)


Westley, P.A.H, T.P. Quinn, and A.H. Dittman. Straying in hatchery produced Pacific salmon and steelhead differs among species, life history forms, and populations. Canadian Journal of Fisheries and Aquatic Sciences 70: 735-746.


Westley, P.A.H, E.J. Ward, and I.A. Fleming. Fine-scale local adaptation in an invasive freshwater fish has evolved in contemporary time*. Proceedings of the Royal Society B: Biological Sciences 280: 20122327.

Westley, P.A.H, C.M. Conway, and I.A. Fleming. Phenotypic divergence of exotic fish populations is shaped by spatial proximity and habitat differences across an invaded landscape. Evolutionary Ecology Research 14: 147-167.


Veinott, G., P.A.H Westley, C.F. Purchase, and L.Warner. Assigning origins in a potentially mixed-stock recreational sea trout (Salmo trutta) fishery. Ecology of Freshwater Fish 21: 541-551.


Westley, P.A.H. and I.A. Fleming. Landscape factors that shape a slow and persistent biological invasion: brown trout in Newfoundland 1883-2010*. Diversity & Distributions 17: 566-579.


Westley, P.A.H. What invasive species reveal about the rate and form of contemporary phenotypic change in nature. American Naturalist 177: 496-509.


Westley, P.A.H., D.E. Schindler, T.P. Quinn, G.R. Ruggerone, and R. Hilborn. Natural habitat change, commercial fishing, climate, and dispersal interact to restructure an Alaskan fish metacommunity. Oecologia 163: 471-484.


Westley, P.A.H., R. Hilborn, T.P. Quinn, G.R. Ruggerone, and D.E. Schindler. Long-term changes in rearing habitat and downstream movement by juvenile sockeye salmon (Oncorhynchus nerka) in an interconnected Alaska lake system. Ecology of Freshwater Fishes 17: 443-454.


Westley, P.A.H†, S.M. Carlson, and T.P. Quinn. Among-population variation in adipose fin size parallels the expression of other secondary sexual characteristics in sockeye salmon (Oncorhynchus nerka). Environmental Biology of Fishes 81: 439-446.

Publications (peer-reviewed technical reports)


1. Westley, P.A.H, D. W. Ings, and I.A .Fleming. 2011. A review and annotated bibliography of the impacts of invasive brown trout (Salmo trutta) on native salmonids, with an emphasis on Newfoundland waters. Canadian Technical Report of Fisheries and Aquatic Sciences 2924, 81 p.

2. Westley, P.A.H and R. Hilborn.  2006.  Chignik Salmon Studies: Investigations of Salmon Populations, Hydrology, and Limnology of the Chignik Lakes, Alaska, During 2005-2006.  Available at:

3. Westley, P.A.H, B.E. Chasco, and R Hilborn.  2005. Chignik Salmon Studies: Investigations of Salmon Populations, Hydrology, and Limnology of the Chignik Lakes, Alaska, During 2004-2005.  Available at: .

Popular literature and Outreach

1. Alaska at the frontlines of biological invasions: A call to arms for salmon-dependent stakeholders to unite. Article in 'Smolt'- Newsletter of Cook Inlet Aquaculture Association.


2. Interview on local CBC news to discuss brown trout in Newfoundland, aired May 24, 2008.  Can be viewed on YouTube at:

3. Westley, P.A.H. 2008. One trout, two trout, brook trout, brown trout: an overview of the wonderfully bizarre biology of salmonids. The Osprey (nature journal of Newfoundland and Labrador) 39: 139-143.

4. Westley, P.A.H. 2008. Life-time angler with passion for salmon and trout biology lands ideal job in Newfoundland. Spawner Magazine. Published by the Salmon Preservation Association for the Waters of Newfoundland. Page 75.

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