Rogers Carnivore Coexistence Lab

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Scientific Publications

To see the abstract of each article, roll your mouse over the authors' names (in blue. If you have trouble accessing a copy of an article, please email Adrian Treves.

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Coming soon: Why use pre-prints?

To reach a larger number of peer scientists before an article goes through the publication process, we present our work in pre-print format. This also has the advantage of reaching the public and policy-makers more quickly. The downside is if we get something wrong (demonstrated by peer-reviewed work or a pre-print shared with us) yet it reached the public and went into effect. We accept that risk because of the pressing public need for scientific information for public policy. We will clearly communicate any corrections if we find an error.

Treves, A., Agan, SW, Langenberg, JA, Lopez-Bao, J.V., Louchouarn, NX. Parsons, DR, Rabenhorst, MF, Santiago-Ávila, FJ, 2024.

Commentary: rebutting Roberts et al.-- gray wolf (Canis lupus) mortality patterns in Wisconsin: a critique. Journal of Mammalogy in press 2024. COMING SOON

Abi R. Fergus, Samuel J. Hermanstorfer, Adrian Treves in review. Combining two non-lethal methods in crossover design randomized experiments. Pre-print posted for pre-publication review at

Peer-reviewed or published with editorial review

Treves A, Khorozyan, I.

in review. Robust inference and errors in studies of wildlife control. Pre-print posted for pre-publication review at

Treves A, Fergus, AR, Hermanstorfer, SJ, Louchouarn, NX, Ohrens, O, Pineda Guerrero, AA. 2024.

Gold-standard experiments to deter predators from attacking farm animals. Animal Frontiers 14(1)"40-52.

After years of research, our conclusions about non-lethal deterrents and how to design randomized, controlled trials with crossover design.

(1) The long-held belief that randomized, controlled trials (RCTs) are impossible in wild ecosystems with working livestock is laid to rest.

(2) Crossover designs reduce most confounding variables between subjects and strengthen inference beyond the gold-standard of RCTs, yet we describe limitations precisely.

(3) Non-lethal methods can be effective in preventing carnivore approaches and attacks on working livestock in fenced pastures or open rangelands. The relationship between approaches and attacks remains uncertain.

(4) Lethal methods of predator control have been subjected to less robust study designs that suggest mixed results including increases in livestock losses.

(5) Non-lethal methods promise the elusive triple-win for wildlife, domestic animals, and livelihoods.

Treves, A. 2023.

Replace the ivory tower with the fire tower. Frontiers in Ecology and the Environment (guest editorial) p.355, doi:10.1002/fee.2676

Treves, A. 2023.

Replace ivory towers with fire towers. Frontiers in Ecology and the Environment (guest editorial) 2023.

Rogers A, Treves A, Karamagi R, Nyakoojo M, Naughton-Treves, L. 2023

Trenches reduce crop foraging by elephants: Lessons from Kibale National Park, Uganda for elephant conservation in densely settled rural landscapes. PLoS One 18(7): e0278501.

Louchouarn NX. 2023

Don’t judge the roar by its echo: Tests of assumptions, tools and policies for human-carnivore coexistence in North America. PhD dissertation, August 2023, Carnivore Coexistence Lab, Nelson Institute for Environmental Studies, University of Wisconsin-Madison.

Hermanstorfer, S. J. 2023

Western Colorado carnivore coexistence: Gold-standard non-lethal deterrent experiments and human-carnivore coexistence in Montrose, Colorado. Master's thesis, June 2023, Carnivore Coexistence Lab, Nelson Institute for Environmental Studies, University of Wisconsin-Madison.

Elbroch L & Treves A. (equal co-authors) 2023.

Why might removing carnivores maintain or increase risks for domestic animals? Biological Conservation 283:110106.

Treves, A., L. M. Elbroch, J. T. Bruskotter, 2024.

Evaluating fact claims accompanying policies to liberalize the killing of wolves, peer-reviewed chapter TBA, In press. Alpha Wildlife Publications, Canada.

Treves, A., Santiago-Ávila, F.J. 2023

Estimating wolf abundance with unverified methods. Academia Biology 1 doi 10.20935/AcadBiol6099 Compressed source documents

from the state (WDNR 2022 population reports, greensheet and Stauffer et al. 2021).

Treves, A.

'Best available science' and the reproducibility crisis. Frontiers in Ecology and the Environment 20(9):495, November 2022.

The infographic below refers to information contained in the above articles.


Treves A, Elbroch, LM, Koontz F, Papouchis CM. 2022.

How should scientific review and critique support policy? PLoS One Comment on Laundre & Papouchis 2020. Click here to access PLoS One pages

and full disclosure for the Comment

Treves, A., Elbroch, L.M. 2022.

Does killing wild carnivores raise risk for domestic animals? Wild Felid Monitor, the newsletter of the Wild Felid Research & Management Association. Summer 2022.

Louchouarn & Treves 2023.

Low-stress livestock handling protects cattle in a five-predator habitat. PeerJ in press, PeerJ 11:e14788

Supplementary Materials for the above article here.

Treves, Darimont, Santiago-Ávila 2022.

calling out irreproducible research underlying the State of Wisconsin's wolf policy set by the Department of Natural Resources.

Also attached to the original article here.

Treves and louchouarn 2022.

recalculating our 2022 model of the Wisconsin wolf population in April 2022 see here

using a different estimate of infant survival. We also enumerate the (now) five cases in which current or former WDNR employees failed to correct, retract, or inform the public of errors or omissions of important information about wolf science that could affect public policy.

Santiago-Ávila FJ, Agan S, W.,, Hinton JW, Treves A. 2022.

Evaluating how management policies affect red wolf mortality and disappearance. Royal Society Open Science 9:210400.

Treves, A., Louchouarn, N.X. 2022.

Uncertainty and precaution in hunting wolves twice in a year. PLoS One, 2022. 17(3): e0259604.

A third-party raised a concern about a typographical error in one of our sources (Thiel et al. 2009), so for due diligence we re-analyzed our data and published a comment here

However our conclusions did not change.

The DNR and NRB documents we cited from 2021.

The Supplementary Information from Treves & Louchouarn 2022.

Santiago-Ávila, F.J. & Treves, A. 2022.Poaching of protected wolves fluctuated seasonally and with non-wolf hunting. Scientific Reports 12:e1738.

Treves, A., F.J. Santiago-Ávila, and K. Putrevu 2021.

Quantifying the effects of delisting wolves after the first state began lethal management. PeerJ, 9:e11666, doc 10.7717/peerj.11666.

Treves A, Paquet PC, Artelle KA, Cornman AM, Krofel M, Darimont CT. 2021.

Transparency about values and assertions of fact in natural resource management. Frontiers in Conservation Science: Human-Wildlife Dynamics, 2:e631998, doc 10.3389/fcosc.2021.631998.

Treves, A., C. Batavia 2021.

Improved disclosures of non-financial competing interests would promote independent review. Academia Letters, 2021. Article 514: p. 1-9.

Agan, S.W., A. Treves, and E.L. Willey 2021.

Estimating poaching risk for the critically endangered wild red wolf (Canis rufus). PLoS One, 2021. 16(5):e0244261. DOI 10.1371.

Agan, S.W., A. Treves, and E.L. Willey 2021.

Majority positive attitudes cannot protect red wolves (Canis rufus) from a minority willing to kill illegally. Biological Conservation 109321. DOI

Louchouarn NX, Santiago-Ávila FJ, Parsons DR, Treves A.2021.

Evaluating how lethal management affects poaching of Mexican wolves. Royal Society Open Science 8 (registered report):: e2003300.

Just Preservation

In 2018, we published our ethic of public trusteeship, non-anthropocentric, multispecies justice that presents a method to give voice to future generations and to nonhumans when decisions are made to allocate or preserve nature. Start here with the original article Treves, A., Santiago-Ávila, F., Lynn, W.S. (equal co-authors) 2018. Just Preservation. Biological Conservation 229: 134-141.

The newest discussion of Just Preservation played out in 2021 in the journal Animal Sentience, which included commentaries by several dozen colleagues and our replies to each: F.J. Santiago-Ávila, A. Treves (equal co-authors), W.S. Lynn, Just preservation, trusteeship and multispecies justice. Animal Sentience 393. This continues our work on trusteeship, legal standing for nonhumans, and future generations, and equitable consideration of nonhumans as members of our moral community.

Treves, A. and N. J. Balster (2021

The effect of extended student hours on performance of students in an interdisciplinary, introductory undergraduate ecology course. North American Colleges and Teachers of Agriculture Journal, in press.

Tshabalala, T., McManus, J., Treves, A., Masocha, V., Faulconbridge, S., Schurch, M., Goets, S., Smuts, B. 2021.

Leopards and mesopredators as indicators of mammalian species richness across diverse landscapes of South Africa. Ecological Indicators 121, 107201.

Masters thesis

by Abi Fergus, M.S. December 2020.

Santiago-Ávila, F.J., R.J. Chappell, and A. Treves, 2020.

Liberalizing the killing of endangered wolves was associated with more disappearances of collared individuals in Wisconsin, USA. Scientific Reports 10:e13881.

Darimont, C.T., Hall, H., Mihalik, I., Artelle, K.A., Eckert, L., Treves A., Paquet, P.

Large carnivore hunting and the social license to hunt. Conservation Biology 35(4):1111-1119.

Carroll, C., Rohlf, D.J., von Holdt, B.M., Treves, A., Hendricks, S.A. 2020

Wolf delisting challenges demonstrate need for an improved framework for conserving intraspecific variation under the Endangered Species Act. Bioscience  biaa125,1-12. doi:10.1093/biosci/biaa125. .

With a podcast from four of the authors to explain the analysis and recommendations, in the wake of 2020 Trump Administration rule to delist the gray wolf nationwide Listen here (55 minutes).

Treves, Louchouarn, Santiago-Ávila. 2020.

Modelling concerns confound evaluations of legal wolf-killing. Biological Conservation. 249:108643,

Treves & Santiago-Ávila. 2020

Myths and assumptions about human-wildlife conflict and coexistence. Conservation Biology 10.1111/cobi.13472.

Treves, A. 2020

Standards of evidence in wild animal research. A report for the Brooks Institute for Animal Rights Law and Policy.

Treves 2020. Elephants and pandemics. Animal Sentience 28(20). URL

Treves, A., Krofel M, Ohrens O and van Eeden 2019

Predator Control Needs a Standard of Unbiased Randomized Experiments With Cross-Over Design. Frontiers in Ecology and Evolution 7:402-413. doi: 10.3389/fevo.2019.00462

Treves, A. 2019

Scientific ethics and the illusion of naïve objectivity. Guest editorial in Frontiers in Ecology and the Environment 7:1.

Treves, A. A. F. J. Santiago-Ávila, V. D. Popescu, P. C. Paquet, W. S. Lynn, C. T. Darimont, K. A. Artelle 2019

Trophy hunting: Insufficient evidence. Letter in Science 366(6464):435.

Ohrens, O., Bonacic, C., Treves, A. 2019. Non-lethal defense of livestock against predators: Flashing lights deter puma attacks in Chile. Frontiers in Ecology and the Environment 17(1):1-7.

van Eeden, L., Eklund, A., Miller, J.R.B.,...17 co-authors... Treves, A. (equal first authors) 2018. Carnivore conservation needs evidence-based livestock protection. PLOS Biology

. 2018. Van Eeden, Treves, Ritchie. The Conversation.

A short popular science summary of the above article.

Treves, A., Artelle, K.A., Paquet, P.C. 2018. Differentiating between regulations and hunting as conservation interventions. Conservation Biology 33(2):472–475. DOI:10.1111/cobi.13211.

Santiago-Avila, F.J., Lynn, W.S., Treves, A. 2018.

Inappropriate consideration of animal interests in predator management: Towards a comprehensive moral code. In Large Carnivore Conservation and Management: Human Dimensions and Governance, ed. T. Hovardos, Taylor & Francis, London.

Ohrens, O., Santiago-Avila, F.J., Treves, A.2019.

The challenges of preventing real and perceived threats to livestock. In Human-Wildlife Interactions: Turning Conflict into Coexistence, eds. B. Frank, S. Marchini, J. Glikman, Cambridge University Press: Cambridge.

Treves, A., K. A. Artelle, C. T. Darimont, W. S. Lynn, P. C. Paquet, F. J. Santiago-Avila, R. Shaw and M. C. Wood 018.

Intergenerational equity can help to prevent climate change and extinction. Nature Ecology & Evolution DOI: 10.1038/s41559-018-0465-y.

Supporting Data. @Burgessart Credit: Jen Burgess @jenburgessart @Burgessart

Credit: Jen Burgess @jenburgessart

Infographic below for 'Hallmarks of science missing from North American wildlife hunting and trapping plans': infographic

Artelle, K.A., Reynolds, J.D., Treves, A. Walsh, J.C., Paquet, P.C., Darimont, C.T. 2018.

Hallmarks of science missing from North American wildlife management. Science Advances. 2018.

short video explaining the findings

Santiago-Avila, F.J., Cornman, A.M., Treves, A. 2018.

Killing wolves to prevent predation on livestock may protect one farm but harm neighbors. PLOS One 13:e0189729 here

Chapron, G. and A. Treves 2016a and b, 2017a and b.

We first showed that Michigan and Wisconsin wolf population growth slowed whenever the government liberalized wolf-killing and the slow-down was proportional to the length of time that culling was liberalized, regardless of how many wolves were killed. Then starting a lively debate, Pepin et al. tried to counter our hypothesis but did not succeed in our opinion. That debate improved our model which strengthened its findings here Strengthening our findings.


a 2017 rebuttal.

Then Stien and Olson and his colleagues tried again.

Those critiques only made our evidence stronger.

Olson et al. in particular weakened their own position. Furthermore, independent findings for Mexican wolves presented by David Parsons in 2014 corroborate the idea that relaxing protections slows population growth more than expected. Furthermore work by Dr. F. J. Santiago-Ávila et al. i 2020, 2022a,b, and N. Louchouarn in 2022 corroborated the finding that reducing protections for wolves increased poaching. These three studies used independent datasets from red wolves in North Carolina, Mexican wolves in southwest USA and Wisconsin wolf survival rates. In sum, the debate is over: blood does not buy goodwill.

Treves A, Rabenhorst MF. 2017.

Risk Map for Wolf Threats to Livestock still Predictive 5 Years after Construction. PLoS ONE:

Lopez-Bao, J.V., Chapron, G., Treves, A. 2017.

The Achilles heel of participatory conservation. Biological Conservation 212: 139-143.

Treves, A., Artelle, K.A., Darimont, C.T., Parsons, D.R. 2017.

Mismeasured mortality: correcting estimates of wolf poaching in the United States. Journal of Mammalogy 98(3): open access at DOI:

Summary and FAQs

Darimont, C.T., Paquet, P., Treves, A., Artelle, K.A., Chapron, G. 2018. Political populations of large carnivores.Conservation Biology 32(3):747-749.

Carroll, C., B. Hartl, G.T. Goldman, D.J. Rohlf, A. Treves, J.T. Kerr, E.G. Ritchie, R.T. Kingsford, K.E. Gibbs, M. Maron, J.E.M Watson. 2017. Defending scientific integrity in conservation policy processes: lessons from Canada, Australia, and the United States. Conservation Biology DOI: 10.1111/cobi.12958

Treves, A., J.A. Langenberg, J.v. López-Bao, M.F. Rabenhorst 2017. Gray wolf mortality patterns in Wisconsin from 1979 to 2012 Journal of Mammalogy 98(1): DOI:10.1093/jmammal/gyw145

Treves, A., Krofel, M., McManus, J. (equal co-authors).2016.

Predator control should not be a shot in the dark. Frontiers in Ecology and the Environment14: 380-388. This article has the highest Altmetric score among articles of the same age in that journal, ranks in the 99th percentile for all research outputs of similar age, and the top 5% for research outputs of all types and ages; see for details.

In a nutshell: