Category Archives: wildlife management

Whither Demography in an Era of Biodiversity Science?

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Biodiversity science has overtaken traditional ecological science in an important way that bears some discussion. Biodiversity science seeks as its main goal to protect species from declining to extinction. It overlaps traditional population and community ecology partly by concentrating on iconic species that are declining in abundance and thus trying to prevent species loss, but the major focus of biodiversity science is on the species composition of communities and ecosystems and trying to understand what factors are driving deleterious changes. This task is difficult to complete currently because of a shortage of research time, person-power and money, so we are driven to select relatively few ecosystems to concentrate our conservation efforts upon. If we cannot do everything for all species, we must choose what to do at the present time, and this releases a cascade of discussions and arguments about what species are ‘flagship’, “umbrella”, or “keystone” in any particular community or ecosystem (Barua 2011).

Now we can go in two directions. We can take the simplest route and say that all species are of equal importance, so the objective of biodiversity science becomes ‘occupancy” – is species X still existing in ecosystem Y? Occupancy can be established in a variety of ways from simple visual sightings, traps, cameras, e-DNA, to electronic sound recording devices. Occupancy for species in a particular ecosystem is a useful parameter for biodiversity studies but it is alpha-level ecology for understanding community or ecosystem dynamics. Determining occupancy every month, year, or decade will provide a start in community and ecosystem understanding but in order to achieve its goal as a science it needs population and community ecology to measure and understand the dynamics underlying occupancy, and this is the more complicated route that we can take. The defining science for this second level of understanding comes from dynamics, both population and community dynamics.

Population dynamics is necessary for determining if a particular species is declining in numbers or biomass, and what the causes are of the observed trends. We are led into demography, the measurement of births, deaths, and movements in animals and the equivalent parameters in plants. But now we run into the most serious problem of determining what parameters are the causal agents of declines in abundance and what procedures will alleviate species declines. If we have many species in our community or ecosystem, the requirements for research are extravagant, given the current workers and dollars currently available for environmental science. All ecologists want to protect biodiversity but how might we best achieve that goal?

We fall back at this point into simple general procedures for biodiversity conservation like designation of national parks or protected areas with the hope that the biodiversity of the designated area will not decline. We do not have the person-power to have frequent occupancy surveys for any national park, much less to have the investigations of why an iconic species is declining in an area. At present we must fall back on ‘umbrella species’ or ‘flagship species’ (Simberloff 1998). There is an extensive literature on this approach to biodiversity conservation and recent reviews, some critical (Tälle et al. 2023), others somewhat more positive (Sumbh and Hof 2022, Clark-Wolf et al. 2024), combining this approach with food web structure (Wang and Zhou 2023).   

Complicating the whole issue of protecting biodiversity is the issue of cryptic species, undescribed species, and rare species that are capable of taxonomic and ecological resolution but only at a large cost and a long timeframe to achieve results (Cheng et al. 2024). Once we are successful in protecting our communities and ecosystems, we immediately face the demographic issue of what affects the abundance of all the species of interest, and then the social and political issue of the funds available for conservation of these species. We need to bring the approaches of biodiversity science and classical demographic ecology together to achieve conservation goals. We can do this only by recognizing that knowing occupancy is not enough to achieve conservation success, and we need to follow up with population and community ecology within the context of food webs so that we can understand trends in abundance and finally propose possible actions for conservation management. We have much to do.

Barua, M. (2011) Mobilizing metaphors: the popular use of keystone, flagship and umbrella species concepts. Biodiversity and Conservation, 20, 1427-1440.doi:10.1007/s10531-011-0035-y.

Cheng, R., Luo, A., Orr, M., Ge, D., Houu, Z.e., Qu, Y., Guo, B., Zhang, F., Sha, Z., Zhao, Z., Wang, M., Shi, X., Han, H., Zhou, Q., Li, Y., Liu, X., Shao, C., Zhang, A., Zhou, X. & Zhu, C. (2024) Cryptic diversity begets challenges and opportunities in biodiversity research. Integrative Zoology, (in press). doi: 10.1111/1749-4877.12809.

Clark-Wolf, T.J., Holt, K.A., Johansson, E., Nisi, A.C., Rafiq, K., West, L., Boersma, P.D., Hazen, E.L., Moore, S.E. & Abrahms, B. (2024) The capacity of sentinel species to detect changes in environmental conditions and ecosystem structure. Journal of Applied Ecology, (in press). doi: 10.1111/1365-2664.14669.

Simberloff, D. (1998) Flagships, umbrellas, and keystones: is single-species management passe in the landscape era? Biological Conservation, 83, 247-257.doi: 10.1016/S0006-3207(97)00081-5.

Sumbh, O. & Hof, A.R. (2022) Can pikas hold the umbrella? Understanding the current and future umbrella potential of keystone species Pika (Ochotona spp.). Global Ecology and Conservation, 38, e02247.doi: 10.1016/j.gecco.2022.e02247.

Tälle, M., Ranius, T. & Öckinger, E. (2023) The usefulness of surrogates in biodiversity conservation: A synthesis. Biological Conservation, 288; e110384. doi: 10.1016/j.biocon.2023.110384.

Wang, Q., Li, X.C. & Zhou, X.H. (2023) New shortcut for conservation: The combination management strategy of “keystone species” plus “umbrella species” based on food web structure. Biological Conservation, 286, 110265.doi: 10.1016/j.biocon.2023.110265.

On Discourse and Evidence

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A major problem that bedevils society as well as science today is the distinction between opinion and evidence. The world is awash in opinions and short of evidence for many questions that fill the news media as well as the scientific literature. In trying to evaluate this statement we should recognize that there are many important issues for which we have no evidence but only beliefs. A current discussion in the news media is whether or not any particular country should spend 2% of its GDP on military expenses. Much hot air follows from these discussions because as an individual person you have no evidence one way or the other for different points of view, spend more, spend less. You will have an opinion that everyone should respect but for this and many issues any evidence that can be cited is vague. Discussion on many issues like this example are important and should be civilized but often are not.

But there are a range of issues for which scientific evidence is available. The first rule of discourse on these issues ought to be that you as a person are allowed to have any opinion you wish but you must be able to present evidence to support your opinions. You should be allowed as a person to proclaim that the Earth is not round but flat and provide the evidence one way or the other. More serious issues with different opinions involve issues like vaccination for a particular disease. On these issues scientists can only advise and provide evidence. But if a vaccine X for example has a complex side effect rate of 1 person in 1000, you could always argue that you are that one person and you are opposed to vaccination X.

How does all this relate to ecological science? First, we should recognize that many of the arguments in the ecological literature are about opinions rather than evidence. In many cases this should lead to more studies of particular problems to gather more evidence. But as we see with climate change research the evidence is accumulating but varies greatly in quality and time span from area to area and from taxonomic group to taxonomic group. We cannot agree whether our research should be focused on the oceans or on land, or on birds rather than mammals or insects. We cannot do everything, and the consequence is that ecological research funding is driven in many directions depending on who is on the committee dispersing funding and what their opinions are. The result is that for large scale problems like climate change we have convinced most people that it is a reality, but we cannot agree on the details of future change. So, we build models with past data and try to project them into the future with uncertain confidence.

The consequence is that the ecological world is awash in opinions in the same way as other parts of society, and in support of opinions the evidence often gets lost. The main problem here is that opinions are generated rapidly while evidence accumulates slowly. We see this more readily in medical science in which the media trumpets treatment X rather than Y with opinions and little evidence. We cannot demand answers to important questions tomorrow when the problem spans years or decades for evidence to accumulate. 

Ecology like every science becomes more complicated with age, and a fisheries biologist trained 40 years ago lives in a different world from one trained today. The accumulated evidence from research changes our list of important questions illustrated well by the reviews of progress in conservation science by Sutherland et al. (2022, 2023) and Christie et al. (2023), in predator- prey dynamics by Sheriff et al. (2020), wildlife management by Hone et al. (2023), and in insect conservation by Saunders et al. (2020). Understanding and solving ecology problems must rely more on evidence and less on opinions.

Christie, A.P., Christie, A.P., Morgan, W.H. & Sutherland, W.J. (2023) Assessing diverse evidence to improve conservation decision‐making. Conservation Science and Practice, 5, e13024.doi. 10.1111/csp2.13024

Hone, J., Drake, A. & Krebs, C.J. (2023) Evaluation Options for Wildlife Management and Strengthening of Causal Inference. BioScience, 73, 48-58.doi: 10.1093/biosci/biac105.

Saunders, M.E., Janes, J.K. & O’Hanlon, J.C. (2020) Moving On from the Insect Apocalypse Narrative: Engaging with Evidence-Based Insect Conservation. BioScience, 70, 80-89.doi: 10.1093/biosci/biz143.

Sheriff, M.J., Peacor, S.D., Hawlena, D. & Thaker, M. (2020) Non-consumptive predator effects on prey population size: A dearth of evidence. Journal of Animal Ecology, 89, 1302-1316.doi: 10.1111/1365-2656.13213.

Sutherland, W.J. & Jake M. Robinson, D.C.A., Tim Alamenciak, Matthew Armes, Nina Baranduin, Andrew J. Bladon, Martin F. Breed, Nicki Dyas, Chris S. Elphick, Richard A. Griffiths, Jonny Hughes, Beccy Middleton, Nick A. Littlewood, Roger Mitchell, William H. Morgan, Roy Mosley, Silviu O. Petrovan, Kit Prendergast, Euan G. Ritchie,Hugh Raven, Rebecca K. Smith, Sarah H. Watts, Ann Thornton (2022) Creating testable questions in practical conservation: a process and 100 questions. Conservation Evidence Journal, 19, 1-7.doi. 10.52201/CEJ19XIFF2753

Sutherland, W.J., Sutherland, W.J., Bennett, C. & Thornton, A. (2023) A global biological conservation horizon scan of issues for 2023. Trends in Ecology & Evolution, 38, 96-107.doi. 10.1016/j.tree.2022.10.005

On Critical Evaluation in Ecology

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Science proceeds by “conjecture-and-refutation” if we agree with Karl Popper (1963). There is a rich literature on science in general and ecological science in particular that is well worth a series of graduate discussions even if it is pre-2000 ancient history (Peters 1991, Weiner 1995, Woodward and Goodstein 1996). But I wish to focus on a current problem that I think is hindering ecological progress. I propose that ecological journals at this time are focusing their publications on papers that present apparent progress and are shedding papers that are critical of apparent progress. Or in Popper’s words, they focus on publishing ‘conjecture’ and avoid ‘refutation’. The most important aspect of this issue involves wildlife management and conservation issues. The human side of this issue may involve personal criticism and on occasion the loss of a job or promotion. The issue arises in part because of a confusion between the critique of ideas or data and the interpretation that all critiques are personal. So, the first principle of this discussion is that I discuss here only critiques of ideas or data.

There are many simple reasons for critiques of experimental design and data gathering. Are the treatments replicated, are the estimates of data variables reliable and sufficient, are proxy variables good or poor? Have the studies been carried out long enough? All these critiques can be summarized under the umbrella of measurement reliability. There are many examples we can use to illustrate these ideas. Are bird populations declining across the globe or locally? Are fisheries overharvesting particular species? Can we use climate change as a universal explanation of all changes in wildlife populations? Are survey methods for population changes across very large areas reliable? The problem is tied into the need for good or bad news that must be filtered to the news media or social media with high impact but little reliability. 

The problem at the level of science is the temptation to extrapolate beyond the limits of the available data. Now we come to the critical issue – how do our scientific journals respond to critical reviews of papers already published? My concern is that in the present time journals do not wish to receive or accept manuscripts that are critical of previously published papers. These decisions are no doubt confidential for journal publishers. There is perhaps some justification for this rejection policy, given that in the few cases where critiques are published on existing papers, the citation score of the original paper may greatly exceed that of the critique. So, conjecture pays, refutation does not.

Journals are flooded with papers and for the better journals I would expect at least a 60-80% rejection rate. For Science the rejection rate is 94%, for Nature 92%, and for the Journal of Animal Ecology 85% of submitted manuscripts are rejected. Consequently, the suggestion that they reserve space for ‘refutation’ is too negative to their publication model. There is little I can suggest if one in caught in this dilemma except to try another less premium journal, and remember that web searches find papers easily no matter where published. If you need inspiration, you can follow Peters (1991) and write a book critique and suffer the brickbats from the establishment (e.g. Nature 354: 444, 12 December 1991).

But if you are upset about a particular paper or series of papers, remember critiques are valuable but follow these rules for a critique:

  1. Keep it short, 5 typed pages should be near maximal length.
  2. Raise a set of major points. Do not try to cover everything.
  3. Summarize briefly the key points you are in agreement with, so they are not confounded in the discussion.
  4. Discuss what studies might distinguish hypothesis A vs B, or A+B vs C.
  5. Discuss what better methods of measurement might be used if funding is available.
  6. Never attack individuals or research groups. The discussion is about ideas, results, and inferences.

Decisions to accept some management actions may have to be taken immediately and journal editors must take that into consideration. Prognostication over accepting critiques may be damaging. But all actions must be continually evaluated and changed once the understanding of the problem changes.

There are too many examples to recommend reading about past and present controversies in ecology, so here are only two examples. Dowding et al. (2009) report a comment on suggested methods of controlling introduced pests on Macquarie Island in the Southern Ocean. I was involved in that discussion. A much bigger controversy in Canada involves Southern Mountain caribou populations which are in rapid decline. The proximate explanation for the decline is postulated to be predation by wolves and thus the suggested management action is shooting the wolves. Johnson et al. (2022), Lamb et al. (2022) and Superbie et al. (2022) provide an entre into this literature and the decisions of what to do now and in the future to prevent extinction of these ungulates. The caribou problem is complicated by the interaction of human alteration of landscapes and the natural processes of predation and food availability. Alas nothing is simple.

All these ecological dilemmas are controversial and the important role of criticism involving evaluations of alternative hypotheses are the only way forward for ecologists involved in controversies. In my opinion most ecological journals are not doing their part is publishing critiques of the conventional wisdom.

Dowding, J.E., Murphy, E.C., Springer, K., Peacock, A.J. & Krebs, C.J. (2009) Cats, rabbits, Myxoma virus, and vegetation on Macquarie Island: a comment on Bergstrom et al. (2009). Journal of Applied Ecology, 46, 1129-1132. doi: 10.1111/j.1365-2664.2009.01690.x.

Johnson, C.J., Ray, J.C. & St-Laurent, M.-H. (2022) Efficacy and ethics of intensive predator management to save endangered caribou. Conservation Science and Practice, 4: e12729. doi: 10.1111/csp2.12729.

Lamb, C.T., Willson, R., Richter, C., Owens-Beek, N., Napoleon, J., Muir, B., McNay, R.S., Lavis, E., Hebblewhite, M., Giguere, L., Dokkie, T., Boutin, S. & Ford, A.T. (2022) Indigenous-led conservation: Pathways to recovery for the nearly extirpated Klinse-Za mountain caribou. Ecological Applications 32 (5): e2581. doi: 10.1002/eap.2581.

Peters, R.H. (1991) A Critique for Ecology. Cambridge University Press, Cambridge, England. 366 pp. ISBN:0521400171.

Popper, K.R. (1963) Conjectures and Refutations: The Growth of Scientific Knowledge. Routledge and Kegan Paul, London. 412 pp. ISBN-13: 978-0415285940.

Superbie, C., Stewart, K.M., Regan, C.E., Johnstone, J.F. & McLoughlin, P.D. (2022) Northern boreal caribou conservation should focus on anthropogenic disturbance, not disturbance-mediated apparent competition. Biological Conservation, 265, 109426. doi: 10.1016/j.biocon.2021.109426.

Weiner, J. (1995) On the practice of ecology. Journal of Ecology, 83, 153-158.

Woodward, J. & Goodstein, D. (1996) Conduct, misconduct and the structure of science. American Scientist, 84, 479-490.

On Conservation Complexities

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It is too often the case that biodiversity problems are managed by single species solutions. If you have too many deer in your parks or conservation areas, start a culling program. If your salmon fishing stocks are declining, cull seals and sea lions. The overall issue confounding these kinds of ‘solutions’ are now being recognized as a failure to appreciate the food web of the community and ecosystem in which the problem is embedded. Much of conservation action is directed at heading back to the “good old days” without very much data about what the ecosystem was like in the “good old days”.

Problems with introduced species top the list of conservation dilemmas, and nowhere are these problems more clearly illustrated than by the conservation dilemmas of New Zealand and Australia. If we concentrate our management efforts on introduced predators or herbivores, we face a large set of conservation issues, well-illustrated by the current New Zealand situation (Leathwick and Byrom 2023, Parkes and Murphy 2003).

New Zealand is a particularly strong case history because we have a good knowledge of its indigenous biodiversity from the time that people colonized these islands, as well as reasonable information about how things have changed since Europeans colonized the country (Thomson 1922). It is in some respects the classic case of biodiversity impacts from introduced species. The introduced species list is large and I can talk only about part of these species introduced mostly in the late 1800s. Seven species of deer were released in New Zealand, along with chamois, hares, rabbits, cats, hedgehogs, three mustelid species, brushtail possums, rats, house mice, along with all the usual farm animals like cattle, horses, and dogs (King & Forsyth 2021). The first concerns began about 100 years ago over ungulate browsing in forests and grasslands. Deer control began about 1930, and over 3 million deer were shot between 1932 and 1954. Caughley (1983) showed that this amount of control did not reduce the impact of browsing and grazing by ungulates in native ecosystems. Control and harvesting efforts decreased in recent years partly from a lack of government funding with the result that deer numbers have rebounded. The recognition of the impact of other pests like rabbits, weasels, and rats led to a focus on poison campaigns. Brushtail possum control with poisons was started to reduce tree browsing damage by the 1970s and gradually increased to reduce TB transmission to domestic livestock by the 1990s. Large scale predator control began in the late 1990s with a focus on rats, stoats (weasels, Mustela erminea), and possums with good success in preventing declines in threatened bird species. All this history is covered in detail in Leathwick and Byrom (2023).

These efforts led to a declaration in 2016 of “Predator Free New Zealand 2050” (PF2050) a compelling promise that would alleviate biodiversity problems by making New Zealand free of possums, mustelids, and rats by 2050, and predator control has thus became the focus of recent conservation action. The 2050 part of the promise was always a worry, since governments in general promise much in advances by that year, but the optimistic view is that predator control will achieve this objective if careful planning is made, adequate funding is available (c.f. Department of Conservation 2021), and well-articulated guidelines for eradication of invasive species are followed (Bomford & O’Brien 1995). The message is that biodiversity goals can be achieved if we move from single species management to a stable system of ecosystem management in the broad sense, including strong research, good public participation and support toward these goals, and that biodiversity conservation will be greatly boosted by thorough consultation with (if not leadership by) the indigenous groups involved.

The New Zealand specific situation cannot be applied directly to all biodiversity concerns, but the New Zealand conservation story and the 12 recommendations given in Leathwick and Byrom (2023) show the necessity of goal definition and coordination between the public, government, and private foundations if we are to maximize the effectiveness of our approach to the biodiversity crisis. Not every conservation issue involves introduced species, but the principle must be: What do we want to achieve, and how are we going to get there?

Bomford, M, & O’Brien, P 1995. Eradication or control for vertebrate pests? Wildlife Society Bulletin 23, 249–255.

Caughley, G. (1983) The Deer Wars: The Story of Deer in New Zealand. Heinemann, Auckland. ISBN: 0868633895.

Department of Conservation (2020). Annual Report. Available at: https://www.doc.govt. nz/nature/pests-and-threats/predator-free-2050/goal-tactics-and-new-technology/tools-to-market/.    See also: PF2050-Limited-Annual-Report-2022.pdf

King, C.M. & Forsyth, D.M. (2021). eds. The Handbook of New Zealand Mammals. 3rd edition. CSIRO Publishing, Canberra. ISBN 978-1988592589.

Leathwick, J.R. & Byrom, A.E. (2023) The rise and rise of predator control: a panacea, or a distraction from conservation goals? New Zealand Journal of Ecology, 47, 3515. doi: 10.20417/nzjecol.47.3515.

Parkes, J. & Murphy, E. (2003) Management of introduced mammals in New Zealand. New Zealand Journal of Zoology, 30, 335-359. doi:10.1080/03014223.2003.9518346.

Thomson, G.M. (1922) The Naturalisation of Animals and Plants in New Zealand. The University Press, Cambridge, England. doi: 10.5962/bhl.title.28093.

Is Ecology Becoming a Correlation Science?

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One of the first lessons in Logic 101 is classically called “Post hoc, ergo propter hoc” or in plain English, “After that, therefore because of that”. The simplest example of many you can see in the newspapers might be: “The ocean is warming up, salmon populations are going down, it must be another effect of climate change. There is a great deal of literature on the problems associated with these kinds of simple inferences, going back to classics like Romesburg (1981), Cox and Wermuth (2004), Sugihara et al. (2012), and Nichols et al. (2019). My purpose here is only to remind you to examine cause and effect when you make ecological conclusions.

My concern is partly related to news articles on ecological problems. A recent example is the collapse of the snow crab fishery in the Gulf of Alaska which in the last 5 years has gone from a very large and profitable fishery interacting with a very large crab population to, at present, a closed fishery with very few snow crabs. What has happened? Where did the snow crabs go? No one really knows but there are perhaps half a dozen ideas put forward to explain what has happened. Meanwhile the fishery and the local economy are in chaos. Without very many critical data on this oceanic ecosystem we can list several factors that might be involved – climate change warming of the Bering Sea, predators, overfishing, diseases, habitat disturbances because of bottom trawl fishing, natural cycles, and then recognizing that we have no simple way for deciding cause and effect and therefore making management choices.

The simplest solution is to say that many interacting factors are involved and many papers indicate the complexity of populations, communities and ecosystems (e,g, Lidicker 1991, Holmes 1995, Howarth et al. 2014). Everyone would agree with this general idea, “the world is complex”, but the arguments have always been “how do we proceed to investigate ecological processes and solve ecological problems given this complexity?” The search for generality has led mostly into replications in which ‘identical’ populations or communities behave very differently. How can we resolve this problem? A simple answer to all this is to go back to the correlation coefficient and avoid complexity.

Having some idea of what is driving changes in ecological systems is certainly better than having no idea, but it is a problem when only one explanation is pushed without a careful consideration of alternative possibilities. The media and particularly the social media are encumbered with oversimplified views of the causes of ecological problems which receive wide approbation with little detailed consideration of alternative views. Perhaps we will always be exposed to these oversimplified views of complex problems but as scientists we should not follow in these footsteps without hard data.

What kind of data do we need in science? We must embrace the rules of causal inference, and a good start might be the books of Popper (1963) and Pearl and Mackenzie (2018) and for ecologists in particular the review of the use of surrogate variables in ecology by Barton et al. (2015). Ecologists are not going to win public respect for their science until they can avoid weak inference, minimize hand waving, and follow the accepted rules of causal inference. We cannot build a science on the simple hypothesis that the world is complicated or by listing multiple possible causes for changes. Correlation coefficients can be a start to unravelling complexity but only a weak one. We need better methods for resolving complex issues in ecology.

Barton, P.S., Pierson, J.C., Westgate, M.J., Lane, P.W. & Lindenmayer, D.B. (2015) Learning from clinical medicine to improve the use of surrogates in ecology. Oikos, 124, 391-398.doi: 10.1111/oik.02007.

Cox, D.R. and Wermuth, N. (2004). Causality: a statistical view. International Statistical Reviews 72: 285-305.

Holmes, J.C. (1995) Population regulation: a dynamic complex of interactions. Wildlife Research, 22, 11-19.

Howarth, L.M., Roberts, C.M., Thurstan, R.H. & Stewart, B.D. (2014) The unintended consequences of simplifying the sea: making the case for complexity. Fish and Fisheries, 15, 690-711.doi: 10.1111/faf.12041

Lidicker, W.Z., Jr. (1991) In defense of a multifactor perspective in population ecology. Journal of Mammalogy, 72, 631-635.

Nichols, J.D., Kendall, W.L. & Boomer, G.S. (2019) Accumulating evidence in ecology: Once is not enough. Ecology and Evolution, 9, 13991-14004.doi: 10.1002/ece3.5836.

Pearl, J., and Mackenzie, D. 2018. The Book of Why. The New Science of Cause and Effect. Penguin, London, U.K. 432 pp. ISBN: 978-1541698963

Popper, K.R. 1963. Conjectures and Refutations: The Growth of Scientific Knowledge. Routledge and Kegan Paul, London. 608 pp. ISBN: 978-1541698963

Romesburg, H.C. (1981) Wildlife science: gaining reliable knowledge. Journal of Wildlife Management, 45, 293-313.

Sugihara, G., et al. (2012) Detecting causality in complex ecosystems. Science, 338, 496-500.doi: 10.1126/science.1227079.

On the Meaning of ‘Food Limitation’ in Population Ecology

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There are many different ecological constraints that are collected in the literature under the umbrella of ‘food limitation’ when ecologists try to explain the causes of population changes or conservation problems. ‘Sockeye salmon in British Columbia are declining in abundance because of food limitation in the ocean’. ’Jackrabbits in some states in the western US are increasing because climate change has increased plant growth and thus removed the limitation of their plant food supplies.’ ‘Moose numbers in western Canada are declining because their food plants have shifted their chemistry to cope with the changing climate and now suffer food limitation”. My suggestion here is that ecologists should be careful in defining the meaning of ‘limitation’ in discussing these kinds of population changes in both rare and abundant species.

Perhaps the first principle is that it is the definition of life that food is always limiting. One does not need to do an experiment to demonstrate this truism. So to start we must agree that modern agriculture is built on the foundation that food can be improved and that this form of ‘food limitation’ is not what ecologists who are interested in population changes in the real world are trying to test. The key to explain population differences must come from resource differences in the broad sense, not food alone but a host of other ecological causal factors that may produce changes in birth and death rates in populations.

‘Limitation’ can be used in a spatial or a temporal context. Population density of deer mice can differ in average density in 2 different forest types, and this spatial problem would have to be investigated as a search for the several possible mechanisms that could be behind this observation. Often this is passed off too easily by saying that “resources” are limiting in the poorer habitat, but this statement takes us no closer to understanding what the exact food resources are. If food resources carefully defined are limiting density in the ‘poorer’ habitat, this would be a good example of food limitation in a spatial sense. By contrast if a single population is increasing in one year and declining in the next year, this could be an example of food limitation in a temporal sense.

The more difficult issue now becomes what evidence you have that food is limiting in either time or space. Growth in body size in vertebrates is one clear indirect indicator but we need to know exactly what food resources are limiting. The temptation is to use feeding experiments to test for food limitation (reviewed in Boutin 1990). Feeding experiments in the lab are simple, in the field not simple. Feeding an open population can lead to immigration and if your response variable is population density, you have an indirect effect of feeding. If animals in the experimentally fed area grow faster or have a higher reproductive output, you have evidence of the positive effect of the feeding treatment. You can then claim ‘food limitation’ for these specific variables. If population density increases on your feeding area relative to unfed controls, you can also claim ‘food limitation of density’. The problems then come when you consider the temporal dimension due to seasonal or annual effects. If the population density falls and you are still feeding in season 2 or year 2, then food limitation of density is absent, and the change must have been produced by higher mortality in season 2 or higher emigration.

Food resources could be limiting because of predator avoidance (Brown and Kotler 2007). The ecology of fear from predation has blossomed into a very large literature that explores the non-consumptive effects of predators on prey foraging that can lead to food limitation without food resources being in short supply (e.g., Peers et al. 2018, Allen et al. 2022).

All of this seems to be terribly obvious but the key point is that if you examine the literature about “food limitation” look at the evidence and the experimental design. Ecologists like medical doctors at times have a long list of explanations designed to sooth the soul without providing good evidence of what exact mechanism is operating. Economists are near the top with this distinguished approach, exceeded only by politicians, who have an even greater art in explaining changes after the fact with limited evidence.

As a footnote to defining this problem of food limitation, you should read Boutin (1990). I have also raved on about this topic in Chapter 8 of my 2013 book on rodent populations if you wish more details.

Allen, M.C., Clinchy, M. & Zanette, L.Y. (2022) Fear of predators in free-living wildlife reduces population growth over generations. Proceedings of the National Academy of Sciences (PNAS), 119, e2112404119. doi: 10.1073/pnas.2112404119.

Boutin, S. (1990). Food supplementation experiments with terrestrial vertebrates: patterns, problems, and the future. Canadian Journal of Zoology 68(2): 203-220. doi: 10.1139/z90-031.

Brown, J.S. & Kotler, B.P. (2007) Foraging and the ecology of fear. Foraging: Behaviour and Ecology (eds. D.W. Stephens, J.S. Brown & R.C. Ydenberg), pp. 437-448.University of Chicago Press, Chicago. ISBN: 9780226772646

Krebs, C.J. (2013) Chapter 8, The Food Hypothesis. In Population Fluctuations in Rodents. University of Chicago Press, Chicago. ISBN: 978-0-226-01035-9

Five Stages of Ecological Research

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Ecological research falls into five broad classes or stages. Each stage has its strengths and its limitations, and it is important to recognize these since no one stage is more or less important than any other. I suggest a classification of these five stages as follows:

  1. Natural History
  2. Behavioural Ecology
  3. Applied Ecology
  4. Conservation Ecology
  5. Ecosystem Ecology

The Natural History stage is the most popular with the public and in some sense the simplest type of ecological research while at the same time the critical foundation of all subsequent research. Both Bartholomew (1986) and Dayton (2003) made impassioned pleas for the study of natural history as a basis of understanding all the biological sciences. In some sense this stage of biological science has now come into its own in popularity, partly because of influential TV shows like those of David Attenborough but also because of the ability of talented wildlife photographers to capture amazing moments of animals in the natural world. Many scientists still look upon natural history as “stamp-collecting” unworthy of a serious ecologist, but this stage is the foundational element of all ecological research.

Behavioural ecology became popular as one of the early outcomes of natural history observations within the broad framework of asking questions about how individuals in a population behave, and what the ecological and evolutionary consequences of these behaviours are to adaptation and possible future evolution. One great advantage of studying behavioural ecology has been that it is quick, perfectly suited to asking simple questions, devising experimental tests, and then being able to write a report, or a thesis on these results (Davies et al. 2012). Behavioural ecology is one of the strongest research areas of ecological science and provides entertainment for students of natural history and excellent science to understand individual behaviour and how it fits into population studies. It is perhaps the strongest of the ecological approaches for drawing the public into an interest in biodiversity.

Applied ecology is one of the oldest fields of ecology since it arose more than 100 years ago from local problems of how organisms affected human livelihoods. It has subdivided into three important sub-fields – pest management, wildlife management, and fisheries management. Applied ecology relies heavily on the principles of population ecology, one level above the individual studies of behavioural and natural history research. These fields are concerned with population changes, whether to reduce populations to stop damage to crops, or to understand why some species populations become pests. All applied ecology heavily interreacts with human usage of the environment and the economics of farming, fisheries, and wildlife harvesting. In a general sense applied ecology is a step more difficult than behavioural ecology because answering the applied problems or management has a longer time frame than the typical three-year thesis project. Applied ecology has a broad interface with evolutionary ecology because human actions can disrupt natural selection and pest evolution can complicate every management problem.

Conservation ecology is the new kid on the block. It was part of wildlife and fisheries management until about 1985 when it was clear to all that some populations were endangered by human changes to the ecosystems of fisheries, forestry, and agriculture. The essential problems of conservation ecology were described elegantly by Caughley (1994). Conservation issues are the most visible of all issues in population and community ecology, and they are often the most difficult to resolve when science dictates one conservation solution that interferes with the dominant economic view of human society. If species of interest are rare the problem is further confounded by the difficulty of studying rare species in the field. What will become of the earth’s ecosystems in the future depends in large part as to how these conservation conflicts can be resolved.

Ecosystem ecology and community ecology are the important focus at present but are hampered by a lack of a clear vision of what needs to be done and what can be done. The problem is partly that there is much poor theory, coupled with much poor data. The critical questions in ecosystem ecology are currently too vague to be studied in a realistic time period of less than 50 years. Climate change is impacting all our current ideas about community stability and resilience, and what predictions we can make for whole ecosystems in the light of a poor database. Ironically experimental manipulations are being done by companies with an economic focus such as forestry but there are few funds to make use of these large-scale landscape changes. In the long term, ecosystem ecology is the most significant aspect of ecology for humans, but it is the weakest in terms of understanding ecosystem processes. We can all see the negative effects of human changes on landscapes, but we have little in the way of scientific guidance to predict the long-term consequences of these changes and how they can be successfully ameliorated.

All of this is distressing to practical ecologists who wish to make a difference and be able to counteract undesirable changes in populations and ecosystems. It is important for all of us not to give up on reversing negative trends in conservation and land management and we need to do all we can to influence the public in general and politicians in particular to change negative trends to positive ones in our world. An array of good books points this out very forcefully (e.g., Monbiot 2018, Klein 2021). It is the job of every ecologist to gather the data and present ecological science to the community at large so we can contribute to decision making about the future of the Earth.

Bartholomew, G. A. (1986). The role of natural history in contemporary biology. BioScience 36, 324-329. doi: 10.2307/1310237

Caughley, G. (1994). Directions in conservation biology. Journal of Animal Ecology 63, 215-244. doi: 10.2307/5542

Davies, N.B., Krebs, J.R., and West, S.A. (2012) ‘An Introduction to Behavioural Ecology.‘ 4th edn. (Wiley-Blackwell: Oxford.). 520 pp.

Dayton, P.K. (2003). The importance of the natural sciences to conservation. American Naturalist 162, 1-13. doi: 10.1086/376572

Klein, Naomi (2021) ‘How to Change Everything: The Young Human’s Guide to Protecting the Planet and Each Other ‘ (Simon and Schuster: New York.) 336 pp. ISBN: 978-1534474529

Monbiot, George. (2018) ‘Out of the Wreckage: A New Politics for an Age of Crisis.’ (Verso.). 224 pp. ISBN: 1786632896

What is the Ratio of Thought to Action in Biodiversity Conservation?

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Many ecologists who peruse the conservation literature will come away with a general concern about the amount of effort that goes into thoughts about how conservation should be done and how much action is currently being carried out to achieve these goals in the field. My premise here is that currently the person-power given to thought greatly exceeds the person-power devoted to actually achieving the broad conservation goal of protecting biodiversity. Let me illustrate this with one dilemma in conservation: should we be concerned predominately with the loss of threatened and endangered species, or should we concentrate on the major dominant species in our ecosystems? Of course, this is not a black-or-white dichotomy, and the first answer is that we should do both. But the economist would suggest that resources are limited, and you cannot do both, so the question should be reworded as to what fraction of resources should go to one or the other of these two activities.

Consider the example of threatened and endangered species. Many of these species are rare numerically at present. In the past they may have been abundant but that is not always the case. The ecologist will know as a universal constant that most species in ecosystems are rare, and because they are rare, they are most difficult to study to answer the simple question why are they rare? Pick your favourite rare species and try to answer this question. For some species under persecution by humans the answer is simple; for most it is not, and ecologists fall back on explanations like the resources they require are not abundant, or their niche is specialized, meaningless statements that can be called panchrestons unless we have infinite time and funds to find out exactly what the limiting resources are, or why their niche is specialized. Now let us make a simple thought experiment that asks: what would happen if all these rare and endangered species disappeared from the world’s ecosystems? The first response would be total outrage that anyone would ask such a terrible question, so it is best not to talk about it. The second would be that we would be outraged if our favorite bird or frog disappeared like the passenger pigeon. The third might be that we should consider this question seriously.

Some community and ecosystem ecologists might wager that nothing would happen to ecosystem dynamics if all the rare and endangered species disappeared. No one of course would admit to such a point of view since it would end their career. At the moment we are in the unenviable state of doing the opposite experiment on the world’s coral reefs which are suffering in an ocean that is acidifying and heating up, pollution that is increasing, and overfishing that is common (Fraser et al. 2019, Lebrec et al. 2019, Romero-Torres et al. 2020). Coral reefs are an extreme example of human impacts on areas of high conservation and economic value such that the entire ecosystem will have to reconstruct itself with corals of greater tolerance to current and future conditions, a future with no clear guess of what positive effects will transpire.

Perhaps the message of both coral reef conservation and terrestrial ecosystem conservation is that you cannot destroy the major species without major consequences. Australia provides a good example of the consequences of altering predator abundance in an ecosystem. The dingo (Canis familaris) has been persecuted because of predation on sheep, and at the same time domestic cats (Felis catus) and red foxes (Vulpes vulpes) have been introduced to the continent. The ecological question is whether the reintroduction of the dingo to places where it has been exterminated will reduce the abundance of cats and foxes, and thus save naïve prey species from local extinction (Newsome et al. 2015). The answer to this question is far from clear (Morgan et al. 2017, Hunter and Letnic 2022) and may differ in different ecosystems within Australia.  

The bottom line is that our original question about rare species cannot be answered. There is much literature on introduced predators affecting food webs, following from Estes et al. (2011) important paper. and now there is much research effort on the roles of apex predators and consumers on ecosystem dynamics (Serrouya et al. 2021). Much of this effort concentrates on the common animals rather than the rare ones with which we began this discussion. Much more action in the field is needed on all conservation fronts since in my opinion the amount of thought we have available now will last field workers for the rest of the century.

Estes, J.A., Terborgh, J., Brashares, J.S., Power, M.E., Berger, J., et al. (2011). Trophic downgrading of Planet Earth. Science 333, 301-306. doi: 10.1126/science.1205106.

Fraser, K.A., Adams, V.M., Pressey, R.L., and Pandolfi, J.M. (2019). Impact evaluation and conservation outcomes in marine protected areas: A case study of the Great Barrier Reef Marine Park. Biological Conservation 238, 108185. doi: 10.1016/j.biocon.2019.07.030

Hunter, D.O. and Letnic, M. (2022). Dingoes have greater suppressive effect on fox populations than poisoning campaigns. Australian Mammalogy 44. doi: 10.1071/AM21036.

Lebrec, M., Stefanski, S., Gates, R., Acar, S., Golbuu, Y., Claudel-Rusin, A., Kurihara, H., Rehdanz, K., Paugam-Baudoin, D., Tsunoda, T., and Swarzenski, P.W. (2019). Ocean acidification impacts in select Pacific Basin coral reef ecosystems. Regional Studies in Marine Science 28, 100584. doi: 10.1016/j.rsma.2019.100584.

Morgan, H.R., Hunter, J.T., Ballard, G., Reid, N.C.H., and Fleming, P.J.S. (2017). Trophic cascades and dingoes in Australia: Does the Yellowstone wolf–elk–willow model apply? Food Webs 12, 76-87. doi: 10.1016/j.fooweb.2016.09.003.

Newsome, TM., Ballard, G.-A., Crowther, M.S., Dellinger, J.A., Fleming, P.J.S., et al. (2015). Resolving the value of the dingo in ecological restoration. Restoration Ecology 23, 201-208.  doi: 10.1111/rec.12186.

Romero-Torres, M., Acosta, A., Palacio-Castro, A.M., Treml, E.A., Zapata, F.A., Paz-García, D.A., and Porter, J.W. (2020). Coral reef resilience to thermal stress in the Eastern Tropical Pacific. Global Change Biology 26, 3880-3890. doi: 10.1111/gcb.15126

Serrouya, R., Dickie, M., Lamb, C., Oort, H. van, Kelly, A.P., DeMars, C., et al. (2021). Trophic consequences of terrestrial eutrophication for a threatened ungulate. Proceedings of the Royal Society B: Biological Sciences 288, 20202811. doi: 10.1098/rspb.2020.2811.

On Rewilding and Conservation

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Rewilding is the latest rage in conservation biology, and it is useful to have a discussion of how it might work and what might go wrong. I am reminded of a comment made many years ago by Buzz Holling at UBC in which he said, “do not take any action that cannot be undone”. The examples are classic – do not introduce rabbits to Australia if you can not reverse the process, do not introduce weasels and stoats to New Zealand if you cannot remove them later if they become pests, do not introduce cheatgrass to western USA grasslands and allow it to become an extremely invasive species. There are too many examples that you can find for every country on Earth. But now we approach the converse problem of re-introducing animals and plants that have gone extinct back into their original geographic range, the original notion of rewilding (Schulte to Bühne et al. 2022).

The first question could be to determine what ‘rewilding’ means, since it is a concept used in so many ways. As a general concept it can be thought of as repairing the Earth from the ravages imposed by humans over the last thousands of years. It appeals to our general belief that things were better in the ‘good old days’ with respect to conservation, and that all we have seen are losses of iconic species and the introduction of pests to new locations. But we need to approach rewilding with the principle that “the devil is in the details”, and the problems are triply difficult because they must engage support from ecologists over the science and the public over policies that affect different social groups like farmers and hunters. Rewilding may range from initiatives that range from “full rewilding” to ‘minimal rewilding’ (Pedersen et al. 2020). Rewilding has been focused to a large extent on large-bodied animals and particularly those species of herbivores and predators that are high in the food chain, typified by the reintroduction of wood bison back into the Yukon after they went extinct about 800 years ago (Boonstra et al. 2018). So the first problem is that the term “rewilding” can mean many different things.

Two major issues must be considered by conservation ecologists before a rewilding project is initiated. First, there should be a comprehensive understanding of the food web of the ecosystem that is to be changed. This is a non-trivial matter in that our understanding of the food webs of what we describe as our best-known ecosystems are woefully incomplete. At best we can do a boxes and arrows diagram without understanding the strength of the connections and the essential nature of many of the known linkages. The second major issue is how rewilding will deal with climate change (Bakker and Svenning, 2018). There is now an extensive literature on paleoecology, particularly in Europe and North America. The changes in climate and species distributions that flowed from the retreat of the glaciers some 10,000 years ago are documented as a reminder to all ecologists that ecosystems and communities are not permanent in time. Rewilding at the present has a time frame with less than necessary thought to future changes in climate. We make the gigantic assumption that we can recreate an ecosystem that existed sometime in the past, and without being very specific about how we might measure success or failure in restoring ecological integrity. 

Pedersen et al. (2020) recognize 5 levels of rewilding of which the simplest is called “minimal rewilding” and the measure of success at this level is the “Potential of animal species to advance self-regulating biodiverse ecosystems” which I suggest to you is an impossible task to achieve in any feasible time frame less than 50-100 years, which is exactly the time scale the IPCC suggests for maximum climatic emergencies. We do not know what a ‘biodiverse ecosystem’ is since we do not know the boundaries of ecosystems under climate change, and we cannot measure what “natural population dynamics” is because we have so few long-term studies. Finally, at the best level for rewilding we cannot measure “natural species interaction networks” without much arm waving.

Where does this leave the empirical conservation ecologist (Hayward et al. 2019)? Rewilding appears to be more a public relations science than an empirical one. Conservation issues are immediate, and a full effort is needed to protect species and diagnose conservation problems of the day. Goshawks are declining in a large part of the boreal forest of North America, and no one knows exactly why. Caribou are a conservation issue of the first order in Canada, and they continue to decline despite good ecological understanding of the causes. The remedy of some conservation dilemmas like the caribou are clear, but the political and economic forces deny their implementation. As conservation biologists we are ever limited by public and governmental policies that favour exploitation of the land and jobs and money as the only things that matter. Simple rewilding on a small scale may be useful, but the losses we face are a whole Earth issue, and we need to address these more with traditional conservation actions and an increase in research to find out why many elements in our natural communities are declining with little or no understanding of the cause.

Bakker, E.S. and Svenning, J.-C. (2018). Trophic rewilding: impact on ecosystems under global change. Philosophical Transaction of the Royal Society B 373, 20170432. doi: 10.1098/rstb.2017.0432.

Boonstra, R., et al. (2018). Impact of rewilding, species introductions and climate change on the structure and function of the Yukon boreal forest ecosystem. Integrative Zoology 13, 123-138. doi: 10.1111/1749-4877.12288.

Hayward, M.W., et al. (2019). Reintroducing rewilding to restoration – Rejecting the search for novelty. Biological Conservation 233, 255-259. doi: 10.1016/j.biocon.2019.03.011.

Pedersen, P.B.M., Ejrnæs, R., Sandel, B., and Svenning, J.-C. (2020). Trophic rewilding advancement in Anthropogenically Impacted Landscapes (TRAAIL): A framework to link conventional conservation management and rewilding. Ambio 49, 231-244. doi: 10.1007/s13280-019-01192-z.

Schulte to Bühne, H., Pettorelli, N., and Hoffmann, M. (2022). The policy consequences of defining rewilding. Ambio 51, 93-102. doi: 10.1007/s13280-021-01560-8.

On Feeding Birds and Other Wildlife

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There is a very large global movement to feed birds and I want to address why this is a human success story and could be an ecological disaster. These two alternatives follow from two divergent views of the role of humans in the world’s ecosystems. The first is the dominant view that humans are the most important species on Earth, and that we can design the world to maximize our wellbeing without concern of the ecological consequences. The second is a view that we are the custodians of the Earth and that our aim must be to conserve the Earth’s biodiversity and protect its ecosystems. The second view is gaining more visibility with the conservation movement, but if it is to become dominant, there are many ecological problems that deserve our attention. One of the most obvious ones is bird feeding. There are at present no global policies on feeding birds and views on feeding are controversial (Baverstock et al. 2019).

Humans feed birds because they like to look at them and because they have a general belief that feeding in winter or severe weather prevents bird deaths (Brock et al. 2021, Clark et al. 2019). If that is correct, we would expect to see that if we had one large area where birds were fed in winter, and another in which birds were not fed, there should be a difference in population size in the two areas in the following spring. I have yet to see any study that shows this differential effect. Consider an alternative hypothesis that feeding does indeed improve bird survival in winter, but this merely feeds more predators that now have a larger prey base, so the improvement is largely in the predator populations.  It is certainly true for some migratory bird species that if they are fed they can overwinter in more northerly areas or in cities and towns, so geographic winter ranges can expand.

Perhaps the most obvious impact of feeding birds and providing water is the transmission of diseases associated with feeding stations and bird baths. Lawson et al. (2017) explored this problem with bird feeding in Great Britain and found emerging diseases over a 25-year period, focusing on protozoan, viral, and bacterial diseases with contrasting modes of transmission. They also considered mycotoxin contamination of food residues in bird feeders, which present a direct risk to bird health. Rogers et al. (2018) described a mortality event in a declining population of band-tailed pigeons in California with a loss off about 18,000 pigeons associated with tricomonosis in a drought in which birds visited artificial water sites like bird baths. Purple et al. (2015) have demonstrated that the protozoan parasite Trichomonas gallinae could persist in bird baths.

There is a certain irony in the general belief that feeding improves the survival of wild birds. I am reminded of an old story about the English ornithologist David Lack who in the 1930s was talking to a local bird club about his long-term study for a life table of the English Robin. He reported from his banding studies that the life expectancy of the robin was about one year. After his talk, an elderly woman came up to him and started beating him over the head with her umbrella. Once she calmed down, she challenged him because she had a robin singing in her back yard for the last 10 years, so she assumed it was the same robin.

There are other consequences of feeding birds. One is the attraction of squirrels to bird feeders, and the subsequent displacement of birds. One study in southern England showed that grey squirrels occupied the feeders nearly half of the time they were in service (Hanmer et al. 2018). Another consequence of feeders is feed spilled to the ground which can attract rats and other less desirable species in urban settings. Many of these problems are not unique to bird feeding. Fležar et al. (2019) used cameras to investigate sites where European brown bears were being artificially fed year-round on plant-food and carrion from road kills in Slovenia. Over one year they detected 23 vertebrate species at the feeding sites in about 68,000 photos, most frequently brown bears, red foxes and European badgers, but also about half of the species coming to the feeding sites were birds. Roe deer also used these bear feeding sites, even though it is technically illegal to feed roe deer in this jurisdiction because deer feeding on corn and other plants materials can lead to fatal metabolic diseases. The key point is that feeding stations can attract a variety of non-target species with largely unknown consequences for the local wildlife community.  

It will take a brave set of ecologists and veterinarians to define and test the critical hypotheses that arise from feeding wildlife of any kind if only because of the vested interests of the bird seed producers along with so many humans who ‘know’ that feeding is ‘good’ for wildlife. The irony of all this in the end is that many people in parts of the Earth suffer from poor nutrition and starvation while in the first world we use agricultural products to feed birds and other wildlife.

Baverstock, S., Weston, M.A., and Miller, K.K. (2019). A global paucity of wild bird feeding policy. Science of The Total Environment 653, 105-111. doi: 10.1016/j.scitotenv.2018.10.338.

Brock, M., Doremus, J., and Li, Liqing (2021). Birds of a feather lockdown together: Mutual bird-human benefits during a global pandemic. Ecological Economics 189, 107174. doi: 10.1016/j.ecolecon.2021.107174.

Clark, D.N., Jones, D.N., and Reynolds, S.J. (2019). Exploring the motivations for garden bird feeding in south-east England. Ecology and Society 24, 26. doi: 10.5751/ES-10814-240126.

Fležar, Urša, Costa, B., and Krofel, M. (2019). Free food for everyone: artificial feeding of brown bears provides food for many non-target species. European Journal of Wildlife Research 65, 1. doi: 10.1007/s10344-018-1237-3.

Hanmer, H.J., Thomas, R.L., and Fellowes, M.D.E. (2018). Introduced Grey Squirrels subvert supplementary feeding of suburban wild birds. Landscape and Urban Planning 177, 10-18. doi: 10.1016/j.landurbplan.2018.04.004.

Lawson, B., Robinson, R. A., and Cunningham, A. A. (2018). Health hazards to wild birds and risk factors associated with anthropogenic food provisioning. Philosophical Transactions of the Royal Society, Biological Sciences 373 (1745): 20170091. doi: 10.1098/rstb.2017.0091.

Purple, K.E. and Gerhold, R.W. (2015). Persistence of two isolates of Trichomonas gallinae in simulated bird baths with and without organic material. Avian Diseases 59, 472-474. doi: 10.1637/11089-041115-Reg.1.

Rogers, K.H., Girard, Y.A., Woods, L.W., and Johnson, C.K. (2018). Avian trichomonosis mortality events in band-tailed pigeons (Patagioenas fasciata) in California during winter 2014–2015. International Journal for Parasitology: Parasites and Wildlife 7, 261-267. doi: 10.1016/j.ijppaw.2018.06.006.