Tag Archives: conservation ecology

On Ecological Imperialism

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It is well known among ecologists that there are more species of almost everything in the tropical regions, and it is also well known that there is rather much more research in the ecosystems of the temperate zone. A recent note in Science 379 (6632) – 8 Feb. 2023 highlights the problems faced by ornithologists in Latin America and the Caribbean trying to carry out research on their local birds. The details are in two papers now published (Soares et al. 2023, Ruelas Inzunza et al. 2023). Both of these papers are a response to a review paper published in 2020 (Lees et al. 2020) which discussed how much was not known about birds in Latin America, but which ignored most of the contributions of Latin American scientists. The red flag arose in part because all the authors of the 2020 paper were based at universities either in the United States or in the United Kingdom. The central criticisms were that the 2020 paper perpetuated an elitist, exclusionary, “northern” approach that has overlooked the knowledge produced by Latin American experts and Indigenous people, partly because these papers were not in English.

    Their case is certainly important and should be a call-to-arms but it should be read with a few minor qualifications. It is certainly not valid to ignore local knowledge both of scientists and indigenous peoples. But this has been going on now for more than 200 years in all areas of biological science, not that history justifies these barriers. Alas Charles Darwin would fall under the knife of this criticism. The funding for ecological research is higher in most European countries as well as North America compared with tropical countries. So we are dealing with economic issues that underlie the scientific funding that is less in Latin America in addition to the global problem that too many governments prefer guns to butter. We recognize these problems, but we can do nothing immediately about them.

    The language issue is much more difficult because it is so clear. There is a long history of this conflict in scientific papers as well as in literature in general. French scientists years ago refused to publish in English, that has changed. Chinese scientists were all educated in Russian but when the tide turned they learned English and started to write scientific papers in English. The problem revolves back to the education system of North American schools that seem to operate on the assumption that to learn a foreign language is very close to being a traitor. Alas students hardly learn to speak and write English but that is another social issue. I think many northern scientists have helped Latin America scientists to assist them in English usage, so it is to me quite obscene to think that someone has a business charging people $600 for a translation. So much of the complaint in the predominance of English scientific papers arises from social issues that are difficult to overcome.

    In the end I am very sympathetic with the inequities raised in these papers and the desire to move forward on all these issues. Ironically the skeleton of the Lees et al. (2020) paper is an excellent roadmap for the analysis of any taxonomic group anywhere is the world, and these papers should be a reminder that similar reviews should be more inclusive of all published literature. Remember always that European or American knowledge is not the only or the best knowledge.

Lees, A.C., Rosenberg, K.V., Ruiz-Gutierrez, V., Marsden, S., Schulenberg, T.S. & Rodewald, A.D. (2020) A roadmap to identifying and filling shortfalls in Neotropical ornithology. Auk, 137, 1-17. doi: 10.1093/auk/ukaa048.

Ruelas Inzunza, E., Cockle, K.L., Núñez Montellano, M.G., Fontana, C.S., Cuatianquiz Lima, C., Echeverry-Galvis, M.A., Fernández-Gómez, R.A., Montaño-Centellas, F.A., Bonaccorso, E., Lambertucci, S.A., Cornelius, C., Bosque, C., Bugoni, L., Salinas-Melgoza, A., Renton, K., Freile, J.F., Angulo, F., Mugica Valdés, L., Velarde, E., Cuadros, S. & Miño, C.I. (2023) How to include and recognize the work of ornithologists based in the Neotropics: Fourteen actions for Ornithological Applications, Ornithology, and other global-scope journals. Ornithological Applications, 125, duac047. doi: 10.1093/ornithapp/duac047.

Soares, L., Cockle, K.L., Ruelas Inzunza, E., Ibarra, J.T., Miño, C.I., Zuluaga, S., Bonaccorso, E., Ríos-Orjuela, J.C., Montaño-Centellas, F.A., Freile, J.F., Echeverry-Galvis, M.A., Bonaparte, E.B., Diele-Viegas, L.M., Speziale, K., Cabrera-Cruz, S.A., Acevedo-Charry, O., Velarde, E., Cuatianquiz Lima, C., Ojeda, V.S., Fontana, C.S., Echeverri, A., Lambertucci, S.A., Macedo, R.H., Esquivel, A., Latta, S.C., Ruvalcaba-Ortega, I., Alves, M.A.S., Santiago-Alarcon, D., Bodrati, A., González-García, F., Fariña, N., Martínez-Gómez, J.E., Ortega-Álvarez, R., Núñez Montellano, M.G., Ribas, C.C., Bosque, C., Di Giacomo, A.S., Areta, J.I., Emer, C., Mugica Valdés, L., González, C., Rebollo, M.E., Mangini, G., Lara, C., Pizarro, J.C., Cueto, V.R., Bolaños-Sittler, P.R., Ornelas, J.F., Acosta, M., Cenizo, M., Marini, M.Â., Vázquez-Reyes, L.D., González-Oreja, J.A., Bugoni, L., Quiroga, M., Ferretti, V., Manica, L.T., Grande, J.M., Rodríguez-Gómez, F., Diaz, S., Büttner, N., Mentesana, L., Campos-Cerqueira, M., López, F.G., Guaraldo, A.C., MacGregor-Fors, I., Aguiar-Silva, F.H., Miyaki, C.Y., Ippi, S., Mérida, E., Kopuchian, C., Cornelius, C., Enríquez, P.L., Ocampo-Peñuela, N., Renton, K., Salazar, J.C., Sandoval, L., Correa Sandoval, J., Astudillo, P.X., Davis, A.O., Cantero, N., Ocampo, D., Marin Gomez, O.H., Borges, S.H., Cordoba-Cordoba, S., Pietrek, A.G., de Araújo, C.B., Fernández, G., de la Cueva, H., Guimarães Capurucho, J.M., Gutiérrez-Ramos, N.A., Ferreira, A., Costa, L.M., Soldatini, C., Madden, H.M., Santillán, M.A., Jiménez-Uzcátegui, G., Jordan, E.A., Freitas, G.H.S., Pulgarin-R, P.C., Almazán-Núñez, R.C., Altamirano, T., Gomez, M.R., Velazquez, M.C., Irala, R., Gandoy, F.A., Trigueros, A.C., Ferreyra, C.A., Albores-Barajas, Y.V., Tellkamp, M., Oliveira, C.D., Weiler, A., Arizmendi, M.d.C., Tossas, A.G., Zarza, R., Serra, G., Villegas-Patraca, R., Di Sallo, F.G., Valentim, C., Noriega, J.I., Alayon García, G., de la Peña, M.R., Fraga, R.M. & Martins, P.V.R. (2023) Neotropical ornithology: Reckoning with historical assumptions, removing systemic barriers, and reimagining the future. Ornithological Applications, 125, duac046. doi: 10.1093/ornithapp/duac046.

Should Empirical Ecology be all Long-term?

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The majority of empirical ecology research published in our journals is short-term with the time span dictated by the need for 1–2-year Master’s degree studies and 3-4-year PhD research. This has been an excellent model when there was little of a framework for researching the critical questions ecologists ought to answer. Much of ecology in the good old days was based on equilibrium models of populations, communities, and ecosystems, an assumption we know to be irrelevant to a world with a changing climate. Perhaps we should have listened to the paleoecologists who kept reminding us that there was monumental change going on in the eras of glaciation and much earlier in the time of continental drift (Birks 2019). All of this argues that we need to change direction from short-term studies to long-term studies and long-term thinking.

There are many short-term ecological studies that are useful and should be done. It is necessary for management agencies to know if the spraying of forest insect pests this year reduces damage next year, and many similar problems exist that can be used for student projects. But the big issues of our day are long term problems, defined in the first place by longer than the research lifespan of the average ecologist, about 40 years. These big issues are insufficiently studied for two reasons. First, there is little funding for long term research. We can find a few exemptions to this statement, but they are few and many of them are flawed. Second, we as research scientists want to do something new that no one has done before. This approach leads to individual fame and sometimes fortune and is the social model behind many of the research prizes that we hear about in the media, the Nobel Prize, the MacArthur Awards, the National Medal of Science, the Kyoto Prize and many more. The point here is not that we should stop giving these awards (because they are socially useful), but that we should take a broader perspective on how research really works. Many have recognized that scientific advances are made by groups of scientists standing on the shoulders of an earlier generation. Perhaps some of the awards in medicine recognize this more frequently than other areas of science. My point is that large problems in ecology require a group effort by scientists that is too often unrecognized in favour of the individual fame model of science prizes.

A few examples may exemplify the need in ecology to support group studies of long-term problems. The simplest cases are in the media every day. The overharvesting of trees continues with little research into the long-term recovery of the harvested area and exactly how the forest community changes as it recovers. We mine areas for minerals and drill and mine tar sands for oil and gas with little long-term view of the recovery path which may stretch to hundreds or thousands of years while our current research program is long-term if it goes for 10 years. Canada has enough of these disturbance problems to fill the leger. The Giant Gold Mine in the Northwest Territories of Canada mined 220,000 kg of gold from 1948 to 2004 when it closed. It left 237 tonnes of arsenic trioxide dust, a by-product for extracting gold. The long-term ecosystem problems from this toxic compound will last for centuries but you might expect it will be much sooner forgotten than subjected to long-term study.

So where are we ecologists with respect to these large problems? We bewail biodiversity loss and when you look at the available data and the long-term studies you would expect to measure biodiversity and, if possible, manage this biodiversity loss. But you will find only piecemeal short-term studies of populations, communities, and ecosystems that are affected. We tolerate this unsatisfactory scientific situation even for ecosystems as iconic as the Great Barrier Reef of eastern Australia where we have a small number of scientists monitoring the collapse of the reef from climate change. The only justification we can give is that “Mother Nature will heal itself” or in the scientific lingo, “the organisms involved will adapt to environmental change”. All the earth’s ecosystems have been filtered through a million years of geological change, so we should not worry, and all will be well for the future, or so the story goes.

I think few ecologists would agree with such nonsense as the statements above, but what can we do about it? My main emphasis here is long-term monitoring. No matter what you do, this should be part of your research program. If possible, do not count birds on a plot for 3 years and then stop. Do not live trap mice for one season and think you are done. If you have any control over funding recommendations, think continuity of monitoring. Long-term monitoring is a necessary but not a sufficient condition for managing biodiversity change.

There are many obstacles interfering with achieving this goal. Money is clearly one. If your research council requests innovation in all research proposals, they are probably driven by Apple iPhone producers who want a new model every year. For the past 50 years we have been able to fund monitoring in our Yukon studies without ever using the forbidden word monitor because it was not considered science by the government granting agencies. In one sense it is not whether you consider science = innovation or not, but part of the discussion about long term studies might be shifted to consider the model of weather stations, and to discuss why we continue to report temperatures and CO2 levels daily when we have so much past data. No one would dream of shutting down weather monitoring now after the near fiasco around whether or not to measure CO2 in the atmosphere (Harris, 2010, Marx et al. 2017).

Another obstacle has been the destruction of research sites by human developments. Anyone with a long history of doing field research can tell you of past study areas that have been destroyed by fire or are now parking lots, or roads, or suburbia. This problem could be partly alleviated by the current proposals to maintain 30% of the landscape in protected areas. We should however avoid designating areas like the toxic waste site of the Giant Gold Mine as a “protected area” for ecological research.

Where does this all lead? Consider long-term monitoring if you can do the research as part of your overall program. Read the recent contributions of Hjeljord, and Loe (2022) and Wegge et al. (2022) as indicators of the direction in which we need to move, and if you need more inspiration about monitoring read Lindenmayer (2018).

Birks, H.J.B. (2019) Contributions of Quaternary botany to modern ecology and biogeography. Plant Ecology & Diversity, 12, 189-385.doi: 10.1080/17550874.2019.1646831.

Harris, D.C. (2010) Charles David Keeling and the story of atmospheric CO2 measurements. Analytical Chemistry, 82, 7865-7870.doi: 10.1021/ac1001492.

Hjeljord, O. & Loe, L.E. (2022) The roles of climate and alternative prey in explaining 142 years of declining willow ptarmigan hunting yield. Wildlife Biology, 2022, e01058.doi: 10.1002/wlb3.01058.

Lindenmayer, D. (2018) Why is long-term ecological research and monitoring so hard to do? (And what can be done about it). Australian Zoologist, 39, 576-580.doi: 10.7882/az.2017.018.

Marx, W., Haunschild, R., French, B. & Bornmann, L. (2017) Slow reception and under-citedness in climate change research: A case study of Charles David Keeling, discoverer of the risk of global warming. Scientometrics, 112, 1079-1092.doi: 10.1007/s11192-017-2405-z.

Wegge, P., Moss, R. & Rolstad, J. (2022) Annual variation in breeding success in boreal forest grouse: Four decades of monitoring reveals bottom-up drivers to be more important than predation. Ecology and Evolution.12, e9327. doi: 10.1002/ece3.9327.

Alas Biodiversity

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One would have to be on another planet not to have heard of the current COP 15 meeting in Montreal, the Convention on Biological Diversity. Negotiators have recently finalised an agreement on what the signatory nations will do in the next 5 years or so. I do not wish to challenge the view that these large meetings achieve much discussion and suggestions for action on conservation of biodiversity. I do wish to address, from a scientific viewpoint, issues around the “loss of biodiversity” and in particular some of the claims that are being made about this problem.

The first elephant in the room which must not be ignored is human population growth. At a best guess there are perhaps three times as many people now on earth as the earth can support. So the background for all biodiversity action is human population size and the accompanying resource demands. Too few wish to discuss this elephant.

The second elephant is the vagueness of the concept of biodiversity. If we take its simple meaning to be ‘all life on Earth’, we must face the fact that we are not even close to having a complete catalogue of life on earth. To be sure we know most of the species of birds and mammals, a lot of the fish and the reptiles, so we have made a start. But look at the insects and you will find guesses of several million species that are undescribed. And we have hardly begun to look at the bacteria, fungi, and viruses.

The consequence of this is loose speech. When we say we wish to ‘protect biodiversity’ what exactly do we wish to protect? Only the birds but not all of them, only the ones we like? Or only the large mammals like the polar bears, the African lion, and the panda? Typically, conservation of biodiversity focuses on one charismatic species and hopes for spill over to others, applying the well-known principles of population ecology to the immediate threat. But ecologists talk about ecological communities and ecosystems, so this raises another issue of how to define these entities and how protecting biodiversity can be applied to them.

Now the third elephant comes into play, climate change. To appreciate this, we need to talk to paleoecologists. If you were fortunate to live in central Alaska or the Yukon 30,000 years ago and you formed a society for the conservation of biodiversity, you would face a vegetation community that was destined to disappear or change dramatically, not to mention species like the mammoths and saber-toothed tigers that no longer exist but we love to see in museums. So there is a time scale as well as a spatial scale to biodiversity that is easily forgotten. Small national parks and reserves may not be a solution to the issue.

So whither biodiversity science? If we are serious about biodiversity change, we must lay out more specific questions as a start. Exactly what species are we measuring and for how long and with what precision? We need to concentrate on areas that are protected from human exploitation, one of the main reasons for biodiversity losses, the loss of habitat due to agriculture, mining, forestry, human housing, roads, invasive pests, the list goes on. We need groups of ecologists to concentrate on the key areas we define, on the key threats affecting each area, how we might mitigate these effects, and once these questions are decided we need to direct funding to these groups. Biodiversity funding is all over the map and often wasted on trivial problems. Biodiversity issues are at their core problems in community and ecosystem ecology, and yet we typically treat them as single species problems. We need to study communities and ecosystems. To say that we as ecologists do not know how to study community and ecosystem ecology would be a start. Studying one fish species extensively will not protect the community and ecosystem it requires for survival. If you need a concrete example, consider Pacific salmon on the west coast of North America and the ecosystems they inhabit. This is not a single species problem. In some river systems stocks are doing well, while in other rivers salmon are disappearing. Why? If we know that at least part of the answer to this question lies in ecosystem management and yet no action is undertaken, is this because it costs too much or what? Why can we spend a billion dollars going to the moon and not spend this money on serious ecological problems subject to biodiversity increases or declines? Perhaps part of the problem is that to get to the moon we do not give money to 10 different agencies that do not talk or coordinate with one another. Part of the answer is that governments do not see biodiversity loss or gain as an important problem, and it is easier to talk vaguely about it and do little in the hope that Nature will rectify the problems.

So, we continue in the Era of Biodiversity without knowing what this means and too often without having any plan to see if biodiversity is increasing or declining in any particular habitat or region, and then devising a plan to ameliorate the situation as required. This is not a 5 year or a 10-year plan, so it requires a long-term commitment of public support, scientific expertise, and government agencies to address. For the moment we get an A+ grade for talking and an F- grade for action.

Dupont-Doaré, C. & Alagador, D. (2021) Overlooked effects of temporal resolution choice on climate-proof spatial conservation plans for biodiversity. Biological Conservation, 263, 109330.doi: 10.1016/j.biocon.2021.109330.

Fitzgerald, N., Binny, R.N., Innes, J., Pech, R., James, A., Price, R., Gillies, C. & Byrom, A.E. (2021) Long-Term Biodiversity Benefits from Invasive Mammalian Pest Control in Ecological Restorations. Bulletin of the Ecological Society of America, 102, e01843.doi: 10.1002/bes2.1843.

Moussy, C., Burfield, I.J., Stephenson, P.J., Newton, A.F.E., Butchart, S.H.M., Sutherland, W.J., Gregory, R.D., McRae, L., Bubb, P., Roesler, I., Ursino, C., Wu, Y., Retief, E.F., Udin, J.S., Urazaliyev, R., Sánchez-Clavijo, L.M., Lartey, E. & Donald, P.F. (2022) A quantitative global review of species population monitoring. Conservation Biology, 36, e13721.doi. 10.1111/cobi.13721.

Price, K., Holt, R.F. & Daust, D. (2021) Conflicting portrayals of remaining old growth: the British Columbia case. Canadian Journal of Forest Research, 51, 1-11.doi: 10.1139/cjfr-2020-04530.

Shutt, J.D. & Lees, A.C. (2021) Killing with kindness: Does widespread generalised provisioning of wildlife help or hinder biodiversity conservation efforts? Biological Conservation, 261, 109295.doi: 10.1016/j.biocon.2021.109295.

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

On Ecological Climate Change Research

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The media world is awash in climate change articles and warnings. When your town is faced with the fourth one-in-100-year-flood or your favourite highway has been washed away, you should perhaps become aware that something is changing rapidly. Ecologists are aware of the problems that climate change is producing, and the question I want to raise here is what kind of research is needed to outline current and future problems and suggest possible solutions. This fact of current climate change means that each of us has something important to do at the individual level to reduce the impacts of climate change, like taking the bus or bicycling. But that is another whole set of social issues that I cannot cover here.

The first thing most scientific organizations want to do when faced with a big problem is to have endless meetings about the problem. This unfortunately eats up much money and produces little understanding except that the problem is complicated and multidimensional. Ecological research on climate change must begin with the axiom that climate change is happening rapidly, and that we as ecological scientists can do nothing about this at the level of climate physics. Given this, what are we to do? The first approach we could take is to ignore climate change and carry on with normal research agendas. This works very well for short term problems on the time scale of 20-30 years. Since this is the research lifespan of most ecological scientists, it is not an unreasonable approach. But it does not help solve the earth’s future problems, and this is not a desirable path to take in science.

There are three broad problems that accompany climate change for ecological science. First, geographical ranges of species will shift. We have from paleoecology much information on some of these changes since the last Ice Age. Data from palaeontology is less useful to planning, given that we have enough problems trying to forecast the next 100 years of change. So, we have major ecological question #1 – what limits the geographical distributions of species? This relatively simple question is greatly confounded by human activities. If we send oil and other chemical pollution out onto a coastal coral reef, we should not be surprised if the local distribution of sea life is affected. For ecologists this class of problems of distribution changes caused by human activities is a very important focus of research. If you doubt this, read about Covid viruses. But there is also a large area of research needed to estimate the possible changes in geographic distributions of organisms that are not immediately affected by human activities. How fast will tree species colonize up-slope in mountains around the globe, and how will this affect the bird and mammals that depend on trees or the vegetation types the trees displace? These changes are local and complex, and we can begin by describing them, but to understand the limiting factors involved in changes in geographical distributions is not easy.

Population ecology addresses the second central question of ecology: what causes changes in the abundance of particular species? While we need answers to this simple question for our conservation and management issues, population ecology is an even bigger minefield for research on the effects of climate change. There is no doubt that climate in general can affect the abundance and changes in abundance of organisms, but the complications lie in determining the detailed mechanisms of explaining these changes in abundance. Large scale climate indicators like ENSO sometimes correlate positively with animal population increases, sometimes negatively, and sometimes not at all in different populations (Wan et al. 2022). Consequently, a changing climate may not have a universal effect on biodiversity. This means we must dive into details of how climate affects our specific population, is it via maximum temperatures?, minimum temperatures?, dry season rainfall?, wet season rainfall? etc., and each of these aspects of weather have many subcomponents – March temperatures, April temperatures, etc. and the search for an explanation can thus become infinite. The problem is that the number of possible explanatory variables in weather dwarfs the number of years of observations of our study species (c.f. Ginzburg and Jensen 4004, Loken and Gelman 2017). The result is that some of the strongest papers with conclusions about the impact of climatic change on animals can be in error (Daskalova. Phillimore, and Myers-Smith 2021). The statistical pitfalls have been discussed for many years (e.g., Underwood and Chapman 2003) but are still commonly seen in the ecological literature today.

A third central question is that each population is embedded in a community of other species which may interact so that we must analyse the changes occurring community and ecosystem dynamics. Changes in biological communities and ecosystems are subject to complications arising from climate change and more because of species interactions which are not easy to measure. These difficulties do not mean that we should stop trying to explain population and community changes that might be related to climate change. What it does mean is that we should not jump to strong conclusions without considering all the alternate possible agents that are changing the earth’s biomes. The irony is that the human caused shifts are easy to diagnose but difficult to fix because of economics, while the pure climate caused shifts in ecosystems are difficult to diagnose and to validate the exact mechanisms involved. We need both strong involvement in diagnosing the major ecological problems associated with climate change, but this must be coupled with modesty in our suggested conclusions and explanations. There is much to be done.

Daskalova, Gergana N., Phillimore, Albert B., and Myers-Smith, Isla H. (2021). Accounting for year effects and sampling error in temporal analyses of invertebrate population and biodiversity change: a comment on Seibold et al. 2019. Insect Conservation and Diversity 14, 149-154. doi: 10.1111/icad.12468.

Ginzburg, L. R. and Jensen, C. X. J. (2004). Rules of thumb for judging ecological theories. Trends in Ecology and Evolution 19, 121-126. doi: 10.1016/j.tree.2003.11.004.

Loken, Eric and Gelman, Andrew (2017). Measurement error and the replication crisis. Science 355, 584. doi: 10.1126/science.aal3618.

Underwood, A. J. and Chapman, M. G. (2003). Power, precaution, Type II error and sampling design in assessment of environmental impacts. Journal of Experimental Marine Biology and Ecology 296, 49-70. doi: 10.1016/s0022-0981(03)00304-6.

Wan, Xinru, Holyoak, Marcel, Yan, Chuan, Maho, Yvon Le, Dirzo, Rodolfo, et al. (2022). Broad-scale climate variation drives the dynamics of animal populations: A global multi-taxa analysis. Biological Reviews 97. (in press).

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.

How Do We Decide Controversial Issues in Conservation?

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While almost everyone favours conservation of plants and animals around the globe, it is far from clear how this broad goal can be disarticulated into smaller issues. Once we have done this the solution of the conservation problem should be simple. But it is not (Sutherland et al, 2021). Take an example of the koala in Australia, cute mid-size marsupials that live in trees and eat leaves. If koalas are to be protected, you must protect forests, but if you protect forests the companies that survive by logging on both private and crown land will be adversely affected. We have an immediate conflict, so how do we decide what to do. One response which we can label have-your-cake-and-eat-it-too suggests that we use some of our forests for logging and protect some forests for ecological reserves. Everyone is now happy, but things unravel. As the human population grows, we need more wood, so over time we would have to log more and more of the forested areas that could support koalas. Conflict now, jobs for loggers vs. conservation of koalas. The simplest solution is to decide all this in economic terms. Logging produces much money; conservation is largely a drain on the taxpayers. To propose that conservation should win, ecologists will pull out David Attenborough to show all the beauties of the forest and to point out that the forest contains many other animals and plants and not just trees for lumber. Stalemate, and social and economic goals begin to override the ecological issue until some compromise is suggested and accepted.

While this kind of oversimplified scenario is common, the whole issue of conservation decision making is fraught with problems and who is going to decide these issues (Christie et al. 2022)? In a democracy in the good old days, you took a vote or a poll and decided to win/lose at >50% of the vote. But this cannot work for critical problems. We have a good example of this problem now with Covid vaccination requirements, and a vocal minority opposed to vaccinations. This now spills over into the issue of whether to wear a face mask or not. In all these kinds of scenarios science delivers a simple decision about the consequences of decision A vs decision B, but the problem is that society can refuse to recognize the scientific results or just prefer decision B with little visible justification. Science is not always perfect, adding further complications. And in the case of the covid virus, the virus can mutate in unexpected ways, complicating prognoses. In the case of protected conservation areas, we can suffer fires, floods, insect outbreaks and any number of events that affect the balance of decision making.

There is a large literature on decision making in conservation (e.g., Bower et al. 2018) and even good advice from the field of psychology about this problem of making decisions (Papworth 2017). The best systematic decision tree I have found is that in Sutherland et al. 2021). Sutherland et al. (2021) compiled a framework that can be used profitably in deciding on the level of evidence assessment (see Table 1 and Figure 1 below from their paper).

Table 1 and Figure 1 from Sutherland et al. (2021)

The Strategic Evidence Assessment Framework. Seven levels of evidence assessment, how to apply them.

Assessment LevelApproach UsedGeneral Database ApplicationApproximate Time to reflect on the evidence
1 No consideration of evidenceContinue with existing practice or make decisions without considering scientific evidencenone
2 Assertion but no independent consideration of evidenceConsultation with others (including experts) that affect decision but are not verified e.g. “we normally do this”, “accepted best practice is to do this”minutes
3Papers reviewed, looking at: Read the title and/or summary points to determine whether action described in the paper is likely to be effective or not. Review effectiveness category e.g. “likely to be beneficial” on action page to decide whether action is likely to be effective or notminutes
4 Read abstract to assess the evidence described in the paper in relation to the local problemTens of minutes- hours
5  Read abstract, key results and conclusion assessing each paper in relation to the decision being madeHours
6 Read the full underlying paper/s. This is likely to affect decisions on study quality, relevance and modificationsHours to days
7Comprehensive assessmentA systematic review of all available literature. Assessed papers summarised as part of new reviewMonths to a year

Figure 1. A framework for considering the appropriate level of effort in decision making. Numbers refer to assessment level (Table 1). For a given decision about an action identify the column with the relevant level of consequence, start at the lowest level (1) and decide whether it would benefit from examining higher levels of evidence. Keep moving up until either the uncertainty in the effectiveness of the action is resolved from examining the evidence (from any platform) or the arrows end. This final number is the level at which the evidence assessment should occur. (From Sutherland et al. 2021 with permission).

Clearly conservation ecologists cannot use the highest assessment level for all issues that arise and must result to triage in many cases (Hayward and Castley 2018). But triage and assessment levels 1 and 2 should be rare in making judgement on what program to adopt. We need to get the science right for all conservation problems.

But this is not enough to get thoughtful political decisions. Some native species can be pests, yet nothing is done to reduce their damage (e.g. horses in North America and Australia, camels and goats in Australia, feral pigs in North America) and the list goes on. Nothing is done because of budget limitations or political concerns about “cute species”. The science of conservation is difficult enough, the social science of conservation is too often out of our control.

Bower, S.D., Brownscombe, J.W., Birnie-Gauvin, K. Ford, M.I. et al. (2018). Making Tough Choices: Picking the appropriate conservation decision-making tool. Conservation Letters 11, e12418. doi: 10.1111/conl.12418.

Christie, A.P., Downey, H., Bretagnolle, V., Brick, C., Bulman, C.R., et al. (2022). Principles for the production of evidence-based guidance for conservation actions. Conservation Science and Practice 4, e579. doi: 10.1111/csp2.12663.

Hayward, M.W. and Castley, J.G. (2018). Triage in Conservation. Frontiers in Ecology and Evolution 5, 168. doi: 10.3389/fevo.2017.00168.

Papworth, Sarah (2017). Decision-making psychology can bolster conservation. Nature Ecology & Evolution 1, 1217-1218. doi: 10.1038/s41559-017-0281-9.

Sutherland, W.J., Downey, H., Frick, W.F., Tinsley-Marshall, P., and McPherson, T. (2021). Planning practical evidence-based decision making in conservation within time constraints: the Strategic Evidence Assessment Framework. Journal for Nature Conservation 60, 125975. doi: 10.1016/j.jnc.2021.125975.

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 Assumptions in Ecology Papers

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What can we do as ecologists to improve the publishing standards of ecology papers? I suggest one simple but bold request. We should require at the end of every published paper a annotated list of the assumptions made in providing the analysis reported in the paper. A tabular format could be devised with columns for the assumption, the perceived support of and tests for the assumption, and references for this support or lack thereof. I can hear the screaming already, so this table could be put in the Supplementary Material which most people do not read. We could add to each paper in the final material where there are statements of who did the writing, who provided the money, and add a reference to this assumptions table in the Supplementary Material or a statement that no assumptions about anything were made to reach these conclusions.

The first response I can detect to this recommendation is that many ecologists will differ in what they state are assumptions to their analysis and conclusions. As an example, in wildlife studies, we commonly make the assumption that an individual animal having a radio collar will behave and survive just like another animal with no collar. In analyses of avian population dynamics, we might commonly assume that our visiting nests does not affect their survival probability. We make many such assumptions about random or non-random sampling. My question then is whether or not there is any value in listing these kinds of assumptions. My response is that this approach of listing what the authors think they are assuming should alert the reviewers to the elephants in the room that have not been listed.

My attention was called to this general issue by the recent paper of Ginzburg and Damuth (2022) in which they contrasted the assumptions of two general theories of functional responses of predators to prey – “prey dependence” versus “ratio dependence”. We have in ecology many such either-or discussions that never seem to end. Consider the long-standing discussion of whether populations can be regulated by factors that are “density dependent” or “density independent”, a much-debated issue that is still with us even though it was incisively analyzed many years ago.  

Experimental ecology is not exempt from assumptions, as outlined in Kimmel et al. (2021) who provide an incisive review of cause and effect in ecological experiments. Pringle and Hutchinson (2020) discuss the failure of assumptions in food web analysis and how these might be resolved with new techniques of analysis. Drake et al. (2021) consider the role of connectivity in arriving at conservation evaluations of patch dynamics, and the importance of demographic contributions to connectivity via dispersal. The key point is that, as ecology progresses, the role of assumptions must be continually questioned in relation to our conclusions about population and community dynamics in relation to conservation and landscape management.

Long ago Peters (1991) wrote an extended critique of how ecology should operate to avoid some of these issues, but his 1991 book is not easily available to students (currently available on Amazon for about $90). To encourage more discussion of these questions from the older to the more current literature, I have copied Peters Chapter 4 to the bottom of my web page at https://www.zoology.ubc.ca/~krebs/books.html for students to download if they wish to discuss these issues in more detail.

Perhaps a possible message in all this has been that ecology has always wished to be “physics-in-miniature” with grand generalizations like the laws we teach in the physical sciences. Over the last 60 years the battle in the ecology literature has been between this model of physics and the view that every population and community differ, and everything is continuing to change under the climate emergency so that we can have little general theory in ecology. There are certainly many current generalizations, but they are relatively useless for a transition from the general to the particular for the development of a predictive science. The consequence is that we now bounce from individual study to individual study, typically starting from different assumptions, with very limited predictability that is empirically testable. And the central issue for ecological science is how can we move from the present fragmentation in our knowledge to a more unified science. Perhaps starting to examine the assumptions of our current publications would be a start in this direction.  

Drake, J., Lambin, X., and Sutherland, C. (2021). The value of considering demographic contributions to connectivity: a review. Ecography 44, 1-18. doi: 10.1111/ecog.05552.

Ginzburg, L.R. and Damuth, J. (2022). The Issue Isn’t Which Model of Consumer Interference Is Right, but Which One Is Least Wrong. Frontiers in Ecology and Evolution 10, 860542. doi: 10.3389/fevo.2022.860542.

Kimmel, K., Dee, L.E., Avolio, M.L., and Ferraro, P.J. (2021). Causal assumptions and causal inference in ecological experiments. Trends in Ecology & Evolution 36, 1141-1152. doi: 10.1016/j.tree.2021.08.008.

Peters, R.H. (1991) ‘A Critique for Ecology.’ (Cambridge University Press: Cambridge, England.) ISBN:0521400171 (Chapter 4 pdf available at https://www.zoology.ubc.ca/~krebs/books.html)

Pringle, R.M. and Hutchinson, M.C. (2020). Resolving Food-Web Structure. Annual Review of Ecology, Evolution, and Systematics 51, 55-80. doi: 10.1146/annurev-ecolsys-110218-024908.