Category Archives: Charley Krebs’ blogs

Is Ecology Becoming a Correlation Science?

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.

In Honour of David Suzuki at his Retirement

David Suzuki is retiring from his media work this year at age 86. If you wish to have a model for a lifetime of work, he should be high on your list – scientist, environmentalist, broadcaster, writer. He has been a colleague of mine at the Department of Zoology, UBC from the time when I first came there in 1970. He was a geneticist doing imaginative and innovative research with his students on the humble fruit fly Drosophila melanogaster. The Department at that time was a beehive of research and teaching, and David was a geneticist breathing fire at the undergraduates taking the genetics course. Many a doctor would probably tell you now that Suzuki’s genetics course was the most challenging in their undergraduate education.

The hierarchy in the Department of Zoology was very clear in the 1970s. First came the physiologists, top of the pack and excellent scientists who turned the spotlight on the Department nationally and internationally. Second came the geneticists, with the DNA revolution full on. At the bottom of the pile were the ecologists causing nothing but trouble about fisheries and wildlife management problems, pointing out a rising tide of environmental problems including climate change. Contrary to what you might conclude from the media, environmental problems and climate change issues were very alive even in the 1970s. But somehow these problems did not get through to governments, and David has been a key person turning this around. In 1979 he began a natural history and science program on the CBC entitled “The Nature of Things” which he then hosted for 43 years. In doing so he began to fill an empty niche in Canadian news affairs between the environmental scientists who had data on what was going on in the environment and what needed attention. Environmental scientists were severely ignored both by industry and the governments of the day who operated on two premises – first, that the most critical issues for Canada were economics and economic growth, and second that environmental issues could largely be ignored or could be solved by promises but no action. Alas we are still inundated with the news that “growth is good”, and “more growth is better”.   

I had relatively little involvement in David’s increasing interest in environmental issues by 1979, but I had written 3 ecology textbooks by then, pushing some of the environmental issues that are still with us, and I became a friend of David’s in the Department. We ecologists could only admire his ability to speak so clearly on the environmental issues of our day and connect these issues with the many travesties of how First Nations people had been sidelined. He pointed out very forcefully the astonishing failure of governments to address these issues. The public which was much less aware of environmental issues in the 1980s is now highly mobilized thanks in great part to all the work David and his colleagues have done in the last 50 years. He has many friends now but still strong enemies who continue to think of the environment as a large garbage can for economic growth. And he, still in his retirement, having achieved so much from his environmental work, bemoans the slow pace of government actions on environmental problems, as does every ecologist I know. His Foundation continues to press for action on many conservation fronts. So, thank you David for all your work and your wisdom over all these many years. You have engineered a strong environmental movement among old and young and I thank you for all that.

https://davidsuzuki.org/

How to Destroy a Research Station

I have had the ‘privilege’ over the last 60 years of watching three ecological field stations be destroyed. Admittedly this is a small sample, against which every ecologist can complain, but I wanted to present to you my list of how to achieve this kind of destruction should you ever be commanded to do so. I will not name names or specific places, since the aim is to develop a general theory rather than to name and pillory specific historical actions and people. I suggest that nine rules are needed to proceed smoothly in this matter if you are given this job.

  1.  Have a clear vision why you wish to destroy an existing station. Do not vacillate. The background may be money, or philosophy of science, or orders from those higher in the echelon, or a personal peeve. Remember you are an administrator, and no one can challenge your wisdom in making major changes or closing the station.  
  2. Speak to none of the current users of the research station. If the research station has a Users Committee, avoid talking to them until after all the decisions are made. A users committee is just an honorary appointment, and it helps if very few of the users are actually people who do research at the station. It is very important that your vision should not be clouded by personnel or research programs currently running at the station. And it is best if the scientists using the station have no information except gossip about the changes that are coming.
  3. Avoid loose talk around your office. If you or your group are paying a visit in the field to the research station before closing it or repositioning its purpose, give out no information to anyone on future courses of action.
  4. Communicate upwards in the hierarchy, never downwards. You must keep all the members of the higher echelons fully informed. Do not dwell on the details of your progress in destruction but emphasize the gains that will flow from this dismantling. Tell fibs as much as you like because no one will question your version of events.
  5. Never read anything about the history of the research station or read any of the papers and reports that have originated there. The key is that you as an administrator know what should be done, and the last consideration is history. Administrators must keep a clear mind, unconcerned with historical trivia.
  6. Let none of the destruction news reach the media lest the public in general might begin to see what is happening. Newspaper and media coverage are rarely flattering to bureaucrats. If possible, line up a sympathetic media person who can talk about the brilliant future of the research station and the wisdom of the decisions you have made.
  7. Take a strong business approach. Do not worry if you must fire people currently running the research station or eject scientists currently working there. Everyone must retire at some point and all business leaders have solid recipes for hiring contractors to take care of any problems with the buildings. No matter what the extra cost.
  8. Sell the research station if you possibly can in order to gain revenue for your yet to be revealed vision. You may talk complete nonsense to explain why you are making major changes or closing the research station because few of your possible critics will be in a position to distinguish nonsense statements from truth. ‘Alternative facts’ are very useful if your decisions are questioned.
  9. Realize that if you have made a mistake in destroying a research station, your employer will not know that for several years. By that time, you will have ascended in the hierarchy of your employment unit for having carried out such a definitive action. And if your co-workers know the poor job you are doing, they will write sterling letters of reference for you to move you to another position in a different department or agency so that the worse the job you have done, the stronger will be the reference letters to recommend you for another job.

There is almost no literature I can find on this topic of administering a field station. If you think field stations are eternal, it may be a sign that you are very young, or you are very fortunate in working for an agency where moving forward is correctly labeled as progress. I have always thought that long-term field research stations were considered sacred but clearly not everyone agrees. Administrators must have something to do to leave their mark on the world for better or worse. All we can do is watch and be alert for emerging symptoms of collapse.

Swanson, F.J. (2015). Confluence of arts, humanities, and science at sites of long-term ecological inquiry. Ecosphere 6 (8), Article 132. doi: 10.1890/ES15-00139.1.

On the Meaning of ‘Food Limitation’ in Population Ecology

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 Climate Change Research Funding

I have grown weary of media and news statements that climate change research should be a priority. At the present time military spending, war, and oil and gas companies seem to be the priority spending of many governments. Climate change research seems to be more focused on the physical sciences in attempts to predict what changes in temperature, rainfall, and sea conditions can be expected if we continue at the present global rates of greenhouse gas emissions. This is all very good, and the IPCC reports are excellent. The people are listening and reacting to the bad news even if all the major western governments are close to ignoring the problem. So where does this leave ecological scientists?

Our first response is that we should mimic the climatologists in predicting what the ecological world will be like in 2050 or 2100. But there is a major problem with this centered around the fact that physics has a whole set of fixed laws that will not change in a thousand years, so that the physics of the atmosphere and the oceans is reasonably understood and by the application of the laws of physics, we can arrive at a reasonable prediction that should be constrained by physical laws. Ecological science is nowhere near that paradigm of predictability because it deals with organisms that can evolve and interactions that can change rapidly when an unexpected invasive species arrives on the scene or humans interfere with ecosystem services. Ecological changes are not driven solely by climate change, a fact it is easy to forget. One consequence of this limitation is that we cannot make any kind of reliable predictions about the state of our ecosystems and the state of the Earth’s biodiversity by 2050 or 2100. We can however, in contrast to the physical sciences, do something about ecological changes by finding the limiting factors for the species under concern, protecting these endangered species and setting aside natural areas protected from human depredation. While we can do this to some extent in rich countries, in poor countries, particularly tropical ones, we have a poor record of protecting the exploitation of national parks and reserves. Think Brazil or the Central African Republic.

But given this protection of areas and funding for threatened species, conservation ecologists still have some very difficult problems to face. First and foremost is the conservation of rare, endangered species. It is nearly impossible to study rare species to discover the limiting factors that are pushing them toward extinction. Second, if you have the information on limiting factors, it is difficult to reverse trends that are determined by climate change or by human disrespect for conservation values.

In spite of these problems, the ecological literature is full of papers claiming to solve these issues with various schemes that predict a brighter future sometime. But if we apply the same rigor to these papers as we do to other areas of ecology, we must treat them as a set of hypotheses that make specific predictions, and try to test them. If we have solutions that are feasible but will require 50 years to accomplish, we should be very clear that we are drawing a long bow. Some statement of goals for the next 5 years would be desirable so we can measure progress or lack of progress.

The screams of practitioners go up – we have no time to test hypotheses, we need action! If we have clear-cut a forest site, or bulldozed shrub habitats, we may have a good idea of how to proceed to restoration. But with a long term view, restoration itself in highly contestable. In particular with climate change we have even less ability to predict with knowledge based on the last 50 year or so. So if you are in a predictive mode about conservation issues, have multiple working hypotheses about what to do, rather than one certain view of what will solve the problem.

This is not a cry to give up on conservation, but rather to trim our certainty about future states of ecosystems. Trying to predict what will happen under climate change is important for the Earth but we must always keep in mind the other critical factors affecting biodiversity, from predators to parasites and diseases, and the potential for evolution. Human destruction of habitats is a key issue we do not control well enough, and yet it may be the most important short term threat to conservation.

All of this leads into the fact that to achieve anything we need resources –people and money. The problem at present is where can we get the money? Governments in general place a low value on conservation and the environment in general in the quest for money and economic growth. Rich philanthropists are useful but few, and perhaps too often they have a distorted view of what to invest in. Improving the human condition of the poor is vital; medical research is vital, but if the environment suffers losses as it is at present, we need to balance or reverse our priorities of where to put our money. I do not know how to accomplish this goal. The search for politicians who have even a grade 1 understanding of environmental problems is not going well. Read Boris Johnson and Vladimir Putin. What is being accomplished now is more to the credit of private philanthropy which has clear goals but may pull in diverse directions. I submit that to date we have not been successful in this pursuit of environmental harmony, but it is a goal we must keep pushing for. E.O. Wilson once said that there was more money spent in New York City on a Friday night on beer than was devoted to biodiversity conservation for the entire world for the year.  This should hardly be a good epitaph for our century.

On Ecological Climate Change Research

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

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?

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?

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.

What Can You Do About the Climate Emergency?

It is very easy to do little in the climate emergency because it is a long-term problem, and many of us will be gone by 2050 when Shell Oil and our government promise Net Zero emissions. Possibly the first thing you should do is find out what “net zero” really means. “Net zero emissions” refers to achieving an overall balance between greenhouse gas emissions produced by us and greenhouse gas emissions taken out of the atmosphere. So clearly it does not mean zero emissions so pollution will still be with us, and all it promises is equality between what goes in and what comes out. If you believe that net-zero will happen, you are living in la-la land, but consider it a scientific hypothesis and if you are young and live to 2050, check the numbers. It means that all the greenhouse gases that are here today will remain and all the problems on our doorstep today will continue – floods, fires, drought, sea level rise, agricultural changes, temperature increases – and if you think none of this will bother you, you can probably buy an inexpensive house in New Mexico and avoid shopping for groceries.

But do not throw your hands up since there are many small things all of us can do to minimize these problems. Here is a partial list:

  1. Drive less, fly less, walk more, get an electric car if you can. Try a bicycle.
  2. Avoid coal, gasoline, and natural gas implements. Sit in the sun, not under a propane heater on the deck.
  3. Put solar panels on your roof if you can. In addition to your windmill generating power.
  4. Put your retirement funds into renewable energy funds, not into oil companies.
  5. Educate yourself and ignore all the dangerous nonsense about climate change that is provided in advertisements, radio, TV, and social media.
  6. Protest against climate nonsense by writing letters, using social media, phoning the stations that allow nonsense to be perpetrated. Your one letter may have minimal effect, but if a million people do the same, someone might listen.
  7. Demand that politicians actually answer questions about climate change action plans. And as they say in Chicago, vote early and vote often.
  8. Nominate Greta Thunberg again for the Nobel Prize. If she does not receive it, request that the Nobel Committee be disbanded and replaced by young people.
  9. Relax and enjoy your life while keeping a lid on your carbon budget.

The climate emergency is not difficult to comprehend. Help the world survive it for your grandchildren.