Tag Archives: conservation

On the Canadian Biodiversity Observation Network (CAN BON)

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I have been reading the report of an exploratory workshop from July 2021 on designing a biodiversity monitoring network across Canada to address priority monitoring gaps and engage Indigenous people across Canada. The 34 pages of their workshop report can be accessed here, and I recommend you might read it before reading my comments on the report:

https://www.nserc-crsng.gc.ca/Media-Media/NewsDetail-DetailNouvelles_eng.asp?ID=1310

I have a few comments on this report that are my opinion only. I think the Report on this workshop outlines a plan so grand and misguided that it could not be achieved in this century, even with a military budget. The report is a statement of wisdom put together with platitudes. Why is this and what are the details that I believe to be unachievable?

The major goal of the proposed network is to bring together everyone to improve biodiversity monitoring and address the highest priority gaps to support biodiversity conservation. I think most of the people of Canada would support these objectives, but what does it mean? Let us do a thought experiment. Suppose at this instant in time we knew the distribution and the exact abundance of every species in Canada. What would we know, what could we manage, what good would all these data be except as a list taking up terabytes of data? If we had these data for several years and the numbers or biomass were changing, what could we do? Is all well in our ecosystems or not? What are we trying to maximize when we have no idea of the mechanisms of change? Contrast these concerns about biodiversity with the energy and resources applied in medicine to the mortality of humans infected with Covid viruses in the last 3 years. A monumental effort to examine the mechanisms of infection and ways of preventing illness, with a clear goal and clear measures of progress toward that goal.

There is no difficulty in putting out “dream” reports, and biologists as well as physicists and astronomers, and social scientists have been doing this for years. But in my opinion this report is a dream too far and I give you a few reasons why.

First, we have no clear definition of biodiversity except that it includes everything living, so if we are going to monitor biodiversity what exactly should we do? For some of us monitoring caribou and wolves would be a sufficient program, or whales in the arctic, or plant species in peat bogs. So, to begin with we have to say what operationally we would define as the biodiversity we wish to monitor. We could put all our energy into a single group of species like birds and claim that these are the signal species to monitor for ecosystem integrity. Or should we consider only the COSEWIC list of Threatened or Endangered Species in Canada as our major monitoring concern? So, the first job of CAN BON must be to make a list of what the observation network is supposed to observe (Lindenmayer 2018). There is absolutely no agreement on that simple question within Canada now, and without it we cannot move forward to make an effective network.

The second issue that I take with the existing report is that the emphasis is on observations, and then the question is what problems will be solved by observation alone. The advance of ecological science has been based on observation and experiment directed to specific questions either of ecological interest or of economic interest. In the Pacific salmon fishery for example the objective of observation is to predict escapement and thus allowable harvest quotas. Despite years of high-quality observations and experiments, we are still a long way from understanding the ecosystem dynamics that drive Pacific salmon reproduction and survival.

Contrast the salmon problem with the caribou problem. We have a reasonably good understanding of why caribou populations are declining or not, based on many studies of predator-prey dynamics, harvesting, and habitat management. At present the southern populations of caribou are disappearing because of a loss of habitat because of land use for forestry and mining, and the interacting nexus of factors is well understood. What we do not do as a society is put these ideas into practice for conservation; for example, forestry must have priority over land use for economic reasons and the caribou populations at risk suffer. Once ecological knowledge is well defined, it does not lead automatically to action that biodiversity scientists would like. Climate change is the elephant in the room for many of our ecological problems but it is simultaneously easy to blame and yet uneven in its effects.

The third problem is funding, and this overwhelms the objectives of the Network. Ecological funding in general in Canada is a disgrace, yet we achieve much with little money. If this ever changes it will require major public input and changed governmental objectives, neither is under our immediate control. One way to press this objective forward is to produce a list of the most serious biodiversity problems facing Canada now along with suggestions for their resolution. There is no simple way to develop this list. A by-product of the current funding system in Canada is the shelling out of peanuts in funding to a wide range of investigators whose main job becomes how to jockey for the limited funds by overpromising results. Coordination is rare partly because funding is low. So (for example) I can work only on the tree ecology of the boreal forest because I am not able to expand my studies to include the shrubs, the ground vegetation, the herbivores, and the insect pests, not to mention the moose and the caribou.  

For these reasons and many more that could be addressed from the CAN BON report, I would suggest that to proceed further here is a plan:

  1. Make a list of the 10 or 15 most important questions for biodiversity science in Canada. This alone would be a major achievement.
  2. Establish subgroups organized around each of these questions who can then self-organize to discuss plans for observations and experiments designed to answer the question. Vague objectives are not sufficient. An established measure of progress is essential.
  3. Request a realistic budget and a time frame for achieving these goals from each group.  Find out what the physicists, astronomers, and medical programs deem to be suitable budgets for achieving their goals.
  4. Organize a second CAN BON conference of a small number of scientists to discuss these specific proposals. Any subgroup can participate at this level, but some decisions must be made for the overall objectives of biodiversity conservation in Canada.

These general ideas are not particularly new (Likens 1989, Lindenmayer et al. 2018). They have evolved from the setting up of the LTER Program in the USA (Hobbie 2003), and they are standard operating procedures for astronomers who need to come together with big ideas asking for big money. None of this will be easy to achieve for biodiversity conservation because it requires the wisdom of Solomon and the determination of Vladimir Putin.

Hobbie, J.E., Carpenter, S.R., Grimm, N.B., Gosz, J.R., and Seastedt, T.R. (2003). The US Long Term Ecological Research Program. BioScience 53, 21-32. doi: 10.1016/j.oneear.2021.12.008

Likens, G. E. (Ed.) (1989). ‘Long-term Studies in Ecology: Approaches and Alternatives.’ (Springer Verlag: New York.) ISBN: 0387967435

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.

Lindenmayer, D.B., Likens, G.E., and Franklin, J.F. (2018). Earth Observation Networks (EONs): Finding the Right Balance. Trends in Ecology & Evolution 33, 1-3. doi: 10.1016/j.tree.2017.10.008.

On Cats and Birds and Policy Gaps

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Many people in western societies like to keep cats as pets, and with that simple observation we run into two problems that require resolution. First, cats are killers of wildlife, particularly birds but also an array of other small prey. Most people do not believe this, because cats are adored and make good, if somewhat disinterested pets. So, my first point might be that if you think cats are not killers, I invite you to keep another cat like a mountain lion for a pet. But we need some data on the kill rate of cats. Before we begin this search, we should note that cats can be kept inside dwellings or in cat runs with no access to birds or other prey. If this is the case, no problem exists for wildlife, and you can skip to the bottom of this blog for one other issue to recognize.

How much mortality can be traced to cats roaming out of doors? This will include normal house cats let out to roam at night, as well as wild cats that have been discarded by their owners into the wild. There is extensive literature on cats killing birds. If you want a brief introduction Greenwall et al. (2019) discuss a nesting colony of Fairy Terns, a threatened species of Australian seabird, along a beach in southwestern Australia. With detailed observations and photographic data, they recorded the complete failure of all 111 nests in this colony with the loss of all tern chicks in the early summer of 2018. The predator was a single desexed feral cat. Many local governments allow the capture of feral cats with the protocol that they are desexed and then released back into the environment. Clearly desexing and release does not remove the problem.

The domestic cat has been spread world-wide, so that the cat problem is not a local one. Li et al. (2021) completed a survey of feral cat kill rates in the eastern part of China and found that the minimum annual loss of wildlife to feral cats was in the range of 2.7-5.5 billion birds, and 3.6-9.8 billion mammals, as well as large numbers of amphibians, reptiles, and fish. In gardens in Western Europe cat predation on ringed birds studied with precise data showed that up to 25% of dead birds were killed by cats, but these data varied greatly among species (Pavisse et al. 2019). For the European Robin which often feeds on the ground 40% of all ringed birds were killed by cats, for the European Greenfinch the figure was 56% of ringed birds killed. These are just two examples of an extensive literature on cat kills going back many years (Calvert et al. 2013).

What can we do about this predation? As with too many conservation issues the answer is simple but difficult to implement: Ban all cats from free-ranging unless they are on a leash and under control. Keep cats in the house or in special cat runs that are confined outdoors. Ban completely stupid programs of catching feral cats, sterilizing them, and releasing them back to the wild to continue their killing. Cats may make marvellous pets but need to be kept indoors. Many people would support these measures but many cat owners would disagree about such measures. Some progress is being made in urban environments in which some suburbs do not permit cats to roam freely.

Feral cats are a serious issue in Australia because they attack many threatened birds and reptiles (Doherty et al. 2019). In this case a federal environmental policy to kill 2 million cats is popular but from a conservation viewpoint still a poor policy. We do not know if killing 2 million cats is too much or too few, and without specific goals for conservation and careful monitoring of bird populations widespread killing my not achieve the goal of protection for threatened species. Eradications of cats on islands is often feasible, but no mainland eradication is currently possible.

As conservation biologists know too well, when humans are the problem, wise policies may not be implemented. So, the second issue and the bottom line might be to consider the human costs of cat ownership. Adhikari et al. (2020) report a highly significant association between the risk of dying from colon cancer and cat ownership. These results are not confounded by sedentary lifestyle, cigarette smoking or socio-economic status. In a similar study Adhikari et al. (2019) found that living with a cat significantly increased the death rate from lung cancer among women. The cause of these associations cannot yet be deciphered but are postulated to result from mycotoxins, toxic secondary metabolites produced by fungi (moulds) in cereal crops used in cat food. Aflatoxin is a mycotoxin that produces well-known chemicals that are seriously toxic to animals and humans.

These kinds of studies of associations arising from surveys can be tossed off by the typical comments ‘these-things-do-not-concern-my cats’ or ‘that there is no proof of the exact cause’ so if you are concerned you might investigate the literature on both mycotoxins and the diseases that cats carry.

It is up to humans to solve human problems, but up to conservation biologists to point out the detrimental effects of household pets and their feral cousins on wildlife. The present situation is a complete policy failure by governments at all levels. Good science is relatively easy, good policy is very difficult.

Adhikari, Atin, Adhikari, A., Jacob, N. K., and Zhang, J. (2019). Pet ownership and the risk of dying from lung cancer, findings from an 18 year follow-up of a US national cohort. Environmental Research 173, 379-386. doi: 10.1016/j.envres.2019.01.037.

Adhikari, Atin, Adhikari, A., Wei, Y. D., and Zhang, J. (2020). Association between pet ownership and the risk of dying from colorectal cancer: an 18-year follow-up of a national cohort. Journal of Public Health 28, 555-562. doi: 10.1007/s10389-019-01069-1.

Calvert, A.M., Bishop, C.A., Elliot, R.D., Krebs, E.A., Kydd, T.M., Machtans, C.S., Robertson, G.J., 2013. A synthesis of human-related avian mortality in Canada. Avian Conservation and Ecology 8: 11. doi 10.5751/ACE-00581-080211.

Doherty, T.S., Driscoll, D.A., Nimmo, D.G., Ritchie, E.G., and Spencer, R. (2019). Conservation or politics? Australia’s target to kill 2 million cats. Conservation Letters 12, e12633. doi: 10.1111/conl.12633.

Li, Yuhang, Wan, Yue, Shen, Hua, Loss, S.R., Marra, P.P., and Li, Zhongqiu (2021). Estimates of wildlife killed by free-ranging cats in China. Biological Conservation 253, 108929. doi: 10.1016/j.biocon.2020.108929.

Greenwell, C.N., Calver, M.C., and Loneragan, N.R. (2019). Cat Gets Its Tern: A Case Study of Predation on a Threatened Coastal Seabird. Animals 9, 445. doi: 10.3390/ani9070445.

Pavisse, R., Vangeluwe, D., and Clergeau, P. (2019). Domestic Cat Predation on Garden Birds: An Analysis from European Ringing Programmes. Ardea 107, 103-109. doi: 10.5253/arde.v107i1.a6.

On Biodiversity Science

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With David Attenborough and all the amazing picture books on biodiversity there can be few people in the world who have not been alerted to the array of beautiful and interesting species on Earth. Until recently the subject of biodiversity, known to First Nations since long, long ago, had not entered the western world of automobiles, industry, farming, fishing, music, theatres, and movies. Biodiversity is now greatly appreciated by most people, but perhaps more as entertainment for western societies and more for subsistence food in less wealthy parts of our world.

There are many different measures of ‘biodiversity’ and when discussing how we should protect biodiversity we should be careful about exactly how this word is being used. The number of different species in an area is one simple measure of biodiversity. But often the types of organisms being considered are less well defined. Forest ecologists attempt to protect forest biodiversity, but logging companies are more concerned only with trees and tree size for commercial use. Bird watchers are concerned with birds and have developed much citizen science in counting birds. Mushroom connoisseurs may worry about what edible mushrooms will be available this summer. But in many cases biodiversity scientists recognize that the community of organisms and the ecosystem that contains them would be a more appropriate unit of analysis. But as the number of species in an ecosystem increases, the complexity of the ecosystem becomes unmanageable. A single ecosystem may have hundreds to thousands of species, and we are in the infant stage of trying to determine how to study these biological systems.

One result is that, given that there are perhaps 10 million species on Earth and only perhaps 10,000 biologists who study biodiversity, where do we begin? The first and most popular way to answer this question is to pick a single species and concentrate on understanding its ecology. This makes are researcher’s life fairly simple. If elephants in Africa are under threat, find out all about the ecology of elephants. If a particular butterfly in England is very rare, try to find out why and how to protect them. This kind of research is very valuable for conservation because it provides a detailed background for understanding the requirements of each species. But the single species approaches lead into at least two quagmires. First, all species exist in a web of other species and understanding this web greatly expands the problem. It is possible in many cases to decipher the effects other species have on our elephants or butterflies, but this requires many more scientists to assist in analysing the species’ food chain, its diseases, its predators and parasites, and that is only a start. The second quagmire is that one of the general rules of ecology is that most species on Earth are rare, and few are common. So that we must concentrate our person-power on the common species because they are easier to find and study. But it is often the rare species that are of conservation concern, and so we should focus on them rather than the common species. In particular, given that only about 10% of the species on Earth have been described scientifically, we may often be assigned a species that does not have any information on its food habits or habitat requirements, its distribution, and how its abundance might be changing over time, a lifetime research program.

The result of this general overview is that the mantra of our day – Protect Biodiversity – begins as a compelling slogan and ends in enormous scientific complexity. As such it falls into the category of slogans like ‘Reduce Poverty’ and ‘Peace on Earth’, something we can all agree on, but the devil is in the details of how to achieve that particular goal.

One way to avoid all these pitfalls has been to jump over the problems of individual species and analyse communities of species or entire ecosystems. The result of this approach is to boil down all the species in the community to a number that estimates “biodiversity” and then use that number in relating ‘biodiversity’ to community attributes like ‘productivity’ or ‘stability’. This approach leads to testing hypotheses like ‘Higher biodiversity leads to greater stability’. There are serious problems with this approach if it is used to test any such hypothesis. First, biodiversity in this example must be rigorously defined as well as stability. The fact that higher biodiversity of butterflies in a particular region is associated with a more stable abundance of these butterflies over time is worthy of note but not of generalization to global communities or ecosystems. And as in all ecological studies we do not know if this is a generalization applicable to all butterfly populations everywhere until many more studies have been done.

A second problem is that this community or ecosystem approach to address ecological questions about biodiversity is not very useful in promoting conservation which boils down to particular species in particular environments. It should force us back to looking at the population ecology of species that are of conservation concern. It is population ecologists who must push forward the main goals of the conservation of the Earth’s biota, as Caughley (1994) recognized long ago.

The practical goals of conservation have always been local, and this constraint is mostly ignored in papers that demand some global research priorities and global ecological rules. The broad problem is that the conservation of biodiversity is a gigantic scientific and political problem that is currently underfunded and in its scientific infancy. At the present too much biodiversity research is short-term and not structured in a comprehensive framework that identifies critical problems and concentrates research efforts on these problems (Nichols et al. 2019, Sutherland et al. 2018). One more important issue for a seminar discussion group. 

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

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

Sutherland, W.J., Butchart, Stuart H.M., Connor, B., Culshaw, C., Dicks, L.V., et al. (2018). A 2018 Horizon Scan of Emerging Issues for Global Conservation and Biological Diversity. Trends in Ecology & Evolution 33, 47-58. doi: 10.1016/j.tree.2017.11.006.

Have We Lost the Plot?

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The decisions we make as a society depend directly on what knowledge we have achieved through our educational system. Two major problems the Earth faces occupy the day – the Covid epidemic and climate change. In both major emergencies, a significant fraction of humanity seems to have completely missed the plot and I would like to ask a few simple questions about why this might be.

The Covid epidemic is indeed a global emergency, and if you do not recognize this you should stop reading here. We have had major human epidemics in the last 1000 years so we might start by asking what knowledge we have garnered from past events. Epidemics occur because a particular disease is transmissible among people, and the three most obvious observations that could be made from previous epidemics are that large groups of people should not congregate, travel should be restricted, and that people should always wear a mask, a point made very clearly in the 1918 flu epidemic. More recent medical studies since the 1940s have shown conclusively that immunity to any particular disease can be achieved by vaccination programs, and many people have been vaccinated over their lifespan to reduce greatly the chance of infection. So, to make the point simple, many people are alive today because of the vaccinations they have received over time.

Vaccine hesitancy at this time with respect to the Covid epidemic has been decreasing, and as more of the population becomes vaccinated, disease incidence should decline. My question is how did many people become educated in our schools about these general points and then join the anti-vaxxers? I do not know the answer to this, but at least part of the answer might be a failure of our education systems.

A second emergency over climate change will probably be with us for a much longer time than the Covid pandemic, so we need to think very clearly about it. The problem in part is that climate change is long term (10-100+ years) and it is difficult to change human behaviour in a short time. Consequently, advances like renewable energy, solar panels on roofs, electric cars, and good insulation in houses need to be pushed by government policies. Since governments are too often concerned only about the next 4 years, and all the good policies will result in rising taxes, there is much talking but little action. Longer term issues like population control are too often swept under the table as too hot to handle. News outlets push panic buttons over reduced birth rates in the world today and translate this into immediate population collapse. Elementary issues of human demography that ought to be part of any curriculum are not understood, and the failure to appreciate the consequences of continued growth seem lost on much of the population. Consequently, part of our current problems involving action on the climate emergency must be laid to poor education about these simple matters.

We have gone through a long period when economics triumphed over ecology and sustainability, but that problem is rapidly being rectified. More people are recognizing that a single country cannot ignore global problems, conservation is strong on the agenda of many governments, although again these issues emit more talk than actions.

I certainly do not know the solution to these current issues but the polarization in the world today is strong enough to prohibit many policies being achieved that would improve and overcome our present emergencies. Unless we can achieve agreement on sustainable goals for all of society these emergencies will continue to build. Thinking that I could fly to Mars and get away from these problems is something even the British royalty recognize as ridiculous.

A few possible ideas:

  1. Call out and protest as much as you can about uninformed pseudo-scientific comments on ecology, economics, medical science, and sustainability. Demand political action on these two global emergencies now.
  2. Improve our education systems to demand a curriculum that addresses current problems of climate change and agriculture, population growth, medical history, disease, and the history of the biosphere.
  3. Get accurate data on global change and Covid from reliable sources.
  4. Never give up. Present scientific truth to counteract nonsense.
  5. And use social media effectively to improve communication of the science that speaks to the solution of these major problems.

Kolata, Gina B. (2019) Flu: The Story of the Great Influenza Pandemic of 1918 and the Search for the Virus that caused it.’ Atria Books: New York. 352 pp. ISBN: 978-0743203982

MacKenzie, Debora (2020) COVID-19: The Pandemic that Never Should Have Happened and How to Stop the Next One. Hachette Books: New York. 304 pp. IBN: 978-0306924248  (Published in North America in 2021 as Stopping the Next Pandemic, 339 pp. ISBN 978-036924224.)

Piketty, Thomas (2021). Time for Socialism: Dispatches from a World on Fire, 2016-2021
Yale University Press: New Haven, Connecticut. 360 pp. ISBN: 978-0300259667

Salamon, Margaret Klein (2020). Facing the Climate Emergency: How to Transform Yourself with Climate Truth. New Society Publishers: Gabriola Island, B.C. Canada. 160 pp. ISBN: 978-0865719415

Blaming Climate Change for Ecological Changes

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The buzz word for all ecological applications for funding and for many submitted papers is climate change. Since the rate of climate change is not something ecologists can control, there are only two reasons to cite climate change as a reason to fund current ecological research. First, since change is continuous in communities and ecosystems, it would be desirable to determine how many of the observed changes might be caused by climate change. Second, it might be desirable to measure the rate of change in ecosystems, correlate these changes to some climate variable, and then use these data as a political and social tool to stimulate politicians to do something about greenhouse gas emissions. The second approach is that taken by climatologists who blame hurricanes and tornadoes on global warming. There is no experimental way to trace any particular hurricane to particular amounts of global warming, so it is easy for critics to say these are just examples of weather variation of which we have measured much over the last 150 years and paleo-ecologists have traced over tens of thousands of years using proxies from tree rings and sediment cores. If we are to use the statistical approach we need a large enough sample to argue that extreme events are becoming more frequent, and that might take 50 years by which time the argument would be made too late to request proper action.

The second approach to prediction in ecology is fraught with problems, as outlined in Berteaux et al. (2006) and Dietze (2017). The first approach has many statistical problems as well in selecting a biologically coherent model that can be tested by in a standard scientific manner. Since there are a very large number of climate variables, the possibility of spurious correlations is excessive, and the only way to avoid these kinds of results is to be predictive and to have a biological causal chain that is testable. Myers (1998) reviewed all the fishery data for predictive models of juvenile recruitment that used environmental variables as predictors and data was subsequently collected and tested with the published model. The vast majority of these aquatic models failed when retested but a few were very successful. The general problem is that model failures or successes might not be published so even this approach can be biased if only a literature survey is undertaken. The take home message from Myers (1998) was that almost none of the recruitment-environment correlations were being used in actual fishery management.

How much would this conclusion about the failure of environmental models in fishery management apply to other areas in ecology? Mouquet et al. (2014) pointed out that predictions could be classified as ‘explanatory’ or ‘anticipatory’ and that “While explanatory predictions are necessarily testable, anticipatory predictions need not be…….In summary, anticipatory predictions differ from explanatory predictions in that they do not aim at testing models and theory. They rely on the assumption that underlying hypotheses are valid while explanatory predictions are based on hypotheses to be tested. Anticipatory predictions are also not necessarily supposed to be true.” (page 1296). If we accept these distinctions, we have (I think) a major problem in that many of the predictive models put forward in the ecological literature are anticipatory, so they would be of little use to a natural resource manager who requires an explanatory model.

If we ignore this problem with anticipatory predictions, we can concentrate on explanatory predictions that are useful to managers. One major set of explanatory predictions in ecology are those associated with range changes in relation to climate change. Cahill et al. (2014) examined the conventional hypothesis that warm-edge range limits are set by biotic interactions rather than abiotic interactions. Contrary to expectations, they found in 125 studies that abiotic factors were more frequently supported as setting warm-edge range limits. Clearly a major paradigm about warm-edge range limits is of limited utility.

Explanatory predictions are not always explicit. Mauck et al. (2018) for example developed a climate model to predict reproductive success in Leach’s storm petrel on an island off New Brunswick in eastern Canada. From 56 years of hatching success they concluded that annual global mean temperature during the spring breeding season was the single most important predictor of breeding success. They considered only a few measures of temperature as predictor variables and found that a quadratic form of annual global mean temperature was the best variable to describe the changes in breeding success. The paper speculates about how global or regional mean temperature could possibly be an ecological predictor of breeding success, and no mechanisms are specified. The actual data on breeding success are not provided in the paper, even as a temporal plot. Since global temperatures were rising steadily from 1955 to 2010, any temporal trend in any population parameter that is rising would correlate with temperature records. The critical quadratic relationship in their analysis suggests that a tipping point was reached in 1988 when hatching success began to decline. Whether or not this is a biologically correct explanatory model can be determined by additional data gathered in future years. But it would be more useful to find out what the exact ecological mechanisms are.

If the ecological world is going to hell in a handbasket, and temperatures however measured are going up, we can certainly construct a plethora of models to describe the collapse of many species and the rise of others. But this is hardly progress and would appear to be anticipatory predictions of little use to advancing ecological science, as Guthery et al. (2005) pointed out long ago. Someone ought to review and evaluate the utility of AIC methods as they are currently being used in ecological and conservation science for predictions.

Berteaux, D., Humphries, M.M., Krebs, C.J., Lima, M., McAdam, A.G., Pettorelli, N., Reale, D., Saitoh, T., Tkadlec, E., Weladji, R.B., and Stenseth, N.C. (2006). Constraints to projecting the effects of climate change on mammals. Climate Research 32, 151-158. doi: 10.3354/cr032151.

Cahill, A.E., Aiello-Lammens, M.E., Fisher-Reid, M.C., Hua, X., and Karanewsky, C.J. (2014). Causes of warm-edge range limits: systematic review, proximate factors and implications for climate change. Journal of Biogeography 41, 429-442. doi: 10.1111/jbi.12231.

Dietze, M.C. (2017). Prediction in ecology: a first-principles framework. Ecological Applications 27, 2048-2060. doi: 10.1002/eap.1589.

Guthery, F.S., Brennan, L.A., Peterson, M.J., and Lusk, J.J. (2005). Information theory in wildlife science: Critique and viewpoint. Journal of Wildlife Management 69, 457-465. doi: 10.1890/04-0645.

Mauck, R.A., Dearborn, D.C., and Huntington, C.E. (2018). Annual global mean temperature explains reproductive success in a marine vertebrate from 1955 to 2010. Global Change Biology 24, 1599-1613. doi: 10.1111/gcb.13982.

Mouquet, N., Lagadeuc, Y., Devictor, V., Doyen, L., and Duputie, A. (2015). Predictive ecology in a changing world. Journal of Applied Ecology 52, 1293-1310. doi: 10.1111/1365-2664.12482.

Myers, R.A. (1998). When do environment-recruitment correlations work? Reviews in Fish Biology and Fisheries 8, 285-305. doi: 10.1023/A:1008828730759.

 

On Culling Overabundant Wildlife

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Ecologists have written much about the culling of wildlife from an ecological and conservation perspective (Caughley 1981, Jewell et al. 1981, Bradford and Hobbs 2008, Hampton and Forsyth 2016). The recommendations for culling as a method for reducing overabundant wildlife populations are typically scientifically well established and sensitive to animal welfare. The populations chosen for culling are classified as ‘overabundant’. But overabundant is a human-defined concept, and thus requires some form of social license to agree about what species, in which conditions, should be classified as ‘overabundant’. The problem of overabundance usually arises when humans make changes that permit a species to become so numerous locally that it is having an adverse effect on its food supply, its competitors, or the integrity of the ecosystem it occupies. Once overabundance is recognized, the management issue is to determine which methods should be used to reduce abundance to a suitable level. Culling is only one option for removing wildlife, and animals may be captured and moved elsewhere if that is possible or sterilized to prevent reproduction and further increase (Liu et al. 2012, Massei and Cowan 2014).

All these policy issues are subject to open public debate and these debates are often heated because of different belief systems. Animal rights advocates may push the assumption that we humans have no rights to kill any wildlife at all. News media often concentrate on the most stringent views on controlling populations that are overabundant, and public discussion becomes impossible. Two aspects need to be noted that are often lost in any discussion. First is the cost of alternatives in dollars and cents. As an example, most ecologists would agree that wild horses are overabundant on open range in western United States (Davies et al. 2014, Rutberg et al. 2017) but the question is what to do about this. Costs to reduce horse populations by capturing horses and penning them and feeding them are astronomical (the current situation in western USA, estimated at $25,000 per animal) but this method of control could be done if society wishes to spend money to achieve this goal. Culling would be much cheaper, but the killing of large animals is anathema to many people who speak loudly to politicians. Fertility control methods are improving with time and may be more acceptable socially, but costs are high and results in population reduction can be slow in coming (Hobbs and Hinds 2018). Models are essential to sort out many of these issues, whether it be the projected costs of various options (including doing nothing), the expected population trajectory, or the consequences for other species in the ecosystem.

The bottom line is that if overabundant wildlife populations are not reduced by some means, the result must be death by starvation or disease coupled with extensive damage to other species in these ecosystems. This type of “Plan B” is the second aspect not often considered in discussions of policies on overabundant species. In the present political scene in North America opposition to culling overabundant wildlife is strong, coherent discussion is rarely possible, and Plan B problems are rarely heard. Most overabundant wildlife result from human actions in changing the vegetation, introducing new species, and reducing and fragmenting wildlife habitats. Wishing the problems will go away without doing anything is not a feasible course of action.

These kinds of problems in wildlife management are soluble in an objective manner with careful planning of research and management actions (Hone et al. 2017). Ecologists have a moral duty to present all scientific sides of the management of overabundant species, and to bring evidence into the resulting social and political discussions of management issues. It is not an easy job.

Bradford, J.B., and N.T. Hobbs. 2008. Regulating overabundant ungulate populations: An example for elk in Rocky Mountain National Park, Colorado. Journal of Environmental Management 86:520-528. doi: 10.1016/j.jenvman.2006.12.005

Caughley, G. 1981. Overpopulation. Pages 7-19 in P.A. Jewell S. Holt, and D. Hart, editors. Problems in Management of Locally Abundant Wild Mammals. Academic Press, New York. ISBN: 978-0-12-385280-9

Davies, K. W., Collins, G. & Boyd, C. S. (2014) Effects of feral free-roaming horses on semi-arid rangeland ecosystems: an example from the sagebrush steppe. Ecosphere, 5, 127. doi: 10.1890/ES14-00171.1

Hampton, J. O., and D. M. Forsyth. 2016. An assessment of animal welfare for the culling of peri-urban kangaroos. Wildlife Research 43:261-266. doi: 10.1071/WR16023

Hobbs, R.J. and Hinds, L.A. (2018). Could current fertility control methods be effective for landscape-scale management of populations of wild horses (Equus caballus) in Australia? Wildlife Research 45, 195-207. doi: 10.1071/WR17136.

Hone, J., Drake, V.A. & Krebs, C.J. (2017) The effort–outcomes relationship in applied ecology: Evaluation and implications BioScience, 67, 845-852. doi: 10.1093/biosci/bix091

Jewell, P. A., Holt, S. & Hart, D. (1982) Problems in Management of Locally Abundant Wild Mammals. Academic Press, New York. 360 pp. ISBN: 978-0-12-385280-9

Liu, M., Qu, J., Yang, M., Wang, Z., Wang, Y., Zhang, Y. & Zhang, Z. (2012) Effects of quinestrol and levonorgestrel on populations of plateau pikas, Ochotona curzoniae, in the Qinghai-Tibetan Plateau. Pest Management Science, 68, 592-601. doi: 10.1002/ps.2302

Massei, G. & Cowan, D. (2014) Fertility control to mitigate human–wildlife conflicts: a review. Wildlife Research, 41, 1-21. doi: 10.1071/WR13141

Rutberg, A., Grams, K., Turner, J.W. & Hopkins, H. (2017) Contraceptive efficacy of priming and boosting doses of controlled-release PZP in wild horses. Wildlife Research, 44, 174-181. doi: 10.1071/WR16123

On the Loss of Large Mammals

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The loss of large mammals and birds in the Pleistocene was highlighted many years ago (Martin and Wright 1967, Grayson 1977, Guthrie 1984 and many other papers). Hypotheses about why these extinctions occurred were flying left and right for many years with no clear consensus (e.g. Choquenot and Bowman 1998). The museums of the world are filled with mastodons, moas, sabre-tooth tigers and many other skeletons of large mammals and birds long extinct. The topic has come up again in a discussion of these extinctions and a prognosis of future losses (Smith et al. 2018). I do not want to question the analysis in Smith et al. (2018) but I want to concentrate on this one quotation that has captured the essence of this paper in the media:

“Because megafauna have a disproportionate influence on ecosystem structure and function, past and present body size downgrading is reshaping Earth’s biosphere.”
(pg. 310).

What is the evidence for this very strong statement? The first thought that comes to mind is from my botanical colleagues who keep reminding me that plants make of 99% of the biomass of the Earth’s ecosystems. So, if this statement is correct, it must mean that large mammals have a very strong effect on plant ecosystem structure and function. And it must also imply that large mammals are virtually immune to predators, so no trophic cascade can occur to prevent plant overgrazing.

I appreciate that it is very difficult to test such a statement since evolution has been going on for a long time before humans arrived, and so there must have been a lot of other factors causing ecosystem changes in those early years. Humans have a disproportionate love for biodiversity that is larger than us. So, we revel in elephants, tigers, bears, and whales, while at the same time we pay little attention to the insects, small mammals, most fish, and plankton. Because of this size bias, we are greatly concerned with the conservation of large animals, as we should be, but much less concerned about what is happening to the small chaps.

What is the evidence that large mammals and birds have a disproportionate influence on ecosystem structure and function? In my experience, I would say there is very little evidence for strong ecosystem effects from the collapse of the megafauna. DeMaster et al. (2006) evaluated a proposed explanation for ecosystem collapse caused by whaling in the North Pacific Ocean and concluded that the evidence was weak for a sequential megafauna collapse caused by commercial whaling. Trites et al. (2007) and Wade et al. (2007) supported this conclusion. Citing paleo-ecological data for Australia, Johnson (2010) and Rule et al. (2012) argued in another evaluation of ecosystem changes that the human-driven extinction of the megafauna in Australia resulted in large changes in plant communities, potentially confounded by climate change and increases in fire frequency about 40K years ago. If we accept these controversies, we are left with trying to decide if the current losses of large mammals are of similar strength to those assigned to the Pleistocene megafauna, as suggested by Smith et al. (2018).

If we define ecosystem function as primary productivity and ecosystem structure as species diversity, I cannot think of a single case in recent studies where this idea has been clearly tested and supported. Perhaps this simply reflects my biased career working in arctic and subarctic ecosystems in which the vast majority of the energy flow in the system rotates through the smaller species rather than the larger ones. Take the Great Plains of North America with and without the bison herds. What aspect of ecosystem function has changed because of their loss? It is impossible to say because of human intervention in the fire cycle and agricultural pre-emption of much of the landscape. It is certainly correct that overgrazing impacts can be severe in human-managed landscapes with overstocking of cattle and sheep, and that is a tragedy brought on by economics, predator elimination programs, and human land use decisions. All the changes we can describe with paleo-ecological methods have potential explanations that are highly confounded.

I think the challenge is this: to demonstrate that the loss of large mammals at the present time creates a large change in ecosystem structure and function with data on energy flow and species diversity. The only place I can see it possible to do this experimentally today would be in arctic Canada where, at least in some areas, caribou come and go in large numbers and with relatively little human impact. I doubt that you could detect any large effect in this hypothetical experiment. It is the little chaps that matter to ecosystem function, not the big chaps that we all love so much. And I would worry if you could do this experiment, the argument would be that it is a special case of extreme environments not relevant to Africa or Australia.

No one should want the large mammals and birds to disappear, but the question of how this might play out in the coming 200 years in relation to ecosystem function requires more analysis. And unlike the current political inactivity over the looming crisis in climate change, we conservation biologists should certainly try to prevent the loss of megafauna.

Choquenot, D., and Bowman, D.M.J.S. 1998. Marsupial megafauna, Aborigines and the overkill hypothesis: application of predator-prey models to the question of Pleistocene extinction in Australia. Global Ecology and Biogeography Letters 7: 167-180.

DeMaster, D.P., Trites, A.W., Clapham, P., Mizroch, S., Wade, P., Small, R.J., and Hoef, J.V. 2006. The sequential megafaunal collapse hypothesis: testing with existing data. Progress in Oceanography 68(2-4): 329-342. doi:10.1016/j.pocean.2006.02.007

Grayson, D.K. 1977. Pleistocene avifaunas and the Overkill Hypothesis. Science 195: 691-693.

Guthrie, R.D. 1984. Mosaics, allelochemics and nutrients: An ecological theory of late Pleistocene megafaunal extinctions. In: Quaternary Extinctions: A Prehistoric Revolution ed by P.S. Martin and R.G. Klein. University of Arizona Press Tucson.

Johnson, C.N. 2010. Ecological consequences of Late Quaternary extinctions of megafauna. Proceeding of the Royal Society of London, Series B 276(1667): 2509-2519. doi: 10.1098/rspb.2008.1921.

Martin, P.S., and Wright, H.E. (eds). 1967. Pleistocene Extinctions; The Search for a Cause. Yale University Press, New Haven, Connecticut. 453 pp.

Rule, S., Brook, B.W., Haberle, S.G., Turney, C.S.M., Kershaw, A.P., and Johnson, C.N. 2012. The aftermath of megafaunal extinction: ecosystem transformation in Pleistocene Australia. Science 335(6075): 1483-1486. doi: 10.1126/science.1214261.

Smith, F.A., Elliott Smith, R.E., Lyons, S.K., and Payne, J.L. 2018. Body size downgrading of mammals over the late Quaternary. Science 360(6386): 310-313. doi: 10.1126/science.aao5987.

Trites, A.W., Deecke, V.B., Gregr, E.J., Ford, J.K.B., and Olesiuk, P.F. 2007. Killer whales, whaling, and sequential megafaunal collapse in the North Pacific: a comparative analysis of the dynamics of marine mammals in Alaska and British Columbia following commercial whaling. Marine Mammal Science 23(4): 751-765. doi: 10.1111/j.1748-7692.2006.00076.x.

Wade, P.R., et al. 2007. Killer whales and marine mammal trends in the North Pacific – a re-examination of evidence for sequential megafaunal collapse and the prey-switching hypothesis. Marine Mammal Science 23(4): 766-802. doi: 10.1111/j.1748-7692.2006.00093.x.

Three Approaches to Ecology

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I ask the question here why ecology is not appreciated as a science at a time when it is critical to the survival of the existing world. So the first question we need to answer is if this premise is correct. I offer only one example. A university zoology department has recently produced a discussion paper on its plans for faculty recruitment over the next 15 years. This document does not include the word “ecology” in any of its forward planning. Now it is probably not unusual for biology or zoology departments in major universities to downplay ecology when there is so much excitement in molecular biology, but it is an indicator that ecology is not a good place to put your money and reputation as you await a Nobel Prize. So if we can accept the initial premise that ecology is not appreciated, we might ask why this situation exists, a point raised long ago by O’Connor (2000). Here are a few thoughts on the matter.

There are three broad approaches to the science of ecology – theoretical ecology, empirical ecology, and applied ecology. These three areas of ecology rarely talk to each other, although one might hope that they could in future evolve into a seamless thread of science.

Theoretical ecology deals with the mathematical world that has too often only a tangential concern with ecological problems. It has its own journals and a whole set of elegant discussions that have few connections to the real world. It is most useful for exploring what might be if we make certain mathematical assumptions. It is without question the most prestigious part of the broad science of ecology, partly because it involves elegant mathematics and partly because it does not get involved in all the complexities of real-world ecological systems. It is the physics of ecology. As such it carries on in its own world and tends to be ignored by most of those working in the other two broad areas of ecology.

Empirical ecology has set itself the task of understanding how the natural world works at the level of individuals, populations, communities and ecosystems. In its pure form it does not care about solving practical ecological or environmental problems, but its practitioners assume probably correctly that the information they provide will in fact be useful now or in the future. It seeks generality but rarely finds it because all individuals and species differ in how they play the ecological game of survival. If it has a mantra, it is “the devil is in the details”. The problem is the details of empirical ecology are boring to politicians, business people, and to much of the television generation now operating with a 7 second or 140 character limit on concentration.

Applied ecology is where the action is now, and if you wish to be relevant and topical you should be an applied ecologist, whether a conservation biologist, a forester, or an agricultural scientist. The mantra of applied ecologists is to do no harm to the environment while solving real world problems. Applied ecologists are forced to put the human imprint into empirical ecology, so they are very much concerned with declining populations and extinctions of plants and animals. The main but not the sole impact of humans is on climate change, so much of applied ecology traces back to the impacts of climate change on ecosystems, all added to by the increasing human population with its rising expectations. But applied ecologists are always behind the environmental problems of the day because the issues multiply faster than possible solutions can be evaluated. This ought to make for high employment for applied ecologists but in fact the opposite seems to be happening because governments too often avoid long-term problems beyond their 4-year mandate. If you do not agree, think climate change.

So, the consequence is that we have three independent worlds out there. Applied ecologists are too busy to apply the successful paradigms of empirical ecology to their problems because they are under strict time limits by their line managers who need to suggest immediate action on problems. They must therefore fire off solutions like golf balls in all directions, hoping that some might actually help solve problems. Empirical ecologists may not be helpful for applied ecologists if they are overwhelmed by the details of their particular system of study and are constrained by the ‘publish or perish’ mentality of the granting agencies.

Finally, we lay on top all this a lack of funding in the environmental sciences for investigating and solving both immediate and long-term ecological problems. And I am back to my favourite quote in the ecological literature:

“Humans, including ecologists, have a peculiar fascination with attempting to correct one ecological mistake with another, rather than removing the source of the problem.” (Schindler 1997).

What can we do about this? Three things. Pressure our politicians to increase funding on long-term environmental problems. This will provide the person-power to find and test solutions to our known problems. Vote with your ballot and your feet to improve sustainability. And whether you are young or old strive to do no harm to the Earth. And if all this is too difficult, take some practical advice not to buy a house in Miami Beach, or any house near the beach. Do something for the environment every day.

 

O’Connor, R.J. (2000) Why ecology lags behind biology. The Scientist 14(20):35. (October 16, 2000).

Schindler, D.W. (1997) Liming to restore acidified lakes and streams: a typical approach to restoring damaged ecosystems? Restoration Ecology 5:1-6

 

On Evolution and Ecology and Climate Change

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If ecology can team up with evolution to become a predictive science, we can all profit greatly since it will make us more like physics and the hard sciences. It is highly desirable to have a grand vision of accomplishing this, but there could be a few roadblocks on the way. A recent paper by Bay et al. (2018) illustrates some of the difficulties we face.

The yellow warbler (Setophaga petechia) has a broad breeding range across the United States and Canada, and could therefore be a good species to survey because it inhabits widely different climatic zones. Bay et al. (2018) identified genomic variation associated with climate across the breeding range of this migratory songbird, and concluded that populations requiring the greatest shifts in allele frequencies to keep pace with future climate change have experienced the largest population declines, suggesting that failure to adapt may have already negatively affected population abundance. This study by Bay et al. (2018) sampled 229 yellow warblers from 21 locations across North America, with an average of 10 birds per sample area (range n = 6 to 21). They examined 104,711 single-nucleotide polymorphisms. They correlated genetic structure to 19 climate variables and 3 vegetation indices, a measure of surface moisture, and average elevation. This is an important study claiming to support an important conclusion, and consequently it is also important to break it down into the three major assumptions on which it rests.

First, this study is a space for time analysis, a subject of much discussion already in plant ecology (e.g. Pickett 1989, Blois et al. 2013). It is an untested assumption that you can substitute space for time in analyzing for future evolutionary changes.

Second, the conclusions of the Bay et al. paper rest on an assumption that you have adequate data on the genetics involved in change and on the demography of the species. A clear understanding of the ecology of the species and what limits its distribution and abundance would seem to be prerequisites for understanding the mechanisms of how evolutionary changes might occur.

The third assumption is that, if there is a correlation between the genetic measures and the climate or vegetation indices, one can identify the precise ‘genomic vulnerability’ of the local population. Genomic variation was most closely related to precipitation variables at each site. The geographic area with one of the highest scores in genomic vulnerability was in the desert area of the intermountain west (USA). As far as I can determine from their Figure 1, there was only one sampling site in this whole area of the intermountain west. Finally Bay et al. (2018) compared the genomic vulnerability data to the population changes reported for each site. Population changes for each sampled site were obtained from the North American Breeding Bird Survey data from 1996 to 2012.

The genetic data and its analysis are more impressive, and since I am not a genetics expert I will simply give it a A grade for genetics. It is the ecology that worries me. I doubt that the North American Breeding Bird Survey is a very precise measure of population changes in any particular area. But following the Bay et al. paper, assume that it is a good measure of changing abundance for the yellow warbler. From the Bay et al. paper abstract we see this prediction:

“Populations requiring the greatest shifts in allele frequencies to keep pace with future climate change have experienced the largest population declines, suggesting that failure to adapt may have already negatively affected populations.”

The prediction is illustrated in Figure 1 below from the Bay et al. paper.

Figure 1. From Bay et al. (2018) Figure 2C. (Red dot explained below).

Consider a single case, the Great Basin, area S09 of the Sauer et al. (2017) breeding bird surveys. From the map in Bay et al. (2018) Figure 2 we get the prediction of a very high genomic vulnerability (above 0.06, approximate red dot in Figure 1 above) for the Great Basin, and thus a strongly declining population trend. But if we go to the Sauer et al. (2017) database, we get this result for the Great Basin (Figure 2 here), a completely stable yellow warbler population for the last 45 years.

Figure 2. Data for the Great Basin populations of the Yellow Warbler from the North American Breeding Bird Survey, 1967 to 2015 (area S09). (From Sauer et al. 2017)

One clue to this discrepancy is shown in Figure 1 above where R2 = 0.10, which is to say the predictability of this genomic model is near zero.

So where does this leave us? We have what appears to be an A grade genetic analysis coupled with a D- grade ecological model in which explanations are not based on any mechanism of population dynamics, so that the model presented is useless for any predictions that can be tested in the next 10-20 years. I am far from convinced that this is a useful exercise. It would be a good paper for a graduate seminar discussion. Marvelous genetics, very poor ecology.

And as a footnote I note that mammalian ecologists have already taken a different but more insightful approach to this whole problem of climate-driven adaptation (Boutin and Lane 2014).

Bay, R.A., Harrigan, R.J., Underwood, V.L., Gibbs, H.L., Smith, T.B., and Ruegg, K. 2018. Genomic signals of selection predict climate-driven population declines in a migratory bird. Science 359(6371): 83-86. doi: 10.1126/science.aan4380.

Blois, J.L., Williams, J.W., Fitzpatrick, M.C., Jackson, S.T., and Ferrier, S. 2013. Space can substitute for time in predicting climate-change effects on biodiversity. Proceedings of the National Academy of Sciences 110(23): 9374-9379. doi: 10.1073/pnas.1220228110.

Boutin, S., and Lane, J.E. 2014. Climate change and mammals: evolutionary versus plastic responses. Evolutionary Applications 7(1): 29-41. doi: 10.1111/eva.12121.

Pickett, S.T.A. 1989. Space-for-Time substitution as an alternative to long-term studies. In Long-Term Studies in Ecology: Approaches and Alternatives. Edited by G.E. Likens. Springer New York, New York, NY. pp. 110-135.

Sauer, J.R., Niven, D.K., Hines, J.E., D. J. Ziolkowski, J., Pardieck, K.L., and Fallon, J.E. 2017. The North American Breeding Bird Survey, Results and Analysis 1966 – 2015. USGS Patuxent Wildlife Research Center, Laurel, MD. https://www.mbr-pwrc.usgs.gov/bbs/

On Caribou and Hypothesis Testing

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Mountain caribou populations in western Canada have been declining for the past 10-20 years and concern has mounted to the point where extinction of many populations could be imminent, and the Canadian federal government is asking why this has occurred. This conservation issue has supported a host of field studies to determine what the threatening processes are and what we can do about them. A recent excellent summary of experimental studies in British Columbia (Serrouya et al. 2017) has stimulated me to examine this caribou crisis as an illustration of the art of hypothesis testing in field ecology. We teach all our students to specify hypotheses and alternative hypotheses as the first step to solving problems in population ecology, so here is a good example to start with.

From the abstract of this paper, here is a statement of the problem and the major hypothesis:

“The expansion of moose into southern British Columbia caused the decline and extirpation of woodland caribou due to their shared predators, a process commonly referred to as apparent competition. Using an adaptive management experiment, we tested the hypothesis that reducing moose to historic levels would reduce apparent competition and therefore recover caribou populations. “

So the first observation we might make is that much is left out of this approach to the problem. Populations can decline because of habitat loss, food shortage, excessive hunting, predation, parasitism, disease, severe weather, or inbreeding depression. In this case much background research has narrowed the field to focus on predation as a major limitation, so we can begin our search by focusing on the predation factor (review in Boutin and Merrill 2016). In particular Serrouya et al. (2017) focused their studies on the nexus of moose, wolves, and caribou and the supposition that wolves feed preferentially on moose and only secondarily on caribou, so that if moose numbers are lower, wolf numbers will be lower and incidental kills of caribou will be reduced. So they proposed two very specific hypotheses – that wolves are limited by moose abundance, and that caribou are limited by wolf predation. The experiment proposed and carried out was relatively simple in concept: kill moose by allowing more hunting in certain areas and measure the changes in wolf numbers and caribou numbers.

The experimental area contained 3 small herds of caribou (50 to 150) and the unmanipulated area contained 2 herds (20 and 120 animals) when the study began in 2003. The extended hunting worked well, and moose in the experimental area were reduced from about 1600 animals down to about 500 over the period from 2003 to 2014. Wolf numbers in the experimental area declined by about half over the experimental period because of dispersal out of the area and some starvation within the area. So the two necessary conditions of the experiment were satisfied – moose numbers declined by about two-thirds from additional hunting and wolf numbers declined by about half on the experimental area. But the caribou population on the experimental area showed mixed results with one population showing a slight increase in numbers but the other two showing a slight loss. On the unmanipulated area both caribou populations showed a continuing slow decline. On the positive side the survival rate of adult caribou was higher on the experimental area, suggesting that the treatment hypothesis was correct.

From the viewpoint of caribou conservation, the experiment failed to change the caribou population from continuous slow declines to the rapid increase needed to recover these populations to their former greater abundance. At best it could be argued that this particular experiment slowed the rate of caribou decline. Why might this be? We can make a list of possibilities:

  1. Moose numbers on the experimental area were not reduced enough (to 300 instead of to 500 achieved). Lower moose would have meant much lower wolf numbers.
  2. Small caribou populations are nearly impossible to recover because of chance events that affect small numbers. A few wolves or bears or cougars could be making all the difference to populations numbering 10-20 individuals.
  3. The experimental area and the unmanipulated area were not assigned treatments at random. This would mean to a pure statistician that you cannot make statistical comparisons between these two areas.
  4. The general hypothesis being tested is wrong, and predation by wolves is not the major limiting factor to mountain caribou populations. Many factors are involved in caribou declines and we cannot determine what they are because they change for area to area, year to year.
  5. It is impossible to do these landscape experiments because for large landscapes it is impossible to find 2 or more areas that can be considered replicates.
  6. The experimental manipulation was not carried out long enough. Ten years of manipulation is not long for caribou who have a generation time of 15-25 years.

Let us evaluate these 6 points.

#1 is fair enough, hard to achieve a population of moose this low but possible in a second experiment.

#2 is a worry because it is difficult to deal experimentally with small populations, but we have to take the populations as a given at the time we do a manipulation.

#3 is true if you are a purist but is silly in the real world where treatments can never be assigned at random in landscape experiments.

#4 is a concern and it would be nice to include bears and other predators in the studies but there is a limit to people and money. Almost all previous studies in mountain caribou declines have pointed the finger at wolves so it is only reasonable to start with this idea. The multiple factor idea is hopeless to investigate or indeed even to study without infinite time and resources.

#5 is like #3 and it is an impossible constraint on field studies. It is a common statistical fallacy to assume that replicates must be identical in every conceivable way. If this were true, no one could do any science, lab or field.

#6 is correct but was impossible in this case because the management agencies forced this study to end in 2014 so that they could conduct another different experiment. There is always a problem deciding how long a study is sufficient, and the universal problem is that the scientists or (more likely) the money and the landscape managers run out of energy if the time exceeds about 10 years or more. The result is that one must qualify the conclusions to state that this is what happened in the 10 years available for study.

This study involved a heroic amount of field work over 10 years, and is a landmark in showing what needs to be done and the scale involved. It is a far cry from sitting at a computer designing the perfect field experiment on a theoretical landscape to actually carrying out the field work to get the data summarized in this paper. The next step is to continue to monitor some of these small caribou populations, the wolves and moose to determine how this food chain continues to adjust to changes in prey levels. The next experiment needed is not yet clear, and the eternal problem is to find the high levels of funding needed to study both predators and prey in any ecosystem in the detail needed to understand why prey numbers change. Perhaps a study of all the major predators – wolves, bears, cougars – in this system should be next. We now have the radio telemetry advances that allow satellite locations, activity levels, timing of mortality, proximity sensors when predators are near their prey, and even video and sound recording so that more details of predation events can be recorded. But all this costs money that is not yet here because governments and people have other priorities and value the natural world rather less than we ecologists would prefer. There is not yet a Nobel Prize for ecological field research, and yet here is a study on an iconic Canadian species that would be high up in the running.

What would I add to this paper? My curiosity would be satisfied by the number of person-years and the budget needed to collect and analyze these results. These statistics should be on every scientific paper. And perhaps a discussion of what to do next. In much of ecology these kinds of discussions are done informally over coffee and students who want to know how science works would benefit from listening to how these informal discussions evolve. Ecology is far from simple. Physics and chemistry are simple, genetics is simple, and ecology is really a difficult science.

Boutin, S. and Merrill, E. 2016. A review of population-based management of Southern Mountain caribou in BC. {Unpublished review available at: http://cmiae.org/wp-content/uploads/Mountain-Caribou-review-final.pdf

Serrouya, R., McLellan, B.N., van Oort, H., Mowat, G., and Boutin, S. 2017. Experimental moose reduction lowers wolf density and stops decline of endangered caribou. PeerJ  5: e3736. doi: 10.7717/peerj.3736.