Monthly Archives: November 2013

On Charles Elton

Charles Elton was the Father of Animal Ecology and many young ecologists do not learn very much about him. He founded the Bureau of Animal Population at Oxford in 1932, and much of the history of his research group is captured in Peter Crowcroft’s book “Elton’s Ecologists” (1991). I was fortunate to spend the winter of 1960-61 at the Bureau while I was completing my Ph.D. at UBC with Dennis Chitty. It was Dennis’ last year at the Bureau, having gone there in 1935 when he had just finished his undergraduate work at the University of Toronto.

The Bureau of Animal Population or BAP, as all connected with it came to call it, had been born in January 1932 and by 1934 Oxford University had guaranteed funds for its core costs for five years with 3-4 scientific staff and a very few assistants. Survival as a unit depended on working on numerous applied projects, and the species receiving attention included Canadian snowshoe hares, Canadian lynx, muskrat, beaver, lemmings, European rabbits, squirrels, voles and the wood mouse. The Bureau was the home of the newly created Journal of Animal Ecology, of which Charles Elton was the first editor.

Charles Elton was a proper English gentleman, a gentle soul who had a coterie of first class ecologists in the BAP. The BAP was very nearly the world centre for ecology from the 1930s to the 1960s when ecology began its great growth around the world, so everyone who was interested in population ecology considered it equivalent to Mecca for science. Every day there was tea in the BAP in 1960, when we all took time to interact with the other postdocs and graduate students in the BAP, a total group of perhaps 15-20. Once a week Mr. Elton (as he never did a Ph.D.) would preside over tea around a table in the BAP Library and give out any news of the week to the staff and students. On most days he wore a tie and a sport coat in the best English tradition, and signed his letters as “Elton”. In 1960 he was compiling a species list for Wytham Woods, a 390 ha forest reserve belonging to the University. He felt strongly that one had to know all the species in a community before you could understand how it operated. So one could see him day after day pinning insects in trays. He was always very serious, and the only joke I ever heard him tell was about how he could never understand Americans. He had gone to the New World after the War, perhaps 1947 or 1948 and was visiting a famous American scientist. They had to get up at 0700 in the morning and rush to work without a proper English breakfast, and so at 0800 they arrived in the professor’s office, and then Elton said he was told ‘now you can relax’. It was not the proper English way to start the day and he could never understand the rush-rush style of the New World.

Charles Elton founded the Journal of Animal Ecology in 1931, now one of our leading journals. In the early days he did much of the reviewing and accepting of papers for the Journal. He had an amusing tale of the classic papers of A.J. Nicholson (1933) on the balance of animal populations. He received this very long paper and he could not find anyone who would agree to review it so he did it himself. He confessed to us one day at tea that he found he could not understand anything in the paper, so he decided it must be very brilliant so he published it immediately. Alas those days are gone.

There were of course no electronic machines even in 1960 and Elton did all his writing by pen and paper. He had just finished the now famous book “The Ecology of Invasions” and his secretary who typed all his work pointed out to me that he never changed a word from what he first wrote. No need for revisions and revisions. He was of course like a god to all of us young ecologists, and so we were very fortunate that this was the year in which he was teaching his Animal Ecology course to Oxford undergraduates. All of us graduate students and postdocs went along, as it was only a series of 14 lectures in the best Oxford tradition. The classroom was full in the first lecture, which was one of the worst lectures I have ever attended. We were rather stunned that such a great man could lecture so hopelessly, mumbling in a monotone, showing slides but almost never referring to them, every mistake in the book. We realized then that greatness could occur in many dimensions and his skill was as a writer. Classroom attendance fell like an exponential and by the fifth lecture no one was left in the classroom but we of the BAP.

Elton organized the BAP as a small research unit and did not believe that any research unit should exceed more than a handful of scientists who interacted all the time over a small subset of problems. In the early days much of the research was on cyclic populations of rodents and fur bearers in Europe and North America, but it moved to insects and broader problems after the War.

Oxford was a strange place to a North American in 1960. Too many of the professors were at odds with one another, jostling for fame we all presumed. It was impossible not to have many enemies within and outside the walls of Oxford, and we as students never quite knew why some were praised and others reviled. Perhaps ideas were confounded with personalities, and no one thought that you could respect a scientist but disagree with his or her view of science. But much was at stake then, and when you were King of Oxford you were king of the hill. Now 50 years later we have many kings of science all around the world, and I hope that Oxford has changed.


Crowcroft, P. 1991. Elton’s Ecologists: A History of the Bureau of Animal Population. University of Chicago Press, Chicago. 177 pp.

Elton, C. S. 1958. The Ecology of Invasions by Animals and Plants. Methuen, London. 181 pp.

Nicholson, A. J. 1933. The balance of animal populations. Journal of Animal Ecology 2:132-178.


Science and Money

Why do we the public support science? The general answer is that science produces products we like, improves our possibilities of a healthy life, and increases wealth. A less general answer is that science informs us about how the Earth works and how the Earth fits into the universe. Most people would agree that science should not provide us with ethical judgments or define good and evil. The result of this dichotomy between science and ethics in the broad sense is that scientists live in a divided world. Each scientist has definite views on what is good for society and what is evil, and these views can differ among scientists in different cultures. But as a scientist he or she cannot use scientific information to define good and evil and therefore to advise governments about what actions to take in particular problems. All this is very vague until you bring it into the arguments of our time – abortion, gay marriage, the death penalty for criminals, nuclear power, fish farming in the ocean, tar-sands oil, fracking – the list goes on.

Scientific information is vital to the decisions made on all these issues. Consider fracking for oil and gas. One scientific question is: Does fracking contaminate the water table? Does fracking release the greenhouse gas methane to the atmosphere? Given adequate scientific information, governments and the public may support or ban fracking, and to support or ban is not a scientific issue but an ethical one. Public opinion of course is affected by scientific findings, and the job of the scientist is to make these findings precise and accurate. But to do that requires money.

The result of all this is that governments and the public have developed a ranking system of the sciences. At the top of the totem pole are physics and chemistry (and their associated engineering sciences) because their findings and products are typically thought to be very useful – cars, computers, IPhones, medical drugs. Not only are they useful but they make lots of money for many people. Geology is also somewhere near the top of the pole because it produces oil and minerals, but suffers somewhat from being responsible for earthquakes and tsunamis. Somewhat lower on the totem pole are the biological sciences. Molecular biology is closely akin to chemistry and offers medical promises so it is high on the totem pole. Biochemistry and physiology follow closely, but they are somewhat suspect unless they promise that their results can be applied to human wellbeing. Near the bottom of the totem pole are the ecologists who describe how the web of life works on Earth and how it has been affected by human actions. The top position of ecology goes to natural history, and bird watching brings much happiness to many people. TV programs like those of David Attenborough bring images of many areas and species that few will be able to visit or see. Descriptive ecology fares slightly less well because it seems harmless to most people but is unable to generate money in any useful manner. Conservation ecology sits at the bottom of the ecology heap, falling into the dark side because it continually points out problems of what humans have done or are doing to life on earth, to ecosystem processes that are essential to a healthy environment. Only climate scientists are lower on the totem pole than ecologists because they are always talking about the coming train wreck of climate change, with the ethical implication that we the public should be doing something by changing our habits.

The results are that funding for scientific work follows the totem pole. Ecologists fare poorly along with organismal biology with the result that we do not have an inventory of life on Earth or an adequate understanding of how most of the Earth’s ecosystems operate. Climate scientists are perhaps fortunate because the gathering of climate data has been extensive because people need weather information to drive to work or plant their crops. Consequently, even though it is at the lower end of the totem pole climate science has much data to utilize, although many do not like the ensuing message. I suspect many governments of the day would like to close down all the weather stations to save money as well as to avoid further negative findings.

There is unlikely to be any move soon in the relative positions on the totem pole for the different sciences. We ecologists live in a trickle down world where some funding sieves through to the lower layers of biology, partly by accident and partly because there are some who think that we should know more about our Earth’s biological heritage.

Fortin, J.-M. and D. J. Currie. 2013. Big science vs. little science: How scientific impact scales with funding. PLoS ONE 8:e65263.

Haufe, C. 2013. Why do funding agencies favor hypothesis testing? Studies in History and Philosophy of Science Part A 44:363-374.

May, R. M. 1997. The scientific wealth of nations. Science 275:793-796.


The 7 Generation Rule

Some of the First Nations people of northern Canada believe that we are stewards of the Earth, and for their particular area the land must be managed within a time horizon of 7 generations, approximately 200 years. If we are serious about sustainability, we need to ask for each situation how the impact of this or that environmental decision will track for the next 7 generations. It is quite clear to anyone who listens to any of the news media that we are not at present even living by a 1 Generation Rule. The guide of governments and corporations of virtually all developed nations is economic growth, producing societies that are more and more inequitable, the rich 1% and the poor 99%. The environment is almost never mentioned. What might we do if we lived by the 7 Generation Rule?

The first item to question might be the transportation system of the world and the use of fossil fuels. All is well you might argue, gasoline and diesel are cheap, we carry on. But if we think of future generations we might worry about whether increasing CO2 is causing climate change; the naïve belief that burning fossil fuels has nothing to do with climate change means that we do not believe any of the laws of physics. There is yet another problem somewhere on the 7 Generation horizon – fossil fuels are a non-renewable resource. At some point sooner or later we will run out of fossil fuels, or as an economist would say fossil fuels will not run out but will get very expensive. How far will you be driving in 7 Generations if the price of gasoline is $10,000 a litre? Round that to $40,000 a gallon if you calculate in those units.

But if I cannot drive my car on gasoline, surely someone will invent a car that runs on solar power. Technology will save us. This is akin to a religious belief for many people, and it might be true. If it is, then we can leave the coal, oil, and natural gas in the ground, which is what we ought to plan in any event if we live by the 7 Generation Rule. It is good to be an optimist but it is also good to have a Plan B.

There is one more problem that might be even more important than driving our cars – the provisioning of food. The demand for food in the world today grows at a rate exceeding the rate of food production. No problem, you say, we have plenty of food as long as we continue to neglect one-third of the people on Earth that are undernourished and as long as we operate with the 1 Generation Rule. There are several ways of solving this problem but many of the suggestions are quite impossible. We can become more vegetarian in our diets, and that would be good. But we cannot develop more farmland because virtually all of it is in use. We can increase the productivity of our crops by genetic means, but we cannot compensate for losses in soil fertility and erosion. Fertilizer which is essential to modern agriculture could be problematic. Nitrogen fertilizer is now made largely from fossil fuel (natural gas) and phosphate fertilizer comes entirely from phosphate rocks which are being mined but are also a non-renewable resource. What does our 4th or 5th generation do when phosphate runs out? Might we consider recycling starting now to prepare for the 7th generation?

Ecologists fight now with minimal funding to describe and protect the biodiversity of the Earth, which might be useful already to generation 3, while governments spend much more money subsidizing the fossil fuel industry that is destroying the Earth. There is little money left for environmental protection. How is your government tracking toward a sustainable planet? What Generation Rule are they following? Ask yourself these questions the next time you vote.

Diamond, J. 2011. Collapse: How Societies Choose to Fail or Survive. Penguin Books, London. 608 pp. ISBN: 9780241958681

McKibben, B. 2013. Oil and Honey: The Education of an Unlikely Activist. Times Books, New York. 272 pp. ISBN: 9780805092844

On Alpha-Ecology

All science advances on the back of previous scientists. No advances can be made without recognizing problems, and problems cannot bet recognized without having completed a great deal of descriptive natural history. Natural history has been described by some physicists as ‘stamp-collecting’ and so has been condemned forever in the totem pole of science as the worst thing you could possibly do. Perhaps we would improve our image if we called natural history alpha-ecology.

Let us start with the biggest problem in biology, the fact that we do not know how many species inhabit the earth (Mora et al. 2011). A minor problem most people seem to think and little effort has gone into encouraging students to make a career of traditional taxonomy. Instead we can sequence the genome of any organism without even being able to put a Latin name on it. Something is rather backwards here, and a great deal of alpha-biology is waiting to be done on this inventory problem. Much of taxonomic description considers low-level hypotheses about evolutionary relationships and these are important to document as a part of understanding the Earth’s biodiversity.

In ecology we have an equivalent problem of describing the species that live in a community or ecosystem, and then constructing the food webs of the community. This is a daunting task and if you wish to understand community dynamics you will have to do a lot of descriptive work, alpha ecology, before you can get to the point of testing hypotheses about community dynamics (Thompson et al. 2012). Again it is largely a bit of detective work to see who eats whom in a food web, but without all this work we cannot progress. The second part of community dynamics is being able to estimate accurately the numbers of organisms in the different species groups. Once you dig into existing food web data, you begin to realize that much of what we think is a good estimate of abundance is in fact a weak estimate of unknown accuracy. We have to be careful in analysing community dynamics to avoid estimations based more on random numbers than on biological reality.

The problem came home to me in a revealing exchange in Nature about whether the existing fisheries data for the world’s oceans is reliable or not (Pauly, Hilborn, and Branch 2013). For years we have been managing the oceanic fisheries of the world on the basis of fishing catch data of the sort reported to FAO, and yet there is considerable disagreement about the reliability of these numbers. We must continue to use them as we have no other source of information for most oceanic fisheries, but there must be some doubt that we are relying too much on unreliable data. On the one hand, some fishery scientists argue with these data that we are overexploiting the ocean fisheries, but other fishery scientists argue that the oceanic fisheries are by and large in good shape. Controversies like this confuse the public and the policy makers and tell us we have a long way to go to improve our alpha-ecology.

I think the bottom line is that if you wish to test any ecological hypothesis you need to have reliable data, and this means a great deal of alpha-ecology is needed, research that will not get you a Nobel Prize but will help us understand how the Earth’s ecosystem operates.

Mora, C., et al. 2013. How Many Species Are There on Earth and in the Ocean? PLoS Biology 9:e1001127.

Pauly, D., R. Hilborn, and T. A. Branch. 2013. Fisheries: Does catch reflect abundance? Nature 494:303-306.

Thompson, R. M., et al. 2012. Food webs: reconciling the structure and function of biodiversity. Trends in Ecology & Evolution 27:689-697.

Bandwagons in Ecology

Scientists are like most people in their attraction to bandwagons. Often this is good, since some parts of any particular science may move more quickly than others, creating a bandwagon for scientists building a career. But sometimes this is detrimental in diverting efforts and money from one aspect of a science to another. All would be fine if the older parts of a science were thoroughly understood, and the new bandwagon opened up the solutions to critical problems.

So what does all this have to do with ecology? Ecology has been one example of a science beset by one bandwagon after another during the past 50 years. Many of these bandwagons were relatively harmless because they started with the promise to solve all problems and ended up contributing a small bit of understanding to the subject as it matured. I am thinking now of energy flow in the 1950s, systems ecology and density dependence in the 1960s, competition theory in the 1970s, and mathematical modelling from the 1980s onward. Other examples could be added to this list. At the moment we have two bandwagons that deserve some discussion – climate change and evolutionary ecology.

Climate change is one of the three most critical problems of our day and so it is understandable that much is written about it. Consequently it appears on all grant and scholarship applications as a relevant field. The problem is twofold. First of all, we should not take weather, the ecological side of climate, as the universal explanation for everything that is changing without considering alternative hypotheses for change. If the geographical distribution of a species is expanding toward the poles, climate change is only one of several possible reasons for this. The factors limiting geographic ranges are multiple and have been studied less well than any ecologist would like. We need to keep in mind that there are other ecological problems out there that are not directly tied in with climate change, and these need to be pursued as well. If you want an example, consider the problem of biological control of invasive species.

Evolutionary ecology is a second bandwagon and I fear it is tilting the entire focus of ecological research. The reason is quite clear – technological advancements in genetic studies. Much of science is driven by technological advances and that is good, but again it should not mean that we ignore other unresolved problems. In particular evolutionary ecology has the great potential to describe the world in great detail without necessarily adding any critical insights. In many cases it is stamp collecting and it reminds me of the saying that “Nero fiddles while Rome burns”. Should we as ecologists be concerned more about the practical problems of our day, or about simply understanding nature? There is no reason of course not to do both, and different individuals have talents in different areas of science. But some ecologists might feel as I do that ecological questions are poorly served by much of evolutionary ecology. I listen to many evolutionary ecologists telling us that their work is solving some ecological question when it is obvious that this is a leap of faith with little substance.

I think we need to ask as ecologists what are the problems we wish to solve, and if we could ever decide on a list of these problems, we could ask where we currently sit in solving these problems. It causes a great focus of the mind to look at a practical problem and ask what ecologists are doing about it. At the moment I am in the Philippines at the International Rice Research Institute, and I am overwhelmed by the ecological questions that interface with sustainable rice cropping in Southeast Asia, of pests and beneficial animals and plants, of migratory birds, of chemical poisons and their impacts on non-target species, the list goes on. The assumption at the moment seems to be that plant breeding and genetics will conquer all problems, but we ought to have a Plan B to look at the community and ecosystem dynamics that centre on a rice paddy, and how that might interface with the changing varieties of rice that are produced. We would be more humble if we moved away from genetic determinism to consider that there are other issues, currently ignored, that only ecologists can solve.

Bandwagons will always occur in science, but we should be careful that not everyone follows the pied pipers of the moment.

The Hippocratic Oath and Ecology

“Physician, do no harm” (Hippocrates, Greece, 5th century BC) is one of the classic ethical standards of medicine. Of course as medical science has progressed, treatments that were once considered to be beneficial are now known to cause harm, so one has to apply these standards to the time and place of action. How does all of this apply to ecology and environmental science?
All science is or should be evidence-based and the job of the ecologist is to examine and measure the evidence about how the biological world works, how natural populations, communities, and ecosystems operate and continue to exist. Given that evolution is the background to all these operations, in the long term individual species will come and go and change the dynamics we now describe. At the level of basic or “blue-sky” research, ecologists run into few ethical issues. But at the level of applied ecology, we become the ‘physicians of the world’ because we must assess the problems that arise in the natural world from the actions or inactions of humans. Consequently when ecologists investigate problems caused by mining, logging, aquaculture, or agriculture, and the associated issues caused by population growth, we have an ethical responsibility captured by the Hippocratic Oath.
In many situations ecologists and environmental scientists do well, laying out the issues, the science behind the measured effects, and the best predictions they can make about future changes. Climate change science is the best current example. But in many areas the conclusions of our best ecologists and environmental scientists crash head on into the economic train that drives 95% of decision making at the political and business levels. This is the key point where the Hippocratic Oath must enter if we wish to behave ethically. We cannot allow companies or the government to carry out environmental policies that are harmful to the populations, communities and ecosystems of the Earth without our voices being heard. This does not permit us to fabricate evidence or extrapolate beyond what is known. It does permit us to say what is not known and needs investigation, and that the policy of “what you do not know cannot hurt you” is stupidity squared. None of this endears us to the business community or the government bent on economic growth at all costs.
We can hope that this is changing, albeit slowly. Politicians and oil companies now at least talk about ‘sustainability’ while pushing ahead. But if more wealth is gained at the expense of the Earth we are lost in the long term. A major problem for ecologists is that operational changes are made in forestry, agriculture and mining with little thought to their consequences for biodiversity and ecologists are left to pick up the pieces later. If you wish an immediate example, fracking for oil and gas is more than enough. This is not an intelligent way to operate if we wish to be stewards of the Earth. So in every bit of ecology we do, we need to keep the Hippocratic Oath in mind, and do our best to stop harming the Earth.
And at the political level, we could take the radical step of asking that every Minister of the Environment ought to be trained in environmental science and ecology, and understand the environmental problems of the Earth.