Tag Archives: climate change

Biodiversity Science

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Protecting biodiversity is a goal of most people who value the environment. My question is what are the goals of biodiversity science and how do we achieve them? Some history is in order here. The term ‘biodiversity’ was coined in the 1980s as the complete biosphere including all species and ecosystems on Earth. The idea of cataloguing all the species on Earth was present many decades before this time, since the origin of the biological sciences. By the 1990s ‘biodiversity conservation’ became a popular subject and has grown greatly since then as a companion to CO2 emissions and the climate change problem. The twin broad goals of biodiversity science and biodiversity conservation are (1) to name and describe all the species on Earth, and (2), to protect all species from extinction, preventing a loss of biodiversity. How can we achieve these two goals?

The first goal of describing species faces challenges from disagreements over what a species is or is not. The old description of a species was to describe what group it was part of, and then how different this particular species was from other members of the group. In the good old days this was primarily based on reproductive incompatibility between species, if no successful reproduction, must be a new species. This simple common-sense view was subject to many attacks since some organisms that we see as different can in fact interbreed. Lions and tigers breed together and are an example, but if their interbred offspring are sterile, clearly, they are two different species. But many arguments arose because there was no data available for 99% of species to know if they could interbreed or not. The fallback position has been to describe the anatomy of a potential species and its relatives and judge from anatomy how different they were. Endless arguments followed, egged on by naturalists who pointed out that if the elephants in India were separated by a continent from elephants in Africa, clearly, they must be different species defined by geography. Many academic wars were fought over these issues.

Then in 1953 the structure of DNA was unravelled, and a new era dawned because with advances in technology of decoding genes we could describe species in a completely new way by determining how much DNA they had in common. But what is the magic percentage of common DNA? Humans and chimpanzees have 98.6% of their DNA in common, but despite this high similarity no one argues that they are the same species.

Despite this uncertainty the answer now seems much simpler: sequence the DNA of everything and you will have the true tree of life for defining separate species. While this was a dream 20 years ago, it is now a technical reality with rapid sequencing methods to help us get criminals and define species. Problem (1) solved?

Enter the lonely ecologist into this fray. Ecologists do not just want names, they wish to understand the function of each of the ‘species’ within communities and ecosystems, how does all this biodiversity interact to produce what we see in the landscape? For the moment we have approximately 10 million species on Earth, but somewhere around 80% of these ‘species’ are still undescribed. So now we have a clash of biodiversity visions, we cannot describe all the species on Earth even on the time scale of centuries, so we cannot achieve goal (1) of biodiversity science in any reasonable time. We have measured the DNA sequence of thousands of organisms that we can capture but we cannot describe them formally as species in the older sense. Perhaps it is akin to having all the phone numbers in the New York City phone book but not knowing to whom the numbers belong.

But the more immediate problem comes with objective (2) to prevent extinctions. Enter the conservation ecologist. The first problem is discussed above, we ecologists have no way of knowing how many species are in danger of extinction. We must look for rare or declining species, but we have complete inventory for few places on Earth. We must concentrate on large mammals and birds, and hope that they act as umbrella species and represent all of biodiversity. When we do have information on threatened species, for the most part there is no money to do the ecological studies needed to reverse declines in abundance. If there is money to list species and give a recovery plan on paper, then we find there is no money to implement the recovery plan. The Species-At-Risk act in Canada was passed in 2002 and has generated many recovery plans mostly for vertebrate species that have come to their attention. Almost none of these recovery plans have been completed, so in general we are all in favour of preventing extinctions but only it if costs us nothing. By and large the politics of preventing extinctions is very strongly supported, but the economic value of extinctions is nearly zero.

None of this is very cheery to conservation biologists. Two approaches have been suggested. The first is Big Science, use satellites and drones to scan the Earth every year to describe changes in landscapes and from these images infer biodiversity ‘health’. Simple and very expensive with AI to the rescue. But while we can see largescale landscape changes, the crux is to do something about them, and it is here that we fail because of the wall of climate change that we have no control over at present. Big Science may well assist us in seeing patterns of change, but it produces no path to understanding food webs or mediating changes in threatened populations. The second is small-scale biodiversity studies that focus on what species are present, how their numbers are changing, and what are the causes of change. Difficult, possible, but very expensive because you must put biologists in the field, on the ground to do the relevant measurements over a long-time frame. The techniques are there to use, thanks to much work on ecological methods in the past. What is missing again is the money. There are a few good examples of this small-scale approach but without good organization and good funding many of these attempts stop after too few years of data.

We are left with a dilemma of a particular science, Biodiversity Science, that has no way of achieving either of its two main objectives to name and to protect species on a global level. On a local level we can adopt partial methods of success by designating and protecting national parks and marine protected areas, and by studying only a few important species, the keystone species of food webs. But then we need extensive research to determine how to protect these areas and species from the inexorable march of climate change, which has singlehandedly complicated achieving biodiversity science’s two goals. Alas at the present time we may have another science to join the description of economics as a “dismal science” And we have not even started to discuss bacteria, viruses, and fungi.

Coffey, B. & Wescott, G. (2010) New directions in biodiversity policy and governance? A critique of Victoria’s Land and Biodiversity White Paper. Australasian Journal of Environmental Management 17: 204-214. doi: 10.1080/14486563.2010.9725268.

Donfrancesco, V., Allen, B.L., Appleby, R., et al. (2023) Understanding conflict among experts working on controversial species: A case study on the Australian dingo. Conservation Science and Practice 5: e12900. doi: 10.1111/csp2.12900.

Ritchie, J., Skerrett, M. & Glasgow, A. (2023) Young people’s climate leadership in Aotearoa. Journal of Peace Education, 12-2023: 1-23. doi: 10.1080/17400201.2023.2289649.

Sengupta, A., Bhan, M., Bhatia, S., Joshi, A., Kuriakose, S. & Seshadri, K.S. (2024) Realizing “30 × 30” in India: The potential, the challenges, and the way forward. Conservation Letters 2024, e13004. doi: 10.1111/conl.13004.

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

On Ecology and Medicine

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As I grow older and interact more with doctors, it occurred to me that the two sciences of medicine and ecology have very much in common. That is probably not a very new idea, but it may be worth spending time on looking at the similarities and differences of these two areas of science that impinge on our lives. The key question for both is how do we sort out problems? Ecologists worry about population, community and ecosystem problems that have two distinguishing features. First, the problems are complex and the major finding of this generation of ecologists is to begin to understand and appreciate how complex they are. Second, the major problems that need solving to improve conservation and wildlife management are difficult to study with the classical tools of experimental, manipulative scientific methods. We do what we can to achieve scientific paradigms but there are many loose ends we can only wave our hands about. As an example, take any community or ecosystem under threat of global warming. If we heat up the oceans, many corals will die along with the many animals that depend on them. But not all corals will die, nor will all the fish and invertebrate species, and the ecologists is asked to predict what will happen to this ecosystem under global warming. We may well understand from rigorous laboratory research about temperature tolerances of corals, but to apply this to the real world of corals in oceans undergoing many chemical and physical changes we can only make some approximate guesses. We can argue adaptation, but we do not know the limits or the many possible directions of what we predict will happen.

Now consider the poor physician who must deal with only one species, Homo sapiens, and the many interacting organs in the body, the large number of possible diseases that can affect our well-being, the stresses and strains that we ourselves cause, and the physician must make a judgement of what to do to solve your particular problem. If you have a broken arm, it is simple thankfully. If you have severe headaches or dizziness, many different causes come into play. There is no need to go into details that we all appreciate, but the key point is that physicians must solve problems of health with judgements but typically with no ability to do the kinds of experimental work we can do with mice or rabbits in the laboratory. And the result is that the physician’s judgements may be wrong in some cases, leading possibly to lawyers arguing for damages, and one appreciates that once we leave the world of medical science and enter the world of lawyers, all hope for solutions is near impossible.

There is now some hope that artificial intelligence will solve many of these problems both in ecological science and in medicine, but this belief is based on the premise that we know everything, and the only problem is to find the solutions in some forgotten textbook or scientific paper that has escaped our memory as humans. To ask that artificial intelligence will solve these basic problems is problematic because AI depends on past knowledge and science solves problems by future research.

Everyone is in favour of personal good health, but alas not everyone favours good environmental science because money is involved. We live in a world where major problems with climate change have had solutions presented for more than 50 years, but little more than words are presented as the solutions rather than action. This highlights one of the main differences between medicine and ecology. Medical issues are immediate since we have active lives and a limited time span of life. Ecological issues are long-term and rarely present an immediate short-term solution. Setting aside protected areas is in the best cases a long-term solution to conservation issues, but money for field research is never long term and ecologists do not live forever. Success stories for endangered species often require 10-20 years or more before success can be achieved; research grants are typically presented as 3- or 5-year proposals. The time scale we face as ecologists is like that of climate scientists. In a world of immediate daily concerns in medicine as in ecology, long-term problems are easily lost to view.

There has been an explosion of papers in the last few years on artificial intelligence as a potentially key process to use for answering both ecological and medical questions (e.g. Buchelt et al. 2024, Christin, Hervet, and Lecomte, 2019, Desjardins-Proulx, Poisot, & Gravel, 2019). It remains to be seen exactly how AI will help us to answer complex questions in ecology and medicine. AI is very good in looking back, but will it be useful to solve our current and future problems? Perhaps we still need to continue training good experimental scientists in ecology and in medicine.  

Buchelt, A., Buchelt, A., Adrowitzer, A. & Holzinger, A. (2024) Exploring artificial intelligence for applications of drones in forest ecology and management. Forest Ecology and Management, 551, 121530. doi: 10.1016/j.foreco.2023.121530.

Christin, S., Hervet, É. & Lecomte, N. (2019) Applications for deep learning in ecology. Methods in Ecology and Evolution, 10, 1632-1644. doi: 10.1111/2041-210X.13256.

Desjardins-Proulx, P., Poisot, T. & Gravel, D. (2019) Artificial Intelligence for ecological and evolutionary synthesis. Frontiers in Ecology and Evolution, 7. doi: 10.3389/fevo.2019.00402.

The Ecological Outlook

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There is an extensive literature on ecological traps going back two decades (e.g. Schlaepfer et al. 2002, Kristan 2003, Battin 2004) discussing the consequences of particular species selecting a habitat for breeding that is now unsuitable. A good example is discussed in Lamb et al. (2017) for grizzly bears in southeastern British Columbia in areas of high human contact. The ecological trap hypothesis has for the most part been discussed in relation to species threatened by human developments with some examples of whole ecosystems and human disturbances (e.g. Lindenmayer and Taylor 2020). The concept of an ecological trap can be applied to the Whole Earth Ecosystem, as has been detailed in the IPCC 2022 reports and it is this global ecological trap that I wish to discuss.

The key question for ecologists concerned about global biodiversity is how much biodiversity will be left after the next century of human disturbances. The ecological outlook is grim as you can hear every day on the media. The global community of ecologists can ameliorate biodiversity loss but cannot stop it except on a very local scale. The ecological problem operates on a century time scale, just the same as the political and social change required to escape the global ecological trap. E.O. Wilson (2016) wrote passionately about our need to set aside half of the Earth for biodiversity. Alas, this was not to be. Dinerstein et al. (2019) reduced the target to 30% in the “30 by 30” initiative, subsequently endorsed by 100 countries by 2022. Although a noble political target, there is no scientific evidence that 30 by 30 will protect the world’s biodiversity. Saunders et al. (2023) determined that for North America only a small percentage of refugia (5– 14% in Mexico, 4–10% in Canada, and 2–6% in the USA) are currently protected under four possible warming scenarios ranging from +1.5⁰C to +4⁰C. And beyond +2⁰C refugia will be valuable only if they are at high latitudes and high elevations.

The problem as many people have stated is that we are marching into an ecological trap of the greatest dimension. A combination of global climate change and continually increasing human populations and impacts are the main driving factors, neither of which are under the control of the ecological community. What ecologists and conservationists can do is work on the social-political front to protect more areas and keep analysing the dynamics of declining species in local areas. We confront major political and social obstacles in conservation ecology, but we can increase our efforts to describe how organisms interact in natural ecosystems and how we can reduce undesirable declines in populations. All this requires much more monitoring of how ecosystems are changing on a local level and depends on how successful we can be as scientists to diagnose and solve the ecological components of ecosystem collapse.

As with all serious problems we advance by looking clearly into what we can do in the future based on what we have learned in the past. And we must recognize that these problems are multi-generational and will not be solved in any one person’s lifetime. So, as we continue to march into the ultimate ecological trap, we must rally to recognize the trap and use strong policies to reverse its adverse effects on biodiversity and ultimately to humans themselves. None of us can opt out of this challenge.

There is much need in this dilemma for good science, for good ecology, and for good education on what will reverse the continuing degradation of our planet Earth. Every bit counts. Every Greta Thunberg counts.

Battin, J. (2004) When good animals love bad habitats: ecological traps and the conservation of animal populations. Conservation Biology, 18, 1482-1491.

Dinerstein, E., Vynne, C., Sala, E., et al. (2019) A Global Deal For Nature: Guiding principles, milestones, and targets. Science Advances, 5, eaaw2869.doi: 10.1126/sciadv.aaw2869..

IPCC, 2022b. In: Skea, J., Shukla, P.R., et al. (Eds.), Climate Change 2022: Mitigation of Climate Change. Contribution of Working Group III to the Sixth Assessment Report of theIntergovernmental Panel on Climate Change. Cambridge University Press. doi: www.ipcc.ch/report/ar6/wg3/.

Kristan III, W.B. (2003) The role of habitat selection behavior in population dynamics: source–sink systems and ecological traps. Oikos, 103, 457-468.

Lamb, C.T., Mowat, G., McLellan, B.N., Nielsen, S.E. & Boutin, S. (2017) Forbidden fruit: human settlement and abundant fruit create an ecological trap for an apex omnivore. Journal of Animal Ecology, 86, 55-65. doi. 10.1111/1365-2656.12589.

Lindenmayer, D.B. and Taylor, C. (2020) New spatial analyses of Australian wildfires highlight the need for new fire, resource, and conservation policies. Proceedings of the National Academy of Sciences 117, 12481-124485. doi. 10.1073/pnas.2002269117.

Saunders, S.P., Grand, J., Bateman, B.L., Meek, M., Wilsey, C.B., Forstenhaeusler, N., Graham, E., Warren, R. & Price, J. (2023) Integrating climate-change refugia into 30 by 30 conservation planning in North America. Frontiers in Ecology and the Environment, 21, 77-84. doi. 10.1002/fee.2592.

Schlaepfer, M.A., Runge, M.C. & Sherman, P.W. (2002) Ecological and evolutionary traps. Trends in Ecology & Evolution, 17, 474-480.

Wilson, E.O. (2016) Half-Earth: Our Planet’s Fight for Life. Liveright, New York. ISBN: 978-1-63149-252-5.

Back to Nature vs. Nurture

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The ancient argument of ‘nature’ versus ‘nurture’ continues to arise in biology. The question has arisen very forcefully in a new book by James Tabery (Tabery 2023). The broad question he examines in this book is the conflict between ‘nature’ and ‘nurture’ in western medicine. In a broad sense ‘nature’ is discussed as the modern push in medicine to find the genetic basis of some of the common human degenerative diseases – Parkinson’s, dementia, asthma, diabetes, cancer, hypertension – to mention only a few medical problems of our day. The ‘nature’ approach to medicine in this book is represented by molecular genetics and the Human Genome Project. The ‘nurture’ approach to treating these medical conditions is via studying health outcomes in people subject to environmental contamination, atmospheric pollution, water quality, chemicals in food preparations, asbestos in buildings, and other environmental issues including how children are raised and educated. The competition over these two approaches was won very early by the Human Genome Project, and many of the resources for medicine over the last 30 years were put into molecular biology which made spectacular progress in diving into the genome of affected people and then making great promises of personalized medicine. The environmental approach to these medical conditions received much less money and was not viewed as sufficiently scientific. The irony of all this in retrospect is that the ‘nature’ or DNA school had no hypotheses about the problems being investigated but relied on the assumption that if we got enough molecular genetic data on thousands of people that something would jump out at us, and we would locate for example the gene(s) causing Parkinson’s, and then we could alter these genes with gene therapy or specific pharmaceuticals. By contrast the ‘nurture’ school had many specific hypotheses to test about air pollution and children’s health, about lead in municipal water supply and brain damage, and a host of very specific insights about how some of these health problems could be alleviated by legislation and changes in diet for example.

So, the question then becomes where are we today? The answer Tabery (2023) gives is that the ‘nature’ or molecular genetic “personalized medicine” approach has largely failed in achieving its goals despite the large amount of money invested because there is no single or small set of genes that cause specific diseases, but many genes that have complex interactions. In contrast, the ‘nurture’ school has made progress in identifying conditions that help decrease the occurrence of some of our common diseases, realizing that the problems are often difficult because they require changes in human behaviour like stopping smoking or improving diets.

All this discussion would possibly produce the simple conclusion that both “nature” and “nurture” are both involved in these complex human conditions. So, what could this medical discussion tell us about the condition of modern ecological science? I think two things perhaps. First, it is a general error to use science without hypotheses. Yet this is too often what ecologists do – gather a large amount of data that can be measured without too much prolonged effort and then try to make sense of it by applying hypotheses after the fact. And second, technology in ecology can be a benefit or a curse. Take, for example, the advances in vertebrate ecology that have come from the ability to describe the movements of individual animals in space. To have a map of hundreds of locations of an individual animal provides good natural history but does not address any specific hypothesis. Contrast this approach with that of Studd et al. (2021) and Shiratsuru et al. (2023) who use movement data to test important questions about kill rates of predators on different species of prey.

Many large-scale ecological approaches suffer from the same problem as the ‘nature’ paradigm – use ‘big science’ to measure many variables and then try to answer some important question for example about how climate change is affecting communities of plants and animals. Nagy et al. (2021) and Li et al. (2022) provide excellent examples of this approach. Schimel and Keller (2015) discuss what is needed to bring hypothesis testing to ‘big science’. Lindenmayer et al. (2018) discuss how conventional, question-driven long-term monitoring and hypothesis testing need to be combined with ‘big science’ to better ecological understanding. Pau et al. (2022) give a warning of how ‘big science’ data from airborne imaging can fail to agree with ground-based field studies in one core NEON grassland site in central USA.

The conclusion to date is that there is little integration in ecology of the equivalent of “nature” and “nurture” in medicine if in ecology we match ‘big science’ with ‘nature’ and field studies on the ground with ‘nurture’. Without that integration we risk in future another negative review in ecology like that provided now by Tabery (2023) for medical approaches to human diseases.

Lindenmayer, D.B., Likens, G.E. & 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.

Li, D., et al. (2022) Standardized NEON organismal data for biodiversity research. Ecosphere, 13, e4141.doi:10.1002/ecs2.4141.

Nagy, R.C., et al. (2021) Harnessing the NEON data revolution to advance open environmental science with a diverse and data-capable community. Ecosphere, 12, e03833.doi: 10.1002/ecs2.3833.

Pau, S., et al. (2022) Poor relationships between NEON Airborne Observation Platform data and field-based vegetation traits at a mesic grassland. Ecology, 103, e03590.doi: 10.1002/ecy.3590.

Schimel, D. & Keller, M. (2015) Big questions, big science: Meeting the challenges of global ecology. Oecologia, 177, 925-934.doi: 10.1007/s00442-015-3236-3.

Shiratsuru, S., Studd, E.K., Majchrzak, Y.N., Peers, M.J.L., Menzies, A.K., Derbyshire, R., Jung, T.S., Krebs, C.J., Murray, D.L., Boonstra, R. & Boutin, S. (2023) When death comes: Prey activity is not always predictive of diel mortality patterns and the risk of predation. Proceedings of the Royal Society B, 290, 20230661.doi.

Studd, E.K., Derbyshire, R.E., Menzies, A.K., Simms, J.F., Humphries, M.M., Murray, D.L. & Boutin, S. (2021) The Purr-fect Catch: Using accelerometers and audio recorders to document kill rates and hunting behaviour of a small prey specialist. Methods in Ecology and Evolution, 12, 1277-1287.doi. 10.1111/2041-210X.13605

Tabery, J. (2023) Tyranny of the Gene: Personalized Medicine and the Threat to Public Health. Knopf Doubleday Publishing Group, New York. 336 pp. ISBN: 9780525658207.

Belief vs. Evidence

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There is an interesting game you could enter into if you classified the statements you hear or read in the media or in ecological papers. The initial dichotomy is whether or not a statement is a BELIEF or EVIDENCE BASED. There is a continuum between these polar opposites so there can easily be disagreements based on a person’s background. If I say “I believe that the earth is round” you will recognize that this is not a simple belief but a physical fact that is evidence-based. Consequently we use the word ‘belief’ in many different ways. If I say that “Aliens from outer space are firing ray guns to cause flooding in California and Australia”, it is unlikely that you will be convinced because there is no evidence of how this process could work.

If we listen to the media or read the news, you will hear many statements that I or we ‘believe’ that speed limits on streets should be reduced, or that certain types of firearms should be prohibited. The natural response of a scientist to such statements is to ask for what evidence is available that such actions will solve problems, and if there is no evidence, we deal only with opinions or beliefs. If  you lived several hundred years ago, you would be told that “malaria” was a disease caused by “bad air” coming from swamps and rivers, since there was no evidence at the time about microorganisms causing disease. So in a broad sense historical progress was made by people looking for ‘evidence’ to temper and test ‘beliefs’.

How does all this relate to ecological science? I would add the requirement to papers that state some conclusions in ecology journals to also state the beliefs the paper rely on to reach its conclusions, in addition to stating clear hypotheses and alternative hypotheses. Consider the simple case of random sampling, a basic requirement in all statistical methods. But almost no paper states what statistical population is being sampled, and if it does often the study plots are not placed randomly. The standard excuse to this is that our results apply to a large biome, and it is not physically possible to sample randomly, or that we get the same results whether we sample randomly or not. Whatever the excuse, we need to recognize this as a belief or an assumption, a less damning scientific term. And if this assumption is not accepted it is possible to sample other areas or with other methods to test if the evidence validates the assumption. Evidence can always be improved with enough funding, and this replication is exactly what many scientists are doing daily.

Until recently most scientists believed that CO2 was good for plants, and so the more CO2 the better. But the evidence provided was based on simple theory and short term lab experiments. Reich et al. (2018) and Zhu et al. (2018) pointed out that this was not correct when long-term studies were done on C3 plants like rice. So this is a good illustration of the progress of science from belief to evidence. And over the past 50 years it has become very clear that increased CO2 increases atmospheric temperature with drastic climatic and biodiversity consequences (Ripple et al. 2021). The result of these scientific advances is that now there is an extensive amount of scientific research giving the empirical evidence of climate change and CO2 effects on plants and animals. Most people agree with these broad conclusions, but there are people in large corporations and governments around the world who deny these scientific conclusions because they believe that climate change is not happening and is of little consequence to biodiversity or to daily life.

It is quite possible to ignore all the scientific literature about the consequences of climate change, CO2 increase, and biodiversity loss but the end result of passing over these problems now will fall heavily onto your children and grandchildren. The biosphere is screaming the message that ignorance will not necessarily lead to bliss.

Reich, P.B., Hobbie, S.E., Lee, T.D. & Pastore, M.A. (2018) Unexpected reversal of C3 versus C4 grass response to elevated CO2 during a 20-year field experiment. Science, 360, 317-320.doi: 10.1126/science.aas9313.

Ripple, W.J., Wolf, C., Newsome, T.M., Gregg, J.W., Lenton, T.M., Palomo, I., Eikelboom, J.A.J., Law, B.E., Huq, S., Duffy, P.B. & Rockström, J. (2021) World Scientists’ Warning of a Climate Emergency 2021. BioScience, 71, 894-898.doi: 10.1093/biosci/biab079.

Shivanna, K.R. (2022) Climate change and its impact on biodiversity and human welfare. Proceedings of the Indian National Science Academy, 88, 160-171.doi: 10.1007/s43538-022-00073-6.

Watson, R., Kundzewicz, Z.W. & Borrell-Damián, L. (2022) Covid-19, and the climate change and biodiversity emergencies. Science of The Total Environment, 844, 157188.doi: 10.1016/j.scitotenv.2022.157188.

Williams, S.E., Williams, S.E. & de la Fuente, A. (2021) Long-term changes in populations of rainforest birds in the Australia Wet Tropics bioregion: A climate-driven biodiversity emergency. PLoS ONE, 16.doi: 10.1371/journal.pone.0254307.

Zhu, C., Kobayashi, K., Loladze, I., Zhu, J. & Jiang, Q. (2018) Carbon dioxide (CO2) levels this century will alter the protein, micronutrients, and vitamin content of rice grains with potential health consequences for the poorest rice-dependent countries. Science Advances, 4, eaaq1012 doi: 10.1126/sciadv.aaq1012.

Should Empirical Ecology be all Long-term?

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Alas Biodiversity

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

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

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

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

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

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

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

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

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

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

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

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

Is Ecology Becoming a Correlation Science?

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

On Climate Change Research Funding

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

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

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

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

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

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

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

On Ecological Climate Change Research

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

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

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

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

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

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

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

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

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

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