Botkin Chapter 3: The Wolves and Moose of Isle Royale

Sharon is busy with schoolwork this week and has asked me to pinch-hit on the virtual pub book discussion blog.  If this is your first visit to this blog, the best place to start — and to introduce yourself — is:

Rather than provide a summary of Chapter 3, as I did with Chapter 2, I am reprinting a recent post by Dr. Ralph Maughan, an expert on wolves and a strong proponent of their reintroduction in former habitats: The reason for doing this is that Maughan is commenting on the very focus of Botkin’s 3rd chapter, which starts with the story of his (Botkin’s) experiences researching the predator/prey relationships between moose and wolves on Isle Royale, an island in Lake Superior near the Canadian border that is over 200 square miles in size and contains 45 lakes of its own. What has made the island particularly interesting for this type of study is that it has never been heavily influenced by people; moose first arrived there from the mainland only about 100 years ago; and wolves didn’t arrive (except for a failed National Park Service attempt to establish them with zoo animals in the late 1940s) until the lake froze over more than 50 years later — after the moose had a half-century to build their population without a major predator to inhibit their reproduction.

As with Chapter 2, Botkin uses this story and others — sandhill cranes, Canadian lynx and microbes — to examine the “balance of nature” as it has been defined mathematically and as actually observed, to examine the differences between the two. Both writers cite the work of long-time Isle Royale researcher and wildlife ecologist, Rolf Peterson, but they seem to come to different conclusions as to why that work is important:

I would like to thank Dr. Maughan for permitting me to repost his work here. Of the 72 comments on this topic on his blog, a significant number were my own in sometimes heated response to many of his regular commenters. Tree and Matthew will understand. It was my second visit there, and both times I seemed to stir up the natives with my thoughts and opinions — mostly because of the old “anonymous commenter” discussion. I think JZ first sent me there and after I went and got a similar reaction, he said he was just joking. Maybe it was Derek, but you will see the result if you read the comments, too.

I will leave it up to actual readers to determine their own thoughts on these two perspectives, but I’ll repeat Botkin’s analogy of “resilient stability” in regards to the “balance on nature” argument by his comparison of a person with a drink building a house of cards on a train: the house of cards will collapse with a bump or large vibration, but the drink will only slosh around before it returns to its former level. One is a fragile balancing act, and one is resilient. For me at least, that provides a clear metaphor for this discussion. Now for Maughan’s post:

Should the declining inbred wolves of Isle Royale N.P. be augmented?

Should there be genetic rescue (outside wolves brought in)-

For many years the wolves and moose of Isle Royale National Park in Lake Superior have shown that wolves do not wipe out their prey. When wolves become abundant enough that the disappearance of prey seems probable, the wolves die back.

On the other hand, when wolves have declined to few in number, the moose population expands and begins to decimate its prey — the moose-edible vegetation of the island.

This rough balance has existed ever since wolves colonized the island one hard winter. In 1949 a pair of wolves walked over to the island on the frozen lake. The pair found an island overrun with moose. The moose themselves had migrated to the island 40 years earlier.

The wolf population expanded, of course, and brought the moose number in check (and more). Then the wolves began to starve off and the cycle began.

The moose prefer aspen, and they do well eating it. However, they mostly wiped that out before the wolves came.  Ever since, they have relied primarily on the less nutritious balsam fir and lichens.

Both the moose and the wolves are also subject to inbreeding. It is especially a problem for the wolves, all of which descended from the original pair. So, in addition to the cyclic malnutrition when the moose population drops too low, the wolves have been seen to suffer from increasing genetic defects. One of these is poor reproduction even when there is enough food.

Down to just 8 wolves, they seem doomed without outside genes from new wolves. There have been up to 50 wolves at a time on the island, although many scientists think a stable number is about 25. It should be noted that there have always been wide fluctuations around this “mean.” The eight wolves seem to have gained a brief reprieve with the birth of 2 or 3 pups in 2013 after several years with none. Nevertheless, it is hard to see how the unaugmented population can survive much longer. It is less and less likely that the lake will freeze and wolves from Minnesota, Michigan or Wisconsin find their way to the island.

The wolves and their relationship to the moose and the vegetation have been studied since 1958. Dr. Rolf Peterson, in particular, is the person most closely associated with the studies. He would like to see some genetic rescue. Dr. Dave Mech, however, who is another avid student of the island’s wolves is reported to want to first let natural events play out.

With the wolf population so low, we would now expect the moose population to be expanding. It is. However, it is increasingly suffering from tick infestation. This is a problem for moose in general during winters, but Isle Royale has seen warmer winters as the climate changes. This makes the effects of the bloodsucking  arachnids more severe.

Rolf Peterson recently sent out the following letter.

The National Park Service is interested to receive your input on the pending decision regarding the future management of wolves on Isle Royale.  Please send your input to the following email address: (note the “underscore” between ISRO and Wildlife)

The Park Service is considering three options:  (1) do nothing, even if wolves go extinct; (2) allow wolves to go extinct (if that is what they do), and then introduce a new wolf population; or (3) conserve Isle Royale wolves with an action known as genetic rescue by bringing some wolves to the island to mitigate inbreeding.

While expressing your view, consider providing as much detail on the reasons for your preference, as the Park Service believes the reasons for your view are as important as your view.  If you have any questions on the process or anything relating to providing input, please do not hesitate to ask me.

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Botkin Chapter 2: Why the Elephants Died

Sharon is busy with schoolwork this week and has asked me to pinch-hit on the virtual pub book discussion blog (my choices are Kat Anderson or Hugh Raup for book #2). If this is your first visit here, the best place to start — and to introduce yourself — is:

The focus of this chapter is the disparity between computerized model predictions of wildlife populations over time, their actual populations, and why these numbers are usually so different from one another. This is an important distinction because most of our fish and game management objectives are based on the former, inaccurate, “balance of nature” computerized numbers, as are many of our Endangered Species population estimates. Botkin also provides a brief parallel history of ecology as a science, which concludes with one of my favorite quotes in the whole book:

“Many believe that ecology is still a “young” science, but in comparison to most modern sciences, it is not young but simply retarded.”

This chapter examines the truth to that statement by comparing elephants in Africa, anchovies in Peru, salmon in the Pacific Northwest (where I had the pleasure of working under Botkin and participating in the first of his studies of that animal), and whales in the ocean, with fruit flies in a jar. Botkin’s examples and thoughts are clearly presented and described in well written English, with little use of Latin, metrics, or acronyms; i.e., “Plain English.” As a result, almost anyone with a basic education and good reasoning skills can follow his logic, arguments and conclusions.

The chapter opens with the story of one of the world’s first protected wildlife populations, the elephants of Tsavo; a large 5,000-square mile national park in Kenya, Africa dedicated to the survival of African big game animals. The park was created in 1948, 65 years ago, largely through the efforts of a single man, David Sheldrick, for the protection of declining African elephant and rhinoceros populations from their principal predators: human ivory and meat hunters. The primary purpose for protecting these animals was to attract tourists to the park in order to view them. Within 10 years the elephant herd had increased to 36,000 animals and the landscape had become largely denuded of vegetation. By the mid-1960s it was decided that 3,000 of the animals needed to be shot, in order to preserve the habitat. This idea was overturned and the decision was made to “let nature take her course” and allow the elephants and vegetation to achieve a “naturally balanced” “carrying capacity.” A prolonged drought in 1969-1970 contributed to the destruction of most of the remaining vegetation and an estimated 6,000 elephants starved to death.

Botkin uses this story to illustrate the difference between a theoretical balance of nature, and a balance created by people; i.e., roughly the difference between shooting 3,000 elephants and letting 6,000 elephants starve to death. A third alternative is also considered – that Tsavo was simply too small to contain that many animals and that they needed to migrate from one area to another during times of drought or other stressors. Following Sheldrick’s death in 1977, poachers again entered the preserve and by the 1980s the herd had been reduced to 6,000 animals. Today it stands at about 12,000 – far less than the 36,000 that had once lived there under Sheldrick’s management practices.

The elephants of Tsavo are used as a beginning point to examine other human attempts to manage the environment to achieve a desired number of animals. The wildly fluctuating populations of Peruvian anchovies, Pacific sardines, Atlantic menhadens, and several other commercial fisheries are provided as examples where harvest levels were established in attempts to stabilize populations, and failed; typically resulting in abrupt declines in the targeted species. These failed attempts at controlling natural populations of desired fisheries were based on scientific models. This is an important consideration because much of the world’s food supply – particularly in poorer countries – is provided by fish.

This is the principal theme of Chapter 2: the consistent failure of scientific predictive models to accurately estimate wild animal populations, and the reason that Botkin concludes ecology is “retarded” when compared to other modern sciences. He begins in 1838 with Pierre-Francois Verhulst’s simulation of natural populations with the invention of the S-shaped logistic growth curve, which results in a conceptual “carrying capacity” for the environment. Laboratory experiments in the 20th century replicated this model with certain insects and with bacteria, thereby seeming to prove its utility for wild fish and elephants. Alfred Lotka, an early mathematical ecologist, used fruit flies, bananas, and aquariums to fine-tune this equation, and was able to maintain stable populations of these animals in controlled environments. This artificially regulated number of insects was termed a “density-dependent” population, Lotka’s equation was named the “logistic” model, and Botkin cites a paper written in 2010 that examines this potential phenomenon in regards to wild elephants.

As Botkin next explains that, although the logistic equation is considered an “ecological formula,” its mechanical basis can be compared to “a collection of identical colliding balls” with “a certain rate of destruction” and “capable of identical rates of division.” In using this equation to consider a herd of elephants there is no differentiation between bulls, calves, or breeding cows, for example, just a total number, as with the box of identical balls. This idealized balance of population numbers cannot (“has never been observed to”) occur in nature, of course, and Botkin describes the logistic equation as “something from [Lotka’s] imagination, not from actual observation” – as occurred with the world fisheries or the Tsavo elephants.

Following the widespread adoption of Lotka’s work in the field of ecology, Botkin goes on to describe how it has evolved into a simple calculation that is exactly ½ as large as predicted carrying capacity: the “maximum-sustained-yield” population. To complicate the picture further is the fact that it is impossible to accurately measure many wild populations in the first place – Botkin uses Arctic crabs and the television show Deadliest Catch as his example, and comes to the conclusion that the maximum-sustained-yield concept is “fundamentally flawed.”

The Marine Mammal Act of 1972 tried to overcome this broken model with a new concept: “the optimum sustainable population,” and Botkin was hired to help develop this idea. His approach was overturned, however, and a panel of University of Washington scientists recommended a return to the failed logistic model and reinstituted the ideas of “maximum productivity” and “carrying capacity.” This led to Botkin’s conclusion that the scientists had reverted to a belief in the disproved myth of the “balance of nature,” and subsequently led to his book Discordant Harmonies, and ultimately to this present work:

“Thus even today, in both law and in practice, the scientific conservation of endangered marine species continues to be based on the idea that nature undisturbed is constant and stable . . .”

In other words, the management of endangered whales is based on computerized mathematical formulas developed with fruit flies in an aquarium, and not on actual observations by people:

“An irony is that it seems that everybody talks about how complex nature is . . . but we are content to formalize nature in about as simple and simplistic a way as possible.”

2 Questions:

Botkin argues that it is important that people – and particularly scientists – must accept the contradictions between fact (“observations”) and theory (“computer models”). Agree or disagree?

He further argues that this acceptance will lead to a “deeper level of thought” and allow us to find a “true harmony of nature.” Does that even sound plausible or necessary?

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Botkin Chapter 1: The View from the Marsh

Since Botkin did a great job in the introduction of summarizing the whole book, I thought we could go chapter by chapter through the book, and end up with the introduction to see if we missed anything.

So here are some tickler questions:

1) Is there something you strongly agree/disagree with?

2) Is there something that surprised you?

3) Does one of Dan’s stories remind you of a story of your own?

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Botkin Introduction: O’Neill’s Critique of the Ecosystem Idea

Dr. O'Neill

Dr. O’Neill

As I read it, the key question in Botkin’s book is found on page xii on my Kindle..

“Whatever the scientist’s knowledge of the dynamic, changing properties of nature, the formal representations of these remove such considerations in most cases…whether or not environmental scientists know about geological time and evolutionary biology, their policies ignore them. It is strange, ironic and contradictory.”

My feeling is that the locus of these contradictions may lie in the competitive forces among scientific disciplines and ideas, competing for power and funding. When I was a graduate student at Yale, I shared an office with the Hubbard Brookians..they had a way of looking at the world measuring N P and K and applying systems thinking. But there were others around who simply observed or did experiments in manipulating organisms. At the time, these were all equally legitimate approaches to studying Nature- in our mind, if not those of the Big Funding Agencies, such as NSF. Since I had taken History of Science, I understood their (the BFA’s) peculiar attraction to the Mathematical and Abstract; and understood that the rest of us were little more than flower collectors in the hierarchy of Science. And that was OK with us, because our science was more of an interplay or conversation with Nature.. in population genetics we looked for mathematical equilibria but never found them. Ideas, measure, more ideas, more measurement, that was our conversation with Nature. We used math to explore ideas, but we did not have external ideas that we tested Nature against.

I do think the term “ecosystem” can be helpful; if we had talked about “prairies” in the past , the “prairie ecosystem” is handy because it denotes all the critters, plant, animal, insects, fungus, bacteria, viruses, water, soil- all of which we had studied before. We had studied them and their interactions with the environment, but at the time we might call ourselves “wildlife or fish biologists” “plant physiologists” “soil scientists” “entomologists” or “silviculturists”(applied vegetation ecologists). We even had a course at Yale called “genecology” but what else would creatures adapt to other than the environment? My point is that we were all studying things and their relation to the environment, which would make us all “ecologists” I guess. Except that our language was not about equilibria, attractors, functions, etc. These are all abstractions that came from systems theory. It’s legitimate, I think to question how helpful these abstractions have been and continue to be.

But don’t believe me. Check out this 2001 piece.. Is It Time to Bury the Ecosystem Concept? (With full military honors, of course) by Robrt V. O’Neill. The ideas he raised in 2001 are as current as the “ecosystem integrity” requirement in the 2012 Planning Rule.

The term ecosystem was coined by Tansley in 1935. But as Botkin (1990) points out, the underlying concept goes back at least to Marsh (1864). Nature was viewed as relatively constant in the face of change and repaired
itself when disrupted, returning to its previous balanced state. Clements (1905, 1916) and Elton (1930) offered plant and animal succession as basic processes that permitted relative constancy by repairing damage.
Forbes (1925) described the northern lake as a microcosm, a relatively closed, self-regulating system, an archetypic ecosystem.

Science emerged from the Second World War with a new paradigm, Systems Analysis (e.g., Bode 1945), which seemed uniquely suited for this ‘‘balance of nature’’ concept, and fit well with earlier work on the stability of interacting populations (Nicholson and Bailey 1935). Systems Analysis dealt with complex systems as interconnected components with feedback
loops (Hutchinson 1948) that stabilized the system at a relatively constant equilibrium point. Systems Analysis can be seen underlying E. P. Odum’s (1953) definition of the ecosystem as a ‘‘. . . natural unit that
includes living and nonliving parts interacting to pro duce a stable system in which the exchange of materials between the living and nonliving parts follows circular paths . . . .’’

The machine analogy, inherent in Systems Analysis, became a central paradigm for many ecologists (Odum 1971, Holling 1973, Waide and Webster 1976). The paradigm offered a practical approach to the enormous
complexity of natural systems (Teal 1962, Van Dyne 1969). The paradigm helped harness the power of the computer in ecosystem models (Olson 1963). The paradigm permitted a holistic view of system properties
such as nutrient cycling (Webster et al. 1974). The familiarity of the machine analogy facilitated the communication of ecological concepts to the public.
If the ecosystem concept has held such a central place in ecology and been so productive of new ideas, why call it into question? The simple fact is that the ecosystem is not an a posteriori, empirical observation
about nature. The ecosystem concept is a paradigm (sensu Kuhn 1962), an a priori intellectual structure, a specific way of looking at nature. The paradigm emphasizes and focuses on some properties of nature, while ignoring and de-emphasizing others. After a half century of application, the paradigm is showing some rust. Limitations in the concept are becoming more apparent and leading to a vigorous backlash toward ecosystem concepts in particular, and ecology in general.

One more story. When I was working for the FS in R&D we did a review of one of the Research Stations. One of the administrators there was a big aficionado of systems thinking. One of the scientists had done this fascinating study (to me) of how fish move around in streams and discovered something very useful that hadn’t been known before. I thought it was great work. This administrator, though, felt that “organismal biology is passe, it’s all about systems, now.” In arguing that this scientist should be better appreciated, I stated “you can’t understand systems without understanding their components” to which he replied “oh, yes you can.. it’s about flows among boxes and you don’t need to understand what’s in them.”

So you may say “this guy was off the wall, and not in the mainstream.” Well, that could be true. Still the reason I’m telling the story is that to point out that people can cross the line from systems being 1) one way of conceiving of how nature works, to 2) the best way of conceiving how nature works, to 3) how nature works. And somewhere along that line, the empiricism that “science” claims as its basis for legitimacy gets left behind.

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Botkin’s Title: Where Did It Come From?

moon and naut shell

This video explains it.

Video | Posted on by | Leave a comment

Forest Policy Pub Book Club: Introductions


To be honest, I’ve never been in a book club before. So this is an adventure. I thought we should start by introducing ourselves and telling our stories or, at least, some stories. One of the great things about Dan’s book is the stories he tells, and what a great story-teller he is. When you tell stories, say Dan’s triggers yours and yours triggers someone else’s and so on. This is a different kind of thing, and much more right-brained, than what we usually do on the blog. Not that we won’t challenge Dan’s knowledge claims, nor each other’s, through this process. But today let’s take a break from all that and listen to each other’s stories.

Why are stories important? This came across my email a while back from Ronna Detrick, who, I think, said it beautifully.

We live in a world of stories. Childhood fairytales shape our dreams and hopes. Family legends, imparted over kitchen table conversation, at reunions, and during road-trips, build our memory and craft our beliefs. Historical narratives inform our understanding of culture, politics, our larger world. Film, music, literature, and poetry mysteriously and continuously speak to our deepest heart – communicating truths we implicitly know and others we long to grasp.

Stories serve the way in which we are able to make sense of our world, our relationships, our behaviors, everything. They are how we speak of our circumstances, our deepest emotions, and our biggest questions; how we create and apply meaning. And they connect us to one another, bridging differences in language and perspective, time and place, past and future.

Most of us acknowledge that it’s less about a particular story and more about story, itself. It is the device, the vehicle, the means through which we express, listen, and even participate in our own life and others’. We admit (and even enjoy) that most stories, when told over and over again, not only shift and morph over time, but take on a life of their own.

“The fish gets a little bigger, the storm gets a little wilder, the love gets a little stronger, our bravery or disappointment gets a little exaggerated in the telling over time. There is creative tension in story. When we hear it, when we read it, when we speak it, when we write it, we filter words through our own experiences and our need for meaning. We shape the tale to reinforce our understanding of how life is. ~ “Christina Baldwin

This is what we love about them. This is why we tell them. This is why we live our lives within them. This is the power of story.

So, in the form of introduction, please say something about yourself and tell us one story about your relationship to the ideas of the balance of nature or systems ecology, and how they developed.

I’ll do a brief example:
I am a forester and plant geneticist/evolutionary biologist; I worked for the Forest Service for 32 years; I am running for Vice President of the Society of American Foresters; and I am a part-time theology student at Iliff.

When I was a freshman at Yale, I took two courses that were key to my future in forestry, and to my understanding of Nature. One was with Alison Richard, called “Primate Population Adaptations”. Another was “Man and the Environment” (no, I am not kidding; this was 1973 and we were one of the first classes after coeducation at Yale College). As I recall, Dr. Richard was the first, last and only female professor I had for the rest of my college career, which ended with a Ph.D. in Genetics in 1982. This experience (and reading gender studies of science) helped me understand the difference between the aspirations of “objective science” and the down and dirty reality of how it’s produced.

If your first framing of empirical, observational science is adaptation and evolution, as in the Primate class, the idea of a steady state is .. well..very odd. Evolution is change. Equilibrium is “not change” or change such that the results are still somehow “contained” in some abstract sense.

Probably the most important class I took in my college career was the next year at UCLA (I had run out of money so had to stay home and work full-time). It was called “History of American Science” and introduced me to the historical conceptions of science.. Like “applied science” was looked down upon because the upper classes focused on “basic” and could afford to study things without direct outcomes. Or the very real, continuing idea of Physics Envy. Having this class early, before most of my scientific training, helped me understand why different disciplines had more power and funding than others. I was able to watch funding for “science fads” flow and ebb across the forest science community and look at how the community and its different populations (disciplines) competed and evolved.

It is odd to me that history and philosophy of science are not required classes for trainee scientists. Also, remember that in those days “environmental science” was not separate. There was just “science,” and you applied it to whatever issue you had.

Much later one of my colleagues said to me “you’re not a conservation geneticist, you’re an exploitation geneticist!” because she disagreed with my ideas of What Should Be Done. But the key historical fact I’m trying to focus on here was that in the 70’s, there was just plain old genetics, silviculture, physiology, entomology, pathology, ecology, wildlife biology, range ecology, hydrology, fish biology, etc. At least that’s my memory.

Tell us a little about yourself, and what’s your story?

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