Revisiting Vellend 2010

In a paper published in The Quarterly Review of Biology in 2010, Mark Vellend proposed a new conceptual framework for Community Ecology, drawing inspiration from Population Genetics. Vellend proposed that, like in the case of genetic variation in populations, biological communities too can be thought of being governed by four main higher-level processes – selection, drift, speciation and dispersal. Organising our understanding around these processes, Vellend believed, would allow us to clarify the similarities and differences among the plethora of concepts, ideas and theories that have been developed in Community Ecology. Six years after the paper was published, I spoke to Vellend about his motivation for writing this paper and the book that followed from it.

Citation: Vellend, M. (2010). Conceptual synthesis in community ecology. The Quarterly Review of Biology, 85(2), 183-206.

Date of interview: 6th September 2016 (via Skype)


Hari Sridhar: What was your motivation to write this paper?

Mark Vellend: Yeah, so, during my PhD, which started about a decade earlier in 1999, I got quite interested in the parallels between population genetics and community ecology, in terms of the theoretical processes that are at work. And the first way in which that interest manifested was to look for correlated patterns of diversity – so species diversity and genetic diversity– because if the same attributes of habitat patches, like their size and heterogeneity and isolation, might influence one level of diversity, then if these parallels are really important, they should also influence the other level of diversity. And then slowly, I got more and more interested in, not necessarily putting the two bodies of theory together, but simply borrowing from one body of theory and using some of those concepts for the other. And population genetics is quite well known for having quite a coherent body of theory. All the models relate to one another in a pretty simple way. And community ecology is famous for having theories that are just all over the place; very, very difficult to relate one to the next. And so, slowly, I got interested in whether or not we might clean up theoretical community ecology using this structure. And so it built for a number of years, and then I thought I had a sort of complete synthetic framework to put forward. So that’s how it came to be.


HS: When, approximately, did you start sort of putting this together? Was that around 2010, when this was published, or earlier?

MV: It was definitely quite a bit earlier. I just need to turn around and check the date of something. There was this meeting and I have a poster of it on my wall. Give me one second. Okay, so in 2007, there was a symposium at Harvard University celebrating the 40th anniversary of the publication of The Theory of Island Biogeography. And I was invited to give a presentation there. And I’ve always been struck by very strong parallels between the theory of island biogeography and the mainland-island model for population genetics. And so, this particular theme became one component of my presentation there, and also an important component of a book chapter that was in a book that came out of that symposium. The book chapter was published in 2009. But the meeting itself was in 2007. And so, certainly in 2006, in 2007, that would have been the time when I started thinking seriously about doing this. And so, it was probably three to four years before that paper finally came out.


HS: What was the title of the book?

MV: Yeah, I’m going to turn around again and go to my shelf. Jonathan Losos and Bob Ricklefs were the editors. And it’s called something along the lines of ‘The Theory of Island Biogeography at 40’. No, it’s just called The Theory of Island Biogeography Revisited. Losos and Ricklefs were the editors. And anyway, there’s a whole bunch of chapters on different things, and my chapter was co-written with John Orrock, who was a post-doc at the National Center for Ecological Analysis and Synthesis, at the same time I was there. We worked on it together, and then afterwards, I developed things further on my own.


HS: I think you cite that chapter in this paper

MV: Yes, I certainly should have.


HS: How did you get interested in studying plant communities for your PhD? Were you interested in comparing and contrasting species diversity and genetic diversity even at that time?

MV: It developed early on in my PhD. I did not begin my PhD with that particular interest. I was really interested, and I’m still quite interested, in how the history of people’s use of landscapes can have a long-term impact on biodiversity and the composition of plant communities and other communities. So that’s what I started doing, and I did that for my PhD. But then I added this element of, not only how the plant community itself changes, but how genetic variation within species changes, particularly in the context of post-agricultural forests compared to forests that were never converted to agriculture. But I think I can look back further and find motivation. When I was an undergrad in the early 1990s, I worked as a field assistant on a project at McGill University and it was led by Graham Bell and Martin Lechowicz and Marcia Waterway. Graham Bell, for many years, has been talking about communities of different species, using very similar terms to the way you would describe interactions among genotypes in a population. So, you know, talking about selection among species, and random drift and so on. And at the time, I didn’t really think too much about it. But I think that principle stuck with me. And then, early in my PhD, there were several people in my cohort, by which I mean the group of us who started all at the same time, and several of those people were doing molecular genetics, population genetics. And so, right from the outset, I was frequently exposed to this; we had these discussions where it frequently came up. And so the two different fields were constantly present in these discussions we were having, and I think it flowed from there.

Field project team that Mark Vellend worked with as an undergraduate at McGill University. From left to right: Mark Vellend, Robin Naidoo (now senior scientist with WWF), Martin Lechowicz (professor at McGill University, now retired), Tanya Handa (now professor at Université du Québec à Montréal), Eliot McIntire (now Research Scientist with Natural Resources Canada) and Charles Mercier (current whereabouts unknown) © Mark Vellend

HS: How long did writing this paper take? Did you do it all at one go, or was this something that happened over an extended period of time?

MV: That’s a good question. In looking at old files, it looks like it took roughly six months – the second half of 2008 essentially – to produce the first version of a manuscript ready for submission. Although no doubt there were lots of hand-written notes and ideas and outlines that preceded that, and there was the prior book chapter that already contained a sketch of the core idea. As you know, it’s a long paper that covers a lot of ground, and I didn’t have a sabbatical of any kind, so I was working on it while I had all the other normal things I needed to do. So, from notes to manuscript it must have taken at least a year to write a draft of it.


HS: Had other people before you suggested organising community ecology in this fashion, or was what you were proposing completely new?

MV: Other people had definitely thought about this. The person who I think was the most explicit in noting these parallels and even describing them a little bit was Janis Antonovics, who is a plant ecological geneticist. He’s written down something that he called the ‘ecological geneticist’s creed’. And one of the points in that creed is that, and I’m going to paraphrase, the processes underlying species diversity and genetic diversity are essentially the same. And he didn’t develop that in great detail and pursue that very far, but, certainly, it had been noted. And other people – so Bob Holt, famous theoretician, and also Joan Roughgarden – have noted the same thing.


HS: I wanted to spend some time talking about the specific terms that you’ve used in, in the paper itself, for instance, the definitions you provide in a table. Were these definitions you came up with for the paper?

MV: Yeah, I’m just gonna pull up that table because it might help to actually look at it. I came up with them for the paper, but really they were for the most part, simply taking definitions from evolutionary biology or population genetics and inserting different words. So, when you talk about selection, it happens in population genetics when two different alleles confer differential fitness to the individuals that have them. And so then, at a community level, selection is simply when the average fitness of the individuals of two different species is different, in which case one of them will increase in relative abundance and the other one will decrease in relative abundance. And so they are really just slight modifications from what you would get in population genetics. But this paper definitely forced me to, in writing it all down, be more explicit and precise in making these definitions. So if you read Antonovics and others, they know the parallels, but then if you really want to take them further, then you have to start to be more precise in how you define things.


HS: Do you remember if you spent a lot of time thinking about the framing of these definitions and making sure you got it right?

MV: Definitely. I suppose I’ll leave to others the question of whether I got it right! But some of them were pretty simple. And then others…definitely, I think the trickiest ones are things like fitness and selection. The key element that I wanted to retain is that it’s still an individual-level phenomenon. So as soon as you talk about selection at the level of whole species, some people will immediately think about group selection- somehow, species are doing things for the good of the species. But that’s not really it at all. It’s the same way that different individuals in a population can share the same allele. Selection and fitness are still properties of the individual. And then, of course, we can calculate averages and variance and so on among the individuals, but it was important to make sure that it was still the individuals.


HS: In a figure in the paper you describe the black box of community ecology. Do you remember if you considered other terms instead of black box or was that the obvious term?

MV: Wow, I can’t really answer that question. I don’t remember whether I thought about different terms for that particular box. I can certainly tell you that, over the years – because I gave many presentations in which I presented versions of this paper, and they evolve over time – I’ve definitely sketched out many different ways in which you can communicate the basic point: that community ecology would benefit from being sort of cleaned up, But in that particular case, I don’t know if I had different terms for the box.


HS: Can we go over the list of people you have acknowledged, just to get a sense of how you knew them at that point and how they helped?

MV: Sure.


HS: J. Goheen

MV: So, there are several people that fall in the same category here. Jake Goheen and Chris Harley and Jon Shurin and Diane Srivastava were all community ecologists at the University of British Columbia where I was, at the time. And so we had taught a graduate seminar together and talked about these things. And so they were present when I was initially using this concept in a teaching context and helped with refining it and feedback and so on.


HS: R. Holt

MV: Bob Holt. Yeah, he and I had been in a working group or maybe two together and had cross paths in various places. And he, in different contexts, had presented this, and I can’t remember in what form I would have shared ideas, whether I sent him the manuscript, or whether we just talked about it over coffee or something, but he definitely helped in conversations, pinning down some of the parallels, and maybe where the parallels breakdown between disciplines.


HS: J. Levine

MV: Jonathan Levine. Yeah, when I was a post-doc in Santa Barbara – we weren’t at the same institution, but his university at the time – University of California, Santa Barbara- was nearby – I went frequently over to his lab meetings during that year; this was 2004-2005. And after that, we kept in touch, and especially when it comes to coexistence theory and negative frequency dependent selection, which is the key process underlying coexistence, various times we’ve had conversations where he helped me clarify my thinking on that.


HS: J. Losos

MV: Jonathan Losos. He invited me to give this presentation at the island biogeography meeting, which prompted me to do the initial sketch of these ideas, and he also was the editor on the very first paper I had on this topic in The American Naturalist -Island biogeography of genes and species. And so, at a couple of key points along the way, he provided important encouragement.


HS: J. Orrock

MV: John Orrock. We were post-docs together and then we worked on figuring out some ways in which models from population genetics, specifically concerning mutations, can speak to the question of species invasion. When a mutation arises in a population, what determines its success? It’s the same question as when a few individuals of a new species arise…anyway, he was a co-author on the initial sketch of these ideas


HS: R. Ricklefs

MV: Bob Ricklefs. He was a co-organizer of the meeting at which I first presented this. And then he had been at the University of British Columbia, visiting a couple times, I can’t remember if he read a draft of this, but certainly we had chances to discuss it.  And getting feedback from him was really important because he’s also a huge name, in terms of people who have put ecology and evolution together.


HS: J Roughgarden

MV: Joan Roughgarden. So we met at Stanford when I was there visiting. It was a short visit, but she, at the time, was working on something that had certain similarities. She’s published it now. I think it’s in a philosophy of biology journal, where she talks about similar things. But even though it wasn’t long, certainly being able to chat for a little about it was quite helpful.


HS: Do you remember when this was?

MV: My visit to Stanford?


HS: Yes

MV: I believe it was in 2007.


HS: M. Whitlock

MV: Mike Whitlock is a professor at the University of British Columbia. He’s a theoretical population geneticist. And he is also an extremely rigorous thinker. And so, as I was exploring some of these ideas, you get very enthusiastic about these parallels, and he definitely was helpful in pointing out when they broke down -a bitter dose of reality sometimes. But yeah, helpful in the theoretical population genetics


HS: A. Agrawal

MV: Anurag Agrawal. He invited the submission to The Quarterly Review of Biology. I’m not sure if I mentioned to him that it had been rejected or not. I can’t remember what the whole story was, but he basically invited the submission.


HS: J. Wiens

MV: John Wiens. He was the editor-in-chief of The Quarterly Review of Biology at the time, and provided quite a thorough review of the first submission of the paper, which helped me make some important improvements.


HS: And then you thank the ‘UBC community ecology class of 2008’

MV: That was with the other community ecologists that I mentioned. We ran this seminar in which each of us did, let’s say, a quarter of the semester. And all of the students in that class were basically among the first to be on the receiving end, as students of me teaching using this framework. And teaching is a wonderful way to figure out whether or not what you’re saying makes sense or not. And so they were too many to list one by one, but there was maybe a dozen students in that class.


HS: Was this a graduate class?

MV: Yeah, so it was Master’s, PhDs, maybe there’s a couple post-docs in there.


HS: And do remember if you incorporated feedback from the class into the ideas you were developing?

MV: Oh, for sure. I mean, you don’t keep track as these discussions go  – who said what – but certainly as you present it, often in the nature of the questions they pose, you can figure out what it is that you’re not explaining very clearly, or which concepts aren’t as closely parallel as you thought. It’d be impossible to say specifically, but definitely those interactions help you improve your thinking when you work on a synthetic paper like this.


HS: In your email, you mentioned that this was rejected by two journals. I wanted to ask you a little more about the path of this paper through peer-review and how difficult it was?

MV: Yeah, so when I was a graduate student, papers in The American Naturalist were often revered. I mean, many of the synthetic papers in American Naturalist have had a major and lasting impact. And to me this was, of anything I’d ever written, a perfect fit for The American Naturalist. It combined ecology and evolution, and it was synthetic. And so, I submitted it first to The American Naturalist, and it was read by three editors, and they didn’t send it for review. They felt that it didn’t really say anything that they hadn’t already thought about. And so, I can only speculate, but it made me wonder if sometimes the real experts on a topic are necessarily the best to evaluate whether something synthetic is going to be useful. I don’t doubt that they had thought about things that way, but until somebody writes it down, in a way that that can be broadly diffused, then just because it’s in somebody’s mind, doesn’t mean it shouldn’t be published. And so that was quite disappointing, I must say. And then I submitted a proposal to Ecology Letters, it must have been one of their Concepts & Syntheses papers. The proposal was accepted, which just means that they will allow you to submit a full manuscript, but then when I submitted the full manuscript…I can’t remember if I did a proposal or if I just went straight to the manuscript…either way, it was the same thing, where one, maybe a couple of editors, read it and they just didn’t think it was novel enough. Like the perspective wasn’t new; they’d also thought about things that way. The wording was very different, but the thrust of the rejections were more or less the same: these particular editors had already thought about things that way and so it didn’t strike them as pushing things forward enough, or novel enough.

In going through old files while checking this transcript, I actually realized that a proposal to Annual Review of Ecology, Evolution and Systematics was also rejected.  It was a few pages, not a whole manuscript, and I don’t have any record of the rejection itself, just a memory.  But I have the proposal itself and it was definitely submitted and rejected.  Based on the dates on files, it was before the other submissions.


HS: What happened at The Quarterly Review of Biology?

MV: It was based on invitation, although…I think, again…I really should have prepared more for this interview! But I can’t remember whether or not I had spoken with Anurag about the rejections. It didn’t come out of the blue, in the sense that he knew I was working on it. And maybe I’d thought about submitting there, because it’s a fairly long paper, and there aren’t that many journals that will publish longer papers. And so we had some interaction, and it generated an invitation, but the invitation didn’t come spontaneously. I probably put in some sort of inquiry, if you will. Sort of the same way it works at Ecology Letters, where you put in a proposal, even though it wouldn’t have been formal at Quarterly Review, and then if they think it has merit, they will proceed to invite a full manuscript. And then it went through normal review. And there were definitely critical comments and feedback of various sorts – useful ones. And that feedback, definitely, had a certain impact on the paper, although it’s essentially the same paper as it was to begin with, and eventually it was published.


HS: Did you change anything between the submissions to The American Naturalist/Ecology Letters and The Quarterly Review?

MV: The length limit at The Quarterly Review is quite a bit longer than the other journals. So, the paper has a big theoretical section, but it also has an empirical section, where it walks through empirical examples of each of these processes. The initial submissions had less of that, because I had to pare that down in order just to get it under the length limit. And so the empirical portion was definitely beefed up before submission to Quarterly Review, because they allow this longer format of articles, but otherwise, any of the changes would have been of a pretty minor nature.


HS: In hindsight, I’m wondering if, in a sense, if it was a good thing that this was published at Quarterly Review and not American Naturalist or Ecology Letters. It allowed you to flesh out the paper a lot more…

MV: Yeah, that’s definitely a possibility. Of course, all we can do is look back with our 20-20 hindsight and wonder. But certainly, in retrospect, maybe it was a good thing that I was forced, if you will, into a situation where I did have this longer format, because it could well be that the impact has been greater because it went into far more detail on the empirical side. Yeah, that’s totally possible. And it’s also possible that it took a little bit longer than it would have to get noticed by as many people because Quarterly Review doesn’t have the same readership as the other journals that I mentioned.


HS: How was this paper received when it was published? Did it immediately attract a lot of attention?

MV: Well, I actually sent it myself to a pretty long list of people that I knew in the field. So, if it had been published in The American Naturalist, I could have counted on everybody just seeing it. And whether or not they loved it or hated it, they would have at least known about it. Whereas, I know if other people are like me, that not as many people check the Table of Contents or get alerts from Quarterly Review. And so, I sent it to quite a number of people, and I think they shared it with their students. And for those people that I sent it to, they had no choice but to notice it. And I got quite a bit of feedback, from some people who really liked the perspective and even some who had picked it up in its shorter form from the book chapter, and were using it in their teaching. Because I think ultimately one of its utilities, one of its major contributions, is in helping people teach community ecology. And then, I definitely got some feedback that were along the lines of the kinds of things I heard from the editors of American Naturalist and Ecology Letters: “Sure, I thought about it that way, so not all that much new here.”And other people who were just thrilled because they’d never been able to figure out how things fit together and they thought that this really now helped them do that. It was a year or two later when I think a much broader audience started to either read it or notice it. Certainly, if you look at citations or just in talking to people, it seemed to me that it took a year or two before I frequently got comments from people when I’ve run into them at meetings about that particular paper.


HS: Do people still continue to talk about it when they meet you?

MV: Yeah, definitely. When I cross paths with people, it’s not uncommon that that particular paper is mentioned as one that they particularly value.


HS: It’s been cited a few hundred times already [> 1000 times at the time of publication of this interview], not to mention the impact it has had on teaching. At the time when you were working on this paper, and around the time it was published, did you anticipate at all that it would have such a big impact on the field? Did you think that this would sort of change the field in an important way?

MV: That’s a difficult question, because you sort of hope so, but from experience, all of us have a considerable degree of uncertainty when we put something out there. Just as one example, the first paper I published on this broad topic, it was in American Naturalist. It’s called Island biogeography of genes and species. Maybe I was just an overly enthusiastic grad student, but I thought, “Wow, this is incredible”, and everybody’s going to talk about it, and so on.  And then it was sort of well received, but it certainly didn’t change the field or have a huge impact. And so, in this case, I felt confident that it would prompt a lot of people to read it, because I was going to send it to them. And that it would prompt them to think about how they teach and how they view community ecology. But you also have major doubts that it’s gonna have the impact you hope. You see it both ways: you hope, but there’s lots of uncertainty as to the degree to which people will pick it up and enjoy it and find it useful.

HS: It’s only been six years since this paper was published, but have your views on the ideas presented in this paper evolved or do they remain, broadly, the same?

MV: I think it remains broadly the same. Certainly, feedback from certain people has got me thinking. And, as I wrote the book, which is essentially an expansion of this paper, when you hit certain topics, you really have to stop and think about how they fit together. So, one example is disturbance as a process. It’s a tricky one because it involves a change in community size. One of the classic definitions of disturbance is a reduction in biomass; basically, killing things. So you have a reduction in community size, which ought to influence drift, and then it’s clearly a force of selection. And so I would say that, to the extent that my thinking has changed, it’s been on particular topics and how they connect and how they fit, rather than on the big picture. With regard to the big picture: I still think the framework fits.


HS: Would you say the framework fits all theories and ideas that are usually listed under community ecology?

MV: Yes. It definitely fits any type of theory aiming to explain the diversity and composition of species that interact either on the same trophic level or via competition of some sort. It’s more difficult to weave in, for example, food web interactions, which most definitely falls under the broader umbrella of community ecology. And in one sense, there’s no difficulty actually in incorporating those, because it’s just an expansion of the items that are part of your community, or part of your population. So, instead of just having things that compete or facilitate one another, you have things that eat one another, but the same principles of selection – the different forms of selection – still apply. One of the big differences is that, for example in food web ecology, some of the patterns that they aim to explain are themselves processes. So they’ll have a pattern, which is something like ‘connectedness’, but those connections are feeding interactions. So it’s a type of pattern that is quite different than anything you’ll see in population genetics, or in what you can think of as horizontal community ecology. And so the fit is imperfect. And certainly I’ve had feedback from food web ecologists that they don’t feel like it helps them a whole lot, which is a fair criticism.


HS: Can you tell us a little more about how the idea of the book came about?

MV: Certainly. Not long after the paper was published, some of the people who were particularly enthusiastic about it, suggested, almost right away, that it should become a book. So the seed of the idea was planted very shortly after the paper came out. And from talking to other people who’ve written books, I realised it’s quite an investment of time and energy. I knew that you’d have to have a sabbatical break or some sort of break from your regular tasks to be able to complete a book like that. So I didn’t really think about it too much for a while, but then when my sabbatical was coming up, which was in 2014-15, then I had an opportunity to do it. There are different options for how you can take advantage of that time on sabbatical. I was really on the fence for quite some time. I was not at all convinced that I wanted to spend so much time and energy doing one particular project. And then something, I couldn’t even say what exactly, just sort of tipped me over the edge to saying, you know, this should be done. And I guess part of it is just that, in my own experience, or my own impression is that books can have a longer lasting impact than anyone paper can. And I thought, “Well, here’s my chance maybe to have a broader and maybe more long lasting impact.”

Book writing (2015) © Mark Vellend

HS: And if you don’t mind sharing this with us, can you tell us who the people were who pushed you to write the book?

MV: Oh, I mean, one that comes to mind is Diego Vázquez, who’s an ecologist in Argentina; he definitely mentioned that.  I remember early on, Kyle Harms, who’s a forest ecologist, mentioned he was using it in teaching. I don’t think he ever said I should write a book. But that kind of comment gets you thinking about that. And students who found the framework useful said similar things. But otherwise, I couldn’t say exactly who mentioned those things.


HS: In the paper you provide “a very general theory of community ecology: species are added to communities via speciation and dispersal, and the relative abundances of these species are then shaped by drift and selection, as well as ongoing dispersal, to drive community dynamics.”Do you remember how you came up with this, and whether it went through many iterations?

MV: Again, I couldn’t say really; it’s been quite some time. But once you have all the elements in place – the definitions in that table and the basic framework– really, those sentences that you readjust summarize the core idea. And whenever you do that, it definitely takes some effort to find the right words to say it in just a few sentences. But whether or not it took a really long time, I couldn’t say. Writing is a funny endeavour because, sometimes it just takes a really long time to get the words out just the way you want them, and other times they just sort of come out, and you look at it and you think, “Wow, I have no better way to say it than that.” So there’s a major element of stochasticity in how much time it can take to do that.


HS: Apart from the book did this paper have any other sort of direct influence on your career and the course that your research took?

MV: Yeah. I’ll answer the two parts of your question separately. I think it probably did influence my career because – I’ll just give you one specific example -an award I got last year from the Canadian government, which is quite a major award. The proposal I put in didn’t really have to do with that paper, but they take quite seriously reviews from external reviewers. I’ll never know who they were, but they find well-known people in the field to evaluate your application. And several of those specifically mentioned that paper as a major contribution that, in their opinion, spoke to the worthiness of my candidacy for this award. So I’ll guess that things like that, definitely, have been influenced by that paper, and maybe your standing in the field, if you will. My own research – that’s sort of interesting. It hasn’t really influenced my own research all that much. Perhaps, to some degree, the way that I now discuss or describe the processes that are acting in the communities that I study. I’m not sure that I study them in any different way because of that perspective. And, sort of interestingly, that almost feeds back to the initial reasons for rejection: the editors maybe didn’t see any way in which it would alter the way they did their research. Maybe that’s true! Maybe it alters the way we think about things but the impacts on research might take longer to manifest, and might be more subtle. But in the immediate term, perhaps it doesn’t have a huge impact on me; maybe it does on others


HS: I wanted to ask you about others. When you look at empirical papers that cite this particular paper, do you see your framework and ideas reflected in the way research was done?

MV: I see it reflected in the way that people describe and interpret their results. It might not change the way they go about generating data. I think the most specific example I can give is microbial ecologists.  I did not anticipate this, but many of them quite enthusiastically adopted this framework to describe the processes that influence microbial diversity. And part of the reason, I think, is that microbial biologists have, for a very long time, worked on evolutionary questions. And the kinds of data they generate – using sequencing, for example, they have huge amounts of data on the diversity and composition of communities over space and time. But they don’t know a whole lot about the natural history of each organism. And so they’re often unable to really have discussions about predation and competition and disturbance, in very specific terms, which are all what I call low-level processes, but they can certainly see and test for signatures of what I call higher-level processes, like selection and drift and dispersal. So you definitely see it reflected in the way the discussion and introduction sections of those kinds of studies are written.


HS: What is the name of the award that you got?

MV: Oh, it’s called a Steacie fellowship. That was the name of a well-known chemist: Edgar William Richard Steacie. EWR were his initials, so it’s the EWR Steacie Memorial fellowship.


HS: Another point you make in the paper is that, while there’s empirical evidence for all four higher-level processes, most of the emphasis has been on studies of selection. Do you see that balance shifting more recently, where people are paying more attention to the other processes as well?

MV: I think there’s definitely been a shift in the field, but I don’t think it was sparked by my paper. Hubbell’s book on Neutral Theory was definitely the spark that got people thinking more about drift and speciation and dispersal, to some degree. And then, as the term meta-community became widespread, that was a major impetus to get people studying dispersal in more detail. So, I don’t think my paper was the reason behind that at all. I would actually go as far to say that my paper is almost like a response to that. So you have neutral theory pushing people over there, meta-community theory pushing people over there, classic niche theory pushing people over there, and it’s quite difficult for a student to see how all of those things relate to one another. My goal was to do that. To see how do those things all fit together. And in the end, they fit together via a common framework that really isn’t that complicated. It’s not as complicated as we’ve typically made it out to be.


HS: Do you have a sense of how widely the framework you propose is being used in community ecology courses, at least in the US? Have teachers started framing their community ecology courses based on this?

MV: Oh, yes. I mean, I couldn’t say how many, but certainly, I’ve had multiple people tell me that they use this framework. Whether it’s for a whole course or whether it’s for the community ecology section, for example, of a broader ecology course, I don’t always know or remember. But my sense is that there are there are quite a few people out there that use the framework in teaching community ecology, both at the undergrad level and at the grad level. I think it’s made it onto quite a few reading lists, where people hand their students a list of 10 and 20 papers to get them started on figuring out what community ecology is all about.  I think it’s made it onto quite a few lists of that nature.


HS: What about in your own teaching? Is it a paper that you discuss in class?

MV: It is. So the paper itself is an optional read, but the concepts are front and centre in the lectures that I give. And then some simulations that I’ve developed in the book originate from simulations I’ve been using in class for quite some time. For example, using some very simple computer code in R, students can see how selection and drift interact; and then they see how dispersal interacts. And I think it’s been really effective because they can change parameters themselves – e.g., let’s just have a slightly higher proportion of individuals moving between these two habitat patches and see how that influences diversity and the competitive balance in those communities. And so, I definitely use it in teaching myself, but the paper itself maybe hard going for early level undergrads.

Isla Myers-Smith‘s copy of the book after field-use © Cameron Cosgrove

HS: Have you ever read the paper after it was published?

MV: Oh, you know what, it’s funny, because this conversation has made me think that it’s time that I read it again. Certainly, within the year or two after it was published, if people were sending feedback or had questions, I would have been prompted to go through it again. But it’s been several years certainly since I would have read through the paper itself. Well, no, I must have read through it when I started writing the book actually, but books take a while. So that would have been two years ago. I probably haven’t read the whole paper – beginning to end – in at least a couple years.


HS: Would you count this paper as a favourite among all the papers you have published?

MV: Definitely.  There are different criteria for judging papers, but originality, if that’s a criterion, it’s certainly one of the more original things that I’ve contributed.


HS: What would you say to a student who is about to read this paper today? What should he or she takeaway from this, six years after the papers published? And would you add any caveats they should keep in mind when reading this?

MV: I have just published the book in which I go into quite a bit more detail. For example, there were things that were overlooked and needed more attention; they have now received more attention. I guess my answer is that I would not want to direct a student’s thinking before they read the paper or book. I would not want to suggest what they should take away from it. But I want to know what they take away from it, and then we can perhaps have a discussion that prompts them to see things from additional perspectives. But this is what I’ve put down, and they should read it using whatever lens they have on the field. And I’d love to know what they take away from it. But that’s up to them.



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