Congratulations, Alberta! Uh, I think….

Well, the CBC has called it:  Alberta has elected a Progressive Conservative government.

I never thought that I would be vaguely happy to hear that my home province has elected a Tory government;  they’ve been in power for over 40 years now and I would have been happy to see a change, but I am glad that the homophobic, racist, anti-science Wildrose party hasn’t taken power.   And the fact that Alberta has now elected its first female premier is something that shouldn’t be ignored or minimised.

What I’m still upset about, though, is that Alberta has decided to put the Wildrose party in second place, crushing the Liberals and NDP by comparison.  Danielle Smith wouldn’t be out of place running along side Sarah Palin, and now she’s forming the official opposition in Alberta?

For shame.

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Hurrying to go to better things.

Over at Why Evolution is True, Jerry Coyne addresses an article by Andrew Riggio, in which Riggio questions the thought processes of a man named Paul Lord who thanked God for saving him from a tornado that struck in Oklahoma, killing several (including 3 young girls) but sparing him.  Noticing that (unsurprisingly) the comments have exploded into a sprawling mess, Coyne pulls out a few for special attention including this one:

Kleb  •  22 hrs ago
Wooooooow. Bitter much? The author’s argument presupposes that from God’s point of view death is bad. People of “true faith”, as his last sentence mentions, are equally grateful to God for His providence in death as in life. Look at the great heroes in Christianity. When they died they weren’t bawling and begging God to spare them, they were profoundly relieved to be joining Him and, at the same time, deeply grateful for the ride they had been on in this world. From a Christian perspective, then, there is no inconsistency here. The survivor is grateful for the life God has given him here, as he should be, but that doesn’t mean he isn’t also looking forward to meeting the Lord.

Whoa!  Should we be grateful to God for asking 6 million Jews to join Him during the Holocaust?

Seeing this reminded me of something I read a while ago in John Aberth’s great book Plagues in World History.  Faced with the crushing mortality and morbidity of the First Plague (the Justinian plague that ran from roughly 541 to 750 CE), Christian preachers responded the only way that they could, from the pulpit.  In reply to the bubonic scourge, Aberth notes this particular line of thinking:

By the seventh century, sermon cycles were being compiled to be recited on a regular basis whenever plague struck a region as part of the Church’s now standard response to urge its flock to repent in the face of God’s wrathful chastisement;  this at least is the overarching theme of four homilies composed at this time in Toledo, Spain, which, as expected, are replete with quotations from the Old Testament.  Yet, one sermon, the third in the series, adopts a strikingly different tone by employing the carrot rather than the stick […].  In a remarkable passage, one that seems to be inspired by the New Testament, in particular the letters of St. Paul, the preacher now dangles the promise of immortality during the Christian afterlife or resurrection in order to help his listeners conquer their fear of imminent death from the “groin disease”:

But what should we say?  You who take fright at this blow (not because you fear the uncertainty of slavery, but because you fear death, that is, you show yourselves to be terrified), oh that you would be able to change life into something better, and not only that you could not be frightened by approaching death, but rather that you would desire to come to death.  When we die, we are carried by death to immortality.  Eternal life cannot approach unless one passes away from here.  Death is not an end, but a transition from this temporary life to eternal life.  Who would not hurry to go to better things?  Who would not long to be changed more quickly and reformed into the likeness of Christ and the dignity of celestial grace?  Who would not long to cross over to rest, and see the face of his king, whom he had honored in life, in glory?  And if Christ our king now summons us to see him, why do we not embrace death, through which we are carried to the eternal shrine? For unless we have made the passage through death, we cannot see the face of Christ our king.

(emphasis mine)

Kleb the commenter has one thing right:  a belief that death is acceptable and even preferable to this life, whether from God’s point of view or from the worshipper’s, is certainly not a new phenomenon.  I wonder at the power this line of thinking must have held when those who contemplated it were faced with a disease that can kill 60-90% of those it infects and may have wiped out as many as 25 million during the Justinian plague alone.  In the mean time, though, I’ll be thankful for the efforts of modern medicine and science which have brought the plague to its knees. (Even if we are on the verge of squandering that advantage and resurrecting the plague’s power through antibiotic abuse, but that’s an entirely different post).

Now that we’ve found Nemo, it’s time to save his friends.

Nemo! Picture by Peter E. Lee, used under a CC license.

There’s many things that we don’t know about the ocean, and most people won’t find that too surprising, but what might be surprising is the extent of our ignorance in some areas.  This is the subject of a recent paper by McClenachan et al. in the journal Conservation Letters, entitled “Extinction risk and bottlenecks in the conservation of charismatic marine species.”  The problem, they contend, is that while the oceans are currently undergoing a massive loss of biodiversity, the full picture of that loss cannot be seen because we’re not even close to knowing how many species have been lost or how many are currently threatened with extinction.

As a small first step to dealing with that issue, the authors of this paper examine marine charismatic species as a way of estimating the threat to some of those species and identifying possible impediments to their conservation.  What are charismatic species, you ask?  They’re species with widespread appeal that receive more attention and funding for conservation (some might say too much attention and funding);  it’s suggested that charismatic species can raise awareness and drive conservation goals.  In this paper, the authors leverage the fame and fortune of marine charismatic species by arguing that because they receive special attention and are – theoretically – at the least risk of extinction even when threatened, charismatic species can serve as an estimate of the lower boundary of the probability of extinction.

The paper takes an unusual and amusing tack to do so.  In their own words: “we summarize the extinction risk of 1,568 species within 16 families of well-known marine animals represented in the 2003 Academy Award-winning movie, Finding Nemo“.  It’s not many papers in which you get to start “with all major characters, as defined by those with credited speaking parts”, including all species in their taxonomic families which included invertebrates, bony fishes, elasmobranchs (sharks and rays), birds, and turtles.  You can follow through the paper for the exact details of how they constructed the lists, but they then took that list of species and evaluated their risk of extinction.  Following that, they evaluated the potential bottlenecks to conservation among the various families. There’s a lot of fascinating detail in this paper, but I’ll skip to the punchline and let you read the rest for yourself.  For me, the most interesting aspect of this paper is captured by Figure 3:

Here we see that many species, especially in the invertebrates and fishes, are almost entirely ignored either in terms of scientific effort (here the number of papers published on the species), status evaluation (are they endangered?), or conservation assignment (listed by CITES, the species-protection for threatened species).  The authors point out that the charismatic species like turtles and birds receive the bulk of the attention at every level, while conservation bottlenecks arise in the other families.

The focus on Finding Nemo is an amusing hook, but though the paper deliberately trades-off rigor for rhetorical power, the argument that the authors make is a clear and important one.  In calling for a greater focus on the marine biodiversity being lost before our eyes, McClenachan and her coauthors make a great point and deserve the attention that they received for publishing this paper*.

Having said all that, there’s a lot left that to talk about regarding the issues that this paper raises.  In particular, I’m struck by the elements of economics, social psychology, and sociology that would interact with the conclusions of the authors’ work.  The fact is that conservation is, and probably always will be, a finite resource (limited in part by money, and in part by scientific personnel) that must be spread about the overwhelming number of species that are likely to be threatened.  This isn’t to say that the status quo is right, or that we shouldn’t strive to improve it – quite the contrary.  But even in the best of all possible worlds, the fallible human beings tasked with the goal of saving these species (scientists, polticians, the general public) are going to exhibit biases of cognition and simple attention that may make it difficult to drum up support for, say, many of the invertebrates in the paper’s list of species.

Take another look at Figure 3 and try to imagine reasons why some of these species might be relatively ignored.  Off the top of my head, a few potential hypotheses jump to mind.  Physical features of the species’ in question may play a role;  for instance, simple preference for anthropomorphism could account for some of the attention paid to the charismatic megafauna.  It’s easier to care if you can imagine the animal talking but it’s a lot easier to imagine a talking shark or clownfish than a talking shrimp (and don’t discount preference for neoteny;  this paper by Mark Estren looks like a good read on that subject).  Or, even something as obvious as size could play a role:  it’s a lot harder to find or pay attention to many of these marine invertebrates than the turtles or sharks or birds that they compete with.

Even the sheer number of species involved could be important, in more than one way.  On one hand, the fact that there are 6 species of turtles compared with 536 species of invertebrates in this paper seems relevant, as the sheer effort involved in finding and cataloguing the marine invertebrates is daunting.  On the other hand, I’m also reminded of work in economics by people like Barry Schwartz and Sheena Iyengar on the paradox of choice:  when confronted with too many options, people are unhappy and find making decisions difficult, even though they claim that they prefer to have more choices available to them.  The number of possible targets in marine invertebrates for conservation efforts could drain the will of politicians, the attention of the public, and the interest of new grad students selecting a species to work on.

I wouldn’t be surprised at all to hear that the people involved in conservation work have thought about some or even all of these issues, but I would not hesitate to recommend a multidisciplinary approach to this problem.  The answer to the call put out by McClenachan and her co-authors has to include a sober analysis that maximizes the efficiency of the resources we have while, perhaps, searching for innovative new ways to increase those resources.  Here’s one thought:  maybe it’s time for a Kickstarter for conservation – a ConservationStarter?  If the idea of a charismatic species is to reach its fullest expression, I can imagine that it might be in the form of directly crowdfunded conservation efforts targeted at particular species.

Whatever we do, the authors of the paper make it clear that we have to do something, and that our actions have to start with knowledge.  There’s simply too much out there that we don’t know;  we may have found Nemo, but we don’t know very much about his friends and we’re in danger of losing them too.

—————

Loren McClenachan, Andrew Cooper,, Kent Carpenter, and Nick Dulvy (2012). Extinction Risk and Bottlenecks in the Conservation of Charismatic Marine Species. Conservation Letters, 5:73-80.

* My post isn’t especially timely when it comes to this paper, as it was published in January, and others have written about it before me;  a good example is the fine folks over at Southern Fried Science or the Scientific American blogs.  However, the paper was only recently brought to my attention by the good folks in Bill Sherwin’s lab, and the discussion we had about it inspired this blog post.

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Do you know this bird?

Google has failed me (even the reverse image search, which failed rather spectacularly), and since it worked so well last time, I thought I’d ask the readers once again:  any idea what species this bird hails from?

It was spotted while at the walk-in aviary near Canberra, surrounded by dozens of species of parrots, cockatoos, finches, and more.  But there was no picture on the board identifying it, and I forgot to ask on the way out.  So, if you know this bird let me hear about it in the comments!  A prize will be given for the first correct answer*.

* The prize is a lie.  There is no prize.

Update:  pete chimes in below with what I think is the correct answer:  the Rose-ringed Parakeet (Psittacula krameri) found in Africa and Asia according to Wikipedia.  The internet FTW!

The Rose-ringed parakeet, with a colour mutation to blue...

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Rejection Watch Vol. 1, Supplementary Online Material

Photo by Troy C. Boucher Photography, used under a CC license.

Submissions for Rejection Watch have dropped off, which doesn’t really surprise me;  the traffic on this blog isn’t quite strong enough to sustain a feature like that (yet!), though I don’t regret the attempt.  The submissions I did get were fantastic and if anyone out there still wants to send me material, I’ll be happy to resurrect it whenever they do.  In the mean time, though, a couple of relevant posts from around the web have cropped up in the last day or so, and I feel like they make great supplementary reading for those of you reeling from academic rejection.

Rosie Redfield (she of debunking-#arseniclife-fame) over at RRResearch posts her rage over a crappy review of her postdoc’s paper:

We finally (after two months) got the reviews back for the postdoc’s manuscript about DNA uptake bias.    It’s a rejection –  the reviews were quite negative.  The first reviewer was very unfair; they didn’t find any fault with the methods or data or analysis, but they attacked our brief discussion of the functional evolutionary context of uptake bias.  This is all too common for my papers.  The reviewer is so hostile to the idea that bacteria might take up DNA for food that they don’t focus on the science.  Because the paper was rejected we don’t get to do an official response to the reviews, so I’m relieving my frustration by responding to them here.

She goes on to do a detailed, blow-by-blow response to the objections of the two reviewers.  The whole thing is a great read, even if you’re not in this field;  the feeling of ‘oh, that happened to you too?’ is too good to pass up.

Meanwhile, over at The Bug Geek Crystal has found a new pit of despair:

So you know that I handed my draft manuscript in to my advisor last week.  He sent back a document covered in red ink. Then my labmates pointed out all the dumb things I did, and showed me all the cool things I COULD have done but didn’t.

My advisor, a real funny guy, said, “You should make a new graph about the revision process,” and I was all, “Ha ha ha that’s so funny.”

The graph she makes is pretty awesome, but one of the things that struck me was that even the most well-meaning revisions from people close to you (advisors, labmates, colleagues you respect) can cut deeper than the blunt hammerings of an anonymous reviewer with a grudge.  I think that this is because it inspires different emotions, rage for reviewers and despair for labmates.  When we have a personal relationship with those who have dripped red ink on our work, it’s hard to avoid  the attack on your sense of self:  this person knows me, and didn’t think my work was perfect, so there must be something wrong with me.  I should have done better, screwed up.  These are people that you (usually) like, that you want to look smart in front of.  Contrast Crystal’s feelings of despair with Rosie’s feelings of rage;  when anonymous reviewers trash our material, unless we think that they’re right we can work up a really good mad and use it as fuel to revise. In the academic setting, it feels to me like rage is a more productive emotion, a provocation to defiant action (‘I’ll show you Mr. Anonymous Reviewer who will never read this paper again!’) while despair has a soporific effect that leaves us drained and dragging ourselves through the revisions.

Of course, this is just a sweeping generalisation based on my own experience that is almost certainly wrong in some fashion.  But hey, maybe you can leave an anonymous review?  Then I’ll show you.

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A quick follow-up to DolphinGate: cool science!

One of the things that surprised me about the weekend’s DolphinGate flap was that most of the mainstream articles I read were acting as though the paper that they were wildly misinterpreting was providing the first evidence for dolphins engaging in same-sex behaviours.  The Huffington Post is a great example of this;  here, for example, is the first paragraph of their story:

Dolphins were once humorously alluded to as “gay sharks” on an episode of “Glee,” but a new study suggests that bisexuality and even homosexuality among the marine mammals may be very much a reality.

I’ll admit that this caught me off guard, because I was under the impression that the dolphin same-sex play was a well-established fact.  How was everyone reporting this as a big surprise?  Yet when I spent a few minutes poking around on Google Scholar, I couldn’t immediately find a good reference for when this was first talked about in the literature.

Thankfully, the internet provides all!  Justin Gregg of the Dolphin Communication Project mentioned my piece on Twitter and, seeing an opportunity to get to the bottom (?) of this, I reached out to ask him if he knew when these behaviours were first observed.  Justin came through in a big way:

1948!  I can’t get at the actual paper (#icanhazpdf?), but the abstract is a gem of scientific understatement:

A summary of observations of captive porpoises is given. Vision and hearing are well-developed, and vocalizations are produced in the form of “jaw-snapping,” whistling, and barking. In captivity the porpoise shows a diurnal sleep cycle. There is a stable dominance hierarchy. Both homosexual and heterosexual behavior has been observed, as well as masturbation in the male. An instance of live birth is described. The play of the porpoise is complex and goes on for long periods. They manifest many and definite fears. The homologies of these types of behavior with that of other mammals is discussed. When various characteristics of porpoise behavior are considered and compared, the animal may be located at many different points in a scale of phylogenetic complexity. Although no information is available on problem-solving in this form, other types of behavior place the porpoise at a place in the developmental scale between the dog and the chimpanzee.

I’ll have to own up to my own foibles here, though, because the conversation may have gone a little down hill after that.

[Insert your own jokes here….]

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Science journalism blows it, dolphin rape edition.

A few weeks ago I got into a discussion on Twitter with Ananyo Bhattacharya, online editor of Nature News and writer for The Guardian’s science section, after he put out a call asking for ways to improve science journalism. During that conversation, I argued that one way to do this is to create a culture of journalism that values scientific knowledge and expertise as a core value[1]. Ananyo seemed unimpressed with my viewpoint, and suggested that the main point of science journalism was to pry into the dark corners and root out biases, fraud, and the like in science. He views scientific communication and scientific journalism as two distinct things (and thinks that journalists doing ‘PR for science’ is ‘drippy’). Indeed, when asked directly during a Royal Institute forum on science journalism whether journalists should read the original papers behind the stories that they write, he dismissed the idea:

“If the question is ‘must a good science journalist read the paper in order to be able to write a great article about the work’ then the answer is as I said on Tuesday ‘No’. There are too many good science journalists who started off in the humanities (Mark Henderson) – and some who don’t have any degrees at all (Tim Radford). So reading an academic research paper cannot be a prerequisite to writing a good, accurate story … So I stick to the answer I gave to that question on the night – no, it’s not necessary to read the paper to write a great story on it (and I’ll also keep the caveat I added – it’s desirable to have read it if possible).”

He further suggests, in the same comment (original source), that if journalists had to read original papers than no one could report on particle physics[2].

I’m not going to try and hide my bias here: I don’t like Ananyo’s viewpoint on this. I don’t think that it will lead to good writing, either of the communication or journalistic variety, but more importantly I think that forcing journalists to read the papers before they write an article might have stopped stupid @#$@ like what happened today from happening at all.

The story: I received an e-mail this morning from Dr. Bill Sherwin, a member of the Evolution and Ecology Research Centre (E&ERC) here at my current institution, the University of New South Wales. Bill is one of the authors on a new paper coming out in the Proceedings of The Royal Society (B), entitled ‘A novel mammalian social structure in Indo-Pacific bottlenose dolphins (Tursiops sp.): complex male-male alliances in an open social network’. The paper is a nice little exploration of the characteristics of social networks in dolphins found in Western Australia; in essence, they were testing whether two hypotheses about the nature of these social networks were tenable given the data they’ve observed. In particular, they tested whether dolphins show signs of engaging in ‘community defence’, where higher order alliances of dolphins form to patrol and defend a larger community range, similar to chimpanzees, or if it follows a ‘mating season defence’ model where male groups shift their defence to smaller ranges or sets of females when it’s mating season. The comparison to terrestrial species with complex social cognition (such as primates and elephants) is an interesting one, because it provides yet more insight into the relationship between the development of complex cognitive faculties and social relationships.

So far, so good. Bill gave a simple explanation of the paper in an email that he was sent out to the E&ERC this afternoon:

We put out a paper that said “dolphin male alliances are not as simple as other species”, but it has stirred up quite a lot of interest, because somewhere in it, the paper mentioned “bisexual philopatry”, which when translated out of jargon means  “males stay near where they were born, AND females stay near where they were born” – nothing more or less than that.

‘Quite a lot of interest’ is one way to put it. ‘Idiots crawling out of the woodwork’ is another. Here’s the headlines of four stories that were written about this paper:

Dolphins ‘resort to rape’: Dolphins appear to have a darker side, according to scientists who suggest they can resort to ‘rape’ to assert authority. [The Telegraph]

Male dolphins are bisexual, US scientists claim. [news.com.au]. (Note that this is an Australian website, and Bill is Australian).

Male bottlenose dolphins engage in extensive bisexuality. [zeenews.com]

And by far the best of the lot (guess who it’s from?):

The dark side of Flipper: He’s sexual predator of the seas who resorts to rape to get his way. [That’s right, The Daily Mail].

……..

Are you kidding me? If the ‘writers’ of these articles had read the paper, they would have noticed that it contains nothing about the sexual behaviour of the dolphins they studied, bisexual or otherwise, aside from brief mentions of the possible consequences of social networks on reproductive success. It certainly didn’t mention anything about bisexual behaviour, homosexual behaviour, or rape. Now, it’s well known that dolphins engage in homosexual behaviours, and I’ve seen papers arguing that they use sexual coercion as well (Rob Brooks confirms this). But these topics have nothing to do with this paper at all. Even a cursory glance through the original source would have killed these headlines – and the first few paragraphs of the Mail story – which aren’t just a miscommunication but border on outright fabrication. The articles themselves are weird mixes of sensationalist headline with a regurgitated paraphrasing of the much better Discovery News piece that they are treating as the primary source. Here’s the problem, though: it’s Discovery News that makes the original mistake about ‘bisexual philopatry’, interpreting it as bisexual behaviour (hot male dolphin-on-dolphin action, as it were). A reporter who had read the original source could have corrected that mistake fairly easily, or could even have been driven to ask further questions. Without that, however, the press cycle grinds mercilessly forward to Flipper the bisexual rapist.

For my part, I was happy to see that James Randerson’s informal survey of science and health writers showed that many of them do read the original papers. And the kind of people who write things about science that I trust, whether they’re professionally trained in science or not, are not the sort of people who do boneheaded things like this. Ananyo might retort that ‘asking questions’ is enough (he suggested as much in his comment above). Matt Shipman said much the same thing in the piece that Ananyo was commenting on. Yet of all people, Ananyo should be wary of this answer, with his focus on investigative science journalism. A scientist writing an email or doing a phone interview can tell you just about anything that you want to hear; a press officer can write a terrible press release; a wire service will probably distort what comes down the line. But a scientific paper is the One, True Source. It is a public record of what was done, and it is the first and best place to start for answers about a study or a scientific topic[3].

Don’t mistake my criticism of Ananyo’s position of reading scientific papers as a general attack on scientific journalism. I think that there’s a lot of great science journalism out there, and that there are even more great science journalists and communicators. Despite the perennial swirl of internet discussion on the topic, I don’t actually think that the whole field is hopelessly broken like some seem to. I just happen to believe that scientific papers, the products of our time and energy as researchers, form an integral part of the process of talking about science (and it’s part of the reason for my support for Open Access publishing). And I think that disgraceful trainwrecks like the reporting on Bill’s paper are a perfect illustration of the need for these papers to be a part of that process.

[Update: Rob Brooks has also discussed this issue over at TheConversation].

——-

[1] Because of Twitter’s space constraints, this was misconstrued to mean that I was agitating for all science journalists to have a Ph.D. in a scientific discipline. Though I wouldn’t be upset if this happened, that’s not what I meant: it is more than possible to have a deep love and knowledge of science without having a degree in a scientific field. Hell, Carl Zimmer probably knows more about viruses and evolutionary biology than I do, and his only training is an undergraduate degree in English. My argument is only that having scientific training increases the probability of a writer or journalist having a good grasp on how science works, not that it’s the only way for that to happen. I will continue to argue, though, that those having a love of science (professional or amateur) will, on average, produce better science writing and science journalism than those who don’t.

[2] He also claimed that most of the people asking journalists to read papers are biologists and medical people, who write easier-to-understand papers. I would have to turn this back on him: if biology and medical papers are so easy to understand, why shouldn’t journalists read them every time?

[3] Yes, there’s no guarantee that what is written in the paper is true. But the chances of detecting fraud are essentially zero if you don’t read the paper to begin with, and if you’re a journalist looking to catch the next Stapel, chances are that you’ll have to wait for the scientific community to find him and tell you about it anyways.

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Group selection, again. Yay.

I was amused to see that David Sloan Wilson took a weird poke at Dawkins, got thrashed by Jerry Coyne, and didn’t like it.  In fact, I was going to leave this as a link post, but while searching for a link to Coyne’s piece (Wilson can’t seem to figure out how to embed links to anything but his own blog in his posts) I came across a post by a blogger who calls him/herself “The Verbose Stoic”.  This piece is problematic on several points, but discussing this is going to take some space so I’ll do it here instead of a comment on Verbose Stoic’s blog;   from here on, I’m going to refer to him/her as VS.

VS starts off by questioning “examples”:

 What I want to talk about is the objections that Coyne raises against Wilson’s group selection theory:

Dawkins’s argument against the efficacy of group selection was that this form of selection is usually unsuccessful because groups are vulnerable to subversion from within by those selfish replicators. That is, “cheating” replicators that are “good” for individuals but bad for the group as a whole will tend to propagate themselves. Yes, altruism may help groups propagate, but altruistic groups are susceptible to invasion by cheaters unless the “altruism” is based on kin selection or individual selection via reciprocity.

That’s the main one, but he goes on to fill in more later:

Dawkins’s (and my) beef with group selection as a way to evolve traits that are bad for individuals but good for groups is that this form of selection is inefficient, subject to subversion within groups, and, especially, that there’s virtually no evidence that this form of selection has been important in nature.

Let me deal with the two minor ones before getting back to the main event. Starting with the last one, we can see that it’s a bad argument, because what Coyne is doing here is saying that one of the reasons to reject the examples Wilson’s giving of cases where group selection has been important in nature is … that you haven’t found examples of cases where it has been important in nature. Except, perhaps, for the specific cases Wilson is citing. You can’t in any way reasonably claim that the fact that you haven’t found examples of it yet means that you can dismiss this proposed example.

Look, Wilson isn’t citing any specific cases of group selection occurring in nature, mostly because there aren’t any.  When I say that, I mean that Wilson hasn’t been able to demonstrate that a trait arose because of group selection and not kin selection or natural selection or any other evolutionary force.  Wilson’s argument is that (1) group selection (a.k.a. “new” group selection or multi-level selection) is something different than any other variety of selection, and (2) that it is responsible for the evolution of traits such as altruism.  But (1) group selection reduces mathematically to inclusive fitness (follow the links in my previous post), and so (2) is trivially true.  Sure, it arose by “group selection”, but that’s an empty statement.  Wilson’s ‘proposed example’ is a theoretical model that was dealt with when he proposed it nearly 40 years ago (Wilson, 1975), and though it’s been refuted dozens of times since, he keeps holding on to it and insisting that he’s already won.   I’ll quote at length from West et al. (2007) to drive home the point:

It has since been shown that kin selection and new group selection are just different ways of conceptualizing the same evolutionary process. They are mathematically identical, and hence are both valid (Hamilton, 1975; Grafen, 1984; Wade, 1985; Frank, 1986a, 1998; Taylor, 1990; Queller, 1992; Bourke & Franks, 1995; Gardneret al., 2007). New group selection models show that cooperation is favoured when the response to between group selection outweighs the response to within-group selection, but it is straightforward to recover Hamilton’s rule from this. Both approaches tell us that increasing the group benefits and reducing the individual cost favours cooperation. Similarly, group selection tells us that cooperation is favoured if we increase the proportion of genetic variance that is between-group as opposed to within-group, but that is exactly equivalent to saying that the kin selection coefficient of relatedness is increased (Frank, 1995a). In all cases where both methods have been used to look at the same problem, they give identical results (Frank, 1986a; Bourke & Franks, 1995; Wenseleers et al., 2004; Gardner et al.,2007).

VS also isn’t happy about “efficiency”:

The first one is also a pretty bad argument when you look at evolution. The argument is that Wilson’s proposed solution would be inefficient, but it seems to me that one of the main thrusts of evolution is that it can indeed be — and often is — inefficient but as long as it works, that’s not a problem. When has it become a criteria for evolutionary explanations that it achieve maximal or even reasonable efficiency. To go down that route would risk re-introducing a need for a designer, to ensure that the mechanisms stayed efficient. That can’t be what Coyne wants. But, again, why is efficiency even a factor? Why would you sort evolutionary arguments by efficiency? Being more or less efficient isn’t a hallmark of evolutionary mechanisms, so if two mechanisms are proposed but one is more efficient than the other that says absolutely nothing about which one is more likely to be true.

Efficiency is a perfectly fine criterion to use, though the term is a little vague as used here.  Assuming that group selection is different from inclusive fitness (which it’s not):  if group selection results in an very slow rate of change in gene frequencies or a lower probability of fixation compared to inclusive fitness, then inclusive fitness is more ‘efficient’ and is more likely to be the cause of a trait fixating in a population.  At least, that’s how I would use the term;  I don’t want to put words in Dr. Coyne’s mouth, though I think that my view here is consistent with his usage and with the literature I’ve reviewed.  In other contexts, I’ve also seen ‘efficiency’ used to say that group selection wouldn’t actually the enhance relative fitness of altruism vs ‘cheating’ (which isn’t a great term for this, as I discuss below), which ends up in the same place.
In any case, VS seems to be confusing efficiency (whether Dawkins / Coyne would use it the way I do or not) with design.  Adaptations are often very badly designed, such as the case of the amazing recurrent laryngeal nerve, but that doesn’t say anything about how fast (or with what probability) genes for those adaptations spread through populations as a result of natural selection.  Even if group selection works the way that Wilson thinks it does, reasoning from the published theoretical models it’s easy to see why it would be an extremely inefficient process with its cycles of groups / reproduction as compared to overlapping generations with continual selection pressures.
VS finally goes onto what he thinks is the biggest error that Coyne makes:

That leaves us with the main complaint: cheaters. The main issue here is that there is an issue raised against the individual selection explanations of altruism as well, even kin and reciprocal altruism and it is … cheaters. Cheaters will benefit if they can get away with it, and so those individuals will prosper and those who are altruistic will be outstripped, and so altruism is not self-sustaining at the individual level. To get around this, the proponents of evolutionary explanations for altruism end up appealing to cheater detection mechanisms […]

Additionally, it seems to me that group selection can actually get this without having to apply specific cheater detection mechanisms. After all, group selection would imply that the relevant competing entity is the group. Thus, if a group has a significant percentage of people who are altruistic, then it outperforms groups that don’t. Thus, if you have a group where this happens and where too large a percentage of the group are cheaters, then that group will cease to get those benefits and be outcompeted and presumably eventually exterminated by the groups where that does not happen. Thus, group selection here becomes self-sustaining; if you are above or at the magical percentage that means you benefit from being altruistic, you benefit over other groups as long as it stays there, but if it ever drops below that your group may well collapse and your individuals, then, all lose. Note that we would still see cheater detection mechanisms emerge because they are mechanisms that make the group stable and so less likely to fall below that percentage and collapse.

It seems like VS might be on the verge of confusing old and new school group selection, as the bolded statements (my emphasis) suggest.  West et al.’s paper has a great figure that shows the difference between the two:

In the text of their article, they point out that “[a]nother way of looking at this is that the new group selection approach looks at the evolution of individual characters in a group structured population, whereas the old group selection approach looks at the evolution of group characters”.  VS’s own words make him sound like a disciple of Wynne-Edwards, which would be unfortunate since Wynne-Edwards was decisively crushed by George Williams in the 1960s.  But even if he’s just poorly recapitulating Wilson’s models, VS is still wrong on the evolution of altruism.  There are a number of possible explanations for altruism, including inclusive fitness, but I don’t want to get into a long conversation on how altruism might have evolved because I would have research and then write an inconveniently long book to do so.

Having said that, Coyne’s use of “cheating” (even in quotations) is a little unfortunate, because it overlaps with the literature on Prisoner’s Dilemma  and cooperation.  Cooperation and altruism are not the same concept (again, see West et al. for a good breakdown of the different concepts, or any text on social evolution);  altruism might be a subset of cooperation, depending on how you define the terms, but usually altruism comes at a cost to the altruist while cooperators do not necessarily pay a cost to cooperate.  “Cheating detectors” is more appropriate for a conversation about cooperation than altruism  (see also Figure 2 of this paper), which makes the rest of VS’s argument difficult to respond to.  I think that Coyne is using ‘cheating’ to refer to individuals who receive the benefit of altruistic acts without paying the price of altruism, but that’s not the same as cheating in models of cooperation.  (Honestly, ‘cheating’ isn’t a great word on a lot of grounds, including confusion with other areas such as signalling and an implication of conscious intent where none is necessary).

Returning to the posts that started this digression:  my honest belief is that this group selection debate should have been over years ago, but I will still support Wilson’s right to continue trying to make his case.  If he’s going to attack people like Dawkins and Coyne, though, he’d better learn to be prepared for them to hit back.  And though it’s unlikely that either of them will ever read this post, I’d like to tell them that they’re not alone.

P.S. Can I take this opportunity to point out a further example of Wilson claiming that people agree with him when they don’t?  If you read the end of Wilson’s second piece, he says:

For readers who are up for a challenge and want to learn more about the theoretical basis and empirical evidence for group selection from someone other than myself, I recommend Steven A. Frank’s “Natural Selection. III. Selection vs. Transmission and Levels of Selection (Journal of Evolutionary Biology, 2011). For Frank, it goes without saying that natural selection is a multilevel process and that the group level is often a significant evolutionary force.

I’ve actually read that paper.  In it, Frank once again points out that kin selection and group selection are the same thing:

The equivalence of r and Hamilton’s formal theory of kin selection establishes the exact equivalence of multilevel group selection and kin selection.

And then, after a long analysis, he compares the usage of the two methods in a section entitled (tellingly): Reasons to favour kin selection over group selection.  It contains exactly what the title says.  If you can get it and you like technical discussions of evolutionary biology, I urge you to read the paper yourself.  If you don’t, then just do me a favour and don’t accept Wilson’s claims about this paper at face value.

——–

David Sloan Wilson. A theory of group selection. Proceedings of the National Academy of Sciences, 72 (1):143–146, 1975.

S. A. West, A. S. Griffin, and A. Gardner. Social semantics: altruism, cooperation, mutalism, strong reciprocity and group selection. Journal of Evolutionary Biology, 20(2): 415–432, 2007.

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Nerd fight!

I got into a bit of a scientific nerd fight today online with @hylopsar, and I thought that I might make a record of our duel for the internets to see.  It’s good for a laugh, so go check it out:

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