Monday, July 1, 2013

What not to do (and what to do) at conferences


Post written by Kiyoko Gotanda and Ben Haller (equal contributors)

This year, the annual joint congress of the Society for the Study of Evolution (SSE), the American Society of Naturalists (ASN), and the Society for Systematic Biology (SSB) was held in gorgeous Snowbird, Utah. For those who don’t know, Snowbird and Salt Lake City was where the 2002 Winter Olympics was held, and Snowbird is at approximately 7,000 feet elevation. This proved amusing when running between buildings (see #2) as people tried to keep up conversations while gasping for air.

So, instead of just rambling on about all the great talks, we thought we’d take this opportunity to opine about what NOT to do at conferences, and then some counter-examples of what TO do at conferences. Please keep in mind that the what-not-to-do’s are not specific to Evolution 2013, but rather, are an amalgamation of things we’ve noticed over the years. The things TO do at a conference are from this year, and all in all, the conference was a rousing success.

 Snowbird, Utah! Not a bad setting for a conference...

Without further ado, what NOT to do at conferences

1) DON’T forget who is in the audience. Conferences are attended by everyone, and we mean EVERYONE. That’s undergraduates who are just learning about the big, bad academic world, hot-shot profs, graduate students suffering from imposter syndrome, post-docs trying to figure out if they want to stay in academia – and even more diverse folk. At Evolution 2013, there was an artist-in-residence who was making watercolor sketches of the talks she attended. Seriously EVERYONE is there! Gearing your talk just towards your immediate colleagues can be a bit frustrating for others if they aren’t part of that in-group; more importantly, it can deprive you of the cross-fertilization of ideas that comes from talking to people outside your specific field.

However, Richard Lenski gave the SSE Presidential Address, and he knew who was in his audience: EVERYONE! Lenski has been running a bacterial evolutionary experiment for over 25 years, and it has provided remarkable insights into many aspects of evolution. In this talk, most remarkably, it appears that fitness peaks are never really reached. The usual assumption is that if a fitness peak exists, a population will evolve toward that peak asymptotically, making an ever-closer approach to some optimum phenotype over time. But after 50,000 generations in Lenski’s experiment, the approach to the peak does not appear to follow a hyperbolic (i.e. asymptotic) trajectory; rather, it follows a power law, rising without an asymptotic limit (although at an ever-slower rate of increase), and the evidence strongly suggests that this will continue indefinitely. In other words, there IS NO OPTIMUM PHENOTYPE. There is no maximum fitness. This could have profound implications for the underpinnings of evolutionary theory. These results, which could have been shown in a system-specific, jargon-laden manner, were instead shown in a way that swept the whole audience off its feet. That’s how to do a talk!

An American Robin yelling in a jargon-laden manner. That's a no-no


2) DON’T pick talks that require you to run around like a chicken with its head cut off – easier said than done at these massive conferences! The Ecological Society of America is clocking in at close to 3K people each year (!) and Evolution is hitting 1.5-2K. With this many attendees, there have to be a bunch of concurrent sessions. Space limitation is limited, and so conferences are often held between several buildings. You can sprint (literally) between sessions –although this is a bit harder at 7000 feet elevation! – but you miss the questions, and even the beginnings of some talks, and you miss the continuity and flow that comes from a series of talks in a single well-organized session. But there’s another option! You can pick one session and enjoy all of the talks. We both saw complete sessions that impressed us with the quality of speakers and the science presented.

If we could fly, it would be easier to get around the conference!


For Kiyoko, this was one of the evolutionary ecology sessions. Using tropical frogs, Justin Touchon showed that phenotypic plasticity could provide the link between alternative phenotypic states – in this case, where frogs lay their eggs. Alex Badaeyev argued that network connectivity is the key to linking micro- and macro-evolution, and he demonstrated this using the diversification of avian plumage as his system. This was appropriate as his network relied on dietary requirements which are linked to plumage. Corlett Wood showed how we need to consider covariance between heritability and selection when using the breeder’s equation. In general, people estimate heritability and selection separately, but she argued that heritability and selection can vary due to the environment, which means that environmental factors could be causing heritability and selection to co-vary. The model looked at what happens when a covariance metric is included in the breeder’s equation, and the model showed covariance can strongly affect the variance, but not the mean. Katie McGhee demonstrated how paternal and maternal effects can interact to affect behaviour in sticklebacks, and Alison Egge spoke about cold tolerance and plasticity, and Oriol Lapiedra showed that arboreal feeding evolved from terrestrial feeding in birds on islands, and then the arboreal-feeding island species dispersed to the mainland. In other words, arboreal feeding in birds occurs on mainlands only by virtue of its evolution in island species! Just how cool is a session like that?

For Ben, it was a speciation session. Roman Yukilevitch showed a new method of testing whether speciation had occurred in sympatry or allopatry, and used his method to show that perhaps 5% of Drosophila species had resulted from sympatric speciation – a rather earth-shaking result, given the scepticism in some quarters that sympatric speciation is an important force in evolution. Göran Englund gave a very interesting talk about size-specific predation by pike driving sympatric divergence in whitefish. The evolutionary trajectory of the whitefish had a very strong scent of the “adaptive dynamics” perspective: very repeatably, they appeared to first converge on a branching point (of small body size), and then branch into two persistent ecomorphs. Disappointingly, to me, most of this divergence is apparently plasticity, not adaptive evolution (which is perhaps why it does not progress to full speciation, I would speculate); nevertheless, a fascinating talk. Bjørn Ostman (well-known as the moderator of the Carnival of Evolution) then gave a talk on resource specialization and the evolution of trade-offs; I quite enjoyed this, since I think trade-offs and the balance between generalists and specialists is an important and interesting area of theory. Liliana Lettieri finished with a talk on how female choice is a force for diversification in stickleback; this is an important area for empirical researchers to test, given the theoretical importance of magic traits and other mechanisms for assortative mating in producing divergence. At this point I confess that I switched sessions, but that’s because my session was open in the last slot. I went to a remarkable talk by Gideon Bradburd on separating the effects of geographic isolation (i.e. isolation-by-distance) and ecological isolation (i.e. selection against migrants and hybrids), involving a new method: BEDASSLEing your data. It looked quite powerful, and it was striking to me that after the talk ended, nobody got up and left; the whole audience stayed to hear Bradburd’s answers to questions. That’s a good talk. So, OK, switching sessions can be good; but maybe only when your “native” session is over!

3) DON’T spend all day everyday in sessions. If you spend all your time in sessions and don't allow time to explore your surrounding environment, you will burn-out mentally (and physically!). Thank goodness this wasn’t a problem at Evolution 2013. With Snowbird as a backdrop, there were many things to do besides go to presentations and posters. Kiyoko went on the birding trip to Antelope Island (in the Great Salt Lake), which was a rousing success. The bus was full of birders! It was led by Sarah Knutie, Sabrina McNew, Leo Gustaffson, all from the Clayton-Bush lab from the University of Utah. We had a wonderful tour of the local wildlife, including sightings of Say’s Phoebe (NOT a sage grouse), Avocets, California and Franklin Gulls, Western Meadowlarks, Yellow-headed Blackbirds, Loggerhead Shrikes, pronghorn antelope, mule deer, bison, and even two Burrowing Owls! Ben went wandering off photographing flowers in the mountains above Snowbird with some old friends, which also worked out quite nicely (see photos).


Antelope Island



Flowers Ben saw on his alpine hike in Snowbird


4) DON’T be a wallflower and not talk to people and network. Conferences can be intimidating, and meeting people to ask questions or find out more about their work can be a daunting task. The ASN has been trying to facilitate networking by holding mixers where professors and students can mingle and talk. Their session this year was a great success, and was well-attended by both grad students and by “scary professors” (as Keewi calls them). Everyone seemed to be enjoying themselves and sharing ideas and theories.

This year, we shared a condo with our good friends Jurri and Sarah, and the condo included a full kitchen and a dining-room table. This proved useful; we did a grocery store run and cooked all of our meals in our suite. We invited a bunch of people over for lunch or dinner and discussed science while enjoying yummy home-cooked food courtesy of Keewi (Ben’s wife) and Rebecca (Jurri’s wife). Collaborations were born, ideas were conceived, and everyone left with a very full tummy! Conference life doesn’t get much better than this. We plan to continue this tradition at future conferences; if you want to have a geeky science dinner with us, drop us a line!


 Snowbird!
There’s more we could write about what to do and what not to do when it comes to conferences, but it’s time to look at some pretty pictures from Evolution 2013 and Snowbird.

By the way, May 25–29, 2014, will be the first joint congress of the Canadian Society of Ecology and Evolution, the Society of Canadian Limnologists, and the Canadian Society of Zoology. It will be in Montreal, which is a wonderful city to visit. Information can be found here: http://www.genomesbiomes.ca . One month later, Evolution 2014 will be in Raleigh, NC. We’re planning on being at both, so we’ll see you there!

And, here's some more pretty pictures!








Sunday, June 23, 2013

The Living Dead: Darwin's finches and museums.


“Darwin’s finches are dull to look at, not only in their orderly ranks in museum trays, but also when they hop about the ground or perch in the trees of the Galapagos, making dull unmusical noises.”
(David Lack 1947)

I have seen thousands of Darwin’s finches hopping about in the wild and handled many hundreds of them from capture in mist nets to release back into nature – surely something that only a few hundred human beings have every done. So why should I feel awe when faced by 168 of them arrayed before me in their orderly ranks in museum trays? Surely I shouldn’t – and yet that is precisely the feeling I had when handling collections in the Harvard University Museum of Comparative Zoology during the filming of a documentary about selection imposed by humans.

The Living Dead

Perhaps my awe stemmed from the fact that the museum where I work, the Redpath Museum at McGill University, does not have any Darwin’s finches – despite the fact that McGill was where Peter and Rosemary Grant started their celebrated studies and where I have worked on Darwin’s finches for more than a decade*.  Amazing but true – and the main reason is that the Galapagos National Park Service does not allow export of finches (or anything else for that matter). I can see why, of course – because otherwise every one of the tourists would want their own preserved finch to take home as a souvenir. Hell, I would want one too. But surely this shouldn’t apply – within reason – to scientists. In fact, old museum collections from before these draconian rules were put into place have yielded outstanding insights into what has happened to finches from an evolutionary and conservation perspective – as evidenced in this cool paper by Ken Petren. It seems to me that the value of museum collections, including new ones, is very high and the potential impact on finch populations in Galapagos is vanishingly small. In fact, I would bet that the number of Darwin’s finches killed by taxis driving on roads in Galapagos every year exceeds the number of Darwin’s finches in all of the world’s museum collections to date.

Or perhaps it was because the collection had some amazing specimens, including a lot of finches collected in 1929, some collected in 1891, and one that might have been collected before Darwin. As hard as that may be to believe, curator Jeremiah Trimble told me the one specimen came from the private collection of an individual who died around that time – and people certainly did visit the Galapagos for centuries before Darwin. Or maybe this finch saw Darwin back in 1835 and maybe just narrowly avoided a blast from his shotgun. Sadly, however, sufficient information to resolve its collection date and location does not exist for this specimen. It was also fascinating to see the early days of confusion regarding species identification (before David Lack finally convinced everyone in 1947) play themselves out in additions and crossings-out on yellowing bits of paper tied to the feet of finches who were born and died before World War II. Ah the good old days of Geospiza strenuaGeospiza nebulosa, and up to 67 others? In fact, a number of the specimens did not yet have the new (already more than a half-century old) established scientific names on them. And how cool was it to see those old English island names, some of which are so much cooler than the modern Spanish ones? Wouldn’t it be great to say I work on Indefatigable Island rather than the mundane Santa Cruz Island?

Beak diversity in the seed eating ground finches

Or perhaps it was just because museums – particularly old ones – have the aura of mystery about them that, while not replacing the excitement of seeing wild animals doing their thing in nature, augments and expands our sense of wonder and discovery. Perhaps I might – simply in the course of rummaging through these simultaneously sad and yet somehow hopeful rows of feathers and beaks with their vacantly staring cotton eyes – find some new species or have some new realization. As you might imagine, this didn’t happen while looking at these well studied birds. Personal discoveries did, however, happen several times while browsing the public displays. Check out this sampling, starting with my own initially exciting thoughts and then any cold hard reality that came later courtesy of that killjoy Google.

1. The spookfish “has two corneal openings in each eye and uses tiny mirrors to see what is above and below at the same time.” What the f***? I had no idea. How did that evolve? And this was just one fish among a number of others that I did know about but hadn’t seen in the flesh, even in jars: the gulper eel with a mouth many times larger than its body, the tripod fish that stands high above the sediment on absurdly long and stiff fins, and many more. (Google later informed me that the brownsnout spookfish is indeed the only fish with mirrors in its eyes. In addition, the spookfish group as a whole is the same as the amazing barreleye fishes, which I already knew about but still haven’t seen in the flesh. Barreleyes are absolutely amazing  – check out this NGS video. And you can see a tripod fish in action here.).

Gulper eel

2. How about the Cotton Pygmy Goose? We normally think of geese as big, certainly bigger than ducks. So I was rather shocked to be browsing at random along those old-school museum style rows and rows of bird specimens mounted in glass cases and labeled with nothing more than a name and a location. Heightening the effect, this goose was, probably coincidentally, mounted alongside an Oriole that was almost as big. No kidding, I even have the picture to prove it. I had no idea such a thing existed. Why should one goose evolve to be so tiny when all other geese are so much bigger? (Google notes that the Cotton Pygmy Goose is actually a duck – however, it is still the smallest waterfowl in the world, so it remains pretty cool.). 


Death match or epic rap battle? Maroon Oriole v. Cotton Pygmy Goose

3. And then I saw the incredibly colorful and beautiful Tangara tanagers – another amazing example of sexual selection driving extravagant colors – or is it? I read in the text beside the display that the sexes in this group are identical, whereas most other traits that evolved through sexual selection are more exaggerated in one sex (usually males) than the other. So either sexual selection is driving male color and females are simply being dragged along genetically (they do share many of the same genes after all) or they are used for species recognition (but are these birds really so clueless as to need this much color divergence) or maybe it is somehow naturally selected (but why would it be so different – seemingly at random – among species)? I suspect that this topic has been well studied but, at that the moment standing in front of that display, it was fun to make the “discovery” myself. (Limited success on Google suggests that the topic actually hasn’t studied in much detail.)


Tangara Tanagers - a mystery to me.

And on and on and on – one amazing critter after another, a reasonable number of which were totally new to me. Although none of these things were my discoveries in the real sense, they still felt like it in the moment. (Of course, none of these things were discovered by their European “discoverers” either – having been seen by indigenous people long before.) When I get home I can’t wait to kick my kids’ butts in our dinner game “what did I learn today that was new”, I can’t wait. Until then, I just have to say:


"Darwin’s finches are exciting to look at, not only when they hop about the ground or perch in the trees of the Galapagos, but also in their orderly ranks in museum trays, making no noises whatsoever.” (Andrew Hendry 2013)


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For some of our work on Darwin's finches, look here.

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Lest it seem I am disparaging the Redpath Museum and aggrandizing the MCZ, I would note the latter’s great pride in their Laborador Duck, one of about 40 in the world and “one of the museum’s great treasures.” The truth is that the Redpath Museum also has a Labrador Duck, albeit not a male. Our duck even graced the cover of a book about the quest to see all 40+ specimens - probably because, while not beautiful, it has a certain je ne sais quoi.


The Labrador Duck outside my office in the Redpath Museum, McGill University.

Friday, June 7, 2013

Academic dispersal

My name is Ben Haller.  I was a graduate student once, but I have reached my next life stage.  It is time to disperse.  During my juvenile stage, I worked on my PhD with Andrew at McGill; protected from predators, and optimally oxygenated as he fanned his nest, I flourished and grew.  But now I have hatched; I passed my defense on June 3rd, moved from Montreal to New York on June 4th, and submitted my final thesis on June 6th.  Now it’s time for me to start foraging for myself.

My thesis, "The role of heterogeneity in adaptation and speciation", is still grinding its way through McGill’s bureaucracy, but I’ve posted it online at the link given.  At 309 pages, it’s a bit of a long read, so I’ll give a quick summary here!



Chapter 1 is a broad introduction to foundational concepts, ranging from adaptation and fitness landscapes to gene flow, reproductive isolation, and magic traits.  One of the key ideas presented is the idea of “squashed stabilizing selection”, a combination of stabilizing selection and negative frequency-dependent selection (Fig. 1).

Figure 1. Types of selection: (a) directional, (b) stabilizing, (c) stabilizing and directional, (d) disruptive, and (e) “squashed stabilizing selection”. Top panels show the fitness function: fitness as a function of phenotype. Bottom panels show the effect on a population’s phenotypic distribution, before (solid line) and after (dotted line) selection.  See thesis for details.

Chapter 2, “Solving the paradox of stasis: Squashed stabilizing selection and the limits of detection”, is currently in revision at Evolution.  In it, I use an individual-based model to show various reasons why stabilizing selection might be difficult to detect even when it’s present, perhaps explaining the low rate of detection of stabilizing selection in Kingsolver et al. (2001).  This might help to resolve the long-standing “paradox of stasis”, since it suggests that stabilizing selection might be common in nature despite its infrequent detection.  Furthermore, my results suggest that squashed stabilizing selection might be quite common in nature, which might be quite important in explaining patterns of standing genetic variation and speciation (Fig. 2).

Figure 2. Absolute frequency histograms of linear selection gradients, β, and quadratic selection gradients, γ, observed for the modeled population.  White bar areas represent non-significant selection, black bar areas represent significant selection.  The left column summarizes model realizations without competition, showing that stabilizing selection is detected much more frequently than disruptive selection in that scenario.  The right column summarizes model realizations with negative frequency-dependent intraspecific competition, showing that disruptive selection is more commonly detected in that scenario. Overall, significant selection is rarely detected, as in studies in natural populations.

Chapter 3, “Evolutionary branching in complex landscapes”, is in press now at the American Naturalist.  It uses an individual-based model to examine the effects of complex spatial environmental heterogeneity on evolutionary branching in asexual populations.  Among other results, we find a novel “refugium effect” that promotes branching in complex landscapes that combine steep gradients and patchy heterogeneity (Fig. 3).
Figure 3. A complexly heterogeneous landscape. In the steep gradient on the right side of the landscape (blue), refugia are generated by the spatially continuous patchiness (green).  These refugia can provide evolutionary stepping-stones that promote adaptation to the harsh conditions of the gradient, which might otherwise be too extreme to allow evolutionary branching to occur.  This “refugium effect” appears to promote branching in complexly heterogeneous landscapes.

Chapter 4, “A tale of two morphs: Modeling plant-pollinator interactions, reproductive isolation, and local adaptation in parapatry”, is currently in review at Evolution.  It examines the idea that heterostyly, floral syndrome that involves two different floral morphs with different, and reciprocal, reproductive organ positions, might allow reproductive isolation to develop between parapatric populations, promoting adaptive divergence and speciation (Fig. 4).  We show that this outcome does appear plausible, but that other interesting outcomes can also occur, including strongly asymmetric gene flow that might promote the development of dioecy from heterostyly.

Figure 4. An illustration of the hypothesized mechanism of reproductive isolation between populations of heterostylous plants, as a result of the combination of precise pollen transfer and reciprocal herkogamy.  Differential organ positioning between the populations is the result of adaptation to different local pollinators, and results in decreased pollen flow between the differently adapted flowers.

Chapter 5 presents some concluding remarks and a discussion of future research directions.  I use a flowchart to show how the preceding chapters fit together into an overall conceptual framework (Fig. 5).

Figure 5. A flowchart depicting the chain of causation that propagates heterogeneity upward from its initial ecological causes (squashed stabilizing selection and environmental heterogeneity) into heterogeneity among individuals, populations, and species.



My defense went pretty smoothly, all things considered.  I was quite nervous when I started talking – so nervous that people in the audience could tell I was nervous, which I can usually conceal!  But I settled in, and except for some difficult questions from one member of the committee, I felt fairly confident through most of it.  As Andrew told me repeatedly in the days before my defense, there really isn’t anything to worry about; if you’re not ready to defend, your advisor or your committee or your thesis examiners will tell you so.  By the time you get the green light to defend, you’re ready.  So don’t sweat it.  I’ve heard some people say that they actually enjoyed their defense, and were sad when it was over; I can’t say that I shared that feeling, though!

So what’s next?  I’ll be starting a postdoc in Montpellier this fall, working with Luis-Miguel Chevin at CEFE/CNRS, assuming I can convince the French government to issue me a visa.  We’re interested in looking at questions having to do with G matrices, multidimensional niche spaces, phenotypic plasticity, and lots of other cool stuff.  Stay tuned for details!  Will I fall prey to selection against migrants, providing a cautionary tale to those contemplating their own dispersals?  Or will I survive, and reproduce my ideas, showing that gene flow can have positive effects on adaptive divergence in changing environments?  Will I reach reproductive maturity, and start spawning new graduate students of my own?  Or will senescence take its toll, as my papers collect dust but not citations, and potential mate-collaborators leave me for more dominant scientists?  Only time will tell!

Until the fall, I’m more or less homeless; my wife and I plan to road-trip around the U.S., visiting friends and going to the Evolution 2013 conference in Snowbird, Utah.  Maybe I’ll see you there!

Canyonlands National Park in Utah, not so very far from Snowbird.  If you attend Evolution 2013, take some extra days (or weeks!), rent a car, and see the Southwest!
(Photo credit: Ben Haller)

Thursday, June 6, 2013

Carnival of Evolution #60

Carnival of Evolution #60 is now up!  Our contribution this month is from Hendry Lab member Kiyoko Gotanda, making us all envious with her adventures in the Galápagos.  As usual, there’s a wide variety of other nifty entries about evolution, so check it out!

The theme this month was 60th anniversaries, since it’s Carnival #60.  2013 is also the 60th anniversary of the discovery of the structure of DNA by Watson, Crick, and Franklin, so...


Monday, May 27, 2013

Predictable unpredictability and 25 stickleback biologists in the field.

Ecology and evolutionary biology – and their intersection (evolutionary ecology, eco-evolutionary dynamics) – strive for inferences that are robust, consistent, and stable. For instance, investigators commonly explore how organisms differ between habitats (different “ecotypes”) and how these ecotypes have different ecological effects. Once the differences are discovered and the patterns established, we would expect them to be at least relatively consistent through time: year after year after year low-predation guppies should look like a low-predation guppies and high-predation guppies should look like high-predation guppies. With such consistency, we can draw robust general conclusions about the way the world works. Without such consistency, what do we have beyond a series of snap-shots with little generality to connect them?

Sampling the Misty Inlet Stream.

I spent last week in the field on Vancouver Island helping to start some new projects on the threespine stickleback that live in lake and stream environments. This is a system where the patterns are well established, robust, consistent, and stable. For instance, lake stickleback are always shallower-bodied than are stream stickleback – and surely 15 years of studying the same stickleback populations should allow a researcher to draw many such generalities.  

Dieta shows how it's done.

One of this year’s projects, led by PhD student Dieta Hanson, seeks to understand how differences in breeding time contribute to reproductive isolation (i.e., low gene flow) between adjacent lake and stream populations. In particular, previous observations suggested that stream populations start to breed earlier than adjacent lake populations, which should mean that the two ecotypes will show reduced interbreeding and thus restricted gene flow. With this prior experience as her motivation, Dieta has been repeatedly sampling a variety of lake and stream stickleback populations so as to assess changes through time in the number of reproductive males and females, which will allow her to calculate overlap in breeding time and thus the potential restriction on gene flow. This past week, a group of us joined Dieta to sample the Beaver, Misty, and Robert’s watersheds.

The Misty crew hard at work

I had asserted that stream fish breed earlier than lake fish not only to Dieta but also to everyone else on the crew, including stickleback savants Katie Peichel and Rowan Barrett. Much to my surprise, our Misty Lake sample turned up a fair number of reproductively mature individuals but the Misty Inlet stream sample turned up none – not one gravid female. After having not-so-subtly pitched myself as the resident expert on stickleback, I had to start back-tracking to somehow limit the damage to my reputation caused by this new observation. I think I failed because the crew started poking fun at me (or rather with me) – not just for incorrect stickleback assertions but for my tendency to misplace the truck keys in grocery stores, for my failure to have brought any soap or shampoo or towel, for my affinity for hot chocolate, and so on. However, the next day we sampled the Misty Inlet stream again and found a number of gravid females – so perhaps I rehabilitated my reputation somewhat (at least the stickleback part), especially after winning the pool for predicting how many stickleback we would catch (I was only off by one). Overall, however, it was becoming clear that the dramatic difference in breeding time that I had come to expect might not be strong and consistent.

Rowan and Katie: are they laughing at me or with me?

After working on the Misty and Beaver lake-stream pairs, both located on northern Vancouver Island near Port McNeill, we drove south for a couple of hours to Robert’s Lake Resort near Campbell River. Here we were to continue our lake-stream work, starting with the Robert’s pair, and here we would be joined by our collaborators from the University of Texas at Austin, including team leader Dan Bolnick. On the night that everyone arrived, all of us (one-quarter of a hundred people!) gathered in a room for some discussion, introductions, and a pep talk from Dan. Dan went over safety and security issues, of course, but – more importantly – imparted to us his considerable wisdom regarding the local stickleback populations, especially that in Robert’s Lake itself where he had been working for 13 years. “I have never seen a stickleback breeding in Robert’s Lake before June 6” was the phrase I most remember – perhaps because the next morning I walked down to the dock at the lake and saw two breeding males. One of them had babies already, which meant that it must have started breeding at least a week (and perhaps two) earlier. This means that breeding must have started in Robert’s Lake more than three weeks before the previous earliest date. Now it was Dan’s turn to try to salvage his reputation as the local stickleback guru, and together we converged on the best way to do so – we started trading stories about how unpredictable things could be.

Dan: “It is amazing how some years the stickleback are small and other years they are large.”
Andrew: “Ah, yes indeed Dan, and how in some years they have incredible nuptial color and some years much less so.”
Dan: “Right you are Andrew, and some years they breed early and sometimes late.”

I am sure you can see here how we had cleverly shifted from our earlier statements of confidence in generalities to statements showing our rich knowledge of variation and exceptions, knowledge that could only be achieved through our long and detailed experience in these systems. Next year, and in those that follow, I will start from this new position and thus never be wrong again.

Dan's other project - the search for assortative mating in stickleback.

The entire crew only overlapped at Robert’s Lake for one day but we did have time for a farewell drink of Scotch (we now “own” a one square-foot plot of land at Laphroaig – and it even has a stream flowing through it) over which we could argue on topics from the trivial to the important. And argue we did, well past when we should have been in bed – but it was somehow appropriate as it was under a very similar situation in the same building and over the same beverage that Dan and I first met many years ago. Up to that point, we had both been working on stickleback in the same lakes for a number of years but hadn’t realized it. So we converged on Robert’s Lake in 2006, played a Hendry-lab verus Bolnick-lab Ultimate Game (I can’t remember who won but I do know that we asked for funding for an Ultimate trophy in our NSF grant.), and hatched several collaborations.


Robert's Lake Hendry v Bolnick ultimate tourney, 2006.

Sadly, field work is now over for me and I am writing this post in the Campbell River airport. But I can rest easy knowing that the field work will continue unabated without me, although I am not sure the tarps will be set so elegantly anymore. Matt will still be wading right to the top of his waders – and sometimes fearlessly beyond. Suzanna and Elena will still be tucking their jackets inside their waders and cooking amazing dinners. Katie will still be making hundreds of stickleback babies and organizing data by headlamp while in her sleeping bag. Rowan will still be the red canoe that every photo needs. Dieta will still be two steps ahead of everyone else. And Carol will be there to film it all. And –most importantly – large bags of nuts will still be three for five dollars at Sayward Junction. Have some for me!

Be the red canoe, Rowan.
24 of 25 - with a stand in for Duncan.

Friday, May 17, 2013

The Nature of Natural History: Kelowna CSEE 2013


Every few years a group of ecologists or evolutionary biologists experiences existential angst about the decline of natural history knowledge. This angst is wholly justified when many biologists no longer take the time to experience how the organisms they study actually live in nature. At best (and this is better than most), many biologists run off to the field for a day, stop at a bridge over a stream or along a forest trail, quickly collect their samples, and then run back to the lab to extract the DNA, run PCRs, and genotype their critters. Organism reduced to molecules. And yet we can’t possible hope to understand how organisms have evolved and how they fit into the polity of nature without careful observation and experimentation IN NATURE. After all, genotypes do not directly experience selection, nor do genotypes have ecological effects: instead it is phenotypes that experience selection and that have effects. And these phenotypes evolve and have effects through interactions with the environment that usually cannot be discerned without careful observation and study.

A five minute walk from the conference center.

This year, the Canadian Society for Ecology and Evolution held their annual meeting – organized by Jason Pither – in Kelowna, British Columbia. The overall theme was Range Margins in a Rapidly Changing World and I was in a symposium organized by Root Gorelick and Kevin Judge on – you guessed it – natural history. My own talk was about the role of photography as a way to bring readers and listeners a bit of a feel for the natural history of the systems under study. I told two stories through data and pictures. One story was about the interaction between bears and salmon and how the former drive evolu-tion in the latter.  The second story was about how humans are altering the evolution of Darwin’s finches in Galapagos.  I ended my talk by pointing out that, yes, photography was expensive but that anyone could set themselves up well from scratch for only about $3,000. The talk after me was about the role of illustration (as opposed to photography) toward the same goal. Lyn Baldwin pointed out that – in contrast to cameras – you can set yourself up well for illustration with a pencil costing less than a dollar. I don’t think that is really fair though as some pencils cost considerably more than a dollar.

The stories I told through data and photos.

The Natural History symposium was fascinating and yielded many curious tidbits about organisms that reminded me of the game I play with my kids at dinner “what did I learn that was new today.” (I did a similar post a few years ago based on a symposium in Leuven.) If I were home today, here is what I would tell my kids:

  1. Many cactuses have a large and extravagant cephalium which – according to Root Gorelick – has no function and could well be maladaptive. Of course, I would then tell my impressionable kids that, no, Darwin was not wrong, instead Root must be, but that, yes, biologists still have no clear explanation of what benefit the cephalium brings.
  2. Jumping spiders, those charismatic midgets with forward facing eyes that leap on their prey, have crazy mating colors and displays. I will also tell my kids how Wayne Maddison, when he was a kid in Ontario, discovered that two species found on sand dunes have incredibly different strategies for building their “nests.” Wayne told the audience that he feared that he would go to his grave before he published this observation and so I here pledge that, should this be so, my kids (one of whom loves spiders) will take up the study and published it (without mentioning Wayne of course).
  3. Feather mites are tiny (often microscopic) mites that cling to the feathers of birds. Heather Proctor explained how these mites show an incredible diversity of forms - even on the same birds – and have a number of crazy features. Males apparently grapple with each other for access to females by trying to throw each other off the feather – just like tiny sumo wrestlers on a tight-rope. And some groups show handed-ness, with the individuals on one wing of the bird showing a different body coiling pattern than individuals on the other wing of the bird.
  4. Like a number of other insects, male Mormon crickets give nuptial gifts to females – in this case, little “cheese balls” that the female munches on while the male mates with them. Cheese balls appear to be quite costly for the males to produce and they are an important part of the diet for females. This leads to so-called “sex role reversal,” where males become the limiting sex during mating and females compete intensely for males. When asked during the question period whether he had ever tasted these cheese balls, Darryl Gwynne admitted proudly that he had and that they weren’t actually that bad.
  5. Hump-winged grig males, like Mormon crickets, provide a nuptial gift for females but not a yummy cheese ball. Rather, they offer up their wings for the females to munch on while mating. Kevin Judge showed an amazing video of grigs mating, with the female on the back of the male munching vigorously away at the male’s wings (and seemingly trying to get at other male parts too), while the male simultaneously tried to keep her wandering mouth under control with his legs while crimping her abdomen with a structure that looked like a small staple remover (and at least once it removed part of the female’s abdomen).

That is just a small selection of natural history tidbits from the symposium – my apologies for any inaccuracies which I am sure the above hyperlinks can clear up. After sitting through this great symposium, I happened to see Sally Otto, who had some binoculars around her neck. “Have you been bird watching” I cleverly asked, and she proceeded to tell me about a nearby lake with nesting Avocets and many other great birds. How could I not take my own advice, and that of everyone in the symposium? So I ran off to take some photos of the natural history of Kelowna. Amazing stuff. In just a few walks, I saw 45 different bird species, including one – the Pygmy Nuthatch – that I don’t think I have seen before. The conditions were great and I was able to get some good photos, which I show below (many more are here).

So that is it for CSEE 2013 – now I am off to Vancouver Island for field work on stickleback. After that, it is time to get geared up for CSEE 2014, which we are hosting in Montreal. It will be the first ever joint meeting of CSEE, the Canadian Society of Zoologists (CSZ), and the Society of Canadian Limnologists (SCL). The meeting title is “GENOMES TO/AUX BIOMES” and it is supported equally by the three societies and by four Montreal Universities (McGill, UQAM, Condordia, and Universite de Montreal) – we hope to see you there. Montreal has some natural history too!

Breeding Avocets.

The omnipresent chipmunk

A robber fly (thanks for the ID Bob).

Common Flicker - the red-shafted flavor.


The ever-cute California Quail

A Wilson's Phalarope takes flight.


Thursday, May 9, 2013

Life after death: When does your phenotype expire?



Throughout an organisms’ life, the expression of genes, regulated by the biotic and abiotic environment, gives rise to traits that determine how fast it can run or how tall it can grow. Many traits also affect species interactions; for example, are you fast enough to outrun predators? Do you look tasty to herbivores? Most traits (e.g., running speed) cease to be important once an organism dies, but some traits linger and have “afterlife” effects on the environment. A prominent example of afterlife effects can be found in decomposing plant material, which is a crucial part of nutrient cycling. Microbes and fungi are critical to many stages of nutrient cycling, such as the mineralization of organic matter and the nitrification of NH4+, which plants cannot use, to NO3-, which is usable by plants. However, microbes and fungi can be “picky eaters” in a sense, as they prefer substrates with labile simple sugars instead of defensive molecules such as lignin. Simple sugars have carbon and nitrogen supplies that are easily accessible, while larger, more complex molecules require degradation by energetically-costly enzymes. Therefore, genetic and environmental influences on the chemical composition of plant material can persist after a plant sheds its leaves and affect how quickly its nutrients are cycled.

Afterlife effects aren't a new concept; exposure to herbivores and ozone has been shown to indirectly affect decomposition by altering leaf chemistry. However, we recently documented a new type of afterlife effect by showing that genotypic variation in a focal plant’s neighbors could affect the chemical composition of the focal plants litter. Although we don’t have the mechanism completely nailed down, it appears that focal-plant biomass allocation (putting carbon into roots vs. rhizomes vs. stems, and so on) is affected by neighbor-plant genotypic variation, and that shifts in focal-plant biomass allocation are correlated with focal-plant litter quality (specifically, lignin:N). This type of afterlife effect can also be considered an indirect genetic effect (technically, an interspecific indirect genetic effect because the neighboring plants belonged to different species), through which the expression of genes in one individual affects the phenotype of a different, heterospecific individual.



Solidago altissima, one of the study's focal species, along the TN-NC border.


We (Jen Schweitzer, Joe Bailey, and me) were curious whether genetically­-based afterlife effects were unique. Do they have consequences that, for example, ozone-driven afterlife effects would not? Ultimately, we started to think about ecosystem processes (productivity, nutrient cycling, among many others) and the basic drivers of these processes. We argue that ecosystem processes are the “gene-less products of genetic interactions”, meaning that the plant, animal, and microbial traits that interact to create ecosystem processes all have a genetic basis, although the expression of that genetic basis may change depending on how an organism’s genes interact with the biotic and abiotic environment. So, you may ask – “What does this mean?” We’d argue that the “ecosystem processes are gene-less products” perspective allows us to put ecosystems in an evolutionary framework. We can then ask questions like: How might nitrogen cycling or plant productivity change as natural selection acts on the genes that are the most basic drivers of these processes?

If you’re interested in the research behind this post, head to http://www.plosone.org/article/related/info%3Adoi%2F10.1371%2Fjournal.pone.0053718.

The Null Hypothesis is Always Wrong

 No two populations are identical for any trait. No two communities have the same species composition. No detectable phenotype is ever compl...